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Tympanal organs of geometrid moths: A review of their morphology, function, and systematic importance

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Abstract

The basic components of the tympanal organs of Geometridae are described against the background of the relevant literature. The work is prefaced by a summary of the occurrence of hearing organs within adult Lepidoptera as a whole, and the systematic value of these structures is assessed. A Latin-based nomenclature is proposed to standardize the terminology. An assessment of the morphological variation of these organs within the Geometridae was based on the study of over 230 species representing numerous examples from all the subfamilies. Morphological variation between and within the geometrid subfamilies is discussed.

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... In his revision, Turner described considerable variation in the wing venation of what he called the Oenochroma group (currently Oenochrominae) and the Dichromodes group (all now Epidesmiinae). Cook & Scoble (1992) reviewed the tympanal organs of geometrid moths and suggested the circular form of the lacinia and its orientation parallel to the tympanum as a potential synapomorphy of the robustbodied Oenochrominae s.s. The genera that did not fit the definition were listed as Oenochrominae s.l. and it contained among others Abraxaphantes, Dichromodes and Epidesmia (all now Epidesmiinae). ...
... Minet & Scoble, 1999), but the wing pattern characters are variable. Sterrhinae and Larentiinae share a hammer-headed tip in tympanal organ's ansa (Cook & Scoble, 1992, see also Viidalepp, 2011) and a few other characters (summarized in Hausmann, 2004). McGuffin (1967McGuffin ( , 1987 provided diagnostic larva and pupa characters, but those were based on limited Canadian material. ...
... Contrary to the historical view, gnathos or remnants of gnathos are found in Larentiinae (Hausmann, 2004, Schmidt, 2015. Sterrhinae and Larentiinae share hammer-headed tip in the tympanal organ's ansa (Cook & Scoble, 1992, see also Viidalepp, 2011) and a few other characters (summarized in Hausmann, 2004). McGuffin (1987) provided diagnostic larva and pupa characters, but those were based on limited Canadian material. ...
Article
Our study revises Epidesmiinae, the first new Geometridae subfamily that has been described in 127 years. We studied the morphological characters of representatives from all genera currently classified into Epidesmiinae, and compared those with all other geometrid subfamilies. Epidesmiinae were found to have an Australasian distribution, with one species occurring in the Indo-Malayan realm. They compose a lineage diagnosable by a combination of the following morphological characters: male antennae unipectinate; labial palps elongated (particularly the second segment), vom Raths’s organ with an elliptical invagination; forewing with two areoles; hindwing with one anal vein; gnathos arms fused, granulate or dentate apically; female genitalia with two signa, one stellate, another an elongated and spinose plate. We also present a summary of diagnostic characters of all geometrid subfamilies, which confirm the lack of single unique morphological characters. The limited information on the biology and ecology of Epidesmiinae species are summarized, indicating that some species fly during the day, most adult records are from the Southern Hemisphere summer months and larvae are found on Myrtaceae. We transfer Arcina Walker, 1863 from Oenochrominae s.l. to Epidesmiinae. Epidesmiinae includes 102 species that are now classified into nine genera: Abraxaphantes, Adeixis, Arcina, Dichromodes, Ecphyas, Epidesmia, Phrataria, Phrixocomes and Systatica.
... Geometridae are the second most species-rich family of Lepidoptera, with approximately 24,000 described species (number from Van Nieukerken et al. (2011) updated by the authors) found in all regions except Antarctica. The monophyly of Geometridae is well supported based on distinctive morphological characters (Cook & Scoble, 1992;Scoble, 1992;Minet & Scoble, 1999). In particular, adult members of the family possess paired tympanal organs at the base of the abdomen, while in larvae the prolegs are reduced to two pairs in almost all species, which causes the larvae to move in a looping manner (Minet & Scoble, 1999). ...
... Eight subfamilies are currently recognized in Geometridae . Several recent molecular and morphological studies have attempted to confirm the monophyly or clarify the taxonomy of most of these groups, for instance: Sterrhinae (Holloway, 1997;Hausmann, 2004;Sihvonen & Kaila, 2004;Õunap, Viidalepp & Saarma, 2008), Larentiinae (Holloway, 1997;Mironov, 2003;Viidalepp, 2006Viidalepp, , 2011Hausmann & Viidalepp, 2012;Õunap, Viidalepp & Truuverk, 2016), Desmobathrinae (Holloway, 1996;Hausmann, 2001), Archiearinae (Hausmann, 2001;Young, 2006), Oenochrominae (Holloway, 1996;Scoble & Edwards, 1990;Cook & Scoble, 1992;Hausmann, 2001;Young, 2006), Geometrinae (Cook et al., 1994;Pitkin, 1996;Hausmann, 2001;Ban et al., 2018), Orthostixinae (Holloway, 1997) and Ennominae (Holloway, 1994;Pitkin, 2002;Beljaev, 2006;Young, 2006;Wahlberg et al., 2010;Õunap et al., 2011;Skou & Sihvonen, 2015;Sihvonen, Staude & Mutanen, 2015), but questions remain. An important shortcoming is that our understanding of geometrid systematics is biased towards the long-studied European fauna, whereas the highest diversity of this family is in the tropics, which are still largely unexplored (Brehm et al., 2016). ...
... Oenochrominae and Desmobathrinae are considered the most controversial subfamilies in Geometridae. A close relationship of these subfamilies has been proposed both in morphological (Meyrick, 1889;Cook & Scoble, 1992;Holloway, 1996) and in molecular studies Ban et al., 2018). In early classifications, species of Desmobathrinae and Oenochrominae were classified in the family Monocteniadae (Meyrick, 1889), which is currently considered a junior synonym of Oenochrominae Guenée. ...
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Our study aims to investigate the relationships of the major lineages within the moth family Geometridae, with a focus on the poorly studied Oenochrominae-Desmobathrinae complex, and to translate some of the results into a coherent subfamilial and tribal level classification for the family. We analyzed a molecular dataset of 1,206 Geometroidea terminal taxa from all biogeographical regions comprising up to 11 molecular markers that includes one mitochondrial (COI) and 10 protein-coding nuclear gene regions (wingless, ArgK, MDH, RpS5, GAPDH, IDH, Ca-ATPase, Nex9, EF-1alpha, CAD). The molecular data set was analyzed using maximum likelihood as implemented in IQ-TREE and RAxML. We found high support for the subfamilies Larentiinae, Geometrinae and Ennominae in their traditional scopes. Sterrhinae becomes monophyletic only if Ergavia Walker, Ametris Hübner and Macrotes Westwood, which are currently placed in Oenochrominae, are formally transferred to Sterrhinae. Desmobathrinae and Oenochrominae are found to be polyphyletic. The concepts of Oenochrominae and Desmobathrinae required major revision and, after appropriate rearrangements, these groups also form monophyletic subfamily-level entities. Oenochrominae s.str. as originally conceived by Guenée is phylogenetically distant from Epidesmia and its close relatives. The latter is hereby described as the subfamily Epidesmiinae Murillo-Ramos, Sihvonen & Brehm, subfam. nov. Epidesmiinae are a lineage of "slender-bodied Oenochrominae" that include the genera Ecphyas Turner, Systatica Turner, Adeixis Warren, Dichromodes Guenée, Phrixocomes Turner, Abraxaphantes Warren, Epidesmia Duncan & Westwood and Phrataria Walker. Archiearinae are monophyletic when Dirce and Acalyphes are formally transferred to Ennominae. We also found that many tribes were para-or polyphyletic and therefore propose tens of taxonomic changes at the tribe and subfamily levels. Archaeobalbini stat. rev. Viidalepp (Geometrinae) is raised from synonymy with Pseudoterpnini Warren to tribal rank. Chlorodontoperini Murillo-Ramos, Sihvonen & Brehm, trib. nov. and Drepanogynini Murillo-Ramos, Sihvonen & Brehm, trib. nov. are described as new tribes in Geometrinae and Ennominae, respectively.
... Morphologically the geometrids are best defined by the unique structure of the tympanal organs, particularly the presence of the ansa, found at the base of the abdomen and have their tympanal apertures opening ventro-laterally. These structures are reduced or lost in some of the brachypterous females [6]. The alpha-taxonomy of the Geometridae has been developing progressively, and excellent treatises exist, but these are often geographically limited and not aimed at resolving geometrid phylogeny at a deeper global level. ...
... Their definition relied on general similarities of facies, wing venation and male genitalia structure. Cook and Scoble [6] suggested that the circular form of the lacinia and its orientation parallel to the tympanum in the tympanic bulla was an autapomorphy for these robust Oenochrominae. Holloway [8] noted that these features are not apparent in Sarcinodes, the only Oriental representative of the group. ...
... The Dysphaniini have high geoverdin concentrations, sharing this and a few morphological features with the Geometrinae, suggesting these taxa may be linked. The lack of shared, unique characters has led some authors to challenge the placement of the Dysphaniini in the Geometrinae [6,8,52]. The remaining geometrine tribes are difficult to diagnose, and some genera do not fall readily into any of them, vide e.g. ...
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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. 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. 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.
... The Sterrhinae, a subfamily of the Geometridae moths, or loopers, are often called waves because of the numerous wavy fasciae continuing from forewing to hindwing. They share the typical characteristics of family Geometridae, namely the paired tympanal organs at the base of the adbomen (Cook & Scoble 1992). The monophyly of the subfamily has been challenged (Common 1990, Holloway 1997, Minet & Scoble 1999, Hausmann 2001), but within the Sterrhinae, however, there are several possibly monophyletic groups that are more readily diagnosable . ...
... Larentiinae has been suggested to be the sister group of Sterrhinae based on morphology (Holloway 1997, Fänger 1999. Characters that pull Sterrhinae and Larentiinae together are the distribution of the male secondary sexual organs, structure of signum of the female corpus bursae (Holloway 1997), and features of the tympanal organs (Cook & Scoble 1992). A molecular study by Abraham et al. (2001), based on a very limited taxon sample, did not support the Sterrhinae and Larentiinae relationship. ...
... For the phylogenetic analysis of Sterrhinae suprageneric lineages (II), altogether five geometrid species from three subfamilies were chosen as outgroups based on recent literature on the relationships among Geometridae subfamilies (II, Appendix 1). Archiearinae are a compact group of diurnal geometrids, unusual in lacking the accessory tympanum (Cook & Scoble, 1992). Geometrinae and Larentiinae have been speculated to be closely related to Sterrhinae or sister groups of it (Szocs et al. 1991, Holloway 1997. ...
... Ferguson (1985) regarded the presence of this extension as common in Geometrinae and absent from other subfamilies. In the abdomen, the shape of the ansa of the tympanal organ shows more variation than that given by Cook & Scoble (1992) as a probable synapomorphy of the Geometrinae (other than the Dysphaniini): usually narrow just above base, broader medially and narrowing to apex. This state occurs in the majority of Pseudoterpnini, but exceptions (principally Calleremites, Dindica, Hypobapta, Paraterpna and Psilotagma) have the apex of the ansa broadened as in some Ennominae or some other geometrine subfamilies. ...
... In some Pseudoterpnini, notably certain Australian genera: Heliomystis, Hypobapta, Protophyta and Rhuma, the ansa is broad at the base and often tapering. This was remarked on by Young (2006), who noted similarity to the state given for the Oenochrominae s.l. by Cook & Scoble (1992). ...
Article
The classification of the genera of the widely distributed Old World tribe Pseudoterpnini is reviewed and 34 genera are recognized. Two new generic synonyms (Sterictopsis Warren and Oxyphanes Turner as synonyms of Rhuma Walker), and 21 new or reinstated combinations are established. Representative moths of all the genera are illustrated in colour (upper side and underside), and genitalia of all genera are illustrated in monochrome (162 figures). All the known species and subspecies of Pseudoterpnini are listed (321), together with their synonyms. The Pseudoterpnini and their characters are assessed, and many genera are newly assigned to the tribe. © 2007 Natural History Museum, London. Journal compilation © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 150, 343–412.
... The anterior abdominal musculature also shows a few sexual variations (Fig. 13) in relation to the male sternum II structure, especially distinct in a lateral one of the inner ventrolongitudinals (sensu Kristensen, 2003) of the sternum II: this muscle originates on the anterior margin of sternum II and inserts in the anterior margin of sternum III in female, but in male on the anterior margin of sternum II and in the posterior margin of a triangular expansion (Fig. 13: arrow). The ansa, one of the important components of the tympanal organ of the Geometridae, has a hammer shaped tip, a characteristic of the larentiine moths (Cook & Scoble, 1992). In addition, a few modifications are also seen in the generic level: in the genus Sauris, the male abdomen often has a pair of setal tufts laterally on the sterna II and IV to VII (Dugdale, 1980); in the genus Trichopterigia ( Fig. 12E-F), a posterior 1/3 to 2/5 of the female 7th abdominal segment is well sclerotized circularly. ...
Article
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The monophyly of the tribe Trichopterygini is considered. The currently used Trichopterygini are polyphyletic. The present Trichopterygini are defined by the following autapomorphies: the enlarged metepimeron in both sexes and a pair of triangular expansions on the male abdominal sternum II, and are redescribed mainly based on adult characters. Phylogenetic relationships among the Japanese genera are analyzed cladistically with the inclusion of two European, three Oriental, and one New Zealand genera. Japanese Trichopterygini are composed of 15 genera, of which two are new, one is upgraded from a subgeneric rank, and one is synonymized: Pseudacasis gen. nov., Paratrichopteryx gen. nov., Paralobophora Inoue (upgraded), Lobophorodes Hampson (a senior synonym of Epilobophora Inoue), Lobophora Curtis, Neopachrophilla Inoue, Phthonoloba Warren, Trichopterigia Hampson, Acasis Duponchel, Otoplecta Warren, Esakiopteryx Inoue, Cladara Hulst, Trichopteryx Hübner, Episteira Warren, and Sauris Guenée. Except for Phthonoloba and Lobophorodes, the Japanese genera are classified into four genus-groups: 1, Lobophora genus-group consisting of Lobophora and Neopachrophilla; 2, Trichopteryx genus-group consisting of Cladara, Esakiopteryx, Paralobophora, Paratrichopteryx and Trichopteryx; 3, Acasis genus-group consisting of Acasis, Otoplecta, Pseudacasis and Trichopterigia; 4, Sauris genus-group consisting of Episteira and Sauris. The Japanese genera are also redescribed mainly by the adult characters and a key to them is provided. Brief information on morphology, bionomics, and distribution of the Japanese species are also given together with their genital figures. Two Trichopteryx species, T. miracula Inoue and T. muscigera (Butler), are transferred to the genus Cladara. A new tribe Heterophlebini is established for the genera Carige Walker, Heterophleps Herrich-Schäffer, Naxidia Hampson, Palaeomystis Warren and their relatives.
... Ennominae, Geometrinae, Sterrhinae, Larentiinae, Desmobathrinae, Oenochrominae and Orthostixinae represent the Indian fauna of Geometridae. Morphologically, the family is defined by the presence of structurally unique tympanal organ at the base of the abdomen (Cook & Scoble, 1992).The group is also of immense economic importance as the larvae of many species are important pests of agricultural crops, ornamental plants and forest trees. Some adult Geometrid moths also feed upon lachrymal secretions from the eyes of mammals. ...
Article
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The present communication deals with the first time reporting of seven Geometrid species, Omiza herois (Prout), Crypsicometa homoema Prout, Biston mediolata Jiang, Xue and Han, Borbacha punctipardaria Holloway, Ephemerophila pallescens Inoue, Paramaxates taiwana Yazaki and Berta apopemta Prout from India. For each of the newly reported species, first reference, material examined and distribution is given whereas, first reference and name of the type species is provided for the respective genus.
... The tympanal organ morphology is of systematic value at higher taxonomic level (subfamilies), consequently I suppose its detailed description to be unnecessary in the current article. The structure of the tympanal organs is thoroughly reviewed in Kennel & Eggers (1933) and Cook & Scoble (1992). The males of some species are characterized by paired setal patches (many Geometrinae) or transverse setal comb (many Ennominae) on sternite A3 (Fig. 9C). ...
Article
The dichotomous keys to 106 species from 62 genera of Ennominae (Lepidoptera: Geometridae) of the Baikal region (Irkutskaya Oblast and Buryatia, Russia) is given. The annotated catalogue including synonyms, the details of examined specimens, data on distribution and hostplants with references is provided. Distribution of Macaria artesiaria ([Denis et Schiffermüller], 1775) in the Baikal region is confirmed. One species is newly combined with genus Hypoxystis Prout, 1915: H. reticulata (Sterneck, 1928) comb. nov. The genus name Scardostrenia Sterneck, 1928 is established to be a synonym of Hypoxystis Prout, 1915 syn. n. Some taxonomic aspects of Charissa turfosaria (Wehrli, 1922), Hypoxystis reticulata (Sterneck, 1928), Synopsia strictaria Lederer, 1853 and Autotrichia heterogynoides (Wehrli, 1927) are discussed. New westernmost boundary of the range of Abraxas karafutonis Matsumura, 1925 is established.
... Pale yellowish. Ansa of tympanic organ in both sexes same, hammerheaded shape s m Cook & Scoble (1992). Pair of small sclerotized processes on 1st tergite, 8th sternite of male with weakly sclerotized band triangular, front broad; 8th tergite with cone or funnel-shaped band; 7th and 8th segments of female abdomen simple and membranous.. ...
Article
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The genus Cidaria Treitschke is revised. Eight species of the genus which occur widely in the Palaearctic and northern India, are recognized, of which one, Cidaria luteata sp. nov., is described as new while two subspecies of C. fulvata (Forster) are elevated to species, C. nugata Felder stat. rev., and C. distinctata Staudinger stat. nov.C. ochripennis Prout is proposed as a junior synonym of C. ochreata Staudinger. C. deletaria Hampson is excluded from the genus. All species of Cidaria and their genitalia are described and illustrated. The cladistic analysis of these eight species of Cidaria is carried out using two data matrices, the first comprising morphological characters alone and the second morphological and distributional characters. The most parsimonious cladogram of Cidaria is selected, and its monophyly defined. The character analysis shows that some distribution characters contribute to resolving the in group node. The choice of multiple trees is discussed.
... The family Geometridae is distributed worldwide and one of the three most speciesrich families of the order Lepidoptera, with approximately 21000 known species (Scoble 1995(Scoble , 1999. A distinctive autapomorphy is the presence of the ansa, a unique structure to Geometridae, located in the tympanal organ, near to the base of the abdomen (Cook & Scoble 1992;Scoble 1999). Furthermore, maybe the best-known character is the peculiar movement exhibited by the larvae, also called inchworms, as they seem to measure the ground by curving their bodies into loops (Scoble 1999). ...
Article
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Despite the urban and landscape impact caused by Drymoea veliterna (Druce, 1885) (Lepidoptera: Geometridae) larvae on trees of the genus Croton L. (Euphorbiaceae) in the Neotropics, there is no information about its biology and reproductive traits. In this study, we describe the life cycle and reproductive traits of this species. Its life cycle lasts approximately 56 days, the larvae develop during 24 days on average and the longevity of the females is 7 days on average. Females have an average fecundity of 207 eggs. Regarding their reproductive system, the presence of previtellogenic eggs in the reproductive tract of the females is emphasized. The bursa copulatrix and a highly sclerotized signum is depicted. The number of spermatophores found in the reproductive tract of the females was quantified. Male and female genitalia were schematized.
... However, there are also several areas where such groupings are ill-defined or where there exists much ambiguity over relationships between the groups. The family Geometridae is clearly monophyletic, defined particularly on the unique structure of the abdominal tympanal organs (e.g., Kennel and Eggers, 1933;Cook and Scoble, 1992;Scoble, 1992;Minet and Scoble, 1999). A review of higher classification in the Geometridae is presented by Holloway (1994Holloway ( , 1996Holloway ( , and 1997 and Minet and Scoble (1999). ...
Article
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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.
... possess two wing venational characters usually present in Geometrinae: R separate from Rs and anastomosing with Sc for a short distance in the forewing and R2-5 stalked in the forewing (Young 2006b). Also, the ansa is similar to the geometrine type i.e., narrow at the base, widening mesally and again tapering apically, and is not the more typical tapering ansal morphology characteristic of Oenochrominae s. l. (Cook & Scoble 1992). As in the Geometrinae, the caecum of the aedeagus is long, slender and tapered and cornuti are reduced, tending not to be discrete rods and spines (Young 2006b). ...
Article
The assembly of a DNA barcode library for Australian Lepidoptera revealed that Oenochroma vinaria Guenée, 1858, as currently understood, is actually a mix of two different species. By analyzing DNA barcodes from recently collected specimens and the 150 year-old female lectotype of O. vinaria, we propose a reliable assignment of the name vinaria to one of these two species. A lectotype is designated for Monoctenia decora, a confirmed synonym of O. vinaria, and a new species, Oenochroma barcodificata sp. nov., is described. This species is only known from Tasmania and New South Wales; its biology and immature stages are described in detail.
... Examples of the structures of tympanal organs in Rhodometrini. The base of the ansa is enlarged and bottle-shaped in the majority of Sterrhinae, but the Rhodometrini condition in which the base is concave and the outer margin is free appears unique within Sterrhinae (a superficially similar structure may appear in Ennominae; seeCook & Scoble, 1992, fig. 9B). ...
Article
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. We analysed the phylogeny of Sterrhinae moths based on molecular dataset of 76 species and 11 genes, combined those with morphology and included the results in a global classification framework. Two new tribes are described, Mecoceratini is transferred from Desmobathrinae to Sterrhinae, and 50 other taxonomic changes are proposed. Sterrhinae are a cosmopolitan group of about 3000 species, with the highest species richness in the Neotropics and the lowest in Oceania.
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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.
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Auf 63 Untersuchungsflächen am Südwesthang des Kilimanjaro wurden Zusammenhänge zwischen der Geometriden-Diversität und floristischer Diversität, Vegetationsstruktur und abiotischen Faktoren untersucht. Besonderes Interesse galt der Bergwaldregeneration, die auf 16 Flächen zwischen 2000 und 2350 m Höhe untersucht wurde. Entlang eines Höhentransektes, das sich vom unteren Kulturland bei 1200 m durch die Bergregenwaldstufe bis in die Ericaceenstufe in Höhen von 3700 m erstreckte, wurde die Änderung der Lebensgemeinschaften in Abhängigkeit von Höhenlage und Nutzungsintensität erfasst. Der Fang der Geometriden erfolgte an Leuchttürmen zwischen 19 und 22 Uhr, im zentralen Untersuchungsgebiet auch mit sieben automatischen Lichtfallen. Zum Anlocken der Falter wurden 15 W- Schwarzlichtröhren verwendet. Um die Leuchttürme herum wurden 400 m2 große Untersuchungsflächen ausgewiesen, auf denen alle Gefäßpflanzen nach Schichten getrennt erfasst und die Vegetationsstruktur aufgenommen wurde. Im zentralen Untersuchungsgebiet wurden an Leuchttürmen in 59 Fängen 118 Morphospezies und 2603 Falter aus der Familie Geometridae gefangen, mit den weniger effektiven Lichtfallen in 203 Fängen 77 Morphospezies und 662 Individuen. Die auf den zentralen Untersuchungsflächen aufgenommenen höheren Pflanzen verteilten sich auf 44 Familien und 98 Morphospezies. Die Artenzusammensetzung der Geometridenzönosen wie auch der Gefäßpflanzen änderte sich im Regenerationsverlauf. Die Artenzahl der Gefäßpflanzen nahm auf den Flächen insgesamt, in Baum-, Strauch- und Krautschicht, der Gruppe der Epiphyten und in den artenreichsten Pflanzenfamilien im Regenerationsverlauf von großen Lichtungen über junge und alte Sekundärwälder zu naturnahen Wäldern zu. Die Diversität der Geometridenzönosen, gemessen mit Fishers Alpha und Hurlbert Rarefaction, sowie die mit Chao 1 extrapolierte Gesamtartenzahl nahm dagegen ab. Die Werte von Fishers Alpha lagen bei sehr niedrigen Werten zwischen 3,4 und 17,8. Die Vegetationsstruktur korrelierte nur schwach mit der Zusammensetzung der Geometridenzönosen. Die Deckung der Moosschicht, ein Indikator für feuchte Verhältnisse, zeigte dabei die stärkste Korrelation. Im Höhentransekt wurden auf den Offenland-, Agroforst- und Waldflächen mit 212 Leuchtturmfängen insgesamt 304 Morphospezies und 8468 Einzeltiere aus der Familie der Geometridae gefangen. Wie im zentralen Untersuchungsgebiet war Mimoclystia corticearia (Larentiinae) die häufigste Art, gefolgt von Darisodes oritropha (Ennominae). Häufig waren auch Chiasmia fuscataria (Ennominae) aufgrund eines Massenvorkommens am unteren Waldrand sowie Chloroclystis derasata (Larentiinae). Bei den auf 51 Untersuchungsflächen durchgeführten Vegetationsaufnahmen wurden 451 Gefäßpflanzenarten aus 116 Familien erfasst. Die Gesamtartenzahlen der Gefäßpflanzen auf den Flächen schwankte zwischen 5 und 53. Sie zeigte auf den Wald- und Offenlandflächen keine Korrelation mit der Höhe, während sie auf den Agroforstflächen mit steigender Höhe signifikant zunahm, was auf die extensivere Nutzung höher gelegener Standorte zurückzuführen ist. Die Artenzahlen vaskulärer Epiphyten, der Aspleniaceen und Rubiaceen waren in mittleren Höhen zwischen 2200 und 2400 m am höchsten, während die Diversität der Baum- und Strauchschicht und der Leguminosen maximale Werte auf den unteren Flächen erreichte. Auch die Diversität der Geometriden war auf den unteren Flächen mit Fishers Alpha-Werten von maximal 40,2 am höchsten. Sie fiel im Transekt auf den Offenlandflächen gleichmäßig ab, während auf den Waldflächen im Bereich zwischen 1800 und 3100 m ­ abgesehen von dem Bereich des zentralen Untersuchungsgebietes ­ die Diversität bei Werten zwischen 8,3 und 12,5 annähernd konstant blieb. Die Analyse der Beta-Diversität und der Gesamtdiversität einzelner Höhenbereiche zeigte, das die Geometridenzönosen zwischen 2000 und 3000 m sehr homogen sind. Die Höhenlage erwies sich als „Master“-Variable, die einen starken Einfluss auf die Vegetationszusammensetzung wie auch die Zusammensetzung und Diversität der Geometridenzönosen hatte. In Bezug auf Geometriden sind insbesondere die feuchten Bergregenwälder am Kilimanjaro wenig divers. Dies kann auf die Insellage des Vulkans inmitten trockener Savannen, sein geringes Alter und die insgesamt arme afrikanische Lepidopterenfauna zurückgeführt werden. Neben dem Schutz der Bergregenwälder, der für die Aufrechterhaltung der Phytodiversität des Kilimanjaro wie auch für die Sicherung der Wasserversorgung Nordtansanias unerlässlich erscheint, sollte auch die traditionelle Agroforstwirtschaft erhalten bleiben, da sie eine hohe Diversität sowohl an Geometriden als auch an Pflanzenarten gewährleistet. The relationship between the diversity of geometer moths and floristic diversity, vegetation structure and abiotic site factors was studied at 63 plots on the southwestern slopes of Mt. Kilimanjaro. Forest regeneration was investigated on 16 plots between 2000 and 2350 m a.s.l.. Furthermore, an altitudinal transect was established starting at 1200 m in agricultural land and extending through the montane forest belt into the ericaceous belt at 3700 m. Along this transect, changes in composition and diversity of moth and plant communities were studied in relation to altitude and human influence. Geometer moths were caught manually between 7 and 10 p.m. using light towers. In the central study area, seven light traps were additionally deployed. Moths were attracted by 15W-blacklight lamps. Around the light towers and traps, plots of 400-m2 were established. On these plots, vascular plants were recorded according to vegetation layers and the physiognomy of the vegetation. In the central study area, a total of 118 morphospecies and 2603 individual geometer moths were caught during 59 catches at the light towers. The less effective light traps yielded 63 morphospecies and 662 individuals during 203 catches. 98 plant species were recorded on the same plots. Species composition of the geometer moth and vascular plant communities reflected different forest regeneration stages. In the course of forest regeneration, the number of vascular plant species increased in the tree-, shrub- and herblayers. There was also an increase in the number of species of epiphytes, pteridophytes and Rubiaceae in the course of forest regeneration, while the diversity of geometer moth communities decreased when using Fisher’s Alpha, Hurlbert Rarefaction, or Chao 1 as species estimator. Values for Fisher’s Alpha varied between 3.4 and 17.8. Vegetation structure was only weakly related to species composition of geometer moths, with the moss layer ­ indicating humid site conditions ­ showing the strongest correlations. The plots along the altitudinal transect were divided into open habitat, agroforestry and forest plots, where 212 catches with light towers yielded 304 morphospecies and 8468 individual Geometridae. Overall, the most abundant moth was Mimoclystia corticearia (Larentiinae). It was followed by Darisodes oritropha (Ennominae), Chiasmia fuscataria (Ennominae), a species very common on the lower forest boundary at Machame gate, and Chloroclystis derasata (Larentiinae). A total of 51 vegetation surveys identified 51 taxa and 116 families of vascular plants. Between five and 53 species of vascular plant species were recorded per plot. The phytodiversity of forest and open habitat showed no correlation with altitude, whereas the agroforestry plots were correlated positively with elevation, as cultivation was less intensive at higher altitudes. The diversity of epiphytes, Aspleniaceae and Rubiaceae peaked at mid-elevation between 2200 and 2400 m, whereas the diversity of the tree- and shrublayer and the Leguminosae decreased with altitude. The diversity of geometer moth communities was also highest at low altitudes, where Fisher’s Alpha reached a maximum of 40,2. Fisher’s Alpha decreased steadily in open habitats with elevation, whereas it’s value remained stable in the forest between 1800 m and 3100 m, except for the very heterogeneous central study area between 2100 and 2300 m. Further analyses of beta-diversity and diversity within different altitudinal belts confirmed a very homogeneous composition of moth communities between 2000 and 3000 m. Elevation proved to be a key parameter strongly influencing both the composition of vegetation and moth communities. Additionally, changes in the diversity of Euphorbiaceae, Pteridophyta and woody plants as well as the tree cover explained a small proportion of Geometer moth diversity. The diversity of geometer moths on Mt. Kilimanjaro, and particularly of its wet montane rain forests, is extremely low. This can be explained with the isolation of the forests in the middle of dry savannas, the young age of the volcano as well as the low diversity of African Lepidoptera in general. In addition to the protection of the montane rain forests in order to preserve phytodiversity and a steady water supply in northern Tanzania, emphasis should be given to the conservation of traditional agroforestry practice, as this practice preserves a high diversity of both plant and geometer moth species.
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The Ditrysia, which comprise the great majority of extant Lepidoptera, are divided into thirty superfamilies (instead of the fifteen to twenty usually recognized). Several of them are conceived in a new sense and one is newly proposed: the Choreutoidea. The Dudgeoneidae are reassigned to the Cossoidea, the Sematuridae and Uraniidae to the Geometroidea, while the Anthelidae and Lasiocampidae are placed in the Lasiocampoidea, a superfamily differing from the Bombycoidea mainly in retaining unfused prothoracic coxae in the larva. The Alucitoidea comprise the Tineodidae, Oxychirotidae and Alucitidae, and the Drepanoidea the Epicopeiidae and Drepanidae. Sister group relationships are recognized between the following superfamilies: Yponomeutoidea / Gelechioidea, Cossoidea / Sesioidea, Alucitoidea / Pterophoroidea, Hesperioidea / Papilionoidea and Drepanoidea / Geometroidea. Reliable monophyletic groups are the Mimallonoidea + Lasiocampoidea + Bombycoidea and the Hedyloidea + Rhopalocera. A few clades are suggested at a higher level, e.g. the Apoditrysia, the Obtectomera (based on two autapomorphies: pupa with fixed abdominal segments 1-4, and imago with modified pulvilli) and the Macrolepidoptera (mainly based on the shape of the first axillary sclerite in the forewing base). Several other changes are introduced: the ' Pseudocossinae' are restricted to the Madagascan genus Pseudocossus Kenrick (possibly belonging in the Brachodidae); the Macropiratinae (stat. n.) are regarded as the most primitive Pterophoridae; the Charideinae are transferred from the Zygaenidae to the Thyrididae; within the Thyrididae, the Argyrotypinae are regarded identical with the Siculodinae (syn. n.). Finally, two families are newly proposed within the Apoditrysia: the Simaethistidae (fam. n.) and Whalleyanidae (fam. n.). The superfamily relationships of these two families have not been determined.
Article
The Australian genus Hypsidia Rothschild, currently assigned to the Pyralidae, is reassigned to the Drepanoidea (Drepanidae sensu Minet (1983), 'Cymatophoridengruppe' of Kennel & Eggers (1933)). Baryphanes Turner, a genus previously mIsplaced in the Noctuidae is synonymized with Hypsidia. A new species (Hypsidia grisea) is described. Eggersops Sick, a genus with a single species, is correctly assigned to the Drepanoidea, and is synonymized with Hypsidia. Hypsidia is divided into the monophyletic erythropsalis group and, for convenience, the niphosema 'group', which may or may not be monophyletic. The distribution of the expanded genus is disjunct, with the erythropsalis group confined to rainforests of northeastern Queensland and the niphosema 'group' to southern Western Australia. The reassignment of Hypsidia (in the revised sense) to the Drepanoidea is based largely on the form of the tympanal organs. The structure of these organs is summarized. The position of Hypsidia within the Drepanoidea remains uncertain, but we briefly explore the question taking particular account of the presence of ocelli in the species of the niphosema 'group', the arrangement of the veins in the hindwing, and the clubbed frenulum of the male.
Article
In order to clarify the nature of the auditory organs of butterflies, a series of ablation experiments was performed. Both electrophysiological and behavioural techniques demonstrated the location of the sense organs in Heliconius erato to be at the base of the hindwing, and to be maximally sensitive to frequencies about 1200 counts/sec with a latency of 8 to 12 msec. A comparison with forms demonstrating acoustic behaviour, such as Ageronia feronia, showed that the major differences were: (1) whereas the auditory organs were similarly inflated sac-like structures, they were located at the base of the forewings, and (2) the latency of these much larger organs way only about 3 msec.
Article
Males of the pyralid moth, Syntonarcha iriastis Meyrick, perch on vegetation at the tops of trees and bushes and produce ultrasound while their wings are spread and while sclerites at the end of the abdomen are spread to expose the genitalia. Exposing the genitalia appears to engage the sound-producing mechanism; the male genitalia and eighth abdominal sternite of this species are greatly modified and include a file and scraper and possible resonating areas. Sounds produced are consistent between individuals and comprise pulses which are narrow in frequency range, the first pulse being at about 42 kHz and two following pulses at about 57 kHz. The signal is detectable with an ultrasound ‘bat’ detector from 20 m. Two tettigoniid species (Orthoptera) at the same site produced ultrasonic calls of similar frequencies at the same times as the moth; differences in time-amplitude patterns could be used by orienting moths to recognize conspecifics. Signalling by male S. iriastis is compared with that of other pyralid species in which females are attracted to signalling males. The behaviour of S. iriastis males differs from that of other pyralids in that they do not signal in groups or from a resource attractive to females, and do not possess glands known to produce a ‘calling’ pheromone. It is suggested, because of these differences, that sound production in this species does not function at close range, as argued for the wax moth Achroia grisella (Fabricius), but instead as a long-distance calling signal.
Article
The ears of moths we tested in Canada and Cte d'Ivoire are most sensitive to sounds between 20 and 40 kHz, and much less sensitive to sound over 65 kHz. The insectivorous bats most commonly encountered in these (and other) locations use high intensity, frequency modulated echolocation calls with frequency components in the 20–40 kHz range, making them detectable by the most sensitive tympanate moths up to 40 m away. In Africa bats such as species in the Nycteridae, Megadermatidae, and some in the Hipposideridae, use low intensity calls with high frequency components, and these species are not detectable by moths at over 2 m. The hearing ability of moths may significantly influence the feeding efficiency of bats, and changes in the intensity and frequency components of bat echolocation calls can drastically reduce the range at which bats are detected, and thus the time available to the moths for evasive behaviour (Fig. 4). The use of low intensity, high frequency echolocation calls may constitute a bat counter-maneuver against insects tuned to bat calls.
Article
Two species of neotropical moths,Antaea lichyi Franclemont andHapigia curvilinea Schaus (Notodontidae) from Panamá possess paired, cup-like extensions of their pleural segments, reminiscent of mammalian pinnae, that project laterally from the perimeter of the tympanic membrane. These cups are homologous with the post-spiracular abdominal hoods of typical noctuid moths, and may be related to the orientation of the notodontid ear, where the tympanic membrane faces ventrally toward the mid-line of the moth's body and potentially reduces the reception of incident sound energy. Removal of the cups did not alter the neurologically determined best frequencies of either species, nor did it affect the shape ofH. curvilinea's directional sensitivity or its spatial point of maximal sensitivity. Polar threshold curves to stimulus tones of 30 kHz, however, revealed a reduced overall directionality, and thus indicate that the cups may impart a degree of sound localization ability. Both removal and blockage of the external structures result in reduced sensitivities (i.e. increased thresholds) to frequencies higher than approximately 30 kHz. All individuals ofH. curvilinea tested became deaf to tones over 115 kHz when the cups were ablated or blocked. Neotropical moths are exposed to potentially heavy predation pressure from insectivorous bats that characteristically emit faint and/or high frequency echolocation signals, some in excess of 70 kHz (e.g., foliage-gleaning Phyllostomatidae). In certain notodontids, the development of external auditory adaptations that enhance the reception of faint, high frequency sounds may be an evolutionary response to predation pressures by these bats.
Article
The separate impulses contributed by the A1 and A2 acoustic sense cells in the tympanic organs of the noctuids, Autographa pseudogamma and Noctua c.-nigrum, and by the A1, A2, and A3 sense cells in the tympanic organ of the geometrid, Ennomos magnarius, were identified and counted from oscillograms grams made as the moths were exposed to ultrasonic pulses of different intensities. These data were used to construct curves relating the response/intensity characteristics of the less sensitive acoustic sense cells to that of the most sensitive unit, A1. The A2 sense cells of the noctuids were found to be from 20 to 30 dB less sensitive than A1 at sound frequencies to which these ears are most sensitive. In the geometrid it was found that the A2 sense cell was 15 dB less sensitive than A1 and 12 dB more sensitive than A3. Only traces of the response of the fourth geometrid acoustic sense cell (A4) could be identified at high sound intensities. In both noctuids and geometrids the acoustic sensitivity of A2 relative to A1 remained unchanged when tested at selected ultrasonic frequencies between 28 and 50 kHz. This confirms the conclusion that the ears of these moths are incapable of pitch discrimination over this frequency range. Each of the systems had a dynamic range of 40 to 45 dB, that of the geometrid showing greater range overlap of the four A cells and hence greater capacity for sound intensity discrimination.
Crambidae, premiere partie (Lepidoptera Glossata) Annales de la Sociiti Entomologique de France
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Die Bedeutung der Tyrnpanalorgane der Lepi-dopteren fur die Systematik
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