Systematics and taxonomy of the subfamily Larentiinae (Lepidoptera, Geometridae)

Sektion Hymenoptera

Morpho-functional analysis of the male terminalia in the larentiine moths (Lepidoptera, Geometridae)

The characters of skeleton and musculature of the male genitalia have proven useful for the higher classification of Lepidoptera.

  1. The musculature of the male genitalia was reviewed for the tribe Xanthorhoini and related tribes (Lepidoptera, Geometridae, Larentiinae). The genitalia morphology of males of 11 species was discussed and illustrated, and nine paired and unpaired genital muscles identified. Muscles m1, m2(10), m5(7), m6(5), m7(6), m8(3) and m21 have similar position in all species considered in the paper. Comparative morphology of the male terminalia and position of extensors of the valvae m3(2) and flexors m4 confirmed the previously uncertain separation of Euphyiini and Scotopterygini. Cataclysmini share musculature characters with the tribe Xanthorhoini (Schmidt 2013).
  2. The structure and musculature of the male terminalia were described and illustrated in 11 genera of the tribe Cidariini (Lepidoptera, Geometridae, Larentiinae) from the Holarctic and Oriental regions. Nine genital muscles were identified: m1, m2(10), m3(2), m4, m5(7), m6(5), m7(6), m8(3) and m21. Variation in the insertion of the muscles m1, m3(2), m4, m5(7), m6(5) and m8(3) on the sclerites in several generic groups of the tribe Cidariini was discussed, revealing that the Thera species group does not share some apparently cidariine characters. A comparative analysis of the musculature in the tribes Cidariini and Xanthorhoini questions the sister relationship of these tribes that was suggested by earlier studies (Schmidt 2014a).
  3. The skeleton and musculature of the male terminalia were examined and depicted in seven Palaearctic species from the tribes Eupitheciini Pierce, Melanthiini Duponchel, Perizomini Herbulot and Rheumapterini Herbulot (Lepidoptera, Geometridae, Larentiinae) characterized by the presence of the eupitheciine labides in the male genitalia. Nine genital muscles, typical of the subfamily, were identified: m1, m2(10), m3(2), m4, m5(7), m6(5), m7(6), m8(3) and m21. The places of attachment of the muscles m3(2), m4 and m5(7) to the genital sclerites afford useful characters for the higher classification of this group (Schmidt 2014b).
  4. The tribe Eupitheciini and allies. (4.1) The structure and musculature of the male terminalia were described in six Palaearctic and Indo-Australian eupitheciine species from the genera Eupithecia Curtis, Gymnoscelis Mabille, and Pasiphila Meyrick (Lepidoptera: Geometridae: Larentiinae). The tergal flexors of the valvae, muscles m4, running to the medial part of the transtilla or to the membrane dorsad of the transtilla are identified as a potential synapomorphy for the tribe Eupitheciini. The tergal extensors of the valvae, muscles m3(2), running deep into the costa valvae is a possible synapomorphy for the genus Gymnoscelis. The genus Pasiphila does not share all eupitheciine characters, with the tergal flexors of the valvae, muscles m4, extending from the dorsal parts of the vinculum. The Eupithecia haworthiata species-group differs from other species-groups by the shape of the uncus and the sternal extensors of the valvae, muscles m8(3), with lateral fibres longer than the medial ones (Schmidt 2020b). (4.2) The structure and musculature of the male terminalia are described in three type species of the eupitheciine genera Chloroclystis Hübner, Sigilliclystis Galsworthy, and Ziridava Walker (Lepidoptera: Geometridae: Larentiinae). The following traits are characteristic for all three genera: (1) the tergal flexors of the valvae, muscles m4 running to the medial part of the transtilla or membrane anterior to the transtilla; (2) the sternal extensors of the valvae, muscles m8(3), broadly running to the inner wall of the saccus, with long lateral bundles; (3) the twisted muscles m7(6) running from the lower margin of the saccus. The peculiar structure of the transtilla is described and illustrated (Schmidt 2023).
  5. The skeleton and musculature of the male terminalia were described and illustrated for the tribes Operophterini Packard, Phileremini Pierce and Triphosini Tutt (Lepidoptera, Geometridae, Larentiinae), with Solitaneini Leraut included in the discussion. Nine genital muscles, typical of the subfamily, were identified. The previously uncertain separation of Triphosini from Rheumapterini Herbulot was confirmed, based on the morphology of male terminalia, including the position of extensors and flexors of the valvae. Based on the morphology, a close relationship of the Phileremini, Solitaneini, Triphosini and also Operophterini is suggested (Schmidt 2017).
  6. The skeletomuscular system of the male genitalia and the structure of the female genitalia of Hydriomena impluviata (Denis & Schiffermüller) from Europe were compared with those of A. subochraria and A. uncinata from Australia, suggesting that Anachloris does not belong to the tribe Hydriomenini (Schmidt 2001).

Taxonomic revisions of the larentiine genera (Lepidoptera, Geometridae)

  1. Anachloris Meyrick. The genus Anachloris Meyrick containing the species A. subochraria (Doubleday) and A. uncinata (Guenée) is reviewed, and a new species A. tofocolorata is described and illustrated. Keys to species and distribution maps, as well as information about the biology are provided (Schmidt 2001).
  2. Chaetolopha Warren. The Australasian species associated previously with the geometrid moth genus Chaetolopha Warren are reviewed. Six Australian taxa, including C. oxyntis (Meyrick), C. pseudooxyntis, C. leucophragma (Meyrick), C. decipiens (Butler), C. emporias (Turner) with a synonym Scotocyma pteridophila (Turner), and C. niphosticha (Turner) are assigned to Chaetolopha. Eight Papuan high altitude species associated with Chaetolopha or Tephroclystia (Hübner) in original combination and included later in Chaetolopha are reassigned to a new genus Parachaetolopha. These include P. flavicorpus (Warren), P. coerulescens (Warren), P. turbinata (Prout), P. tafa (Prout), P. ornatipennis (Warren), P. anomala Prout, P. nepenthes (Prout), P. peregrina (Prout), with new species P. petasitruncula, P. ferruginoapex, P. collatichaeta, and P. spinosicornutus, described. A phylogenetic analysis of Chaetolopha confirms the proposal of the new genus Parachaetolopha. One species associated with Eupithecia Curtis in original combination and included later in Chaetolopha is transferred to the genus Tripteridia Warren. Lectotypes are designated for all Chaetolopha and Parachaetolopha species except for C. niphosticha, P. turbinata, P. tafa, P. anomala, P. peregrina and P. nepenthes, where a holotype was indicated in the original description. Descriptions of the genera Chaetolopha and Parachaetolopha, as well as a key to Chaetolopha species, are provided. All the known species are redescribed, the new ones are described, and all species are illustrated. Fern-feeding in Australian Lepidoptera and vertical distribution of Chaetolopha and Parachaetolopha species are discussed (Schmidt 2002).
  3. Scotocyma Turner. The Australasian genus Scotocyma Turner is revised, containing the species S. albinotata (Walker), S. legalis (Warren), S. asiatica Holloway, S. scotopepla Prout, stat. n., S. manusensis Prout, stat. n., S. mimula (Warren), stat. n., and S. miscix Prout. Scotocyma euryochra Turner, S. idioschema Turner, S. ischnophrica Turner, and S. transfixa Turner, are regarded synonyms of S. albinotata. Four species are described as new: S. samoensis, S. sumatrensis, S. rutilimixta, and S. longiuncus. Lectotypes are designated for S. scotopepla, S. manusensis, and S. miscix. All species are illustrated, keys to species and distribution maps are provided. A phylogenetic analysis was performed to test the monophyly of the genus and to examine distribution patterns of the species. A biogeographic discussion is included. The tribal position of the genus is clarified and relationships to closely related genera are discussed (Schmidt 2005, 2006b, 2006c, 2007).
  4. Visiana Swinhoe. (4.1) The study clarifies the taxonomy of the Indo-Pacific species-group Visiana sordidata (Moore), a representative of the geometrid subfamily Larentiinae. Historically, V. sordidata comprised three subspecies: V. s. robinsoni (Prout), V . s. inimica (Prout), and V. s. tamborica (Prout). However, examination of about 80 specimens revealed that all supraspecific taxa should be regarded as distinct species, V. robinsoni, V. inimica, and V.  tamborica. Furthermore, the specimens from Borneo (Malaysia) are assigned to a new species, V. hollowayi. The five Visiana species belong to two different species-groups for which the diagnostic characters are defined. Lectotypes are designated for V. sordidata, V. robinsoni, and V. tamborica. The new species, V. hollowayi, is described and illustrated (Schmidt 2006e). (4.2) A new species V. fuscata is described and illustrated (Schmidt 2009). (4.3) The following specimens from the Papua New Guinea region deposited in the Natural History Museum (London, UK) and Australian National Insect Collection (Canberra, Australia) were examined: V. hyperctenista (Prout) from the Bismarck Archipelago, V. vinosa (Warren) from the eastern part of Papua New Guinea, V. ranensis (Prout), stat. nov. from the Moluccas (Indonesia), and four newly described species V. horista, from Bougainville Island, V. grandivinosa from New Britain, V. variovinosa from the eastern part of Papua New Guinea, and V. morobensis from the Morobe province. The lectotype is designated for V. hyperctenista. Redescriptions of V. hyperctenista, V. vinosa and V. ranensis are provided, the four new species are described, figures of all seven species are shown. The species V. hyperctenista and V. horista are treated as a sister-group, the diagnostic characters are defined. A checklist of the known Papuan species of the genus Visiana is provided (Schmidt 2013). (4.4) Based on the study of morphological characters and DNA barcode (CO1) data, the present review revealed the existence of at least three species of Visiana Swinhoe (Lepidoptera: Geometridae: Larentiinae) in Australia. Visiana brujata (Guenée) is redescribed, and two species V. incertata (Walker), stat. rev. and V. repentinata (Walker), stat. rev. are resurrected from synonymy with V. brujata. Visiana breviaria (Walker), syn. rev., previously cited as a synonym of V. brujata, is now considered a synonym of V. incertata. Visiana brujata and V. incertata show close affinities with the sordidata group of species, whereas V. repentinata belongs to the vinosa species group. Images of adults and genitalia of all types are illustrated and the presence of the gnathos in the genus Visiana is discussed (Schmidt 2015). (4.5) The present study contributes to the knowledge of the Indo-Pacific genus Visiana Swinhoe based on historic specimens collected in Indonesia and deposited in the collections of the Natural History Museum (London, UK) and Naturalis Biodiversity Center (Leiden, the Netherlands). Two new species V. cyclopsensis, from the Cyclops Mountains (West Papua) and V. minahasensis, from Minahasa (Sulawesi) are described and illustrated. These belong to two different species groups. An updated checklist of the genus Visiana, comprising 18 species is included (Schmidt 2018).
  5. Crasiloga Warren. The Australasian genus Crasilogia Warren comprising the species C. dispar Warren, C. flavipennis Warren, C. fulvitincta Joicey & Talbo, C. fumipennis Warren, C. gressitti Holloway, C. simplex (Prout), and two newly described species C. parasimplex and C. diversisimplex, is reviewed based on the museum specimens, including primary types. Lectotype is designated for C. simplex (Schmidt 2020a).
  6. Ziridava Walker. The review summarizes the current state of knowledge of the genus Ziridava Walker. Two new species, namely Z. mauriakra from the Philippines and Z. orchidensis from the Lanyu Island (Taiwan), are described and illustrated. They share the characteristics of the genus Ziridava, including the uncus reduced to a short trident, the long, slender valvae with costa distinctly broadened for basal one-fourth, the broad juxta constricted medially and the saccus rather rectangular with rounded corners in males and the square-shaped papillae anales, the ductus seminalis arising from a lateral lobe of corpus bursa, with a signum comprised of scobinate patches on the dorsal and ventral surface of corpus bursa in females (Schmidt & Tautel, 2022).

Checklists of the Larentiinae (Lepidoptera, Geometridae)

  1. An annotated and illustrated list of the primary type specimens of geometrid moths deposited in the Queensland Museum (Australia, Brisbane) (Lepidoptera, Geometridae).The study is a first step towards making available taxonomic and photographic information on type specimens of geometrid moths (Lepidoptera, Geometridae) deposited in Australian insect collections. The current paper provides an annotated and illustrated list of 62 primary types of geometrid moths deposited in the Queensland Museum insect collection in Brisbane. Included are representatives of 23 Ennominae, 12 Oenochrominae, 11 Geometrinae, 10 Larentiinae, and six Sterrhinae. In order to preserve stability of nomenclature, lectotypes are designated for the following eighteen taxa described by A. J. Turner: Aglossophanes adoxima, Anisodes lechriostropha, A. rhodobapta, Anomogenes morphnopa, Dichromodes lechria, D. loxotropha, D. mesotoma, D. tritospila, Eois trissomita, Epidesmia phoenicina, Gelasma selenosema, Idiochroa rufifrons, Pisoraca sticta, Poecilasthena ischnophrica, Scopula loxographa, Taxeotis pleurostigma, T. spodoides, Tephroclystia aphanes. Photographs of both the upper and underside of each primary type are shown and data from all labels are given. Additionally, for the subfamily Larentiinae, information on paratypes is provided (Schmidt 2012).
  2. List of primary types of the larentiine moth species (Lepidoptera: Geometridae) described from Indonesia – a starting point for biodiversity assessment of the subfamily in the region. The study provides a first inventory of the primary type specimens of larentiine moth species (Lepidoptera: Geometridae) described from Indonesia. The list of species described from Indonesia is arranged alphabetically by the tribe, genus, and species, and presents data on 251 species and subspecies. For each species type status, type locality, depository, and a full reference to the original description are listed. Synonyms with Indonesian type localities are included. The study indicates a large part of the Indonesian geometrid fauna belong to the tribe Eupitheciini (Schmidt 2015b).
  3. Records of larentiine moths (Lepidoptera: Geometridae) collected at the Station Linné in Sweden. The island of Öland, at the southeast of Sweden, has unique geological and environmental features. The Station Linné is a well-known Öland research station which provides facilities for effective studies and attracts researchers from all over the world. Moreover, the station remains a center for ecotourism due to extraordinary biodiversity of the area. The present paper is aimed to support popular science activities carried out on the island and to shed light on diverse geometrid moth fauna of the Station Linné. As an outcome of several research projects, including the Swedish Malaise Trap Project (SMTP) and the Swedish Taxonomy Initiative (STI) conducted at the Station Linné, a list of larentiine moths (Lepidoptera: Geometridae) collected on the territory of the station is presented. Images of moths from above and underside are shown. Of the totally 192 species registered for Sweden, 41 species (more than 21%) were collected in close proximity to the main building of the Station Linné. Malaise trap sampling of Lepidoptera is discussed (Schmidt 2016a).

Biology of the larentiine moths (Lepidoptera, Geometridae)

Larval food plants of Australian Larentiinae (Lepidoptera: Geometridae) – a review of available data. In Australia, the subfamily Larentiinae (Lepidoptera: Geometridae) comprises over 45 genera with about 270 species described so far. However, life histories of the Australian larentiine moths have barely been studied. The paper presents a list of larval food plants of 51 Australian larentiine species based on literature references, data from specimen labels and own observations. Some Australian habitats are shown. Possible relationships among the taxa based on food preference of the larvae are discussed. Additionally, a list of Australasian larentiine species from the genera occurring in Australia and their food plants is presented (Schmidt 2016b).

Peculiarities of the male genitalic characters of larentiine species (Lepidoptera, Geometridae)

  1. Gnathos. One century ago, F.N. Pierce in his well-known monograph on the genitalia of Geometridae (Lepidoptera), divided the family into two major subdivisions, the Gnathoi and the Agnathoi, depending on the presence or absence of the gnathos in males. In his study, Pierce assigned the Larentiinae to the Agnathoi based on the apparent absence of the gnathos in this subfamily. A re-examination of the male genitalic characters of numerous larentiine species representing 14 different tribes provided, contrary to Pierce’s results, evidence for the presence of the gnathos in Larentiinae. Illustrations of the gnathos (or its remnants) in male genitalia of selected species are provided and the value of the uncus and gnathos for inferring phylogenetic relationships is discussed (Schmidt 2014b, 2015).
  2. Socii. The definition of the term ‘socius’ is evaluated. Considering several recent publications, in which it was assumed that the socii were generally lacking in the male genitalia of the larentiine moths of the family Geometridae, the male genitalic characters of a number of larentiine species representing all the European tribes have been re-examined. The study reveals that the socii are widely present in the European Larentiinae. The presence of the socii in Larentiinae is reviewed. Illustrations of the uncus and socii in the male genitalia of selected species are provided (Schmidt 2017b).

Rapid biodiversity assessment. Biodiversity Discovery and Information System

1. IndoBioSys.

(1a). A streamlined collecting and preparation protocol for DNA barcoding of Lepidoptera. 

IndoBioSys is a German-Indonesian research project that is conducted by the Museum für Naturkunde in Berlin and the Zoologische Staatssammlung München, in close cooperation with the Research Center for Biology – Indonesian Institute of Sciences (RCB-LIPI, Bogor). The project is focusing on establishing a vertebrate and invertebrate diversity discovery pipeline and a biodiversity information system in Indonesia. Samples are processed through an integrated sorting pipeline that has been set up and optimized at the Zoologische Staatssammlung in Munich, Germany, and tested at the Museum Zoologicum Bogoriense in Cibinong, Indonesia.

A general collecting and preparation protocol for DNA barcoding of Lepidoptera as part of large-scale rapid biodiversity assessment projects is provided, and a comparison with alternative preserving and vouchering methods. About 98% of the sequenced specimens processed using the present collecting and preparation protocol yielded sequences with more than 500 base pairs. The study is based on the first outcomes of the IndoBioSys (Schmidt et al. 2017c).

(1b). The Mt Halimun-Salak Malaise Trap project – releasing the most species rich DNA Barcode library for Indonesia.

The Indonesian archipelago features an extraordinarily rich biota. However, the actual taxonomic inventory of the archipelago remains highly incomplete and there is hardly any significant taxonomic activity that utilises recent technological advances. The IndoBioSys project was established as a biodiversity information system aiming at, amongst other goals, creating inventories of the Indonesian entomofauna using DNA barcoding. Here, we release the first large scale assessment of the megadiverse insect groups that occur in the Mount Halimun-Salak National Park, one of the largest tropical rain-forest ecosystem in West Java, with a focus on Hymenoptera, Coleoptera, Diptera and Lepidoptera collected with Malaise traps. From September 2015 until April 2016, 34 Malaise traps were placed in different localities in the south-eastern part of the Halimun-Salak National Park. A total of 4,531 specimens were processed for DNA barcoding and in total, 2,382 individuals produced barcode compliant records, representing 1,195 exclusive BINs or putative species in 98 insect families. A total of 1,149 BINs were new to BOLD. Of 1,195 BINs detected, 804 BINs were singletons and more than 90% of the BINs incorporated less than five specimens. The astonishing heterogeneity of BINs, as high as 1.1 exclusive BIN per specimen of Diptera successfully processed, shows that the cost/benefit relationship of the discovery of new species in those areas is very low. In four genera of Chalcidoidea, a superfamily of the Hymenoptera, the number of discovered species was higher than the number of species known from Indonesia, suggesting that our samples contain many species that are new to science. Those numbers show how fast molecular pipelines contribute substantially to the objective inventorying of the fauna giving us a good picture of how potentially diverse tropical areas might be (Cancian de Araujo et al. 2018b).

2. Identification of sawflies and horntails (Hymenoptera, ‘Symphyta’) through DNA barcodes: successes and caveats.

The ‘Symphyta’ is a paraphyletic assemblage at the base of the order Hymenoptera, comprising 14 families and about 8750 species. All have phytophagous larvae, except for the Orussidae, which are parasitoids. This study presents and evaluates the results of DNA barcoding of approximately 5360 specimens of ‘Symphyta’, mainly adults, and 4362 sequences covering 1037 species were deemed of suitable quality for inclusion in the analysis. All extant families are represented, except for the Anaxyelidae. The majority of species and specimens are from Europe, but approximately 38% of the species and 13% of the specimens are of non‐European origin. The utility of barcoding for species identification and taxonomy of ‘Symphyta’ is discussed on the basis of examples from each of the included families. A significant level of cryptic species diversity was apparent in many groups. Other attractive applications include the identification of immature stages without the need to rear them, community analyses based on metabarcoding of bulk samples and association of the sexes of adults (Schmidt et al. 2016).

3. From field courses to DNA barcoding data release for West Papua – making specimens and identifications from university courses more sustainable.

The diversity of insects collected during entomological field courses at the University of West Papua (UNIPA), Indonesia, is studied using DNA barcoding tools. The results were compared with public data available for West Papua in the Barcode of Life Data System. During two training courses in 2013 and 2015, 1,052 specimens of insects were collected at eight sites near Manokwari in northern West Papua. The DNA sequences obtained from these specimens were assigned to 311 Barcode Index Numbers (BINs) and represent species in 27 families of Lepidoptera, Hymenoptera and Coleoptera. Of those BINs, 294 (95%) were new to West Papua. The study suggests that DNA barcoding applied to university courses achieves several goals, including capacity building and hands-on experience in molecular biodiversity assessment. In addition, it can provide valuable biodiversity data that are globally available to researchers for further studies (Cancian de Araujo et al. 2018a).


VIETBIO [Innovative approaches to biodiversity discovery and characterization in Vietnam] is a bilateral German-Vietnamese research and capacity building project focused on the development and transfer of new methods and technology towards an integrated biodiversity discovery and monitoring system for Vietnam.

(4a). Using Malaise traps for collecting Lepidoptera (Insecta)

The present study deals with the potential of employing Malaise traps for collecting butterflies and moths for morphological analysis and presents a protocol for preparing Macrolepidoptera from Malaise trap samples that were preserved in ethanol. About 80 specimens of Lepidoptera, including Nymphalidae, Geometridae, Hesperiidae, Erebidae, Noctuidae, Pyralidae and Tortricidae, were mounted, following the protocol. All specimens with robust wings and contrasting wing patterns were well suited for the study of external morphology, regardless of the family. The specimens used in this study were collected in highland forest areas of central Vietnam with a little known entomofauna, as part of the German-Vietnamese biodiversity project ‚VIETBIO‘. The study offers new methodological approaches in an attempt to make the most of the material that was obtained using Malaise traps (Schmidt et al. 2019).

(4b). Contributions to the biodiversity of Vietnam – Results from VIETBIO inventory work and field training in the Cuc Phuong National Park

The editorial paper in the Collection of the Biodiversity Data Journal reports original results from recent biodiversity recording and survey work undertaken in Cuc Phuong National Park, northern Vietnam, under the framework of the VIETBIO project. The main purpose is to make primary biodiversity records openly available, including several new and interesting findings for this biodiversity rich conservation area (Duwe et al. 2022).