Molecular phylogeny of the subfamilies in Geometridae (Geometroidea: Lepidoptera).

Molecular sequence data from three gene fragments were used to examine critically a provisional phylogenetic classification based on morphological characters of the Geometridae, one of the most species-rich families of moths. The sister group relationship between Geometridae and Drepanidae gained further support from the molecular analysis, which was based on the ND1 mitochondrial gene and the first and second expansion segments of the 28S ribosomal RNA gene. Although the alignment of the second expansion segment contained regions with many gaps, it provided the most resolution of the gene fragments. Parsimony analysis of the combined data resulted in a cladogram in which species belonging to Drepanidae, Larentiinae, and Sterrhinae formed monophyletic groups. The Ennominae did not form a monophyletic group but rather were contained within a broader monophyletic group including Archiearinae, Geometrinae, and Alsophilinae (represented by only one species per group in the present study). The molecular results were used to explore further the relationship between Sterrhinae and Larentiinae, the question as to whether Ennominae actually represent a monophyletic group, and the relationships between Ennominae and some of the other subfamilies.     

Faunal composition of geometrid moths changes with altitude in an Andean montane rain forest

Aim The objective of this study was to describe and interpret the changes in faunal composition in the moth family Geometridae (Lepidoptera) along a small‐scale elevational gradient in a tropical montane rain forest. This gradient was compared with a large‐scale latitudinal gradient in Europe. Location Investigations were carried out in the province Zamora‐Chinchipe in southern Ecuador along a gradient ranging from 1040 to 2677 m above sea level at twenty‐two sites. Methods Moths were sampled with light‐traps in three field periods in 1999 and 2000 and subsequently sorted and determined to species or morphospecies. Results We analysed 13,938 specimens representing 1010 species of geometrid moths. The proportional contribution of subtaxa to the local geometrid fauna changes along the elevational gradient at all systematic levels considered. While proportions of species of the subfamilies Ennominae, Sterrhinae and Geometrinae significantly decrease, the proportion of Larentiinae increases with increasing altitude. Changes also occur within the subfamilies Ennominae and Larentiinae. The host–plant specialist ennomine tribes Cassymini, Macariini and Palyadini completely vanish, and the proportion of the tribe Boarmiini decreases at high altitudes. In contrast, the remaining tribes (mostly comprising polyphagous species) either do not show proportional changes (Azelinini, Nacophorini, Nephodiini, Ourapterygini) or even increase (Caberini, ‘Cratoptera group’). Within Larentiinae, the species proportion of the genus Eois decreases, whereas concomitantly the proportion of Eupithecia increases. There is a remarkable similarity between the altitudinal patterns in Ecuador and those found along the latitudinal gradient in Europe. Main conclusions Species of the subfamily Larentiinae seem to be particularly well‐adapted to harsh environmental conditions, towards both high altitudes and latitudes. They might disproportionately profit from lower predation at higher altitudes. Many changes in the faunal composition can be explained by expected host–plant requirements of the species involved. Our results show that diversity estimates based on taxon ratios which are assumed to be constant must be regarded with caution because such ratios can change rapidly along environmental gradients.     

Systematic position of Lythriini revised: transferred from Larentiinae to Sterrhinae (Lepidoptera, Geometridae)

The tribe Lythriini is a small group of diurnally active geometrid moths consisting of a single Palaearctic genus Lythria with five species. The systematic placement of Lythriini has remained controversial: though traditionally it has been placed into the subfamily Larentiinae, a number of morphological characters link this tribe with the subfamily Sterrhinae. A molecular phylogenetic study was conducted to verify the systematic position of Lythriini, using sequences of both mitochondrial and nuclear genes: elongation factor 1α ( EF-1α ), wingless ( wgl ), 28S rRNA expansion segment D2 ( 28S D2 ), cytochrome oxidase subunit 1 ( COI ) and NADH dehydrogenase subunit 1 ( ND1 ) (a total of 3784 bp). Phylogenetic analysis reliably demonstrated that Lythriini belong to the subfamily Sterrhinae. Therefore, we propose to remove tribe Lythriini from Larentiinae and unite it with Sterrhinae. Moreover, our analysis supports the monophyly of both Sterrhinae and Larentiinae. However, although both morphological data and interspecific genetic distances insinuated that Lythria cruentaria and L. sanguinaria are sister species, the latter formed a clade of sister taxa together with L. purpuraria .     

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.     

Additions to the geometrid fauna (Lepidoptera: Geometridae) of Chile

Three species of geometrid moths are reported from the extreme North of Chile. All three are new for the Chilean fauna: Scopula umbilicata (Fabricius, 1794) (Sterrhinae), Cataspilates grisescens Warren, 1897, and Pero obtusaria Prout, 1928 (Ennominae).     

An overview of the Tasmanian geometrid moth fauna (Lepidoptera: Geometridae) and its conservation status

A brief review of the geometrid fauna of the large island of Tasmania and a simple analysis of its conservation status and threats are presented. The fauna comprises 310 species of which Ennominae contribute slightly less than half the total and Larentiinae one third; 23% of the geometrid fauna is endemic at species level. Mixed eucalypt-rainforest is identified as the richest wet forest habitat in geometrid species. Using distribution data at 10  km resolution, the most widespread and most restricted taxa are identified. The conservation status of Lepidoptera living above 800 m is relatively good. However, coastal species and those associated with herb-rich native grasslands are under some pressure from habitat change. Three species of geometrid moths are listed as threatened in Tasmania’s Threatened Species Protection Act 1995 but several others may qualify for listing.     

Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes

Geometridae is one of the most diverse families within the Lepidoptera, comprising nine subfamilies. Winter moths, which have a unique life history, are found in three subfamilies. To examine the phylogeny of the Geometridae at the subfamily level and determine the evolutionary history of winter moths, we constructed phylogenetic trees for all nine geometrid subfamilies using two mitochondrial and two nuclear gene sequences. Specimens of all subfamilies were sampled from Japan. Simultaneous analyses of the combined data from all genes revealed that the Geometridae comprised two major clades: one with subfamilies Larentiinae and Sterrhinae, and the other with the remaining seven subfamilies. The second clade included the largest subfamily, Ennominae, and the subfamily Archiearinae, which is traditionally considered to be an ancestral lineage of the Geometridae. The Larentiinae+Sterrhinae clade contained one winter moth lineage, and the second major clade consisted of three winter moth lineages, including Alsophilinae, which contains winter moths exclusively. Using a Bayesian inference of divergence times, we estimated that geometrids began to diverge 54 Mya (62-48 Mya), whereas winter moth lineages differentiated from non-winter moth lineages 34-12 Mya, during the global cooling events in the Oligocene and the early Miocene. The adaptation to cool climates may have been a preadaptation that facilitated the winter moth life cycle.     

The role of environment and mid-domain effect on moth species richness along a tropical elevational gradient

Aim The biodiversity of geometrid moths (Lepidoptera) along a complete tropical elevational gradient was studied for the first time. The patterns are described, and the role of geometric constraints and environmental factors is explored. Location The study was carried out along the Barva Transect (10° N, 84° W), a complete elevational gradient ranging from 40 to 2730 m a.s.l. in Braulio Carrillo National Park, Costa Rica, and adjacent areas. Methods Moths were sampled manually in 2003 and 2004 at 12 rain forest sites using light ‘towers’, each with two 15 W ultraviolet fluorescent tubes. We used abundance‐based rarefaction, statistical estimation of true richness (Chao 1), geographically interpolated observed richness and Fisher's alpha as measures of local diversity. Results A total of 13,765 specimens representing 739 species were analysed. All four measures showed a hump‐shaped pattern with maxima between 500 and 2100 m elevation. The two subfamilies showed richness and diversity maxima at either lower (Ennominae) or higher (Larentiinae) elevation than Geometridae as a whole. Among the four environmental factors tested, relative humidity yielded the highest correlation over the transect with the rarefaction‐based richness estimates as well as with estimated true species richness of Geometridae as a whole and of Larentiinae, while rainfall explained the greatest variation of Ennominae richness. The elevational pattern of moth richness was discordant with both temperature and with tree species richness. A combination of all environmental factors in a stepwise multiple regression produced high values of r² in Geometridae. The potential effects of geometric constraints (mid‐domain effect, MDE) were investigated by comparing them with observed, interpolated richness. Overall, models fitted very well for Geometridae as a whole and for Ennominae, but less well for Larentiinae. Small‐ranged species showed stronger deviations from model predictions than large‐ranged species, and differed strikingly between the two subfamilies, suggesting that environmental factors play a more pronounced role for small‐ranged species. We hypothesize that small‐ranged species (at least of the Ennominae) may tend to be host specialists, whereas large‐ranged species tend to be polyphagous. Based on interpolated ranges, mean elevational range for these moths was larger with increasing elevation, in accordance with Rapoport's elevational rule, although sampling effects may have exaggerated this pattern. The underlying mechanism remains unknown because Rapoport's ‘rescue’ hypothesis could not explain the observed pattern. Conclusions The results clearly show that moth diversity shows a hump‐shaped pattern. However, remarkable variation exists with regard to taxon and range size. Both environmental and geometric factors are likely to contribute to the observed patterns.     

Phylogenetic diversity of geometrid moths decreases with elevation in the tropical Andes

Species diversity of geometrid moths (Lepidoptera, Geometridae) has previously been shown to be extremely and constantly high along a continuously forested elevational gradient in the Andes of southern Ecuador. We analysed samples taken from 32 sites between 1999 and 2011 in northern Podocarpus National Park and adjacent areas from 1020 to 2916 m a.s.l. We conjecture that high elevation habitats were historically mostly colonised by species from lower elevations, and that environmental filtering (e.g. through host plant specificity or temperature tolerance) constrained colonisation from lower elevations, which would yield a pattern of elevationally decreasing phylogenetic diversity. We analysed elevational phylogenetic patterns by means of: 1) the nearest‐taxon index (NTI), 2) DNA barcode‐based terminal branch lengths (TBLs) from maximum‐likelihood phylogeny, 3) the subfamily composition of the local assemblages, and 4), the rarefied number of morphologically defined genera per site. We counted a total of 1445 species. NTI values significantly increased with elevation, both in a conventional and a rarefaction approach. TBLs decreased significantly with elevation. Subfamily composition profoundly changed with elevation, particularly expressed as an increased proportion of the subfamily Larentiinae and decreased fractions of Sterrhinae and Geometrinae. The number of genera in equally rarefied species resamples significantly decreased with elevation. We conclude that environmental filtering indeed contributed to an altitudinal decrease in moth phylodiversity, but these constraints prevented only relatively few clades from colonising high elevation habitats.     

Bergmann's rule does not apply to geometrid moths along an elevational gradient in an Andean montane rain forest

Aim Bergmann's rule generally predicts larger animal body sizes with colder climates. We tested whether Bergmann's rule at the interspecific level applies to moths (Lepidoptera: Geometridae) along an extended elevational gradient in the Ecuadorian Andes. Location Moths were sampled at 22 sites in the province Zamora‐Chinchipe in southern Ecuador in forest habitats ranging from 1040 m to 2677 m above sea level. Methods Wingspans of 2282 male geometrid moths representing 953 species were measured and analysed at the level of the family Geometridae, as well as for the subfamily Ennominae with the tribes Boarmiini and Ourapterygini, and the subfamily Larentiinae with the genera Eois, Eupithecia and Psaliodes. Results Bergmann's rule was not supported since the average wingspan of geometrid moths was negatively correlated with altitude (r = ?0.59, P < 0.005). The relationship between body size and altitude in Geometridae appears to be spurious because species of the subfamily Larentiinae are significantly smaller than species of the subfamily Ennominae and simultaneously increase in their proportion along the gradient. A significant decrease of wingspan was also found in the ennomine tribe Ourapterygini, but no consistent body size patterns were found in the other six taxa studied. In most taxa, body size variation increases with altitude, suggesting that factors acting to constrain body size might be weaker at high elevations. Main conclusions The results are in accordance with previous studies that could not detect consistent body size patterns in insects at the interspecific level along climatic gradients.     

Phylogeny and tribal classification of Sterrhinae with emphasis on delimiting Scopulini (Lepidoptera: Geometridae)

Abstract. The phylogenetic relationships of tribes of the geometrid subfamily Sterrhinae (Lepidoptera) were studied, with special emphasis on finding delimiting characters for the tribe Scopulini. Two cladistic analyses were conducted for fifty‐nine species representing all previously recognized Sterrhinae tribes and covering the geographical range of the subfamily. In the first analysis, twelve putative synapomorphies of Scopulini, taken from the literature, were coded for actual specimens in order to test their ability to support the monophyly of the group. The resulting strict consensus cladogram was totally unresolved. In the second analysis, the twelve characters were combined with additional information from the morphology and ecology of adults and immature stages. Analysis of these ninety‐six characters resulted in a well‐resolved cladogram. The tribes were found to be monophyletic, except Cosymbiini and Rhodostrophiini. There are two main lineages within Sterrhinae: Cosymbiini + Rhodometrini + Timandrini and Rhodostrophiini + Cyllopodini + Sterrhini + Scopulini. Aletini and Problepsini lay within the concept of Scopulini. The association of the included Larentiinae taxa with the Cosymbiini + Rhodometrini + Timandrini lineage questions the monophyly of Sterrhinae. A majority of the recovered synapomorphic characters had been recognized previously, but several new phylogenetically informative characters were found, especially from the thorax. No unique characters diagnosing the tribe Scopulini were found, but many homoplastic synapomorphic features were found which diagnose parts of it. All recognized Sterrhinae genera are assigned tentatively to tribes and problematic cases are discussed.     

Molecular phylogeny of Sterrhinae moths (Lepidoptera: Geometridae): towards a global classification

A multigene phylogenetic study was carried out to test current, mostly morphology-based hypotheses on Sterrhinae phylogeny with additional material included from further geographical areas and morphologically different lineages. A maximum likelihood analysis (11 molecular markers and 7665 bp) was conducted on 76 species and 41 genera using iq-tree software. The resulting phylogenetic hypothesis is well resolved and branches have high support values. Results generally agree with earlier hypotheses at tribal levels and support the hypothesis that Sterrhinae comprises two major lineages. Based on the molecular phylogeny and extensive morphological examination, nine tribes are considered valid and the following taxonomic changes are introduced to recognize monophyletic groups: Mecoceratini Guenée, 1858 (= Ametridini Prout, 1910) is transferred from Desmobathrinae to Sterrhinae, and it is considered valid at tribal level new classification ; Haemaleini Sihvonen & Brehm is described as a new tribe and deemed sister to Scopulini + Lissoblemmini; Lissoblemmini Sihvonen & Staude is described as a new tribe and sister to Scopulini; Lythriini Herbulot, 1962 is now a junior synonym of Rhodometrini Agenjo, 1952 syn.n. ; and Rhodostrophiini Prout, 1935 is now a junior synonym of Cyllopodini Kirby, 1892 syn.n. In addition, 48 taxa are transferred from other geometrid subfamilies to Sterrhinae, or within Sterrhinae from one tribe to another, or they are classified into a tribe for the first time, or a new genus classification is proposed. The results demonstrate the limited explanatory power of earlier classifications, particularly at the tribal level. This is probably a result of earlier classifications being based on superficial characters and biased towards the European and North American fauna. The species richness and distribution of Sterrhinae and its constituent tribes are reviewed, showing that the globally distributed Sterrhinae are most diverse in the Neotropics (31% of global fauna). They are species-rich in the Palaearctic (22%), Afrotropics (19%) and Indo-Malay (16%) regions, whereas they are almost absent in Oceania (1%). In terms of the described fauna, the most species-rich tribes are Scopulini (928 species), Sterrhini (876 species) and Cosymbiini (553 species), all of which have a cosmopolitan distribution. Mecoceratiini and Haemaleini are almost entirely Neotropical. Timandrini and Lissoblemmini, by contrast, are absent in the Neotropics. We present a revised classification of the global Sterrhinae fauna, which includes about 3000 putatively valid species, classified into nine tribes and 97 genera. Four genera are of uncertain position within Sterrhinae. Our results highlight the compelling need to include more genera from a global perspective in molecular phylogenetic studies, in order to create a stable global classification for this subfamily. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:zoobank.org :pub:A66F5DDD-06D6-4908-893E-E8B124BB99B1.     

Diversity of geometrid moths (Lepidoptera: Geometridae) along an Afrotropical elevational rainforest transect

Geometrid moths were investigated at 26 sites on 9 elevational levels along an elevational transect at Mt. Kilimanjaro (Tanzania), stretching from the fine‐grained mosaic of small agroforest plots with combined cultivation of trees, shrubs and crops at 1650 m through mountain rainforest to heathland at 3300 m. We sampled moths manually at light between 19 : 00 and 22 : 00 in the rainy seasons of March to May and October to January in the years 2000, 2001 and 2002. Along the transect, the composition of moth communities changed from a domination by Sterrhinae and Ennominae to a dominance of Larentiinae with increasing elevation. Overall, alpha diversity was very low compared to other tropical mountain regions. Fisher's alpha showed a maximum of 30 in the agroforest mosaic at 1650 m and decreased to values around 12 in the mountain rainforest. Communities of geometrid moths within the forest belt were significantly dissimilar from communities outside the forest. The diversity patterns on Mt. Kilimanjaro can be related to the young age, island‐like position and history of the mountain. These factors have led to the formation of a homogeneous upper mountain rainforest habitat which in turn houses homogeneous moth communities with a low diversity compared to habitats at lower elevations. Here, a heterogeneous habitat mosaic allowing the intrusion of savannah species into this former forest habitat may account for an increased diversity. In the heath zone above the forest, climatic conditions are very harsh, permitting only few specialists to thrive in this ericaceous woodland. Edge effects were discernible at the forest–heathland boundary where some moth species from heathland invaded the closed forest. At the boundary between agroforest and a forest mosaic of exotic Acacia and Eucalyptus forest plantations and natural mountain forest, diversity values remained low as the dominant species Chiasmia fuscataria accounted for far higher proportions than other dominant species in any of the other habitats.     

Unique elevational diversity patterns of geometrid moths in an Andean montane rainforest

Alpha‐diversity of geometrid moths was investigated along an elevational gradient in a tropical montane rainforest in southern Ecuador. Diversity was measured using 1) species number, 2) extrapolated species number (Chao 1 estimator), 3) rarefied species number, and 4) Fisher's alpha. When applied to the empirical data set, 1 and 2 strongly depended on the sample size, whereas 3 and 4 were suitable and reliable measures of local diversity. At single sites, up to 292 species were observed, and extrapolation estimates range from 244 to 445 species. Values for Fisher's alpha are among the highest ever measured for this moth family, and range from 69 to 131 per site. In contrast to theoretical assumptions and empirical studies in other regions of the world, the diversity of geometrid moths remained consistently high along the entire gradient studied. Diversity measures correlated with neither altitude nor ambient temperature. The large subfamily Ennominae has previously been assumed to be a group that occurs mainly at low and medium elevations. However, no decline in diversity was found in the study area. The diversity of the other large subfamily, Larentiinae, even increased from the lowest elevations and was highest at elevations above 1800 m. The roles of a decreasing diversity of potential host‐plants, decreasing structural complexity of the vegetation, increasingly unfavourable climatic conditions and possible physiological adaptations in determining herbivore species richness are discussed. A relatively low predation pressure might be an advantage of high‐altitude habitats. The physiognomy of the Andes (folded mountains, large areas at high altitudes) might also have allowed speciation events and the development of a species‐rich high‐altitude fauna. There is evidence that the species‐richness of other groups of herbivorous insects in the same area declines as altitude increases. This emphasises difficulties that are associated with biodiversity indicator groups, and calls for caution when making generalisations from case studies.     

Intestinal Helminthic Infections in Striped Field Mice,Apodemus agrarius,from Two Southern Regions of Korea

The present study was performed to know the infection status of intestinal helminths in a most common species of field mice, Apodemus agrarius, from 2 southern regions of Korea. Total 133 and 103 mice were collected by the mouse trap in Hapcheon-gun, Gyeongsangnam-do and Gurye-gun, Jeollanam-do, respectively, from July 2005 to June 2006. The small intestine of each mouse was resected and longitudinally opened with a pair of scissors. The intestinal contents were washed with 0.85% saline until the supernatant became clear. Helminths were collected with naked eyes or under a stereomicroscope from the sediment of the intestinal content. More than 11 species of helminths (4 nematode spp., 5 trematode spp., and 2 cestode spp.) were recovered. Among these, heligmosomoid nematodes (97.5%) was the most highly and heavily infected species. As the members of trematodes, Plagiorchis muris, Brachylaima sp., Echinostoma hortense, Echinostoma cinetorchis, and unidentified echinostome larvae were found in the small intestines of 35 (14.8%), 12 (5.1%), 6 (2.5%), 1 (0.4%), and 1 (0.4%) mice respectively. Two species of tapeworms, Hymenolepis nana and Hymenolepis diminuta were also detected in 79 (33.5%) and 21 (8.9%) mice, respectively. Conclusively, heligmosomoid nematodes were the most prevalent (dominant) species among more than 11 helminth species detected, and Brachylaima sp. fluke is newly added in the list of intestinal trematodes in Korea.     

渤海东部海域秋季底层游泳动物种类组成及群落多样性

根据2014年9月在渤海东部海域(37°40'—38°20'N、120°00'—121°20'E)进行的游泳动物底拖网调查数据,应用相对重要性指数、生物多样性指数、群落结构多元统计分析等方法对该海域游泳动物种类组成及群落多样性特征进行了分析。结果表明:调查海域共捕获游泳动物54种,以头足类枪乌贼(Loliolus spp.)为绝对优势种;游泳动物资源密度为28.82kg/h和5166尾/h。游泳动物种类Margalef丰富度指数(D)的变化范围为1.99—3.67,Shannon-Wiener多样性指数(H')变化范围为0.89—2.28,Pielou均匀度指数(J')变化范围为0.29—0.69。群落结构多元统计分析表明,渤海东部海域游泳动物群落结构以60%的相似性可划分为3个组群:长岛群岛海域组群(A组群)、龙口海域组群(B组群)和蓬莱海域组群(C组群);ANOSIM分析表明,组群之间差异极显著,两两之间差异亦极显著。受增殖放流影响,C组群资源密度和种类多样性均保持较高的水平,而A、B组群人为活动频繁,过度捕捞严重,对海域生态环境和渔业资源整体结构的破坏较大。