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.     

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.     

Systematic position of the enigmatic African cycad moths: an integrative approach to a nearly century old problem (Lepidoptera: Geometridae,Diptychini)

The systematic position and hierarchical level of the moth taxon Diptychini Janse (Lepidoptera: Geometridae), the cycad moths, has remained controversial. This is partly due to their unique morphological and biological characteristics. To study the systematics, comprehensive molecular analyses of eight genes, in total 6157 bp, were carried out. We used Bayesian inference to construct phylogenetic trees. The first analysis (46 Geometridae and 7 non‐Geometridae taxa, representing all recently recognised Geometridae subfamilies) demonstrated that the Diptychini belong to the Geometridae subfamily Ennominae. The second analysis, focused on the Ennominae (70 taxa, representing 28 of 30 recently recognised Ennominae tribes worldwide), found that the Diptychini are nested well within the Ennominae; it is monophyletic and associated with the complex of southern Hemisphere Nacophorini, refuting many of the earlier hypotheses about Diptychini relationships. The Diptychini are considered tentatively valid at the tribe level, but relationships with the Nacophorini and the Lithinini need further research. The molecular findings were evaluated from a morphological point of view, which are mostly in agreement with the molecular results. The Diptychini genera are illustrated and characterised using morphological and life‐history traits. Within the Diptychini, three genera are considered valid. Durbana Warren (described in 1904) is proposed as a junior synonym of Veniliodes Warren (described in 1894) ( n.syn. ). Monotypic Larentioides Prout is combined with the tribe Lithinini ( n.comb .). Homonymy of Diptychini Mirza (described in 1991) (Pisces: Cyprinidae, Schizothoracinae) with Diptychini Janse (described in 1933) (Lepidoptera: Geometridae, Ennominae) is noted, the former requiring a replacement name.     

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.     

Precursor structures and evolution of tympanal organs in Lepidoptera (Insecta, Pterygota)

 The patterns of scolopal organs and their innervation were studied by the methylene blue method in larvae, pupae and adults of an Yponomeuta species (Yponomeutidae) and of tympanate adult representatives of the Noctuoidea, Geometridae, Drepanidae and Pyraloidea. The studies were focused mainly on the mesothorax, the metathorax and some anterior abdominal segments. In the abdominal tympanal organs of Geometridae, Drepanidae and Pyraloidea, the auditory scolopidia are homologous with the lateral scolopal organs of the first abdominal segment; however, the hearing organs as such evolved independently in the three taxa. The studies confirm that the tympanal organ in the Noctuoidea is derived from the caudal dorsolateral region of the metathorax including its dorsal scolopal organ and the B-cell. The adult scolopal organs are present already in the larvae and are maintained nearly unchanged during metamorphosis to the adult. Only in the Noctuoidea are the three sensory cells of the larval scolopal organs, which become part of the tympanal organs, reduced to one (in Notodontidae) or two (in other Noctuoidea) during metamorphosis. A hypothetical scenario of the evolution of the tympanal organs is outlined. Accepted: 12 March 1997     

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.     

Disentangling determinants of egg size in the Geometridae (Lepidoptera) using an advanced phylogenetic comparative method

We present a phylogenetic comparative study assessing the evolutionary determinants of egg size in the moth family Geometridae. These moths were found to show a strong negative allometric relationship between egg size and maternal body size. Using recently developed comparative methods based on an Ornstein-Uhlenbeck process, we show that maternal body size explains over half the variation in egg size. However, other determinants are less clear: ecological factors, previously hypothesized to affect egg size, were not found to have a considerable influence in the Geometridae. The limited role of such third factors suggests a direct causal link between egg size and body size rather than an indirect correlation mediated by some ecological factors. Notably, no large geometrid species lay small eggs. This pattern suggests that maternal body size poses a physical constraint on egg size, but within these limits, there appears to be a rather invariable selection for larger eggs.     

Subfamily divergence in the multigene family of ribulose-1,5-bisphosphate carboxylase/oxygenase (rbcS) in Triticeae and its relatives

To investigate genetic mechanisms acting on multigene family in plants, we analyzed sequence variation in the rbcS gene of 13 species of Triticeae and one species each of related tribes (Bromeae and Aveneae). A total of 36 rbcS genes were analyzed. Based on dimorphism in the length of intron, the rbcSs of investigated species were classified into two subfamilies A and B. The difference in intron length was caused by an indel of about 200 bp in the middle of the intron. The two subfamilies of rbcS were present in the three tribes, indicating that the divergence of rbcS subfamilies occurred before the split of these tribes. Generally, variation between the two subfamilies of rbcS was larger than that within subfamily, but these two measures were about the same at the tribe level. This result suggested that divergence of the subfamilies of rbcS occurred at about the same time of tribe diversification. The level of nucleotide variation in the exon region between subfamilies was reduced in the Triticeae, but clear change was not detected in the intron sequence. This result suggested that the exon sequences between subfamilies of rbcS were homogenized without affecting the intron sequence in the Triticeae lineage.     

Basic chromosome numbers of terrestrial orchids

The chromosome numbers of forty-one Brazilian species belonging to 11 genera of preferentially terrestrial orchids (subfamilies Cypripedioideae, Spiranthoideae, Orchidoideae, and Vanilloideae) were examined. Previous records for these subfamilies were reviewed in order to identify the ancestral chromosome numbers of terrestrial orchids. The variation observed within the subfamilies Spiranthoideae (2n=28, 36, 46, 48 and 92), and Orchidoideae (2n=42, 44, ca. 48, ca. 80, 84, and ca. 168) was similar to that previously reported in the literature. In the subfamily Spiranthoideae, some species of Prescottia (subtribe Prescottiinae) and some genera of Spiranthinae showed a bimodal karyotype with one distinctively large pair of chromosomes. The analysis of chromosome numbers of the genera in subfamilies revealed the predominance of the polyploid series 7, 14, 21, 28, 42 with a dysploid variation of ±1 in each ploidy level. These results suggest that the basic chromosome number of terrestrial orchids is x₁=7 for the subfamilies Spiranthoideae and Orchidoideae, as well as other Epidendroid orchids, and that the majority of the genera are composed of palaeopolyploids.     

Evolutionary relationships of multiple alpha satellite subfamilies in the centromeres of human chromosomes 13, 14, and 21

Summary Using Southern and in situ hybridization analyses, we have earlier defined four different subfamilies of alpha satellite DNA (designated pTRA-1, -2, -4, and -7), each of which has a unique higher order structure represented almost identically on human chromosomes 13, 14, and 21. Here we present the complete sequence of single isolates of these four subfamilies, representing approximately 12 kb of sequence information. Sequences of the individual 171-bp monomers that constitute these four subfamilies (and a fifth subfamily, Alpha-R1, that is known to be present on chromosomes 13 and 21) were compared both within and between the different clones. The results indicate that, at the level of their primary sequence, the five alpha subfamilies are characterized by structures that are as unrelated to each other as the different alpha subfamilies from other chromosomes. However, sequence comparisons between monomers of these clones indicate the possibility that pTRA-2, -4, and-1 may have arisen, at least in part, from a common ancestral alphoid sequence. We also provide evidence that exchange of pTRA-1 between nonhomologous centromeres and its homogenization throughout the population, perhaps by unequal exchange mechanisms, could have occurred after the divergence of humans and chimpanzees. The evolution of multiple alphoid subfamilies within a single centromere suggests that unequal exchange mechanisms may be restricted to specific domains. This may in turn contribute to some requirement for subregional pairing of sequences along the length of the centromeres of these chromosomes. Offprint requests to: K.H.A. Choo     

Molecular characterization of the mitochondrial cytochrome oxidase I gene of Oestridae species causing obligate myiasis

A 688-bp region of the mitochondrial cytochrome oxidase I gene was sequenced from larvae of 18 species of Oestridae causing obligate myiasis. Larvae belonged to the four Oestridae subfamilies (Cuterebrinae, Gasterophilinae, Hypodermatinae and Oestrinae), which are commonly found throughout the world. Analysis of both nucleotide and amino acid data was performed. Nucleotide sequences included 385 conserved sites and 303 variable sites; mean nucleotide variation between all species was 18.1% and variation within each subfamily ranged from 5.3% to 13.34%. Intraspecific pairwise divergences ranged from 0.14% to 1.59%, and interspecific variation ranged from 0.7% to 27%. Of the 229 amino acids, 76 were variable (60 of which were phylogenetically informative), with some highly conserved residues identified within each subfamily. Phylogenetic analysis showed a strong divergence among the four subfamilies, concordant with classical taxonomy based on morphological and biological features. This study provides the first molecular data set for myiasis-causing Oestridae species, providing an essential database for the molecular identification of these parasites and the assessment of phylogenetic relationships within family Oestridae.     

Retrotransposon Mys was active during evolution of the Peromyscus leucopus-maniculatus complex

Mys is a retrovirus-like transposable element found throughout the genus Peromyscus. Several mys subfamilies identified on the basis of restriction site variation occur in more than one species. The distribution of these subfamilies is consistent with the accepted species phylogeny, suggesting that mys was present in the ancestor of Peromyscus and has been active through much of the evolution of this genus. Quantitative Southern blot analysis was used to examine the variability of subfamilies in P. leucopus and maniculatus. We found that subfamilies with phylogenetically narrow distributions were more variable in copy number both within and between species than subfamilies with a broader distribution. Taken together, our data suggest that mys has undergone multiple rounds of transposition since the peromyscine radiation, and that five subfamilies have been amplified during the evolution of the leucopus-maniculatus species complex. Correspondence to: H.A. Wichman     

Organ-specific expression of different histone H3 and H4 gene subfamilies in developing and adult maize

The steady-state levels of H3 and H4 mRNAs transcribed from three H3 and two H4 multigene subfamilies were studied during germination and in different organs of maize. During germination the five subfamilies are expressed in parallel to DNA synthesis, but a 5-fold difference in the quantity of mRNAs transcribed per gene copy was found from our subfamily to another. In adult plants H3 and H4 mRNA levels are highest in organs containing meristematic tissues but also high in non-proliferating tissues. No strict tissue specificity expression could be detected but some subfamilies show preferential expression in some tissues.     

Cytogenetic aspects of phylogeny in the Bovidae. II. C-banding

Constitutive heterochromatin in the Bovidae, as revealed by C-banding, was mostly located in the centromeric regions. Considerable variation was, however, evident in the size of the C-bands both within and between subfamilies. Some evidence was found for a reduction in the amount of centromeric heterochromatin in bi-armed relative to acrocentric autosomes, and these findings are discussed in relation to karyotype evolution in the group.     

Genetic variation of recent Alu insertions in human populations

The Alu family of intersperesed repeats is comprised of ovr 500,000 members which may be divided into discrete subfamilies based upon mutations held in common between members. Distinct subfamilies of Alu sequences have amplified within the human genome in recent evolutionary history. Several individual Alu family members have amplified so recently in human evolution that they are variable as to presence and absence at specific loci within different human populations. Here, we report on the distribution of six polymorphic Alu insetions in a survey of 563 individuals from 14 human population groups across several continents. Our results indicate that these polymorphic Alu insertions probably have an African origin and that there is a much smaller amount of genetic variation between European populations than that found between other populations groups. Present address: Department of Pathology, Stanley S. Scott Cancer Center, Louisiana State University Medical Center, 1901 Perdido St., New Orleans, LA 70112 Correspondence to: M.A. Batzer     

Phylogenetic relationships of the Asian palm civets (Hemigalinae & Paradoxurinae, Viverridae, Carnivora)

The Viverridae (Mammalia, Carnivora), one of the least studied groups of carnivorans, include two subfamilies of Asian palm civets: Hemigalinae and Paradoxurinae. The relationships between and within these two subfamilies have never been thoroughly tested using an extensive molecular sample set. In this study, we gathered sequences of four genes (two mitochondrial: Cytochrome b and ND2 and two nuclear: beta-fibrinogen intron 7 and IRBP exon 1) for eight of the eleven extant species representing these two subfamilies. The results showed that: (1) the Asian palm civets (Hemigalinae and Paradoxurinae) have a single origin and form the sister-group of the (Genettinae+Viverrinae) clade, (2) the Hemigalinae (including the otter civet Cynogale bennettii) are monophyletic, (3) the Paradoxurinae are monophyletic and (4) the small-toothed palm civet (Arctogalidia trivirgata) is an early offshoot within the Paradoxurinae. Using a relaxed molecular clock analysis, the differentiation of the (Hemigalinae+Paradoxurinae) was inferred to occur in the Late Oligocene/Early Miocene.     

Nucleolar organiser region (NOR) location in karyotypes of Australian ground frogs (family Myobatrachidae)

Nucleolar organiser regions (NORs) were examined in over 90% of the species of Australian ground frogs (familiy Myobatrachidae), including representatives from all twenty currently recognised genera and the three subfamilies. Throughout the family, location of the NOR within the karyotype showed considerable variation yet karyotype morphology showed uniformity. The precise mechanism(s) whereby variation in NOR location evolved while karyotype morphology was unchanged remains uncertain. Comparison of the two major subfamilies showed that the Limnodynastinae had a greater diversity of NOR location than the Myobatrachinae. The limnodynastine genus,Heleioporus, was the only one to show multiple NOR sites in several species. NOR location was particularly stable within most polytypic genera. Differences in NOR location within the remaining polytypic genera (Heleioporus, Limnodynastes, Neobatrachus, Philoria andRanidella) pointed to taxonomic discriminations that were generally consistent with recent proposals based on other criteria.     

A reappraisal of the family Comasteridae A. H. Clark, 1908 (Echinodermata: Grinoidea), with the description of a new subfamily and a new genus

The family Comasteridae is divided into four subfamilies on the basis of syzygy positions in the division series and proximal arm. Use of this character results in major differences in perceived relationships between genera compared with that obtained when traditional subfamilial characters are used. The primary character previously used to determine comasterid subfamilies, cirrus ornamentation, is rejected as unimportant in recognizing taxa any higher than species. Seventeen genera, including a new genus from the tropical western Atlantic, and 99 species are now recognized in the Comasteridae. Four of these genera and five species cannot be placed within the newly diagnosed subfamilies. Characters of generic importance are considered to be pinnule comb form, mouth position and, to a lesser extent, the characters traditionally used to distinguish genera.