We present a new, online, open access portal to the geometrid moths of the world (Lepidoptera: Geometridae). The portal provides access to the global database including data on classification, valid genera and species, synonyms and type localities, and to an online list of valid names. The value of this online facility, inter alia, is demonstrated by using the underlying data to update our understanding of the global and regional species richness of the family. We also review, clarify and list the family-group names (family, subfamily, tribe) of the Geometridae to provide users with a better understanding of the higher category framework when they access the database. Currently, the Geometridae database includes 34,897 available species-group names. Of these, 7,891 are considered synonyms (23% of available names), giving a total of 27,006 valid species-group names, which in turn comprise 23,872 species names and 3,134 subspecies names. The world fauna is classified into 2,019 genera, 92 tribes and 9 subfamilies. Our paper also includes an annotated list of all 202 subfamily, tribe and subtribe names in systematic order. One hundred family-group names (49.5%) are considered synonyms. Although most geometrid species were described in the 19th and 20th centuries, the average number of new species descriptions from 2000–2022 has been fairly constant at around 80 per year, implying there is still much more hidden geometrid diversity yet to be discovered.
Ein neues, frei zugängliches Online-Portal zu den Geometriden der Welt (Lepidoptera: Geometridae) wird hier vorgestellt. Das Portal bietet Zugang zu einer globalen Datenbank mit Informationen zur Klassifikation, zu sämtlichen validen Gattungs- und Artnamen, Synonymen und Typuslokalitäten. Die Bedeutung dieser öffentlich zugänglichen Datensammlung liegt darin, unsere Kenntnisse über den globalen und regionalen Artenreichtum der Familie Geometridae zu verbessern. Außerdem wurden die Namen der höherrangigen Taxa (Familie, Unterfamilie, Tribus) überarbeitet, ihr Status geklärt und aufgelistet, um den Benutzern ein besseres Verständnis der Phylo-genie nach dem augenblicklichen Stand der Forschung zu vermitteln. Derzeit umfasst die Familie der Geometriden 23.872 beschriebene und valide Arten und 3.134 validierte Unterarten, während weitere 7.891 Namen als Synonyme betrachtet werden (23 % der verfügbaren Namen). Die weltweite Fauna ist in 2.019 Gattungen, 92 Triben und 9 Unterfamilien unterteilt. Unsere Arbeit enthält zudem eine kommentierte Liste aller 202 Unterfamilien-, Tribusund Subtribusnamen in systematischer Reihenfolge. 100 dieser Namen (49,5 %) werden als Synonyme betrachtet. Während die meisten Geometriden-Arten im 19. und 20. Jahrhundert beschrieben wurden, liegt die durchschnitt-liche Zahl der neuen Artbeschreibungen von 2000 bis 2022 ziemlich konstant bei etwa 80 pro Jahr, was bedeutet, dass eine bedeutende Zahl von Geometriden noch darauf wartet, entdeckt zu werden.
Introduction
Card indexes of insect species exist in many museums, such as that in the Natural History Museum, London (Gaston et al. 1995; Scoble 1999). Some of the available published printed catalogues have evolved from such card indexes (e.g., Arctiinae: Watson & Goodger 1986; Geometridae: Scoble 1999) or were compiled otherwise (e.g., Poole 1989; Sobczyk 2011). In recent years, several global Lepidoptera species databases have become available online, notably for Papilionoidea (HÄuser et al. 2005), Pyraloidea (Nuss et al. 2022), Bombycoidea (Kitching et al. 2018), Tortricidae (Gilligan et al. 2018) and Gracillariidae (De Prins & De Prins 2006–2022) in addition to a growing number of regional online catalogues (e.g., De Prins & De Prins 2011–2022; Dominick et al. 1983; Holloway 1986–1999; Karsholt & Nieukerken 2013).
Species catalogues vary somewhat in their content, but provide basic taxonomic information to many of the following fields: valid names and synonyms, homonyms, authors, dates, references, type specimens, type localities and distribution. Depending on their comprehensiveness, they also enable access to consolidated data on species richness and how species are arranged in higher taxonomic categories. Ideally, phylogenetic relationships are also reflected. Such extensive and structured content means that species catalogues can provide the foundation for global species richness estimates, and information on patterns in species description over time (Gaston et al. 1995). The number of insect species in the world is unknown, but current estimates rely significantly on verifiable information about the richness of described species (GARCIA et al. 2020). Furthermore, species catalogues may help to identify collection biases and understudied regions of the world, indicating where future field campaigns exploring the biodiversity should take place.
Species catalogues are lacking for most insect groups for several reasons. First, taxonomic information is scattered across a wide spread of journals and other literature. Second, many articles are difficult to source (although the Biodiversity Heritage Library provides a major improvement in accessibility, https://www.biodiversitylibrary.org/). Third, articles may be written in languages that are not widely understood. Fourth, numerous regional classifications may conflict with each other, making a consensus difficult. Fifth, data included in original publications are often superficial and do not allow unambiguous identification, a point especially true for the early literature. Finally, and perhaps most importantly, compiling such diverse and scattered information into a single entity is a time- consuming and labour-intensive task, requiring expert knowledge of a particular taxon, often with few specialists who are usually suffering a high workload.
Apart from our personal interest in the Geometridae, we consider that the family is particularly suitable for an online facility, with more than 23,000 species accounting for nearly 15% of all Lepidoptera species (Nieukerken et al. 2011). Geometrid moths are the second-largest lepidopteran family (after Erebidae) in terms of described species. Furthermore, this global diversity has been summarised in two species catalogues dating back 23 and 15 years, respectively: (1) “Geometrid Moths of the World: A Catalogue” (Scoble 1999) and (2) “Online list of valid and available names of the Geometridae of the World” (Scoble & Hausmann 2007). Geometrid moths have received considerable attention during the last decades, and the use of molecular data, in particular, has advanced our understanding of the family significantly, so both the catalogue and the online list need updating. Before describing the scope and content of the new database (“Online taxonomic facility of geometrid moths”), we summarise the contents of the two earlier works.
Geometrid Moths of the World: A Catalogue (Scoble 1999). The Catalogue is a massive two-volume printed monograph of 1,016 pages, plus a 129-page index, and it is based on the British Museum of Natural History's card index to genera and species of Geometridae housed in the Natural History Museum, London (NHMUK) [previously known as the British Museum (Natural History)]. The Catalogue provides access to the available names of geometrid moths of the World, including approximately 35,000 species-group names (including synonyms). The following data for each species and subspecies were entered (called ‘Fields’ in the Global Taxonomic Facility): Subfamily, Genus, Original genus, Taxon name (species or subspecies), Author, Date, Title of reference, Volume number of reference, Series number of reference, Part number of reference, Page number of original citation, Whether original reference was checked (yes/no), Whether a junior synonym (yes / no), If a junior synonym, of what species/ subspecies, Number of names synonymised, Drawer number in BMNH collection, Depository of type(s), Type status (holotype, syntype(s) etc.), Type sex (if known), Type locality (country, state, place), Zoogeographical region. The printed Catalogue does not include all above-mentioned fields.
Entries are ordered alphabetically by genus and, within each genus, by species. The Catalogue also includes genus names and genus synonyms, citing author(s), date, reference to the original description, and type species. It includes extensive unpublished data extracted from details in the card index, but these were not separated from the published data. For a small number of species, larval foodplant data are provided. The publication also includes a morphological diagnosis of the family, a summary of the patterns of species description in the Geometridae derived from Gaston et al. (1995) and a list of genera in systematic order, following the order in which the collection in the British Museum of Natural History was laid out.
Online list of valid and available names of the Geometridae of the world (Scoble & Hausmann 2007). The global species list is displayed as an Excel spreadsheet derived from the above-mentioned Catalogue (Scoble 1999), containing information on the subfamily, genus, species, author and year of description for all validated geometrid moth species up to 2007, covering 22,951 species. Unlike the Catalogue (Scoble 1999), it does not give information on the original publication, depository of type(s), type status (holotype, syntype(s), etc.), subspecies, junior synonyms, or type locality. This list is now available the new Forum Herbulot website at https://geometroidea.smns-bw.org/.
In the present paper, we: i) announce the forthcoming launch of an online facility on Geometridae, describe its structure and comment on its value and purpose; ii) complement this open access biodiversity data release with an up-to-date review of Geometridae species diversity and subfamily-level phylogeny; and iii) provide an updated systematic list of family-group names. Finally, we present our plans for the future development of the facility.
Material and methods
In order to update the printed and online world catalogues of geometrid moths (Scoble 1999; Scoble & Hausmann 2007), we searched the Zoological Record database ( https://clarivate.com/webofsciencegroup/solutions/webofscience-zoological-record/) for all records published after 2005, using the keyword “Geometridae”, and compared the contents of each publication against the “Online list of valid and available names of the Geometridae of the World” (Scoble & Hausmann 2007). In addition, we included all publications available to us from 1998, which were not covered by the Zoological Record database. In total, data from 462 publications (journal articles, books, short communications and any other form of available and valid scientific publications) were extracted and entered in the Excel table. These 462 literature references are listed in Appendix 2. Additionally, all published taxonomic changes, including new synonymies, new combinations, new statuses, etc., in all hierarchical categories, were incorporated. The cut-off date for the data input was 23 May 2022.
Geographic coverage of the aforementioned 462 publications was examined by scoring the biogeographic region(s) each publication included. If the taxa in the paper were confined to one biogeographic region only, only one region was scored “1”. If the paper covered multiple regions, each covered region was scored “1”. The scoring is listed in Supplementary File 1.
Statistics in this paper (see “A summary of the species richness and systematics of Geometridae”), below, are based on the Excel table we created, which contains 34,897 species-group names of geometrid moths. The sub-section “Phylogeny, classification and species richness by subfamily” is based partially on our literature review and partially on the Excel table (see Supplementary File 1).
The geographic distribution of geometrid moths (Fig. 4) is based on 23,872 valid species names (see Supplementary File 1), excluding the subspecies and synonyms (see Fig. 4A). Biogeo-graphic regions were coded for each species based on its type locality (type locality information was missing for 17 species, which were excluded from the analysis). The biogeographic regions used in the analysis and shown in Fig. 4B are based on the scheme employed in the NHMUK collection and in The Global Lepidoptera Names Index ( https://www.nhm.ac.uk/ourscience/data/lepindex/lepindex/). The details are available in Gaston & Hudson (1994).
The online portal with the graphical user interface (GUI) to access the data was created by computer engineer Hamid Lagha (Mashhad, Iran) using React.js and python programming languages. The portal's interface was designed to enable easy, user-friendly updates (by approved contributors) to reflect new taxonomic changes after its initial publication.
The classification presented under “Family-group classification and names of Geometridae of the world” (Appendix 1) and in a forthcoming “Online list of valid names of geometrid moths” will be merged into the GBIF Backbone Taxonomy (Gbif Secretariat 2021) and into backbone systematics on BOLD (Ratnasingham & Hebert 2007).
Results
“Online taxonomic facility of geometrid moths” and list of valid names of geometrid moths
The new online portal provides access to all available names of geometrid moths of the world. The data are freely available and can be accessed at https://geometroidea. smns-bw.org/. The portal provides front-end access to a back-end Excel table, which includes 34,897 species-group names of geometrid moths. The table includes data from all published systematic literature on geometrid moths accessible to the authors, covering the period from the 10th edition of Systema Naturae (Linnaeus 1758) to 23 May 2022.
The back-end Excel table includes the following fields for all species-group names: Superfamily, Family, Subfamily, Tribe, Genus, Author, Year, Senior species, Senior subspecies, Senior species year, Senior subspecies year, Species, Subspecies, Species author, Species year, Species in author, Original genus, Country, Locality, Elevation, Latitude, Longitude, Junior synonym (yes/no), BIN, ID holotype (BOLD), Type depository, Type status, Type sex, Type country, Type state, Journal, Series, Volume, Part, Page (first), Comments.
Fields are not completed for every species, nor is a fully updated list of genera with synonyms available. As such, the online portal allows users to search the following columns for 34,897 species-group names: Family, Subfamily, Genus, Species, Subspecies and any name, including synonyms. Fig. 1A shows a typical query result from the back-end Excel table, in this case at genus level using the term “Archiearis”. Note that the search also reports a full list of synonyms and type localities. All species and synonyms are hyperlinked to the relevant species-level page (Fig. 1B). The data can be queried by using search terms (quick search and advanced search options are available) or by using a hierarchy search.
Additionally, the online portal contains a downloadable Excel table, which lists all valid and available names of Geometridae of the world. The structure of the Excel table is similar to that of the “Online list of valid and available names of the Geometridae of the world” (Scoble & Hausmann 2007).
The database will be updated by the authors as new information is published on Geometridae, with once-yearly updates of the facility and a list of valid and available names. This approach has the advantage of providing a citable, static version of the database with a date for each update release.
A summary of the species richness and systematics of Geometridae
Our database of geometrid moths provides an ideal source of information for summarising knowledge of the species richness, distribution and systematics of the family globally, demonstrating more widely the value of compiling databases of species-rich taxa. The present world database includes 34,897 available species-group names. Of these, 7,891 are considered synonyms (23% of available names), giving a total of 27,006 valid species-group names, which in turn comprise 23,872 species names and 3,134 subspecies names. This diversity is classified into 2,019 genera and nine subfamilies (see “Phylogeny, classification and species richness by subfamily” for details).
Fig. 1.
Examples of search results from the “Online taxonomic facility of geometrid moths”, available at https://geometroidea.smnsbw.org/. A. Genus-level result using the term “Archiearis”. B. Species-level result using the term “Archiearis infans infans”, or selecting the hyperlink in (A) (black arrow).
Subspecific names are often controversial in systematics. The current version of the “Online taxonomic facility of geometrid moths” includes 3,134 such entries. A robust approach has been taken in checklists of other lepidopteran taxa (e.g., Poole 1989; Nielsen et al. 1996), in which it was proposed that subspecies names should be treated as junior synonyms of the relevant species name. With respect to the Geometridae, Scoble (1999: xx–xxi) argued that “subspecies are inconsistently founded and usually they merely burden the taxonomy without adding anything to our understanding of natural divisions within a species”. The subspecies problem was also discussed by Skou & Sihvonen (2015: 37) in the European context, a continent overwhelmed by subspecies and synonymies. Indeed, in “The Geometrid Moths of Europe” series (Hausmann 2001, 2004; Mironov 2003; Hausmann & Viidalepp 2012; Skou & Sihvonen 2015; MÜller et al. 2019a, 2019b) hundreds of subspecies names were sunk into synonymy (see Hausmann & Sihvonen 2019). We follow Scoble (1999) and encourage taxonomists using the online database to adopt the approach of synonymising subspecies if they do not merit the rank of species unless there are clear and consistent reasons for not doing so.
Fig. 2.
Summary of species richness and classification of geometrid moths (based on data collected until 23.5.2022) and diagnostic morphological characters of the family. A. Summary of species richness and classification—see text for details. B. Geometra papilionaria Linnaeus, 1758 (photo: Pekka Malinen), adult male, the type species of the type genus of Geometridae. C. Geometra papilionaria caterpillar (photo: Tapio Kujala), showing diagnostic loss or reduction of ventral prolegs; prolegs are present only on segments 6 and 10, which causes the characteristic “looping” progression. Geometrid caterpillars also possess subprimary seta L4 on the abdominal segments (not visible on the photo due to the small size of the seta). D–E. Phrataria replicataria Walker, 1866 (photos: Pasi Sihvonen), showing the morphologically diagnostic tympanal organs on the ventral side of the second abdominal segment. D. Specimen with scales, arrows showing tympanal cavity and arched sclerotised rod curved over the tympanum. The ansa, which is visible in E and F, is inside the tympanal cavity and not visible in D. E. Specimen without scales, showing both tympanal organs. Arrows on left side of picture showing tympanal cavity and sclerotised rod. F. Tympanal organ's ansa (arrow) enlarged. (B and C published under CC-BY-NC-SA-4.0 licence; original photos available on Finnish Biodiversity Info Facility https://laji.fi/en; D and E reproduced from Murillo-Ramos (2021), slightly modified)
Phylogeny, classification and species richness by subfamily
Much research on the systematics of the Geometridae has been carried out from the time of Linnaeus onwards, but more needs to be done to improve its current state. Specifically, many if not most genera outside the Palaearctic or Nearctic regions need revision. Many genera as currently accepted are not monophyletic and their systematic position remains unstudied, even for those that have been collected and examined frequently from well-explored habitats and biomes. Moreover, numerous species remain undescribed.
A major impediment to achieving a more comprehensive and robust taxonomy for the Geometridae is the large number of species in the family compared with the limited resources for study—especially in terms of specialists and funding. The situation is further hindered by geographically and methodologically restricted approaches, which has resulted in classifications that are too narrow in outlook, lacking the necessary global overview of the family. In recent years, however, these shortcomings have been addressed, and with international collaboration and better geographic sampling, significant advances have been made. Notably, the higher-level classification has been rendered decidedly more robust. A summary of the main observations on the state of Geometridae systematics follows:
Geometridae are a monophyletic family, being sister to Uraniidae within Geometroidea (Fig. 2A, B). The systematic position and monophyly of the family are supported by molecular and morphological data (e.g., Holloway 1997; Minet & Scoble 1999; Sihvonen et al. 2011; Regier et al. 2013; HeikkilÄ et al. 2015; Rajaei et al. 2015; Kawahara et al. 2019; Murillo-Ramos et al. 2019). The combination of presence of a central rod (‘ansa’) in the cavities of the paired tympanal organs and tympanal organs present on fused abdominal ster-nites 1–2 [Fig. 2D, E, F; for overview of structures see Cook & Scoble (1992)] is diagnostic, whereas in Uraniidae the tympanal organs differ in morphology and location, being on tergites 2–3 in males and on sternite 2 in females. Other geometroid families (Sematuridae, Epicopeiidae, Pseudobistonidae) lack tympanal organs [see Rajaei et al. (2015) for other diagnostic characters] and other taxa with tympanal organs at the base of the abdomen, such as Pyraloidea and Drepanoidea, differ in structural details (Minet & Scoble 1999; Munroe & Solis 1999). Tympanal organs are reduced in size in a few geometrid lineages, particularly in brachypterous females [see examples in MÜller et al. (2019a, 2019b)]. In Geometridae, larval prolegs are typically reduced to two pairs, causing these caterpillars to move by looping (Fig. 2C), whereas uraniid larvae have five pairs of well-developed prolegs (Minet & Scoble 1999). A few Geometridae species have more abdominal prolegs, e.g., species of Archiearinae (Hausmann 2001) and Oenochrominae (Common 1990) and in Ennominae this characteristic is almost exclusively confined to Diptychini as defined by Murillo-Ramos et al. (2019), including, for instance, Afrotropical Callioratis Felder, 1874 (Duke & Duke 1998; Staude 2001; Sihvonen et al. 2015) and some Australian and South American taxa (Parra & Henriquez-Rodriguez 1993; Young 2008). Hausmann (2001: 69) provided a summary for Ennominae. Geometridae larvae have an extra L seta (subprimary L4) on the abdominal segments, which is absent in other families (Dugdale 1961) except in Thyatiridae (Stehr 1987), but mapping of this character needs wider taxon sampling. Externally, both adults and larvae of geometrids are very variable in size, pattern and colour.
Nine subfamilies of Geometridae are usually accepted, and the relationships between eight of them (Sterrhinae, Larentiinae, Archiearinae, Desmobathrinae, Epidesmiinae, Oenochrominae s. str., Geometrinae and Ennominae) are well-established (Fig. 3). These subfamilies are supported by morphological characters (see Murillo-Ramos et al. 2021a) and molecular characters (Murillo-Ramos et al. 2019). The status of the ninth subfamily, Orthostixinae, needs more research, as it may be part of Desmobathrinae (Murillo-Ramos et al. 2021b). The phylogenetic position of the genus Eumelea Duncan & Westwood, 1841 remains uncertain. It has been considered variously as a sister group to Desmobathrinae: Desmobathrini (Holloway 1996), Geometrinae (Beljaev 2008b), or the Ennominae genus Plutodes Guenée, 1858, depending on the analysis employed (Murillo-Ramos et al. 2019); as a result, Murillo-Ramos et al. (2019) treated it as incertae sedis. The most recent study, based on 329 genetic markers, placed Eumelea either as sister to Geometrinae (analysis based on nucleotides) or as sister to Oenochrominae s. str. (analysis based on a study of amino acids) (Murillo-Ramos et al. 2021b). The situation is under review, with a proposal to accept Eumelea as representing a separate lineage and giving it subfamily status (Murillo-Ramos et al., unpublished). If Orthostixinae becomes a junior synonym of Desmobathrinae and Eumelea is given subfamily rank, the total number of valid subfamilies in Geometridae would remain at nine.
Sterrhinae alone (Murillo-Ramos et al. 2019) or Sterrhinae + Larentiinae are considered the sister group to the rest of Geometridae (e.g., Yamamoto & Sota 2007; Sihvonen et al. 2011). With regard to the phylogeny, many taxa, particularly in Desmobathrinae, Oenochrominae s. l., and Ennominae are still misplaced and will need transferring to other subfamilies.
The most recent tribal-level classification based on a study benefitting from global taxon sampling included 1,206 geometrid species and 11 genes (Murillo-Ramos et al. 2019). The resultant phylogenetic analysis improved the systematics of geometrids significantly. The results revealed that many species are misplaced in tribes and in genera; that many genera are non-monophyletic; and that a great deal of research is needed at these taxonomic levels. Subtribal classification is rarely applied in Geometridae systematics, except to Geometrinae (Ban et al. 2018).
Fig. 3.
Summary of Geometridae phylogeny, classification and species richness by subfamily. Non-ultrametric phylogeny and classification based on Murillo-Ramos et al. (2019) and Murillo-Ramos (2021); species richness in each subfamily based on data presented in the current paper. The size of the coloured subfamilies in the tree is representative of the taxon sampling in Murillo-Ramos et al. (2021b), not the species richness. The figured moths (not in scale relative to each other) represent selected species in each subfamily, both nocturnal and diurnal, to highlight the diversity of wing patterns, shapes and colours.
Our understanding of the phylogenetic relationships and classification of geometrids varies greatly geographically and by taxon. The most intensively studied areas are the Nearctic (e.g., Mcguffin 1987; Ferguson 2008; Pohl et al. 2016) and the West Palearctic (e.g., MÜller et al. 2019a, 2019b and references therein), yet much work remains to be done even in these areas. The taxonomic impediment is highest in the tropics, which are known to be the most species-rich areas (Hillebrandt 2004; Brehm et al. 2016). Significant parts of these faunas have not been included in modern taxonomic studies, and often nothing else exists than the original, usually very superficial species and genus descriptions from late 1800s and early 1900s. Exceptions do exist, for instance The Moths of Borneo series (see Holloway 2011 and references therein).
Species richness and geographic distribution of type localities
Species richness of Geometridae in different biogeographic regions of the world is mapped in Fig. 4, a graphic that is based on an analysis of type localities. As regards the biogeographic regions, the highest described species richness is in the Neotropics, with 6,595 species, followed by the Oriental Region with 4,969 described species. The Palearctic Region, specifically the West Palearctic Region, is the most explored and its fauna has been the target of extensive taxonomic studies for several centuries. In total, 4,047 valid species are known from this region, 53% of them from the West Palearctic. The Afrotropical Region has 3,366 described species, followed by the Australian Region with 2,622 species. Of note is that the Oriental Region is considerably smaller in land area than the Nearctic, but the number of described species is much higher: 4,969 in the Oriental and 1,442 in the Nearctic. Madagascar and New Zealand are the smallest in terms of described species, but many of the species are endemic to these islands.
Fig. 5.
Annual pattern of species description in Geometridae from 1758 to 2022. Similar data until 1995, but summarised by decade, were published by Gaston et al. (1995).
Patterns of species description in the Geometridae
The annual pattern of species description in Geometridae (Fig. 5) shows a peak roughly between the years 1860 and 1950, with the highest numbers in the early 1900s. In seven years, the annual number of new species described exceeded 600: 1897 (1,137 species), 1858 (968 species), 1904 (807 species), 1907 (658 species), 1901 (645 species), 1893 (641 species) and 1916 (603 species). Altogether 1,854 new species and subspecies have been described by various authors since 2000, after publication of “Geometrid Moths of the World: A Catalogue” (Scoble 1999). From 2000–2022, the average number of new species descriptions has been 80 per year.
Most productive authors describing Geometridae
Analysis of all published species-level names of the geometrid moths shows that two English lepidopterists, namelyWILLIAMWARREN(1839–1914)andLOUISBEETHOVEN PROUT (1864–1943), described together 8,961 taxa, which is roughly 25% of all described names in the family Geometridae. The top ten authors describing the most species-level taxa in Geometridae are shown in Fig. 6A. All have passed away, most recently French lepidopterist CLAUDE HERBULOT (19.2.1908–19.1.2006) and Japanese lepidopterist Hiroshi Inoue (8.7.1917–2.6.2008).
Fig. 6.
Ten most prolific authors and ten most species-rich genera. A. Ten most prolific authors in Geometridae systematics and number of species-level taxa described by each, including taxa currently considered as synonyms (see also Gaston et al. 1995). Numbers above the bars are absolute numbers of described taxa, with percentages of synonymised names in parentheses; numbers of currently valid taxa are shown on the blue bars. B. Ten most species-rich genera of Geometridae. Number of valid species on y-axis. (Photo of Eupithecia pusillata by Pekka Malinen, published under CC-BY-NC-SA-4.0 licence and available on Finnish Biodiversity Info Facility https://laji.fi/en)
Most species-rich genera in Geometridae
The ten most species-rich genera are shown in Fig. 6B. The three most species-rich genera [Eupithecia Curtis, 1825 (Larentiinae), Scopula Schrank, 1802 (Sterrhinae) and Idaea Treitschke, 1825 (Sterrhinae)] are cosmopolitan. These genera have a large number of small species relative to the Geometridae in general, with a wingspan of usually much less than 30 mm. While Geometridae larvae typically consume fresh leaves, many Eupithecia caterpillars feed on flowers, and most Idaea and numerous Scopula larvae feed on withered plant debris.
Geographic coverage of taxonomic literature during the last two decades (2003–2022)
The geographic coverage of taxonomic literature on Geometridae during 2003–2022 is shown in Fig. 7. The taxonomic literature is uneven between biogeographic regions for that period, and out of 462 papers, the majority deal with the Palaearctic (36%) and Oriental faunas (30%). Together, 353 publications cover the fauna of these regions, which is 66% of the world total. The fauna of China, which includes Palaearctic and Oriental elements, has been intensively studied during 2003–2022; 46 papers focus on this fauna, and Chinese authors have prepared the majority of these. Fifty-four papers (10% of total) deal with the Neotropical fauna, which hosts the highest number of described species. The majority of taxonomic papers were published in Zootaxa (93), Tinea (74) and SHILAP Revista de Lepidopterologia (23).
Fig. 7.
Geographic coverage of taxonomic literature on Geometridae during 2003–2022. Colours align with biogeographic regions shown in Fig. 4. Details of examined literature listed in Supplementary File 1.
Taxa excluded from Geometridae after 1999
After “Geometrid Moths of the World: A Catalogue” (Scoble 1999) was published, three species have been transferred from Geometridae to other families:
Pseudobiston pinratanai Inoue, 1994 was tentatively assigned to Geometridae: Geometrinae (Inoue 1994), but the absence of antero-abdominal tympanal organs, among other characters, raised doubts about its systematic position (StÜning 2001). Based on molecular and morphological data, P. pinratanai was included in a new family, Pseudobistonidae Minet, Rajaei & Stüning, 2015, as sister to Epicopeiidae, both families being part of Geometroidea (Rajaei et al. 2015). This oriental moth is known from North Thailand and North Vietnam, and it is the only species included in Pseudobistonidae.
Cartaletis dargei Herbulot, 2003 was originally placed in Geometridae: Sterrhinae (Herbulot 2003). Recently, it was shown that the moth only superficially resembles true Cartaletis Warren, 1894 [the genus name is currently considered a junior synonym of Aletis Hübner, 1820, see Sihvonen et al. (2020)], and it was transferred to Noctuidae: Agaristinae as Aletopus dargei (Sihvonen et al. 2021).
Probolaea roboginosa Turner, 1943 from Australia: Queensland was originally placed in Geometridae: Larentiinae (Turner 1943). Fletcher (1979) proposed an objective generic replacement name for Probolaea, namely Ecphysis Fletcher, 1979, because Probolaea Turner, 1943 is a junior homonym of Probolaea Meyrick, 1885. The species roboginosa is, however, a noctuoid moth and was assigned to Erebidae: Cato-calinae by Nielsen et al. (1996). This information was overlooked and P. roboginosa was listed as a Larentiinae in Scoble (1999). The original transfer from Geometridae to Erebidae may have been done earlier than Nielsen et al. (1996), but we have not traced such literature. Nielsen et al. (1996: 372, note 613) also explained that the original species name was misspelled as roboginosa, and that it should be robigonosa instead. We note that robigonosa is misspelled also, justifying this by the etymology note in Turner (1943: 106), who wrote that the species name was based on “robiginosus, rusty” and emended it to robiginosa (Iczn 1999, Articles 32.5.1, 33). The classification in Erebidae was also missed by Özdikmen (2009), who proposed a new generic replacement name in Geometridae for Ecphysis Fletcher, 1979, which is a junior homonym of Ecphysis Townes, 1969 in Hymenoptera. Accordingly, this Australian species is currently named Aslihana robiginosa (Turner, 1943) and is assigned to Noctuoidea: Erebidae. DNA barcodes also support this assignment (BIN: BOLD:AAE0689).
Taxa transferred, after 1999, from other families to Geometridae
Crambometra Prout, 1915 with type species C. derelicta Prout, 1915 from South Africa, is a genus originally classified in Ennominae (as Geometrinae) with the note “Affinities very obscure, apparently rather a primitive form, perhaps verging on the Oenochrominae” (Prout 1915: 376). Crambometra was later transferred to Notodontidae (Janse 1920) without explanation, and this was potentially the reason why geometrid researchers overlooked the taxon and it was not included in the relevant geometrid literature anymore, including the geometrid catalogues (Scoble 1999; Scoble & Hausmann 2007). Schintlmeister (2013) re-examined the type species of Crambometra and transferred it and its three species back to Geometridae. KrÜger (2015) treated Pachycnemoides Krüger, 1999 as a junior synonym of Crambometra Prout, 1915 and listed two further genus synonyms established earlier by Kiriakoff (1970). Crambometra was analysed in a multi-gene molecular phylogeny study and found to be nested within Ennominae: Diptychini, within the Callioratis Felder, 1874 and Veniliodes Warren, 1894 complex of genera (Murillo-Ramos et al. 2019). Currently, Crambometra includes 14 species (KrÜger 2015).
Family-group classification and names of Geometridae of the world
A list of Geometridae family-group names is presented in Appendix 1, to provide a higher classificatory context to the online facility. Many annotations are included to allow readers to gain a perspective on the way that the nomenclature and systematic ordering has evolved. The current list of family-group names is based on the “Moths of Borneo” publications (Holloway 1994, 1996, 1997), as well as Fletcher (1979) and has been expanded and updated in line with recent studies, particularly molecular phylogenetic work (e.g., Sihvonen et al. 2011; Õunap et al. 2016; Jiang et al. 2017; Ban et al. 2018; Brehm et al. 2019; Murillo-Ramos et al. 2019; Sihvonen et al. 2020). The morphology of the subfamilies was reviewed by Murillo-Ramos et al. (2021a) and the overall systematic order is based on the latest extensive molecular phylogeny [Murillo-Ramos et al. (2019), presented in full in Brehm et al. (2019; supplement 1)] and supplemented by more focused systematic studies as indicated in the text. We emphasise the point that there are numerous ways to render a phylogenetic hypothesis as a linear classification, and thus encourage the readers to study the original tree-form hypotheses in parallel.
Names of a few tribes that were not included in the molecular analyses are listed, some of them considered valid by Beljaev (2008b), who discussed their relationships based on morphology. The list, which does not include incorrect subsequent spellings, will be updated and refined, and we expect to publish new versions with future updates in the “Online taxonomic facility [and list of valid names] of geometrid moths”.
In Appendix 1, all available family-group names (family, subfamily, tribe, subtribe) are shown in bold. The numbers of subordinated family-group names are given in brackets in the headline of each subfamily. Names in brackets are those given by the original author.
Discussion
What is the real species richness of geometrid moths?
In many insect groups the real species richness is estimated to be considerably higher than the described fauna (ǾDegaard 2000; Garcia et al. 2020). The indirect data available suggests this is the case in Geometridae also. We estimate that the real species richness on the global scale is likely to rise from the current 23,872 valid species to 40,000–50,000 species. We try to justify this estimate by a few examples:
Geometridae have been among the target groups in the Global Campaign of DNA barcoding (iBOL Lepidoptera). Within this project, numerous unnamed genetic clusters have emerged and in Ennominae, for instance, which is the most species-rich geometrid lineage with about 11,100 described species worldwide, the genetic data on BOLD suggests that at least 7,000 additional species await description (see Preface in MÜller et al. 2019a). Available DNA barcode data suggest a large amount of cryptic diversity in Lepidoptera in general (Garcia et al. 2020) and a considerable percentage of tropical species are not even represented yet in databases such as BOLD systems.
Some of the tropical genera might easily be as species-rich as the genera illustrated in Fig. 6B. For example, Brehm et al. (2011) concluded that Eois Hübner, 1818 could easily comprise more than 1,000 species, given the low rate of matches (10%) with described species of material collected in southern Ecuador.
Regional diversity estimates in tropical regions also strongly suggest that a high number of species still need to be described. For example, Brehm et al. (2016) estimated regional species richness of geometrid moths at 2,350 species in an area of only 40 km2 in the southern Andes. This corresponds to more than one third of the richness of the entire Neotropical Region (6,595 species), a far higher percentage than appears likely for a very small area that did not even include elevations below 1,000 m.
Beta diversity along environmental gradients could currently be still underestimated. In montane areas, closely related species (currently regarded as one species) occur at different elevational bands (e.g., Brehm et al. 2003, 2016). Similar examples exist along the latitudinal gradient (e.g., Paknia & Rajaei 2015).
Several recent revisions based on both museum and fresh materials show a clear trend that the actual species richness is considerably higher than historically thought. An integrative revision of the genus Prasinocyma Warren, 1897 (Geometrinae), which focused on Ethiopia only, revealed that just 52% of the 40 Ethiopian species had been described (Hausmann et al. 2016).
The revision of the same genus at a continental scale (Africa) will raise the validated species number from 86 (in Scoble 1999) to more than 300 (Hausmann, unpublished), i.e., by more than 350%. In the Indo-Australian genus Ziridava Walker, 1863 (Larentiinae), the increase was 33% (Schmidt & Tautel 2022); in the Afrotropical Trimetopia Guenée, 1858 (Larentiinae) the increase was 70% (Stadie & Fiebig 2019); and in the Chinese Timandra Duponchel, 1829 (Sterrhinae) the increase was 140% (Cui et al. 2019). On the other hand, there is a tendency of finding synonyms in better explored faunas, for example in Europe (see Conclusions).
Assuming a real number of at least 40,000 geometrid species on our planet, including at least 16,128 undescribed species, and assuming a linear continuation of the current pace of 80 descriptions per year, traditional taxonomic research will need another ca. 200 years to complete the task. However, detection of undescribed species may be fostered by DNA barcoding and descriptions may be accelerated by automated, modern tools of data transfer and manuscript templates.
Future development of the “Online taxonomic facility of geometrid moths”
We wish, in time and depending on resources, to expand the “Online taxonomic facility of geometrid moths” in its current format to a more comprehensive knowledge base. Specifically, it is our long-term aim to:
Expand coverage to include the entire superfamily Geometroidea, namely the families Geometridae, Epicopeiidae, Pseudobistonidae, Sematuridae and Uraniidae.
Supplement the database with photographs of type specimens. Progress is already being made by the project “Geometridae Mundi” (coordinated by Axel Hausmann and Hans Löbel from Germany). It is intended to photograph not only type specimens but selected other material for all species. The number of photographs taken already exceeds 50,000 and covers ca 50% of the described species.
Add information for each taxon on original descriptions, type localities, type specimens and their depositories, and genera and their synonyms, as in Scoble (1999).
Forge links with the data on the BOLD website, particularly to type specimens.
Add, if possible, information on immature stages and their hostplants, global and elevational distribution and habitats.
Add, if possible, photographs of morphological traits (e.g., male and female genitalia, wing length, size).
Conclusions
We have noted already the geographical bias in Geometridae systematics and conclude that there is an evident need to encourage a greater effort on species-rich but understudied regions if we are to make significant progress. Yet how might this be achieved? The universal and frequent plea for funding for taxonomy of the magnitude needed is unlikely to materialise, so finding ways of accelerating progress must rely on the community of geometrid taxonomists. DNA barcoding (Hebert et al. 2003) has been used for rapid, mass description of species of braconid wasps (Sharkey et al. 2021; FErnandez-Triana 2022) and weevils (Riedel et al. 2014; Riedel & Narakusumo 2019), as well as for descriptions and monographs of Geometridae (e.g., Hausmann et al. 2016; MÜller et al. 2019a, 2019b). Although extensive use of DNA barcoding without complementary evidence bears a risk of taxonomic instability [see Meier et al. (2021) for a criticism of a minimalist, barcode-only-based approach to species description], molecular methods are an invaluable tool for developing taxonomic hypotheses, especially for mega-diverse taxa in the tropics. While this approach should be used ideally in conjunction with morphological evidence, the catastrophic loss of biodiversity requires more haste in species description, and thus some relaxation in selecting the perfect methodology. Therefore, we regard an accelerated integrative approach as the best practical option for promoting the taxonomy of geometrid moths in tropical countries, where species diversity is at its highest.
A further approach is to encourage geometrid taxonomists to concentrate on reviews and revisions at the level of genus, sampling as many species as possible. Past examples include studies on the Neotropical Geometrinae (Pitkin 1996), Ennominae (Pitkin 2002), Macariini (Scoble & KrÜger 2002) and Scopulini (Sihvonen 2005) and numerous papers dealing with the Chinese fauna (see Supplementary File 1), which created a framework to help those dealing with species descriptions. Such works would need to be centred at institutions with large collections with significant global coverage, but undertaken in a highly collaborative way among taxonomists, parataxon-omists and collectors in species-rich regions.
Combined, these two approaches might be an effective means of making progress. A collaborative approach is exemplified in this paper, with authors from five countries contributing and benefitting from e-communication, although we note that only one country (Colombia) falls among those regions with high species richness.
If the description of new species is the major challenge in the tropics (e.g., Brehm 2018), that of synonymy is a greater issue in well-studied but relatively species-poor regions. As many as 23% of available names in “Online taxonomic facility of geometrid moths” are synonyms (n = 7,891). This percentage is likely to increase significantly, considering that many of the 3,134 subspecies are likely to be synonymised in future. In Volume 6, Part 1 of the “The Geometrid Moths of Europe” (MÜller et al. 2019a), merely four new species were described, while 111 new synonymies, new statuses and new combinations were proposed, many of the proposed synonymies benefitting from DNA barcode data. The point raised years ago by Scoble (1999: xiv) is still valid, at least for well-known faunas: “We need to take careful account of the magnitude of synonymy besides the description of new species if we are to have … a better understanding of species richness in the Geometridae”.
Finally, we were astonished by how difficult it sometimes is to find taxonomic literature on geometrid moths, despite the fact that we work in an era where funding bodies and institutions are encouraging or requiring open access publishing. While much of the older taxonomic literature is now available via the Biodiversity Heritage Library, even today, some taxonomists publish in journals that are not accessible online. Indeed, we encountered a few cases where even university librarians could not gain access to papers with data we wished to source. Furthermore, a few papers were published in journals lacking peer review and editorial standards. We encourage geometrid taxonomists to publish their manuscripts in peer-reviewed journals that are easily accessible and preferably open access.
Acknowledgements
We express our most grateful thanks to Dr. Amir M. Javadi for designing the new logo of Forum Herbulot and to engineer Hamid Lagha for designing the new website of Forum Herbulot. We thank the IT staff at SMNS, and in particular Wiebke Walbaum for her amazing patience and support. We thank all members of Forum Herbulot for their continuous scientific support and especially for providing us with their publications on geometrid moths. We highly appreciate Donald Hobern'S (GBIF) great help with detecting errors in the “Online taxonomic facility of geometrid moths”. Finally, we are thankful to reviewers Dr. Catherine J. Byrne and Dr. Sei-Woong Choi for their critical and constructive review of the submitted version of this paper.
Funding
Development of the new website of Forum Herbulot, which includes the “Online taxonomic facility of geometrid moths” presented herein, was supported financially by several Herbu-lotians (names in alphabetic order): Gunnar Brehm, Feza Can, AxeL Hausmann, Gyula László, Hossein Rajaei, Malcolm Scoble, Pasi Sihvonen, Peder Skou, Hermann Staude, Manfred Sommerer, Dieter Stüning, Claude Tautel.
References
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Supplementary file:
[Available from: https://doi.org/10.6084/m9.figshare.21747089]
Supplementary File 1: Examined literature between 2003 and 23.5.2022, with biogeographic coverage (Excel file).
Appendices
Appendix 1.
Family-group classification and names of Geometridae of the world.
Family Geometridae Stephens, 1829 (202 names)
Declared as the correct name by the International Commission on Zoological Nomenclature (ICZN 1957, Opinion 450).
Currently validated: 9 subfamilies (see text for details on problems in subfamily classification), 93 tribes and 11 subtribes.
Subfamily Sterrhinae Meyrick, 1892 (Sterrhidae) (23 names)
Although there are several senior synonyms of this name, the name Sterrhinae should be maintained for nomenclatural stability [as proposed by Holloway (1997: 15)]. For justification, see below under the tribe Sterrhini. Validity and order of tribes based on Holloway (1997), Hausmann (2004), Murillo-Ramos et al. (2019) and Sihvonen et al. (2020).
Tribe Mecoceratini Guenée, 1858 (Mecoceridae)
Mecoceras is a junior objective synonym of Ametris. Validated at tribe level and transferred from Desmobathrinae to Sterrhinae by Sihvonen et al. (2020).
= Ametridini Prout, 1910 (Ametridicae), junior synonym (Sihvonen et al. 2020).
Tribe Cosymbiini Prout, 1911 (Cosymbiinae)
Although there are senior synonyms of this name (see below), the name should be retained to maintain nomenclatural stability [as proposed by Holloway (1997: 23)]; see also ICZN (1999, Art. 40.2).
= Ephyrini Guenée, 1858 (Ephyridae), based on a junior homonym of a genus-group name outside Lepidoptera. Invalid according to Iczn (1999, Art. 39).
= Zonosomini White, 1876 (Zonosomatidi), based on Zonosoma, thus emended, senior synonym (Holloway 1997).
= Cyclophorini Moore, 1887 (Cyclophoridae), senior synonym (Holloway 1997). As the name Cyclophoridae is in common usage in Mollusca, based on the genus Cyclophorus, Holloway (1997: 15) proposed that it should be “gracefully conceded” to malacologists.
Tribe Timandrini Stephens, 1850 (Timandridi)
= Calothysanini Herbulot, 1963 (Calothysanini), based on (misinterpreted) genus name Calothysanis, a synonym of Scopula (Scopulini) (Holloway 1997).
Tribe Cyllopodini Kirby, 1892 (Cyllopodidae)
= Micropinini Kirby, 1912 (Micropinidae), junior synonym, based on an unnecessary genus-group replacement name (Holloway 1997).
= Rhodostrophiini Prout, 1935 (Rhodostrophiicae), junior synonym (Sihvonen et al., 2020).
Tribe Sterrhini Meyrick, 1892 (Sterrhidae)
Although senior synonyms of this name exist (see below), Sterrhini should be maintained for nomenclatural stability, as proposed by Holloway (1997: 15) and in accordance with the intention and spirit of the concept of “prevailing usage” of Iczn (1999, cf. Art. 40.2.).
= Goniacidaliini Packard, 1876 (Goniacidaliinae), senior synonym (Hodges et al. 1983).
= Idaeini Butler, 1881 (Idaeidae), senior synonym (Holloway 1997).
= Ptychopodini Pierce, 1914 (Ptychopodinae), junior synonym (Holloway 1997).
Tribe Haemaleini Sihvonen & Brehm, 2020 (Haemaleini)
Tribe Lissoblemmini Sihvonen & Staude, 2020 (Lissoblemmini)
Tribe Scopulini Duponchel, 1845 (Scopulites)
= Acidaliini Duponchel, 1845 (Acidalites). The original Latin word stem Acidal- (with the suffix -ites) needs to be emended to Acidali- (being based on the type genus Acidalia), leading to “Acidaliini” rather than to Acidalini; junior synonym, based on a junior homonym of a genus-group name outside the Lepidoptera (Holloway 1997).
= Aletini Hampson, 1918 (Aletinae), junior synonym (Holloway 1996).
= Problepsini Wiltshire, 1990 (Problepsini), junior synonym (Holloway 1997).
Subfamily Larentiinae Duponchel, 1845 (Larentites) (47 names)
The original Latin word stem Larent- (with the suffix -ites) needs to be emended to Larenti- (being based on the type genus Larentia), leading to “Larentiinae” rather than to Larentinae. Validity and order of tribes mainly follow Holloway (1997), Viidalepp (2011), Hausmann & Viidalepp (2012), Õunap et al. (2016), Brehm et al. (2019) and Murillo-Ramos et al. (2019). Brehm et al. (2019), in their phylogenetic hypothesis, highlighted four lineages, each marked as “unnamed clade”. If further data substantiate these clades, they will need description at the level of tribe, but at this stage they are excluded from the list.
Tribe Dyspteridini Hulst, 1896 (Dyspteridinae)
Dyspteridini were removed from synonymy with Trichopterygini and given tribal rank by Viidalepp (2011), an action con-
firmed by Õunap et al. (2016) and Murillo-Ramos et al. (2019).
Tribe Brabirodini Brehm, Murillo-Ramos & Õunap, 2019 (Brabirodini)
Tribe Trichopterygini Warren, 1894 (Trichopteryginae)
= Lobophorini Tutt, 1896 (Lobophorinae), junior synonym (Hodges et al. 1983).
Tribe Chesiadini Stephens, 1850 (Chesiadi)
The Latin word stem is Chesiad- (genitive of the type genus name Chesias = Chesiadis). Authorship of tribe name “Pierce, 1914” according to Viidalepp (2011).
= Eucestiini Warren, 1894 (Eucestiinae). Based on Eucestia, which is a junior objective synonym of Chesias; junior synonymy of tribe name: Holloway (1997: 97).
= Odeziini Exposito, 1978 (Odeziini), junior synonym (Hausmann & Viidalepp 2012: 442).
Subtribe Aplocerina Viidalepp, 2011 (Aplocerina)
Rank of subtribe adopted here, based on the findings of Murillo-Ramos et al. (2019).
Tribe Chrismopterygini Brehm, Murillo-Ramos & Õunap, 2019 (Chrismopterygini)
Tribe Eupitheciini Tutt, 1896 (Eupitheciinae)
Preferred over the senior name Tephroclystini, following ICZN (1999, Art. 40.2) (prevailing usage of the pre-1961 replacement substitute name of the type genus).
= Tephroclystiini Warren, 1895 (Tephroclystiinae), senior synonym. Based on Tephroclystia, now treated as a junior synonym of Eupithecia (Iczn 1999, Art. 40.2.1).
= Chloroclystini Mironov, 1990 (Chloroclystina), junior synonym, originally proposed as a subtribe of Eupitheciini (Holloway 1997).
Tribe Operophterini Packard, 1876 (Operophterinae)
Holloway (1997) noted that Operophterinae should probably take precedence over Oporiniini as in VIVES Moreno (1994). Following this, Oporiniini is treated here as junior synonym.
= Oporiniini Pierce, 1914 (Oporiniinae), junior synonym (Holloway 1997).
Tribe Triphosini Tutt, 1896 (Triphosidi)
Subordinated under Rheumapterini in Hausmann & Viidalepp (2012), but validated at tribe level by Viidalepp (2011), Schmidt (2015), Õunap et al. (2016) and Murillo-Ramos et al. (2019). The three tribes Triphosini, Phileremini and Rheumapterini resolved as a single cluster in Brehm et al. (2019), but these authors did not feel ready to synonymise the tribes without further study.
Tribe Rheumapterini Herbulot, 1961 (Rheumapterini) See comment under Triphosini.
= Melanippini Bruand, 1846 (Melanippidae), senior synonym. Based on Melanippe, a junior synonym of Rheumaptera. It is proposed not to apply this name, as Rheumapterini is accepted widely.
= Eucosmiini Gumppenberg, 1887 (Eucosminae), senior synonym. It is proposed not to apply this name, as Rheumapterini is widely accepted in the literature and Eucosmiinae was never used after the original description. Emended by reasons of grammar (based on Eucosmia, a junior synonym of Hydria).
Tribe Psaliodini Brehm, Murillo-Ramos & Õunap, 2019 (Psaliodini)
Tribe Cidariini Duponchel, 1845 (Cidarites)
The original Latin word stem Cidar- (with the suffix -ites) needs to be emended to Cidari- (being based on the type genus Cidaria), leading to “Cidariini” rather than to Cidarini.
= Therini Pierce, 1914 (Therinae), junior synonym (Holloway 1997). Based on Thera. Theriini (based on Theria) is a family-group name in Ennominae.
Tribe Scotopterygini Warren, 1895 (Scotopteryginae)
Subordinated under Xanthorhoini in Hausmann & Viidalepp (2012), but treated as a tribe by Viidalepp (2011), Schmidt (2015) and, using molecular data, by Õunap et al. (2016) and Murillo-Ramos et al. (2019).
= Euboliini Duponchel, 1845 (Eubolites), senior synonym, but not applied, as Scotopterygini is now largely accepted in the literature over Euboliini. Emended by reasons of grammar: based on Eubolia, a junior synonym of Scotopteryx.
= Ortholithini Tutt, 1896 (Ortholithinae), senior synonym, but not applied, as Scotopterygini is now largely accepted in the literature over Ortholithinae. Based on Ortholitha, a junior synonym of Scotopteryx.
= Phasianini Gumppenberg, 1897 (Phasianinae). Based on Phasiane, a junior synonym of Scotopteryx. Senior synonym, but junior homonym of a family-group name in common use in Aves (Phasianus) (Holloway 1997: 98).
Tribe Euphyiini Herbulot, 1961 (Euphyiini)
Tribes Euphyiini, Pterocyphini, Cataclysmini, Xanthorhoini and Epirrhoini form a complex of tribes (Brehm et al. 2019), potentially requiring synonymy. More research is needed.
Tribe Pterocyphini Brehm, Murillo-Ramos & Õunap, 2019 (Pterocyphini)
Tribe Xanthorhoini Pierce, 1914 (Xanthorhoinae)
= Psychophorini Hampson, 1918 (Psychophorinae), treated as a junior synonym in Hausmann & Viidalepp (2012). Vives-Moreno (1994) considered Psychophorini a junior synonym of Larentiini. Psychophora Kirby, 1824 has not been included in molecular phylogenetic analysis yet.
Tribe Rhinurini Brehm, Murillo-Ramos & Õunap, 2019 (Rhinurini)
Tribe Larentiini Duponchel, 1845 (Larentites)
Emended by reasons of grammar (see under subfamily name).
= Entephriini Pierce, 1914 (Entephrinae), emended by reasons of grammar (based on Entephria), junior synonym.
Tribe Ennadini Brehm, Murillo-Ramos & Õunap, 2019 (Ennadini)
Tribe Cophoceratini Brehm, Murillo-Ramos & Õunap, 2019 (Cophoceratini)
Tribe Erebochlorini Brehm, Murillo-Ramos & Õunap, 2019 (Erebochlorini)
Subfamily Geometrinae Stephens, 1829 (Geometrinae) (33 names)
Validity and order of tribes mainly based on Pitkin (1996), Holloway (1996), Hausmann (1996a, 1996b, 2001), Ban et al. (2018), Brehm et al. (2019) and Murillo-Ramos et al. (2019). Brehm et al. (2019), in their phylogenetic hypothesis, highlighted four lineages, each marked as “unnamed clade”. These may require formal description at the category of tribe when more data are available.
Tribe Ornithospilini Ban, Jiang, Cheng, Yue & Han, 2018 (Ornithospilini)
Tribe Agathiini Ban, Jiang, Cheng, Yue & Han, 2018 (Agathiini)
Tribe Chlorodontoperini Murillo-Ramos, Sihvonen & Brehm, 2019 (Chlorodontoperini)
Tribe Comibaenini Inoue, 1961 (Comibaenini)
= Euchlorini Herbulot, 1963 (Euchlorini), junior synonym; based on genus name Euchloris, which is a junior homonym of a generic name in Coleoptera.
Tribe Nemoriini Gumppenberg, 1887 (Nemorinae)
The original Latin word stem Nemor- (with the suffix -inae) needs to be emended to Nemori- (being based on the type genus Nemoria) leading to “Nemoriini” rather than to Nemorini.
= Ochrognesiini Inoue, 1961 (Ochrognesiini), junior synonym (Holloway 1996: 151; Hausmann 1996b: 31).
= Synchlorini Ferguson, 1969 (Synchlorini), relations to Comibaenini discussed in Holloway (1996: 196) and Pitkin (1996: 322), but subordinated under Nemoriini in Murillo-Ramos et al. (2019) and Brehm et al. (2019) based on molecular evidence.
Tribe Dysphaniini Warren, 1895 (Dysphaniinae)
= Hazini Guenée, 1858 (Hazidae), junior synonym; based on Hazis, a junior objective synonym of Euschema, junior synonym of Dysphania (Holloway 1996).
= Euschemini Walker, 1862 (Euschemidae), junior synonym; based on Euschema, a junior synonym of Dysphania (Holloway 1996).
Tribe Pseudoterpnini Warren, 1893 (Pseudoterpninae)
Pitkin et al. (2007) commented the taxonomic history of the tribe.
= Terpnini Inoue, 1961 (Terpnini), junior synonym; based on Terpne (misspelled as Terpna), a junior synonym of Geometra, but referring to Terpne auctorum (= Pachyodes) (Hausmann 1996a: 96; Holloway 1996).
= Pingasini Heppner & Inoue, 1992 (Pingasini), junior synonym (Holloway 1996).
Tribe Dichordophorini Ferguson, 1969 (Dichordophorini)
Validity and potential subordination under another tribe awaiting study of molecular data.
Tribe Hemitheini Bruand, 1946 (Hemitheidae)
Reasons for tribal rank were given by Hausmann (1996; 2001). Holloway (1996: 196) proposed a broad concept of the tribe to include Thalerini, Comostolini, Hemistolini, Jodini and Thalassodini, a view supported by recent molecular analyses, e.g., Ban et al. (2018), Brehm et al. (2019) and Murillo-Ramos et al. (2019).
= Thalerini Herbulot, 1963 (Thalerini), given the rank of tribe by Hausmann (1996b; 2001), but synonymised as subtribe Hemitheina (misspelled as Hemitheiti) by Ban et al. (2018).
= Chlorochromini Duponchel, 1845 (Chlorochromites), senior synonym, based on Chlorochroma, a junior objective synonym of Thalera. Chlorochromini is not applied here, as Hemitheini is accepted widely (cf. Hausmann 1996a: 101).
The following taxa might have been treated as synonyms of Hemitheini (see above), but we list them as subtribes. Many further lineages revealed by molecular analysis, but not yet named, are likely to belong here.
Subtribe Heliotheina Exposito, 1978 (Heliothinae)
Based on Heliothea. Considered to belong to Geometrinae: Rhomboristini (as Rhomboristiti) by Holloway (1996: 150, 195), see also Hausmann (1996b: 12; 2001: 110). Treated as Hemitheini by Ban et al. (2018; misspelled as “Heliotheiti”) and Murillo-Ramos et al. (2019) based on molecular evidence.
Subtribe Hemistolina Inoue, 1961 (Hemistolini)
Treated as a tribe by Hausmann (1996b; 2001). Holloway (1996: 196) proposed a wider concept (including Thalerini, Comostolini, Hemistolini, Jodini and Thalassodini). Subordination under Hemitheini proposed by Ban et al. (2018; misspelled as “Hemistoliti”) based on molecular evidence and supported by Murillo-Ramos et al. (2019).
Subtribe Comostolina Inoue, 1961 (Comostolini)
Treated as a tribe by Hausmann (1996b; 2001), related to Jodini (Hausmann 1996b: 41). Holloway (1996: 196) proposed a wider concept (including Thalerini, Comostolini, Hemistolini, Jodini and Thalassodini). Subordination under Hemitheini proposed by Ban et al. (2018; misspelled as “Comostoliti”) based on molecular evidence and supported by Murillo-Ramos et al. (2019).
Subtribe Microloxiina Hausmann, 1996 (Microloxiini)
Originally published on 21.11.1996 at tribe rank (Hausmann 1996). Subordination under Hemitheini proposed by Ban et al. (2018; misspelled as “Microloxiiti”) based on molecular evidence and supported by Murillo-Ramos et al. (2019).
= Hierochthoniini Viidalepp, 1996, published without exact date (= 31.12.1996)
Subtribe Jodina Inoue, 1961 (Jodiini)
Emended by reasons of grammar (based on Jodis with the Latin word stem Jod-). Treated as a tribe by Hausmann (1996b; 2001), suggesting a relationship with Comostolini (Hausmann 1996b: 41). Holloway (1996: 196) proposed a wider concept (including Thalerini, Comostolini, Hemistolini, Jodini and Thalassodini). Subordination under Hemitheini proposed by Ban et al. (2018; misspelled as “Joditi”) based on molecular evidence and supported by Murillo-Ramos et al. (2019).
Subtribe Thalassodina Inoue, 1961 (Thalassodini)
Treated as a tribe by Hausmann (1996b). Holloway (1996: 196) proposed a wider concept (including Thalerini, Comostolini, Hemistolini, Jodini and Thalassodini). Subordination under Hemitheini proposed by Ban et al. (2018; misspelled as “Joditi”) based on molecular evidence and supported by Murillo-Ramos et al. (2019).
Subfamily Archiearinae Fletcher, 1953 (Archiearinae) (2 names)
The analyses of Sihvonen et al. (2011) and Murillo-Ramos et al. (2019) suggest that Archiearinae are the sister group to all Geometridae except the Sterrhinae + Larentiinae lineage.
= Brephinae auct. nec Hübner, 1826 (Brephidae), based on Brephos Ochsenheimer, 1816, a junior homonym of Brephos Hübner, 1813.
Subfamily Desmobathrinae Meyrick, 1886 (Desmobathridae) (2 names)
Based on Desmobathra Meyrick, 1886 (junior synonym of Ozola Walker, 1861). For validity at subfamily rank see Holloway (1996: 152, 159). See Orthostixinae for more information on Desmobathrinae–Orthostixinae relationship.
Tribe Desmobathrini Meyrick, 1886 (Desmobathridae)
Tribe Eumeleini Warren, 1894 (Eumeleinae)
The position and classification of Eumeleini have proven difficult to elucidate. Based on molecular data, Murillo-Ramos et al. (2019) suggested a sister-group relationship with the subfamily Geometrinae, provisionally placing the genus as “incertae sedis”. Recently, Eumelea Duncan & Westwood, 1841 was included in a whole-genome shotgun sequencing analysis and the name-bearing genus Eumelea was reconstructed as sister to Geometrinae (based on nucleotide analysis) or Oenochrominae s. str. (based on amino acid analysis). The authors proposed to raise Eumeleini to subfamily rank, but the publication and names and nomenclatural acts therein are not available for purposes of zoological nomenclature in accordance with Iczn (2012, Arts. 8.2, 8.3), and Murillo-Ramos and collaborators will formalise the new classification in a forthcoming publication.
Subfamily Orthostixinae Meyrick, 1892 (Orthostixidae) (1 name)
Subfamily rank questioned (Holloway 1996: 151), possibly subordinated to Ennominae. Hausmann (1996a, 1996b) recognized the subfamily but noted that the genera Derambila and Ozola seem to link with Myinodes/Eumegethes, Orthostixis/Naxa and Gypsochroa. The validity at subfamily rank was maintained by Hausmann (2001: 70, 89). Tentative molecular information on the genus Naxa suggests subordination under Ennominae (Sihvonen 2011; Murillo-Ramos et al. 2019) despite several contradicting morphological characters. Recently, the name-bearing genus Orthostixis was included in a whole-genome shotgun sequencing analysis and Orthostixis was reconstructed as sister to Ozola (Murillo-Ramos et al. 2021b). Therefore, the authors proposed Orthostixinae as a junior synonym of Desmobathrinae, but the publication and names and nomenclatural acts therein are not available for purposes of zoological nomenclature in accordance with ICZN (2012, Arts. 8.2, 8.3), and Murillo-Ramos and collaborators will formalise the synonymy in a forthcoming publication.
Subfamily Epidesmiinae Murillo-Ramos, Brehm & Sihvonen, 2018 (Epidesmiinae) (1 name)
Epidesmiinae were recognised as a valid subfamily by Murillo-Ramos et al. (2019). Epidesmiinae have an Australasian distribution, with one species occurring in the Indo-Malayan realm. The lineage includes 102 described species that are classified into nine genera. Those are not classified into tribes thus far, and no family-level synonyms have been recognised (Murillo-Ramos et al. 2021a).
Subfamily Oenochrominae Guenée, 1858 (Oenochromidae) (3 names)
Oenochrominae s. str. has been recovered as sister to Geometrinae in molecular phylogenetic analyses (Sihvonen et al. 2011; Murillo-Ramos et al. 2019).
= Lyrceini Meyrick, 1883 (Lyrceini), based on Lyrcea, a junior homonym of a name outside Lepidoptera; junior subjective synonym. Invalid according to ICZN (1999, Art. 39).
= Monocteniini Meyrick, 1889 (Monocteniadae), emended, junior subjective synonym (Holloway 1996: 151).
Subfamily Ennominae Duponchel, 1845 (Ennomites) (93 names)
Validity and order of tribes mainly based on Murillo-Ramos et al. (2019), supplemented by information from Holloway (1994), Pitkin (2002), Beljaev (2008b), Skou & Sihvonen (2015), Jiang et al. (2017) and Brehm et al. (2019). Brehm et al. (2019), in their phylogenetic hypothesis, highlighted four lineages, marked as “unnamed clade”, which may need to be formally described at tribe level when more data become available. Those are not included in the current list.
Tribe Euangeronini Brehm, Murillo-Ramos & Sihvonen, 2019 (Euangeronini)
Type genus Euangerona, previously assigned to Nacophorini (Pitkin 2002).
Tribe Gonodontini Forbes, 1948 (Gonodontini)
See remarks in Holloway (1994: 8, 111). Gonodontini needs to be referred to the Commission for a ruling (ICZN 1999, Arts. 41 and 65.2.), because of the misidentification of the generic type species (see Skou & Sihvonen 2015).
Tribe Gnophini Duponchel, 1845 (Gnophites)
Earlier suggested to fall, probably, under the wide concept of Boarmiini by Holloway (1994: 7), but recent molecular phylogenies have recovered Gnophini as a monophyletic lineage sister to Gonodontini (Murillo-Ramos et al. 2019).
= Aspilatini Duponchel, 1845 (Aspilatites), senior subjective synonym (Holloway: 1994: 5), but based on a misspelled genus group name (Aspitates) (Holloway: 1994: 5). See Skou & Sihvonen (2015) for further information.
= Dasydini Duponchel, 1845 (Dasydites), based on Dasydia Guenée, 1845, a junior objective synonym of Sciadia.
= Sionini Duponchel, 1845 (Sionites). Included in Gnophini by Herbulot (1961–1963), confirmed repeatedly, for instance by Viidalepp et al. (2007), Wahlberg et al. (2010), Skou & Sihvonen (2015) and Murillo-Ramos et al. (2019).
= Angeronini Forbes, 1948 (Angeronini). Considered valid at tribal rank (e.g., by Mcguffin 1981; Viidalepp 1996; Hausmann et al. 2011; Vives Moreno 2014), but included as a junior synonym of Gnophini by Skou & Sihvonen (2015) and Beljaev (2016). Synonymy also confirmed by molecular data (Yamamato & Sota 2007; Sihvonen et al. 2011; Murillo-Ramos et al. 2019).
= Psodini Povolný & Moucha, 1955: 155, [Fig.] 1 (Psodinae). Not an available name because description is absent (Psodini is only mentioned on the figure). Included in Gnophini by Beljaev (2016).
= Diaprepesillini Kuznetzov & Stekolnikov, 1982: 369 (Diaprepesillini). Included in Gnophini by Viidalepp (1996) and Beljaev (2016).
Tribe Odontoperini Tutt, 1896 (Odontoperinae)
See Holloway (1994: 8, 111). Related to Nacophorini s. l. (Murillo-Ramos et al. 2019).
= Crocallini Tutt, 1896 (Crocallidi), junior subjective synonym according to Beljaev (2016). Limited molecular data suggest relationships to Ennomos and Opisthograptis (Õunap et al. 2011). Skou & Sihvonen (2015) classified Crocallis in Ennomini of uncertain association. More research is needed.
= Azelinini Forbes, 1948 (Azelinini). A junior subjective synonym, proposed on the basis of molecular data (Brehm et al. 2019). In the phylogenetic analysis, Azelinini (represented by Pero) nested within Odontoperini (Brehm et al. 2019). Earlier, based on functional morphology of the male genitalia, relationships to Ennomini s. l. and Prosopolophini were suggested by Beljaev (2009).
Tribe Nacophorini Forbes, 1948 (Nacophorini)
Relationship to Odontoperini (= Azelinini [= junior synonym of Odontoperini] suggested by Holloway (1994: 8). This view is further supported by molecular data (Murillo-Ramos et al. 2019). The North American fauna was revised by Rindge (1983).
Tribe Ennomini Duponchel, 1845 (Ennomites)
= Odopterini Stephens, 1850 (Odopteridi), based on Odoptera, an unnecessary replacement name for Ennomos.
= Ourapterygini Bruand, 1846 (Urapteridae). Emended by Forbes (1848), Holloway (1996: 6). Subordinated under Ennomini by Sihvonen et al. (2011), as earlier suggested by morphology (Beljaev 2008a). Ourapterygini was subordinated under Ennomini based also on extensive molecular data (Murillo-Ramos et al. 2019).
= Emplociini Guenée, 1858 (Emplocidae). Emended by reasons of grammar. Junior subjective synonym (Pitkin 2002: 135). = Oxydiini Butler, 1886 (Oxydiidae). Junior subjective synonym (Pitkin 2002: 135).
= Pantherini Moore, 1887 (Pantheridae); based on a junior homonym of a genus-group name outside Lepidoptera (Holloway 1994: 9); invalid according to ICZN (1999, Art. 39).
= Nephodiini Warren, 1894 (Nephodiinae). Junior subjective synonym (Beljaev 2008a), further supported by molecular data (Murillo-Ramos et al. 2019; Brehm et al. 2019).
= Leuculini Hulst, 1896: 249 (Leuculinae), 317 (Leuculidae). Type genus: Leucula Guenée, 1858. Male
genitalia are typical for Ourapterygini (see Pitkin 2002). However, the description of the tribe is based on “Leucula lacteolata Hulst”, a taxon situated outside Geometridae (E. Beljaev, pers. comm.). Leucula subordinated under Ennomini in mole-clar analyses (Murillo-Ramos et al. 2019, Brehm et al. 2019).
= Cingiliini Forbes, 1948 (Cingiliini). Junior subjective synonym (Holloway 1994: 6; pitkin 2002: 135).
Tribe Rumiini Tutt, 1896 (Rumiinae)
Based on Rumia, which is a junior objective synonym of Opisthograptis. Junior subjective synonym of Ennomini according to Leraut (1997). Phylogenetic position still under investigation. In the molecular analysis of Sihvonen et al. (2011), reconstructed as sister to Epirranthis diversata; in the analysis of Õunap et al. (2011), as sister to Crocallis elinguaria.
Tribe Thinopterygini Holloway, 1994 (Thinopterygini)
Tribe Campaeini Forbes, 1948 (Campaeini)
= Metrocampini Tutt, 1896 (Metrocampidae), senior synonym. Based on Metrocampa, which is a junior objective synonym of Campaea. It is proposed not to apply this name, as Campaeini is largely accepted (see Holloway 1994: 6). In Murillo-Ramos et al. (2019), Campaeini lumped into a single cluster with Campaeini, Alsophilini and Prosopolophini.
Tribe Alsophilini Herbulot, 1963 (Alsophilinae)
Treated as a subfamily by Hausmann (2001), despite earlier doubts about subfamily rank and the suggestion of subordination under Ennominae (Holloway 1996: 149). Evidence from molecular analyses, however, supports the rank of tribe under Ennominae (e.g., Sihvonen et al. 2011; Murillo-Ramos et al. 2019). In Sihvonen et al. (2011) and Murillo-Ramos et al. (2019), classified as a tribe and Alsophilini united into a single cluster with Campaeini, Wilemaniini and Prosopolophini.
Tribe Cheimoptenini Kuznetzov & Stekolnikov, 1982: 347 (Cheimoptenini)
Position of tribe unclear. It has been placed in Ennominae (Beljaev 2006) and in Desmobathrinae (Beljaev 2008b), but has not been subjected to studies on molecular phylogeny.
Tribe Wilemaniini Wehrli, 1941 (Wilemaninae [sic!])
The original Latin word stem Wileman- (with the suffix -inae) needs to be emended to Wilemani- (being based on the type genus Wilemania), leading to “Wilemaniini” rather than to Wilemanini. In Murillo-Ramos et al. (2019), Wilemaniini united into a single cluster with Campaeini, Alsophilini and Prosopolophini. Beljaev (2016) listed Wilemaniini as a junior synonym of Prosopolophini.
Tribe Prosopolophini Warren, 1894: 464 (Prosopolophinae)
In Murillo-Ramos et al. (2019), Prosopolophini was united into a single cluster with Campaeini, Alsophilini and Wilemaniini.
= Ligiini Guenée, 1858 (Ligidae), emended, based on Ligia. Senior synonym, but based on a junior homonym of a genus-group name outside Lepidoptera. Invalid according to ICZN (1999, Art. 39).
= Colotoini Wehrli, 1940: 345 (Colotoinae). Synonymy proposed in Beljaev (2016). In the molecular phylogeny of Brehm et al. (2019), the Holarctic Colotois pennaria (Colotoini) grouped next to the Central American species Himeromima aulis Druce, 1892, which was assigned to Prosopolophini.
= Compsopterini Herbulot, 1963 (Compsopterini). The genus Compsoptera Blanchard, 1845 has not been included in a molecular phylogeny yet. Compsoptera was listed under Prosopolophini in Beljaev (2016) and Hausmann & Sihvonen (2019).
= Zamacrini “Meyrick nec Agassis”: cf. Viidalepp (1989: 102) and Zoological Record (1990/1991, 13D: 326): “Apochimini nom. nov. Viidalepp … for Zamacrini Meyrick nec Agassis”. Listed under Prosopolophini in Beljaev (2016).
= Apochimini Viidalepp, 1989: 102 (Apochimini) (an unnecessary replacement name for Zamacrini). Listed under Prosopolophini in Beljaev (2016).
Tribe Onychorini Herbulot, 1963 (Onychorini)
Hausmann & Sihvonen (2019) listed Onychora Meyrick, 1892 in “Genera of uncertain tribus association”. Not included in a molecular phylogeny yet.
Tribe Diptychini Janse, 1933 (Diptychini)
Historically, Diptychini was considered as an intermediate group between Geometrinae [Geometrini] and Larentiinae [Larentiini] by Janse (1933: 2) and as part of Oenochrominae by Prout (1931: 120, in: Prout 1929–1935). For subordination under Ennominae and supposed relationships to Nacophorini and Ourapterygini, see Holloway (1996: 150–151) and Pitkin (2002: 135). In the most recent and most extensive molecular dataset to date, Diptychini is nested within Ennominae and classsified as a tribe, being sister to an unnamed clade and the Campaeini + Alsophilini + Wilemaniini + Prosopolophini complex. It does not show close relationship to Nacophorini (Murillo-Ramos et al. 2019).
= Lithinini Forbes, 1948 (Lithinini). Synonymised with Diptychini by Murillo-Ramos et al. (2019). Close relationship with Caberini according to Abraham et al. (2001; fig. 5b), not confirmed subsequently.
= Pachycnemiini Kirby, 1903 (Pachycnemiidae). The structure of the male and female genitalia fit that of the Lithinini of Rindge (1986). Synonymy confirmed by Murillo-Ramos et al. (2019).
= Epirrhanthini Forbes, 1948 (Epirrhanthini). In a molecular analysis, the group was included in Ennominae (Sihvonen et al. 2011), but tribal status was not given. Relationships to Ennomini and Desmobathrinae were discussed in Hausmann (2001: 99). Treated under Ennominae in Holloway (1994: 7) and under Lithinini in Beljaev (2016). Hausmann & Sihvonen (2019) recognised Epirrhanthini as a valid tribe, listing it after Lithini in the sequence of tribes.
= Lacariini Orfila & Schajovskoy, 1959 (Lacarini), emended, based on Lacaria. For possible synonymy to Lithinini, see Holloway (1994: 7, 92).
Tribe Oenoptilini Brehm, Murillo-Ramos & Sihvonen, 2019 (Oenoptilini)
The genera Neobapta and Oenoptila, currently classified in Oenoptilini, were previously assigned to Caberini (Pitkin 2002).
Tribe Baptini Forbes, 1948 (Baptini)
Closely related to Palyadini (Abraham et al. 2001; fig. 5b) and subordinated under Caberini in Pitkin (2002: 131). In Murillo-Ramos et al. (2019) and Brehm et al. (2019), treated as a tribe and included in a genetic cluster with Theriini.
= Aleucini Djakonov, 1936: 484 (Aleucini). Treated as a junior synonym of Baptini in Hausmann et al. (2011), but more recently classified in Theriini (Skou & Sihvonen 2015; Hausmann & Sihvonen 2019). The name-bearing genus Aleucis Guenée, 1845 has not been included in molecular phylogenetic analysis yet. Aleucini is the senior synonym if included in the concept of Baptini or Theriini.
= Lomographini Wehrli, 1940: 381, 382 (Lomographinae). The original concept was based on the genus Stegania (possibly Abraxini), because of the misidentification of the generic type species. It needs to be referred to the ICZN Commission for a ruling (ICZN 1999, Arts. 41, 65.2.) (E. Beljaev, pers. comm.).
Tribe Theriini Herbulot, 1963 (Theriini)
Falls into the broad conception of Boarmiini according to Holloway (1994: 13, 167). In Murillo-Ramos et al. (2019) and Brehm et al. (2019), treated as a tribe and included in a genetic cluster with Baptini. See Aleucini under Baptini.
= Cheimatobiini Tutt, 1896 (Cheimatobiidi), senior synonym. Based on Cheimatobia Stephens, 1829, which is a junior objective synonym of Theria. It is proposed not to apply this name, as Theriini is widely accepted (see Holloway 1994: 6).
Tribe Palyadini Guenée, 1858 (Palyadae)
Earlier subordination under Baptini rejected by Holloway (1994: 59), but accepted by Abraham et al. (2001: fig. 5b). Pitkin (2002: 132) suggested status as a subtribe of Caberini/Baptini. In Murillo-Ramos et al. (2019) and Brehm et al. (2019), treated at tribe rank, closely related to Plutodini.
Tribe Epionini Bruand, 1846 (Epionidae)
= Hypochrosini Guenée, 1858 (Hypochrosinae). Hypochrosini sensu Holloway (1994). Potential synonymy suggested also by Murillo-Ramos et al. (2019), but there and in Brehm et al. (2019) formally still treated as a separate tribe because of limited taxon sampling.
= Scardamiini Warren, 1894 (Scardamiinae), based on Scardamia. Treated as a junior synonym of Epionini by Beljaev (2016).
= Anagogini Forbes, 1948 (Anagogini), junior synonym (Holloway: 1994: 5). Potential synonymy suggested also by Murillo-Ramos et al. (2019), but there and in Brehm et al. (2019) formally still treated as a separate tribe because of limited taxon sampling.
= Seleniini Tutt, 1896 (Seleniidi), suggested as a junior synonym in Holloway (1994: 9).
= Apeirini Kuznetzov & Stekolnikov, 1982: 358 (Apeirini). Potential synonymy with Epionini suggested (but not formally established) by Murillo-Ramos et al. (2019) as well as by Brehm et al. (2019). Treated as a valid tribe by Sihvonen & Skou (2015), Beljaev (2016) and Hausmann & Sihvonen (2019). Apeirini is a morphologically isolated group (Sihvonen & SKOU 2015). More research and, particularly, more extensive taxon sampling in this complex are needed.
Tribe Drepanogynini Murillo-Ramos, Sihvonen & Brehm, 2019 (Drepanogynini)
Genus Drepanogynis earlier subordinated under a wider concept of Nacophorini.
Tribe Pyriniini Brehm, Murillo-Ramos & Sihvonen, 2019 (Pyriniini)
Unassigned by Pitkin (2002), who suggested relationships with Caberini/Baptini. Sister lineage relationship to Caberini supported in Murillo-Ramos et al. (2019).
Tribe Caberini Duponchel, 1845 (Caberites)
Sister lineage relationship to Pyriniini supported in Murillo-Ramos et al. (2019).
= Erastriini Herrich-Schäffer, 1845 (Erastridae), junior synonym (Holloway 1996: 7; 98). Emended, based on Erastria, with the Latin word stem Erastri-. The original concept referring to Noctuidae: Acontiinae. Classification of the generic type species (see Fletcher 1979) leaves this family-group name applicable within Ennominae (Holloway 1994: 7).
= Brotini Grote, 1882 (Brotiinae). Emended, based on Brotis, with the Latin word stem Brot-, which is a junior homonym to a genus-group name in Noctuidae. Invalid according to ICZN (1999, Art. 39). Junior synonym in the broad concept of Caberini/Baptini of Pitkin (2002: 131).
= Deiliniini Warren, 1894 (Deiliniinae), based on Deilinia, a junior synonym of Cabera (Holloway 1996: 6; 98).
= Catopyrrhini Warren, 1894 (Catopyrrhinae), junior synonym (Holloway 1996: 6; 98).
= Sphacelodini Forbes, 1948 (Sphacelodini), retained as a synonym in the broad concept of Caberini/Baptini of Pitkin (2002: 131). Based on illustrations in Pitkin (2002), this group could be close to Beljaev's concept of Deviniliini (E. Beljaev, pers. comm.). In molecular phylogenies, Sphacelodes vulneraria (Hübner, 1823) grouped within Caberini (Murillo-Ramos et al. 2019; Brehm et al. 2019).
Tribe Deveniliini Beljaev, 1998: 440 (Deveniliini)
Subordinated under Baptini by StÜning (2000), maintained valid as a tribe by Beljaev (2016). Not included in molecular phylogenetic studies so far.
Tribe Cassymini Holloway, 1994 (Cassymini)
The molecular analysis of Murillo-Ramos et al. (2019) revealed two clearly distinct genetic clusters, with Cassymini clustering together with Abraxini, Eutoeini and Macariini.
Tribe Abraxini Warren, 1893 (Abraxinae)
= Zerenini Duponchel, 1845 (Zerenites), based on a junior homonym of a genus-group name outside Lepidoptera; junior synonym (Holloway 1994: 9). Invalid according to ICZN 1999, Art. 39). Abraxini clustered together with Cassymini, Eutoeini and Macariini in the analyses of Murillo-Ramos et al. (2019).
Tribe Eutoeini Holloway, 1994 (Eutoeini)
Eutoeini clustered together with Abraxini and Cassymini in the analyses of Murillo-Ramos et al. (2019).
Tribe Macariini Guenée, 1858 (Macaridae)
The original Latin word stem Macar- (with the suffix -idae) needs to be emended to Macari- (being based on the type genus Macaria), leading to “Macariini” rather than to Macarini. Macariini clustered with Cassymini, Abraxini and Eutoeini in Murillo-Ramos et al. (2019).
= Atomorphini Wehrli, 1953: 642 (Atomorphinae). Originally part of Semiothisinae, treated similarly in Wiltshire (1990), but later classified in Gnophini (e.g., Viidalepp 1996). The type genus Atomorpha Staudinger, 1901 was considered a junior synonym of Isturgia Hübner, 1823 by Skou & Sihvonen (2015) in Macariini. The name-bearing genus Atomorpha has not been included in molecular phylogenetic studies.
= Semiothisini Warren, 1894 (Semiothisinae), junior synonym (HOLLOWAY 1994: 9).
= Fernaldellini Hulst, 1896 (Fernaldellinae), junior synonym (Holloway 1994: 7).
Tribe Boarmiini Duponchel, 1845 (Boarmites)
A very broad concept of this tribe was proposed by Holloway (1994: 167), confirmed by recent molecular analyses (Murillo-Ramos et al. 2019; Murillo-Ramos et al. 2021c). Jiang et al. (2017) presented a detailed molecular analysis of the tribe, identifying 14 monophyletic lineages (potential subtribes) within it.
= Cleorini Duponchel, 1845 (Cleorites), junior synonym (Holloway: 1994: 6). Synonymy further supported by molecular analyses (e.g., Murillo-Ramos et al. 2019, 2021c).
= Fidoniini Duponchel, 1845 (Fidonites), emended by reasons of grammar; based on Fidonia, a junior synonym of Eurranthis (Leraut 1997: 214). Synonymy further supported by molecular analyses (e.g., Murillo-Ramos et al. 2019, 2021c).
= Ascotini Warren, 1893 (Ascotinae), junior synonym (Holloway: 1994: 5). Synonymy further supported by molecular analyses (e.g., Murillo-Ramos et al. 2019, 2021c).
= Bistonini Stephens, 1850 (Bistonidi). Falling into the broad concept of Boarmiini according to Holloway (1994: 13, 167). Synonymy further supported by molecular analyses (e.g., Sihvonen et al. 2011; Murillo-Ramos et al. 2019, 2021c).
= Amphidasini Duponchel, 1845 (Amphidasites), senior synonym. Based on Amphidasis, a junior objective synonym of Biston. It is proposed not to apply this name (neither for a potential subtribe), as Bistonini/Bistonina is widely accepted (see Holloway 1994: 5).
= Hyberniini Duponchel, 1845 (Hibernites), senior synonym. Emended. Based on Hybernia, a junior objective synonym of Erannis. It is proposed not to apply this name, as Bistonini/Bistonina is widely accepted (see Holloway 1994: 5, 7).
= Eubyjini Warren, 1893 (Eubyjinae), based on Eubyja, which is a junior synonym of Biston (Holloway 1994: 7).
= Erannini Tutt, 1896 (Eranniinae), junior synonym (Holloway 1994: 7). Synonymy further supported by molecular analyses (e.g., Murillo-Ramos et al. 2019, 2021c). Emended, based on Erannis, with the Latin word stem Erann-.
= Desertobiini Viidalepp, 1989: 104 (Desertobiini). Subordinated under Boarmiini by Beljaev (2000).
= Dalimini Wehrli, 1940 (Daliminae), junior synonym (Holloway 1996: 150).
= Selidosemini Meyrick, 1892 (Selidosemidae), junior synonym (Holloway 1994: 9). Synonymy further supported by molecular analyses (e.g., Murillo-Ramos et al. 2019, 2021c).
= Braccini Warren, 1894 (Braccinae), junior synonym (Holloway 1994: 167).
= Melanchroiini Hulst, 1896 (Melanchroiinae), junior synonym (Pitkin 2002: 130). Synonymy further supported by molecular analyses (e.g., Murillo-Ramos et al. 2019, 2021c).
= Phaseliini Wehrli, 1941 (Phaseliinae), suggested as falling into the wide concept of Boarmiini. Synonymy further supported by molecular analyses (e.g., Murillo-Ramos et al. 2019, 2021c).
= Melanolophini Forbes, 1948 (Melanolophini), junior synonym (Holloway 1994: 8; Pitkin 2002: 131). Synonymy further supported by molecular analyses (e.g., Murillo-Ramos et al. 2019, 2021c).
= Glaucinini Rindge, 1959: 265 (Glaucinini). Boarmiini synonymy supported by molecular analysis (Murillo-Ramos et al. 2019, 2021c).
= Bupalini Herbulot, 1963 (Bupalini), junior synonym (Holloway 1994: 6; 167). Synonymy further supported by molecular analyses (e.g., Murillo-Ramos et al. 2019, 2021c).
= Milioniini Holloway, 1994 (Milioniini). First mentioned by Inoue (1992), but without description and thus unavailable. Junior synonym (Holloway 1994: 8, 302). Synonymy further supported by molecular analyses (e.g., Murillo-Ramos et al. 2019, 2021c).
Tribe Cystidiini Kuznetzov & Stekolnikov, 1982: 344 (Cystidiini)
Not included in molecular phylogenetic analysis so far. The tribe cannot be subordinated to Boarmiini because the vinculum and tegumen are separated one from another by a deep narrowing. Lateral lobes of anellus in Cystidia completely reduced; in Obeidia, the mentioned structures possibly remained, but homologies are difficult to interpret [cf. Inoue's (1992) revision of Obeidia] (E. Beljaev, pers. comm.).
= Obeidiini Holloway, 1994 (Obeidiini). First mentioned by Inoue (1992), but without description and thus unavailable. Junior synonym (Holloway 1994: 8, 302).
Appendix 2.
List of literature examined for the preparation of the “Online taxonomic facility of Geometridae”, spanning 1998 to 23.5.2022 and only including references not already listed in Scoble (1999) or Scoble & Hausmann (2007).
Aarvik, L. & Bjornstad, A. (2007): Review of the genus Zamarada Moore, 1887 (Lepidoptera: Geometridae) in Tanzania, with description of new species. – Esperiana Memoir 3: 7–57, 398–415.
Agassiz, D. J. L. (2009): The Macrolepidoptera fauna of Acacia in the Kenyan Rift Valley (Part 2 – Description of new species). – Tropical Lepidoptera Research 19 (1): 9–17.
Aguiar, A. M. F. & Karsholt, O. (2006): Lepidoptera - Systematic catalogue of the entomofauna of the Madeira Archipelago and Selvagens Islands. – Boletim do Museu Municipal do Funchal, Supplement 9: 5–139.
Bálint, Z. & Katona, G. (2016): Notes on two Transcaucasian Lepidoptera described by Gusztáv Emich in 1872 and 1873. – Caucasian Entomological Bulletin 12 (1): 139–142. https://doi.org/10.23885/1814-3326-2016-12-1-139-142
Ban, X., Jiang, N., Cheng, R., Xue, D. & Han, H. (2018): Tribal classification and phylogeny of Geometrinae (Lepidoptera: Geometridae) inferred from seven gene regions. – Zoological Journal of the Linnean Society 184 (3): 653–672. https://doi.org/10.1093/zoolinnean/zly013
Beck, K. R. & Karisch, T. (2016): Ein weiterer Beitrag zur Kenntnis der Gattung Mesothisa Warren, 1905 (Lepidoptera, Geometridae, Ennominae). – Lambillionea CXVI (2): 100–104.
Beck, K. R. & Karisch, T. (2008): Beschreibung einer neuen Art der Gattung Achrosis Guenee, [1858] von den Philippinen (Lepidoptera: Geometridae, Ennominae). – Entomologische Zeitschrift 118 (2): 51–52.
Beck, K. R. & Karisch, T. (2011): Zwei neue afrikanische Arten von Zamarada Moore, [1887] (Lepidoptera: Geometridae). – Entomologische Zeitschrift 121 (2): 85–87.
Beljaev, E. (2007): Taxonomic changes in the emerald moths (Lepidoptera: Geometridae, Geometrinae) of East Asia, with notes on the systematics and phylogeny of Hemitheini. – Zootaxa 1584 (1): 55–68. https://doi.org/10.11646/zootaxa.1584.1.2
Beltran, E. M. (2008): Compsoptera aemiliroum Raineri, 1994, a synonym of C. opacaria (Huebner, [1819]) (Lepidoptera: Geometridae, Ennominae). – Boletín de la SEA 42: 297–303.
Bolotov, I. N., Frolov, A. A., Kolosova, Y. S. & Kondakov, A. V. (2014): The male of Sauris mouliniei (Legrand, 1971) comb. n. (Lepidoptera: Geometridae: Larentiinae: Trichopterygini), an endemic Inner Seychelles moth. – Zootaxa 3765 (4): 397–400. https://doi.org/10.11646/zootaxa.3765.4.8
Bolshakov, L. V. & Ismagilov, N. N. (2016): Moths of Republic of Tatarstan. 1. Geometridae (Lepidoptera). – Eversmannia 47: 35–80.
Brehm, G. (2015): Three new species of Hagnagora Druce, 1885 (Lepidoptera, Geometridae, Larentiinae) from Ecuador and Costa Rica and a concise revision of the genus. – ZooKeys 537: 131–156. https://doi.org/10.3897/zookeys.537.6090
Brehm, G. (2018): Revision of the genus Callipia Guenée, 1858 (Lepidoptera, Geometridae), with the description of 15 new taxa. – European Journal of Taxonomy 404: 1–54. https://doi.org/10.5852/ejt.2018.404
Brehm, G., Murillo-Ramos, L., Sihvonen, P., Hausmann, A., Schmidt, B., Õunap, E., Moser, A., Mörtter, R., Bolt, D., Bodner, F., Lindt, A., Parra, L. & Wahlberg, N. (2009): New World geometrid moths (Lepidoptera: Geometridae): Molecular phylogeny, biogeography, taxonomic updates and description of 11 new tribes. – Arthropod Systematics & Phylogeny 77: 457-486. https://doi.org/10.26049/ASP77-3-2019-5
Byrne, C. J. & Wei, N. S. (2012): Kunanyia stephaniae gen. nov. & sp. nov. (Lepidoptera: Geometridae: Ennominae): an unusual and rare diurnal moth from the mountains of Tasmania. – Zootaxa 3503 (1): 25–46. https://doi.org/10.11646/zootaxa.3503.1.2
Can, F. (2009): DNA barcoding confirms species rank for a cryptic geometrid species from Turkey and Bulgaria (Lepidoptera: Geometridae: Sterrhinae). – Zootaxa 2314 (1): 63–68. https://doi.org/10.11646/zootaxa.2314.1.4
Can, F. & Mironov, V. (2006): Perizoma onurcani sp. n. from Turkey (Geometridae: Larentiinae). – Nota Lepidopterologica 28 (3–4): 163–166.
Chang, W. & Wu, S. (2013): Review of the genus Hemistola Warren, 1893 in Taiwan with notes on an unusual conifer-feeding larva and descriptions of three new species (Lepidoptera, Geometridae, Geometrinae). – Zootaxa 3741 (4): 538–550. https://doi.org/10.11646/zootaxa.3741.4.5
Cheng, R., Xue, D. & Jiang, N. (2019): A taxonomic study of the subspecies of Ourapteryx ebuleata Guenée, 1858 (Lepidoptera: Geometridae). – Entomotaxonomia 41 (2): 154–159.
Choi, S. W. & Kim, S. S. (2016): A checklist of the genus Scopula (Lepidoptera: Geometridae) including description of a new species and three newly recorded species from Korea. – Zootaxa 4178 (1): 131–137. https://doi.org/10.11646/zootaxa.4178.1.6
Choi, S. W. & Stüning, D. (2011): Revision of the genus Paraplaneta Warren, 1895 (Lepidoptera: Geometridae, Larentiinae) from Southeast Asia. – Zootaxa 3038 (1): 29–44. https://doi.org/10.11646/zootaxa.3038.1.2
Cock, M. J. W. (2017): A preliminary catalogue of the moths (Lepidoptera except Papilionoidea) of Tobago, West Indies. – Insecta Mundi 585: 1–58.
Covell, C. V. Jr. (2013): Two Mexican Geometridae new to the United States, with a new synonymy. – Tropical Lepidoptera Research 24 (1): 59–60.
Covell, C. V. Jr. (2015): Three New Species of Idaea Treitschke (Geometridae: Sterrhinae) from the Southwestern United States and Northern Mexico. – Journal of the Lepidopterists' Society 69 (4): 317–325. https://doi.org/10.18473/lepi.69i4.a7
Covell, C. V. Jr. & Heppner, J. B. (2017a): Review of the genus Simena from Central America (Lepidoptera: Geometridae: Ennominae). – Lepidoptera Novae 10 (1–2): 13–20.
Covell, C. V. Jr. & Heppner, J. B. (2017b): The new genus Neosimena from Peru and western Brazil, with two new species (Lepidoptera: Geometridae: Ennominae). – Lepidoptera Novae 10 (1–2): 9–12.
Cui, L. & Jiang, N. (2018): Aquilargilla gen. nov., a new genus of Sterrhinae from China with description of two new species (Lepidoptera, Geometridae). – Zootaxa 4514 (3): 431–437. https://doi.org/10.11646/zootaxa.4514.3.8
Cui, L., Jiang, N., Stüning, D. & Han, H. (2018): A review of Synegiodes Swinhoe, 1892 (Lepidoptera: Geometridae), with description of two new species. – Zootaxa 4387 (2): 259–274. https://doi.org/10.11646/zootaxa.4387.2.2
Cui, L., Xue, D. & Jiang, N. (2019a): A review of Timandra Duponchel, 1829 from China, with description of seven new species (Lepidoptera, Geometridae). – ZooKeys 829: 43–74. https://doi.org/10.3897/zookeys.829.29708
Cui, L., Xue, D. & Jiang, N. (2019b): A review of Organopoda Hampson, 1893 (Lepidoptera, Geometridae) from China, with description of three new species. – Zootaxa 4651 (3): 434–444. https://doi.org/10.11646/zootaxa.4651.3.2
Cui, L., Xue, D. & Jiang, N. (2019c): Description of two new species of Rhodostrophia Hübner, 1823 from China (Lepidoptera, Geometridae). – Zootaxa 4563 (2): 337–353. https://doi.org/10.11646/zootaxa.4563.2.7
Cui, L., Xue, D. & Jiang, N. (2020): Two new species of the tribe Rhodometrini Agenjo, 1951 from Sichuan, China (Lepidoptera: Geometridae). – SHILAP Revista de Lepidopterologia 48 (189): 167–172.
Da, W. & Wang, M. (2019): A new species of the genus Timandromorpha Inoue, 1944 (Lepidoptera, Geometridae, Geometrinae) from S. Xizang, China. – Japan Heterocerists' Journal 288: 328–329.
Dey, P., Uniyal, V. P., Hausmann, A. & Stüning, D. (2021): Revision of the genus Prometopidia Hampson, 1902, with description of the new species P. joshimathensis sp. nov. from West-Himalaya and its subspecies P. j. yazakii ssp. nov. from Nepal (Lepidoptera: Geometridae, Ennominae). – Zootaxa 4980 (1): 028–044. https://doi.org/10.11646/zootaxa.4980.1.2
Embacher, G., Murauer, K. & Tarmann, G. M. (2005): Thera variata mugo Burmann & Tarmann, 1983 - syn. n. von T. cembrae Kitt, 1912 - (Lepidoptera: Geometridae). – Nachrichtenblatt der Bayerischen Entomologen 54 (3–4): 73–81.
Erlacher, S. & Erlacher, J. (2016): A systematic revision of the genus Gnophopsodos Wehrli, 1945, with description of two new species (Lepidoptera: Geometridae). – Zootaxa 4169 (3): 435–456. https://doi.org/10.11646/zootaxa.4169.3.2
Erlacher, S. & Erlacher, J. (2017): A new species of Charissa Curtis, 1826 from Europe (Lepidoptera: Geometridae). – Zootaxa 4341 (1): 089–096. https://doi.org/10.11646/zootaxa.4341.1.7
Erlacher, S. & Junghans, C. (2009): On the identity of Psodos perlinii Turati, 1914 (Geometridae: Ennominae). – Nota Lepidopterologica 32 (1): 47–54.
Erlacher, S., Marrero Palma, L. & Erlacher, J. (2017): A systematic revision of Charissa, subgenus Pterygnophos Wehrli, 1951, with description of a new species (Lepidoptera: Geometridae). – Zootaxa 4341 (3): 400–418. https://doi.org/10.11646/zootaxa.4341.3.4
Expósito-Hermosa, A. (2007): Nuevas contribuciones para la fauna de España (Lepidoptera: Geometridae). – SHILAP Revista de Lepidopterologia 35 (138): 269–271.
Expósito-Hermosa, A. (2018a): A new species of the genus Bracca Huebner, [1820] from Sulawesi, Indonesia (Lepidoptera: Geometridae, Ennominae, Boarmiini). – SHILAP Revista de Lepidopterologia 46 (184): 581–584.
Expósito-Hermosa, A. (2018b): A new species of the genus Platycerota Hampson, 1893 from Myanmar (Burma) (Lepidoptera: Geometridae, Ennominae, Baptini). – SHILAP Revista de Lepidopterologia 46 (184): 695–697.
Expósito-Hermosa, A. (2019a): A new species of genus Gonodontis Hübner, (1823) from Indonesia, West of the Timor Island (Lepidoptera: Geometridae, Ennominae, Gonodontini). – SHILAP Revista de Lepidopterologia 47 (186): 233–236.
Expósito-Hermosa, A. (2019b): A new species of the genus Ecliptopera Warren, 1894 from Sulawesi. Indonesia (Lepidoptera: Geometridae, Larentiinae, Cidariini). – SHILAP Revista de Lepidopterologia 47 (185): 189–191.
Expósito-Hermosa, A. (2019c): A new species of the genus Xylinophylla Warren, 1898 from West Papua, Indonesia (Lepidoptera: Geometridae, Ennominae, Gonodontini). – SHILAP Revista de Lepidopterologia 47 (186): 209–211.
EXPÓSITO-HERMOSA, A. (2019d): A new species of the genus Yazakia Warren, 1894 from the island of Sulawesi (Celebes), Indonesia (Lepidoptera: Geometridae, Ennominae, Boarmiini). – SHILAP Revista de Lepidopterologia 47 (185): 49–51.
Expósito-Hermosa, A. (2019e): New contributions for the genus Bulonga Walker, 1859 with description from a new species of Sulawesi, Indonesia and Pseudobulonga Expósito, gen. n. (Lepidoptera: Geometridae, Ennominae, Baptini). – Shilap Revista de Lepidopterologia 47 (187): 475–478.
Expósito-Hermosa, A. (2020): A new species of the genus Synegia Guenee, 1858 from the Moluccas Islands: Ceram-Seram, Indonesia (Lepidoptera: Geometridae, Ennominae, Baptini). – SHILAP Revista de Lepidopterologia 48 (190): 253–255.
Expósito-Hermosa, A. (2020): Una nueva especie del género Naxa Walker, 1856, de las montañas Arfak, Papúa Occidental (Indonesia) (Lepidoptera: Geometridae, Orthostixinae). – SHILAP Revista de Lepidopterologia 48 (192): 689–691.
Expósito-Hermosa, A. (2021): Una nueva especie del género Abraxas Leach, [1815] del Monte Langgaliru, de la isla de Sumba (Indonesia) (Lepidoptera: Geometridae, Ennominae). – SHILAP Revista de Lepidopterologia 49 (196): 691–694.
Expósito-Hermosa, A. & Viidalepp, J. (2011): Xanthorhoe iberica (Staudinger, 1901) sp. bon., de España (Lepidoptera: Geometridae, Larentiinae, Xanthorhoeini). – SHILAP Revista de Lepidopterologia 39 (156): 419–422.
Falck, P. & Hausmann, A. (2020): Scopula villumi Falck & Hausmann, sp. n. from Tenerife, Canary Islands, Spain (Lepidoptera: Geometridae, Sterrhinae). – SHILAP Revista de Lepidopterologia 48 (191): 507–511.
Fazekas, I. (2016): Magyar Eupitheciini tanulmanyok 4. Az Eupithecia catharinae Vojnits, 1969 tipusanyaganak revizioja. Hungarian Eupitheciini studies, No. 4. Revision of the Eupithecia catharinae Vojnits, 1969 type material (Lepidoptera: Geometridae). – Acta Naturalia Pannonica 10: 5–12.
Ferguson, D. C. (2008): Geometroidea, Geometridae (part), Ennominae (part), Abraxini, Cassymini, Macariini. – In: Hodges, R.W. et al. (eds.): The Moths of North America, Fascicle 17.2, 431 pp.; Washington (The Wedge Entomological Research Foundation).
Ferguson, D. C. (2009): A revision of the red-brown caberine geometrids of the southeastern United States (Geometridae: Caberini). – Tropical Lepidoptera Research 19 (1): 35–51.
Ferris, C. D. (2007): Three new species of Eupithecia Curtis from Arizona and New Mexico with discussion of associated species (Lepidoptera: Geometridae: Eupitheciini). – Zootaxa 1516 (1): 49–60. https://doi.org/10.11646/zootaxa.1516.1.5
Ferris, C. D. (2009a): Synaxis triangulata (Barnes & McDunnough) moved to Caripeta Walker (Geometridae: Ennominae). – Journal of the Lepidopterists' Society 63 (3): 164–165.
Ferris, C. D. (2009b): Metanema brunneilinearia Grossbeck misplaced in Synaxis Hulst (Geometridae: Ennominae). – Journal of the Lepidopterists' Society 63 (3): 166–168.
Ferris, C. D. (2010a): A new geometrid genus and species from Southeastern Arizona (Ennominae: Nacophorini). – Journal of the Lepidopterists' Society 64 (3): 147–153. https://doi.org/10.18473/lepi.v64i3.a3
Ferris, C. D. (2010b): A revision of the genus Antepione Packard with description of the new genus Pionenta Ferris (Lepidoptera, Geometridae, Ennominae). – ZooKeys 71: 49–70. https://doi.org/10.3897/zookeys.71.789
Ferris, C. D. & McFarland, N. (2010): A new species of Plataea (Geometridae: Ennominae) from Southeastern Arizona. – Journal of the Lepidopterists' Society 64 (2): 98–102. https://doi.org/10.18473/lepi.v64i2.a5
Ferris, C. D. & Mironov, V. (2007): Replacement name for Eupithecia deserticola (Lepidoptera: Geometridae: Eupitheciini). – The Canadian Entomologist 139: 131–132. https://doi.org/10.4039/n06-067
Ferris, C. D. & Schmidt, B. C. (2010): Revision of the North American genera Tetracis Guenee and synonymization of Synaxis Hulst with descriptions of three new species (Lepidoptera: Geometridae: Ennominae). – Zootaxa 2347 (1): 1–36. https://doi.org/10.11646/zootaxa.2347.1.1
Ferris, C. D. & Schmidt, C. (2011): Pterospoda nigrescens (Hulst), a synonym of Ixala klotsi Sperry (Lepidoptera, Geometridae, Ennominae). – ZooKeys 149: 31–37. https://doi.org/10.3897/zookeys.149.2343
Fischer, H. (2008): Eumera lewandowskii sp. n. eine neue Art der Gattung Eumera Staudinger, 1892 aus Jordanien (Lepidoptera: Geometridae, Ennominae). – Nachrichten des Entomologischen Vereins Apollo, Neue Folge 29 (1–2): 77–79.
Fischer, H. (2010): Epirrhoe balearia spec. nov. - a new geometrid species from the island of Ibiza with new records of geometrids from the Balearic Islands. – Atalanta (Marktleuthen) 42 (1–4): 245–247.
Fischer, T. C., Michalski, A. & Hausmann, A. (2020): Eogeometer vadens gen. n., sp. n., the first member of Macrolepidoptera in Eocene Baltic amber (Lepidoptera, Geometridae). – Mitteilungen der Münchner Entomologischen Gesellschaft 110: 125–126.
Fiumi, G. & Flamigni, C. (2013): Dyscia (Dyscia) govii n. sp. della regione sardo-corsa (Insecta Lepidoptera Geometridae Ennominae). – Quaderno di Studi e Notizie di Storia Naturale della Romagna 38: 191–199.
Fiumi, G., Flamigni, C., Zilli, A. & Hausmann, A. (2013): Le specie del genere Tephronia nella regione sardo-corsa e descrizione di Tephronia nuragica n. sp. (Insecta Lepidoptera Geometridae Ennominae). – Quaderno di Studi e Notizie di Storia Naturale della Romagna 38: 201–216.
Fu, C. M., Sato, R. & Kawakami, Y. (2011): Description of a new species of the genus Scardamia Guenée (Geometridae, Ennominae) from Japan and Taiwan, with taxonomic notes on the related species. – Tinea 21 (5): 259–267.
Garzón-Orduña, I. J. (2019): A new species of Ophthalmoblysis Scoble, 1995 (Geometridae: Ennominae) from Mexico with ‘sleepy’ eyespots. – Zootaxa 4706 (3): 469–476. https://doi.org/10.11646/zootaxa.4706.3.7
Gianti, M. (2016a): A new species of Epirrita from Lebanon (Lepidoptera, Geometridae, Larentiinae). – Doriana 8 (397): 1–5.
Gianti, M. (2016b): Monocerotesa galloi Gianti, sp. n. a new species from China (Lepidoptera: Geometridae). – SHILAP Revista de Lepidopterologia 44 (174): 299–301.
Govi, G. & Fiumi, G. (2005): Macaria ichnusae. Una nuova specie di Geometridae della Sardegna (Insecta Lepidoptera Geometridae Ennominae). – Quaderno di Studi e Notizie di Storia Naturale della Romagna 21: 129–135.
Goyal, T., Kirti, J. S. & Saxena, A. (2018): Taxonomy of genus Agathia Geunee (Lepidoptera: Geometridae) with sescription of a new species from Western Ghats, India. – Indian Journal of Entomology, 80 (3): 951–959. https://doi.org/10.5958/0974-8172.2018.00144.X
Guerrero, J. J., Cuenca, E. D., Barros, D. & Ortiz, A. S. (2020): Redescription and DNA barcoding of diurnal moth Athroolopha latimargo Rothschild, 1914 bona sp., stat. rev. from the southern Iberian Peninsula (Lepidoptera: Geometridae: Ennominae). – Zootaxa 4729 (4): 582–588. https://doi.org/10.11646/zootaxa.4729.4.9
Guerrero, J. J., Hausmann, A. & Ortiz, A. S. (2021): Description of Idaea josephinae sp. n. from the Iberian Peninsula (Lepidoptera: Geometridae). – Zootaxa 4990 (2): 369–377. https://doi.org/10.11646/zootaxa.4990.2.10
Guillermet, C. (2011): Contribution à l'étude des Hétérocères de l'île de La Réunion: description de six nouvelles espèces de Pyralidae, Geometridae, Arctiidae and Oecophoridae (Lepidoptera Heterocera). – Entomologiste 67 (2): 95–104.
Han, H., Expósito-Hermosa, A. & Xue, D. (2009a): A taxonomic study of Epipristis Meyrick, 1888 from China, with descriptions of two new species (Lepidoptera: Geometridae, Geometrinae). – Zootaxa 2263 (1): 31–41. https://doi.org/10.11646/zootaxa.2263.1.3
Han, H., Galsworthy, A. C. & Xue, D. (2005): A revision of the genus Metallolophia Warren (Lepidoptera, Geometridae, Geometrinae). – Journal of Natural History 39 (2): 165–195. https://doi.org/10.1080/00222930310001657865
Han, H., Galsworthy, A. C. & Xue, D. (2009b): A survey of the genus Geometra Linnaeus (Lepidoptera, Geometridae). – Journal of Natural History 43 (13–14): 885–922. https://doi.org/10.1080/00222930802702472
Han, H., Galsworthy, A. C. & Xue, D. (2012): The Comibaenini of China (Geometridae: Geometrinae), with a review of the tribe. – Zoological Journal of the Linnean Society 165 (4): 723–772. https://doi.org/10.1111/j.1096-3642.2012.00826.x
Han, H., Li, J. & Xue, D. (2008): Revision of the genus Xenozancla Warren, 1893 (Lepidoptera: Geometridae: Geometrinae) with an analysis of its distribution pattern. – Acta Entomologica Sinica 51 (3): 315–321.
Han, H., Skou, P. & Cheng, R. (2019): Neochloroglyphica, a new genus of Geometrinae from China (Lepidoptera, Geometridae), with description of a new species. – Zootaxa 4571 (1): 099–110. https://doi.org/10.11646/zootaxa.4571.1.6
Han, H., Stüning, D. & Xue, D. (2007): Epichrysodes gen. n., a new genus of Geometrinae from the West Tianmu mountains, China (Lepidoptera, Geometridae), with description of a new species. – Deutsche Entomologische Zeitschrift 54 (1): 127–135. https://doi.org/10.1002/mmnd.200700012
Han, H., Stüning, D. & Xue, D. (2010): Taxonomic review of the genus Pseudostegania Butler, 1881, with description of four new species and comments on its tribal placement in the Larentiinae (Lepidoptera: Geometridae). – Entomological Science 13: 234–249. https://doi.org/10.1111/j.1479-8298.2010.00379.x
Han, H. & Xue, D. (2008): A taxonomic review of Pachyodes Guenee, 1858, with descriptions of two new species (Lepidoptera: Geometridae, Geometrinae). – Zootaxa 1759 (1): 51–68. https://doi.org/10.11646/zootaxa.1759.1.3
Han, H. & Xue, D. (2009): Taxonomic review of Hemistola Warren, 1893 from China, with descriptions of seven new species (Lepidoptera: Geometridae, Geometrinae). – Entomological Science 12 (4): 382–410. https://doi.org/10.1111/j.1479-8298.2009.00341.x
Han, H. & Xue, D. (2011a): Thalassodes and related taxa of emerald moths in China (Geometridae, Geometrinae). – Zootaxa 3019 (1): 26–50. https://doi.org/10.11646/zootaxa.3019.1.2
Han, H. & Xue, D. (2011b): Fauna Sinica (Insecta Lepidoptera, Geometridae, Geometrinae). Volume 54, 864 pp.; Beijing (Science Press).
Hashimoto, S. (2019): New and newly recorded species of the genus Sauris Guenee (Geometridae: Larentiinae) from Japan, with a taxonomic review of S. hirudinata Guenee. – Tinea 25 (1): 45–65.
Hausmann, A. (2009a): Hypochrosis hannelorae sp. n. from Tanzania (Lepidoptera, Geometridae, Ennominae). – Mitteilungen der Münchner Entomologischen Gesellschaft 99: 91–93.
Hausmann, A. (2009b): New and interesting geometrid moths from Sokotra islands (Lepidoptera, Geometridae). – Mitteilungen der Münchner Entomologischen Gesellschaft 99: 95–104.
Hausmann, A. (2009c): New and interesting geometrid moths from Dhofar, southern Oman (Lepidoptera, Geometridae). – Mitteilungen der Münchner Entomologischen Gesellschaft 99: 111–128.
Hausmann, A. (2009e): New and interesting geometrid moths from the Cape Verde Islands (Lepidoptera: Geometridae). – SHILAP Revista de Lepidopterología 37 (146): 241–247.
Hausmann, A. (2011): An integrative taxonomic approach to resolving some difficult questions in the Larentiinae of the Mediterranean region. – Mitteilungen der Münchner Entomologischen Gesellschaft 101: 73–97.
Hausmann, A. (2013): Integrative revision of the African geometrid genus Dargeia Herbulot, 1977. – Mitteilungen der Münchner Entomologischen Gesellschaft 103: 99–104.
Hausmann, A. (2020a): Revision of the West Palaearctic Idaea nocturna species group. – Mitteilungen der Münchner Entomologischen Gesellschaft 110: 71–80.
Hausmann, A. (2020b): The Lepidoptera of Israel. Volume 3: Geometridae. – Proceedings of the Museum Witt 9: 1–256.
Hausmann, A., Chainey, J., Heard, T., Mc Kay, F. & Raghu, S. (2016a): Revision of the genus Eueupithecia Prout, 1910 from Argentina (Lepidoptera, Geometridae, Sterrhinae). – Zootaxa 4138 (2): 392–400. https://doi.org/10.11646/zootaxa.4138.2.11
Hausmann, A., Haszprunar, G. & Hebert, P. D. N. (2011): DNA barcoding the geometrid fauna of Bavaria (Lepidoptera): successes, surprises, and questions. – PLoS ONE 6: 17134. https://doi.org/10.1371/journal.pone.0017134
Hausmann, A. & Hebert, P. D. N. (2009): Order Lepidoptera, family Geometridae (Part 2): The Geometridae of the UAE revised in the light of mtDNA data. – In: HARTEN, T. VAN (ed.): Arthropod fauna of the UAE. Volume 2, pp. 468–479; Abu Dhabi (Dar Al Ammah).
Hausmann, A., Herbert, P. D. N., Mitchell, A., Rougerie, R., Sommerer, M., Edwards, T. & Young, C. J. (2009): Revision of the Australian Oenochroma vinaria Guenée, 1858 species-complex (Lepidoptera: Geometridae, Oenochrominae): DNA barcoding reveals cryptic diversity and assesses status of type specimen without dissection. – Zootaxa 2239 (1): 1–21. https://doi.org/10.11646/zootaxa.2239.1.1
Hausmann, A. & Huemer, P. (2011): Taxonomic decision as a compromise: Acasis appensata (Eversmann, 1832) in Central Italy—a case of conflicting evidence between DNA barcode and morphology (Lepidoptera: Geometridae). – Zootaxa 3070 (1): 60–68. https://doi.org/10.11646/zootaxa.3070.1.7
Hausmann, A., Leipnitz, M. & Bläsius, R. (2008): Idaea omari Hausmann & Bläsius, sp. n. from Morocco (Lepidoptera: Geometridae, Sterrhinae). – SHILAP Revista de Lepidopterologia 36 (143): 411–416.
Hausmann, A., Miller, M. A., Leipnitz, M. & Bläsius, R. (2007): Idaea nigra Hausmann & Bläsius, sp. n. from La Gomera, Canary Islands, Spain (Lepidoptera: Geometridae, Sterrhinae). – SHILAP Revista de Lepidopterologia 35 (140): 499–505.
Hausmann, A., Parisi, F. & Sciarretta, A. (2014): The geometrid moths of Ethiopia I: tribes Pseudoterpnini and Comibaenini (Lepidoptera: Geometridae, Geometrinae). – Zootaxa 3768 (4): 460–468. https://doi.org/10.11646/zootaxa.3768.4.4
Hausmann, A., Parisi, F. & Sciarretta, A. (2016b): The geometrid moths of Ethiopia II: Prasinocyma (Lepidoptera: Geometridae, Geometrinae). – Zootaxa 4065 (1): 001–063. https://doi.org/10.11646/zootaxa.4065.1.1
Hausmann, A. & Parra, L. (2009): An unexpected hotspot of moth biodiversity in Chilean northern Patagonia (Lepidoptera, Geometridae). – Zootaxa 1989 (1): 23–38. https://doi.org/10.11646/zootaxa.1989.1.
Hausmann, A., Pototski, A. & Viidalepp, J. (2020): Archedontia agnesae gen. n., sp. n., a new sterrhine species from Tadjikistan (Lepidoptera, Geometridae, Sterrhinae). – Zootaxa 4743 (2): 275–279. https://doi.org/10.11646/zootaxa.4743.2.10
Hausmann, A. & Sciarretta, A. (2020): The Geometridae of Ethiopia III: genus Zamarada (Lepidoptera: Geometridae, Ennominae, Cassymini). – Zootaxa 4894 (3): 301–328. https://doi.org/10.11646/zootaxa.4894.3.1
Hausmann, A., Sciarretta, A. & Parisi, F. (2016c): The Geometrinae of Ethiopia II: Tribus Hemistolini, genus Prasinocyma (Lepidoptera: Geometridae, Geometrinae). – Zootaxa 4065 (1): 1–63. https://doi.org/10.11646/zootaxa.4065.1.1
Hausmann, A. & Skou, P. (2008): Order Lepidoptera, family Geometridae. – In: HARTEN, T. VAN (ed.): Arthropod fauna of the UAE. Volume 1, pp. 562–590; Abu Dhabi (Dar Al Ammah).
Hausmann, A., Stadie, D. & Fiebig, R. (2016d): Geometridae Leach, [1815]. – In: HACKER, H. H. (ed.): Systematic and illustrated catalogue of the Macroheterocera and superfamilies Cossoidea Leach, [1815], Zygaenoidea Latreille, 1809, Thyridoidea Herrich-Schäffer, 1846 and Hyblaeoidea Hampson, 1903 of the Arabian Peninsula, with a survey of their distribution (Lepidoptera). – Esperiana 20: 61–138.
Hausmann, A., Sommerer, M., Rougerie, R. & Hebert, P. (2009): Hypobapta tachyhalotaria spec. nov from Tasmania – an example of a new species revealed by DNA barcoding. – Spixiana 32 (2): 161–166.
Hausmann, A. & Viidalepp, J. (2012): The Geometrid Moths of Europe. Volume 3. Subfamily Sterrhinae (II) (Lythriini). Subfamily Larentiinae I (Cataclysmini, Xanthorhoini, Euphyiini, Larentiini, Hydriomenini, Stamnodini, Cidariini, Operophterini, Asthenini, Phileremini, Rheumapterini, Solitaneini, Melanthiini, Chesiadini, Trichopterygini), 743 pp. – In: Hausmann, A. (ed.): The Geometrid Moths of Europe; Vester Skerninge (Apollo Books).
Hausmann, A. & Wildfeuer, J. (2017): Nine new emerald species for the fauna of Yemen, with description of two new taxa in the genus Prasinocyma (Lepidoptera, Geometridae, Geometrinae). – Spixiana 40 (2): 171–180.
Hashimoto, S. (2021): Taxonomic study of the Japanese Trichopterygini (Lepidoptera: Geometridae: Larentiinae), with a proposal of a new tribe Heterophlebini. – Japanese Journal of Systematic Entomology, Monographic Series 6: 1–146.
Heppner, J. B. (2010): The new genus Rindgeria and its species in North America and Central America (Lepidoptera: Geometridae: Ennominae). – Lepidoptera Novae 3 (3): 149–153.
Huemer, P. & Hausmann, A. (2009): A new expanded revision of the European high mountain Sciadia tenebraria species group (Lepidoptera: Geometridae). – Zootaxa 2117 (1): 1–30. https://doi.org/10.11646/zootaxa.2117.1.1
Huemer, P. & Mayr, T. (2015): Eine neue Unterart von Colostygia kitschelti (Rebel, 1934) mit Bemerkungen zum Artkonzept im C. austriacaria-Komplex (Lepidoptera: Geometridae). – Wissenschaftliches Jahrbuch der Tiroler Landesmuseen 8: 66–79.
Infusino, M., Scalercio, S. & Hausmann, A. (2016): Nothocasis rosariae sp. n., a new sylvicolous, montane species from southern Europe (Lepidoptera: Geometridae, Larentiinae). – Zootaxa 4161 (2): 177–192. https://doi.org/10.11646/zootaxa.4161.2.2
Inoue, H. (2007a): A new subspecies of Dysphania discalis (Walker) (Geometridae, Geometrinae) from the Lingga Islands. – Lepidoptera Science 58 (1): 4–6.
Inoue, H. (2007b): A new subspecies of Dysphania transducta (Walker) from Tawitawi Island, Sula Archipelago, Philippines (Geometridae, Geometrinae). – Tinea 20 (1): 9–11.
Jiang, N., Li, X., Hausmann, A., Cheng, R., Xue, D. & Han, H. (2017): A molecular phylogeny of the Palaearctic and Oriental members of the tribe Boarmiini (Lepidoptera: Geometridae: Ennominae). – Invertebrate Systematics 31 (4): 427–441. https://doi.org/10.1071/IS17005
Jiang, N., Sato, R. & Han, H. (2012a): One new and one newly recorded species of the genus Amraica Moore, 1888 (Lepidoptera: Geometridae: Ennominae) from China, with diagnoses of the Chinese species. – Entomological Science 15 (2): 219–231. https://doi.org/10.1111/j.1479-8298.2011.00507.x
Jiang, N., Stüning, D., Xue, D. & Han, H. (2016): Revision of the genus Metaterpna Yazaki, 1992 (Lepidoptera, Geometridae, Geometrinae), with description of a new species from China. – Zootaxa 4200 (4): 501–514. https://doi.org/10.11646/zootaxa.4200.4.3
Jiang, N., Xue, D. & Han, H. (2010): A review of Jankowskia Oberthür, 1884, with descriptions of four new species (Lepidoptera: Geometridae, Ennominae). – Zootaxa 2559 (1): 1–16. https://doi.org/10.11646/zootaxa.2559.1.1
Jiang, N., Xue, D. & Han, H. (2011a): A review of Biston Leach, 1815 (Lepidoptera, Geometridae, Ennominae) from China, with description of one new species. – ZooKeys 139: 45–96. https://doi.org/10.3897/zookeys.139.1308
Jiang, N., Xue, D. & Han, H. (2011b): A review of Ophthalmitis Fletcher, 1979 in China, with descriptions of four new species (Lepidoptera: Geometridae, Ennominae). – Zootaxa 2735 (1): 1–22. https://doi.org/10.11646/zootaxa.2735.1.1
Jiang, N., Xue, D. & Han, H. (2012b): A new species of Arbomia Sato & Wang (Lepidoptera, Geometridae, Ennominae) from Guangxi, Southern China. – Zootaxa 3765 (1): 098–100. https://doi.org/10.11646/zootaxa.3765.1.8
Jiang, N., Xue, D. & Han, H. (2012c): A review of Peratophyga Warren, 1894 in China, with descriptions of two new species (Lepidoptera: Geometridae, Ennominae). – Zootaxa 3478 (1): 403–415. https://doi.org/10.11646/zootaxa.3478.1.36
Jiang, N., Xue, D. & Han, H. (2014a): A new species of Arbomia Sato & Wang (Lepidoptera, Geometridae, Ennominae) from Guangxi, Southern China. – Zootaxa 3765 (1): 98–100. https://doi.org/10.11646/zootaxa.3765.1.8
Jiang, N., Xue, D. & Han, H. (2014b): A review of Luxiaria Walker and its allied genus Calletaera Warren (Lepidoptera, Geometridae, Ennominae) from China. – Zootaxa 3856 (1): 073–099. https://doi.org/10.11646/zootaxa.3856.1.3
KANEKO, T. (2016): A new species of the Sterrhinae (Geometridae) from Akiyoshidai-Plateau, Yamaguchi Prefecture, Japan. – Japan Heterocerists' Journal 278: 68–69.
Kaneko, T. (2018): A new species of the Sterrhinae (Geometridae) from Tsushima Is., Nagasaki Prefecture, Japan. – Japan Heterocerists' Journal 287: 285–286.
Kaneko, T. (2019): A new species of the Sterrhinae (Geometridae) collected by the late Mr Y. Yanagita from Iriomote-jima Is., the Ryukyus. – Tinea 25 (Supplement 1): 69–70.
Kaneko, T. & Fu, C.-M. (2020): Three new species of the subfamily Sterrhinae (Lepidoptera, Geometridae) from Taiwan. – Tinea 25 (3): 183–187.
Kaneko, T. & Funakoshi, S. (2018): A new species of the Geometrinae (Geometridae) from Gifu Prefecture, Japan. – Japan Heterocerists' Journal 287: 299–300.
Karisch, T. (2007): Eine neue Hispophora-Art aus Kenia (Lepidoptera: Geometridae: Ennominae). – Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Entomologie 77: 129–131.
Karisch, T. (2008a): Drei neue Ochroplutodes-Arten aus der Sammlung des Musee Royal de l'Afrique Centrale, Tervuren (MRAC) (Lepidoptera: Geometridae, Ennominae). – Entomofauna 29: 213–219.
Karisch, T. (2008b): Zur Kenntnis von Eupithecia jeanneli Herbulot, 1953 (Lepidoptera: Geometridae, Larentiinae). – Entomofauna 29 (14): 201–208.
Karisch, T. (2010): Geometridae der Expeditionen H. Hoppes nach Bioko. 1. Teil: Desmobathrinae, Geometrinae (Lepidoptera, Geometridae). – Lambillionea 110 (3): 344–352.
Karisch, T. (2011): Geometridae der Expeditionen H. HOPPES nach Bioko 2. Teil: Larentiinae (Lepidoptera, Geometridae). – Lambillionea 111 (1): 71–80.
Karisch, T. (2016): Eine neue afrikanische Metallospora-Art (Lepidoptera, Geometridae, Ennominae). – Lambillionea CXVI (1): 20–31.
Karisch, T. (2017): Zur Kenntnis der Gattung Hypomecis Huebner, 1821 in Ostafrika (Lepidoptera: Geometridae: Ennominae). – Entomofauna 38 (3): 29–48.
Karisch, T. (2020a): Eine neue Chlorodrepana von Uganda (Lepidoptera, Ennominae). – Lambillionea 120 (2): 117–121.
Karisch, T. (2020b): Geometridae der Expeditionen H. Hoppes nach Bioko 4. Teil: Ennominae (ausser Zamarada Moore, 1887) (Lepidoptera). – Lambillionea 120 (2): 122–141.
Karisch, T. & Beck, K.-R. (2014): Beitrag zur Kenntnis der Gattung Mesothisa Warren, 1905 (Lepidoptera, Geometridae, Ennominae). – Lambillionea 114 (3): 187–195.
Karisch, T. & Kramp, K. (2019): A new Scopula from mid-Atlantic Ascension Island (Lepidoptera, Geometridae). – Beiträge zur Entomologie 69 (1): 87–90. https://doi.org/10.21248/contrib.entomol.69.1.087-090
Kemal, M. & Koçak, A. O. (2015): Notes on two species of Geometridae from Hatay Province (S. Turkey) (Lepidoptera). – Miscellaneous Papers 119: 1–6.
Kemal, M. & Uçak, H. (2018): Description of a new species of the genus Wehrliola Strand in East Turkey (Lepidoptera, Geometridae). – Miscellaneous Papers 168: 1–5.
Kemal, M., Yildiz, I., Kizildag, S., Uçak, H. & Koçak, A. Ö. (2018): Taxonomical and molecular evaluation of Apochima Agassiz in East Turkey, with a description of a new genus (Lepidoptera, Geometridae, Ennominae). – Miscellaneous Papers 169: 1–14.
Kim, N. H., Choi, S. W. & Kim, S. S. (2018): Additional report of the genus Asthena (Lepidoptera: Geometridae) from Korea. – Animal Systematics, Evolution and Diversity 34 (2): 92–95.
Kimura, S. & Sato, R. (2017): A new species of the genus Odontopera Stephens (Geometridae, Ennominae) from Hahajima Is. in the Ogasawara (Bonin) Islands, Japan. – Tinea 24 (1): 24–26.
Kirti, J. S. & Goyal, T. (2011): A new species of Plutodes Guenee (Lepidoptera: Geometridae: Ennominae) from Western Ghats of India. – Journal of Applied Bioscience 37 (1): 68–70.
Knyazev, S. A. & Mironov, V. G. (2015): New species of the pugs (Lepidoptera, Geometridae: Eupithecia) for southern part of West Siberia, Russia. – Evraziatskii Entomologicheskii Zhurnal 14 (2): 139–141.
Kobayashi, H. & Wang, M. (2020): A new species of the genus Acrodontis (Geometridae, Lepidoptera). – Tinea 25 (2): 119–122.
Koçak, A. & Kemal, M. (2008): Some nomenclatural notes on the Geometridae of the world (Lepidoptera). – Miscellaneous Papers 138: 8–9.
Kostjuk, A. Ö. (2009): Agriopis budashkini sp. n. – a new species of geometrid moths (Lepidoptera, Geometridae, Ennominae) from Crimea. – Proceedings of the Zoological Society of London 5: 61–64.
Kostjuk, I., Mironov, V. & Viidalepp, J. (2020): Review of the Central Asian species of Phthorarcha Meyrick (Geometridae: Alsophilinae) with description of a new species. – Zootaxa 4861 (2): 232–240. https://doi.org/10.11646/zootaxa.4861.2.4
Kostjuk, I. & Viidalepp, J. (2020): Revision of the genus Dzhugesia Wehrli, 1936 (stat. n.), with description of a new species (Lepidoptera: Geometridae: Ennominae). – Mitteilungen der Münchner Entomologischen Gesellschaft 110: 89–95.
Krüger, M. (2007): Composition and origin of the geometrid fauna (Lepidoptera) of the Sneeuberge, Eastern Cape, with description of new taxa. – Annals of the Transvaal Museum 44 (1): 25–66.
Krüger, M. (2008): Conchylia cymopolia sp. nov. from the Richtersveld, northern Cape Province, South Africa (Lepidoptera: Geometridae: Larentiinae). – Metamorphosis 19 (1): 27–32.
Krüger, M. (2009): New Afrotropical Nacophorini in the genera Pacycnemoides Krueger, Argyrophora Guenee, Microligia Warren, Pseudomaenas Prout and Drepanogynis Guenee (Lepidoptera, Geometridae: Ennominae). – Annals of the Transvaal Museum 46: 1–35.
Krüger, M. (2013): Revision of Acrasia Rogenhofer, 1875 and description of Panhyperochia gen. nov., a new genus of putative naco-phorine geometrids from South Africa (Lepidoptera: Geometridae: Ennominae). – Annals of the Ditsong National Museum of Natural History 3: 99–152.
Krüger, M. (2014): A revision of the Mauna Walker, 1865 and Illa Warren, 1914 group of genera (Lepidoptera: Geometridae: Ennominae: Nacophorini). – Annals of the Ditsong National Museum of Natural History 4: 77–173.
Krüger, M. (2015): Taxonomic remarks on southern African Geometridae putatively assigned to the tribe Nacophorini (Lepidoptera: Geometridae: Ennominae). – Annals of the Ditsong National Museum of Natural History 5: 62–63.
Krüger, M. (2017): Henicovalva gen. n., a new genus of predominantly Afrotropical Ennominae with unusually complex genitalia (Lepidoptera: Geometridae). – African Entomology 25 (1): 42–61. https://doi.org/10.4001/003.025.0042
László, G. M. (2003): New species of the genus Erannis Hübner, [1825] 1816 from the North-West Himalaya and Iran (Lepidoptera, Geometridae). – Acta Zoologica Academiae Scientiarum Hungaricae 49 (2): 153–158.
László, G. M. (2018): Descriptions of new Geometridae species from the Vartian collection with remarks on some poorly known taxa. – In: László, G. M., Lödl, M., Gaal-Haszler, S., Galsworthy, A., Ronkay, G., Ronkay, L. & Varga, Z. (eds.): Fibigeriana. Volume 4. The Vartian Collection. Part IV. Geometridae, pp. 55–71; Budapest (Heterocera Press).
László, G. M. & Stüning, D. (2015): Description of a new Paradarisa Warren, 1894 (Lepidoptera, Geometridae, Ennominae) from Taiwan. – Tinea 23 (2): 118–122.
Lehmann, L. (2011): Lithostege stadiei sp. n. from North Iran (Geometridae: Larentiinae). – Nota Lepidopterologica 33 (2): 249–251.
Leraut, P. (2008a): Contribution to the study of the genus Sciadia Hübner, 1822 (Lepidoptera, Geometridae). – Bulletin de la Société Entomologique de France 113 (2): 177–182. https://doi.org/10.3406/bsef.2008.16517
Leraut, P. (2008b): Une nouvelle espèce du genre Triphosa Stephens, 1829 (Lep. Geometridae). – Bulletin de la Société Entomologique de France 113 (2): 452–454. https://doi.org/10.3406/bsef.2008.3040
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Lévêque, A. (2006): Description d'un nouveau Pero du Mexique et compléments à la connaissance de P. fortunata herissa Lévêque, 2005 (Lepidoptera, Geometridae). – Bulletin de la Société Entomologique de France 111: 345–352. https://doi.org/10.3406/bsef.2006.16336
Lévêque, A. (2007): Description d'une nouvelle espece de Pero du Perou (Lepidoptera, Geometridae, Ennominae). – Bulletin de la Société Entomologique de France 112 (3): 389–394. https://doi.org/10.3406/bsef.2007.16454
Lévêque, A. (2010): Description d'une nouvelle espèce péruvienne du genre Pero Herrich-Schäffer, 1855 (Lepidoptera, Geometridae, Ennominae). – Bulletin de la Société Entomologique de France 115 (1): 51–57. https://doi.org/10.3406/bsef.2010.2831
Lévêque, A. (2015): Second contribution to the elaboration of a taxonomic checklist of geometrid moths from French Guiana: the species of the genus Oospila Warren, 1897. – Antenor 2 (2): 232–246.
Lévêque, A. & Viidalepp, J. (2015): Description of a new Oospila Warren, 1897, from French Guiana (Lepidoptera Geometridae Geometrinae Lophochoristini). – Antenor 2 (1): 142–147.
Lewandowski, S. & Tober, K. (2010): Idaea sordida horus subspec. nov., eine neue Unterart der Gattung Idaea TREITSCHKE, 1825 aus Ägypten. – Atalanta (Marktleuthen) 42 (1–4): 249–250.
Lewis, D. & Covell, C. (2008): Review of the Neotropical genus Cyllopoda (Lepidoptera: Geometridae: Sterrhinae: Cyllopodini). – Tropical Lepidoptera Research 18 (2): 88–101.
Li, H., Jiang, N., Rui, C., Zhang, C. & Han, H. (2018): A review of the subgenus Epicterodes of Arichanna (Lepidoptera, Geometridae, Ennominae), with description of one new species. – Journal of Asia-Pacific Entomology 21 (2): 501–512. https://doi.org/10.1016/j.aspen.2018.02.013
Li, J., Jiang, N. & Han, H. (2012a): Heterophleps inusitata, an extremely rare new moth species from western Yunnan, China (Lepidoptera, Geometridae, Larentiinae). – Zootaxa 3278 (1): 58–60. https://doi.org/10.11646/zootaxa.3278.1.2
Li, J., Xue, D., Han, H. & Galsworthy, A. (2012b): Taxonomic review of Syzeuxis Hampson, 1895, with a discussion of biogeogra-phical aspects (Lepidoptera, Geometridae, Larentiinae). – Zootaxa 3357 (1): 1–24. https://doi.org/10.11646/zootaxa.3357.1.1
Li, X., Xue, D. & Jiang, N. (2017): One new species and one new record for the genus Ninodes Warren from China (Lepidoptera, Geometridae, Ennominae). – ZooKeys 679: 55–63. https://doi.org/10.3897/zookeys.679.12547
Lindt, A., Hausmann, A. & Viidalepp, J. (2018): Review of some species groups of the genus Oospila Warren, with descriptions of nine new species (Lepidoptera: Geometridae: Geometrinae). – Zootaxa 4497 (2): 151–194. https://doi.org/10.11646/zootaxa.4497.2.1
Lindt, A., Lennuk, L. & Viidalepp, J. (2017a): The genus Dioscore Warren, 1907: two new species and analysis of characters spread (Lepidoptera: Geometridae: Geometrinae). – Journal of Insect Biodiversity 5 (16): 1–15. https://doi.org/10.12976/jib/2017.5.16
Lindt, A., Tasane, T., Õunap, E. & Viidalepp, J. (2017b): Five new species of the genus Paromphacodes (Lepidoptera: Geometridae: Geometrinae) from High Andes in Ecuador. – Zootaxa 4303 (3): 395–406. https://doi.org/10.11646/zootaxa.4303.3.5
Lindt, A., Tasane, T. & Viidalepp, J. (2014): Two new emerald moth species from Ecuador, Peru and Bolivia (Lepidoptera: Geometridae, Geometrinae). – Shilap Revista de Lepidopterologia 42 (165): 151–156.
Lindt, A. & Viidalepp, J. (2014): Two new emerald geometrid species of Telotheta Warren from Ecuador and Bolivia (Lepidoptera: Geometridae, Geometrinae, Lophochoristini). – Biodiversity Data Journal 2: e1158. https://doi.org/10.3897/BDJ.2.e1158
Lindt, A. & Viidalepp, J. (2015): Oospila bulava, a new emerald geometrid moth from South America (Lepidoptera, Geometridae, Geometrinae). – Zootaxa 4058 (1): 142–144. https://doi.org/10.11646/zootaxa.4058.1.11
Liu, Z., Due, D., Wang, W. & Han, H. (2013): A review of Psyra Walker, 1860 (Lepidoptera, Geometridae, Ennominae) from China, with description of one new species. – Zootaxa 3682 (3): 459–474. https://doi.org/10.11646/zootaxa.3682.3.7
Makhov, I. (2021): Geometridae (Lepidoptera) of the Baikal region: keys to species and an annotated catalogue. Part 1. Ennominae. – Zootaxa 4962 (1): 1–125. https://doi.org/10.11646/zootaxa.4962.1.1
Makhov, I.A. & Lukhtanova, V.A. (2021): Geometrid moths (Lepidoptera, Geometridae) of the Baikal Region: additions to the species list and results of DNA barcoding. – Entomological Review 101 (8): 1154–1172. https://doi.org/10.1134/S001387382108011X
Mallick, K., Dey R., Bandyopadhyay, U., Mazumder, A., Gayen, S., Ali, M., Nandi Das, G., Raha, A., Kumar Sanyal, A., Kumar Gupta, S., Prasad Uniyal, S., Chandra, K. & Kumar, V. (2022): Taxonomy and ecology of genus Psyra Walker, 1860 (Lepidoptera: Geometridae: Ennominae) from Indian Himalaya. – PLoS ONE 17 (4): e0266100. https://doi.org/10.1371/journal.pone.0266100
Matson, T. A. & Wagner, L. D. (2020): A new Stamnodes from the southwestern United States (Lepidoptera, Geometridae, Larentiinae). – ZooKeys 923: 79–90. https://doi.org/10.3897/zookeys.923.48290
Matson, T. A. (2022): A new monotypic genus from the American Southwest to accommodate “Semiothisa” kuschea (Geometridae: Ennominae). – Zootaxa 5093 (1): 067–074. https://doi.org/10.11646/zootaxa.5093.1.4
Mazel, R. & Varenne, T. (2017): Idaea alyssumata selenensis ssp. n. dans le Midi de la France (Lepidoptera, Geometridae, Sterrhinae). – Revue de l'Association Roussillonnaise d'Entomologie XXVI: 82–88.
Mironov, V. (2013): New species and checklist of Turkish Eupithecia Curtis (Geometridae: Larentiinae). – Zootaxa 3717 (1): 39–52. https://doi.org/10.11646/zootaxa.3717.1.3
Mironov, V. & Galsworthy, A. (2007): The genus Eupithecia (Lepidoptera, Geometridae) in Taiwan: an updated survey. – Transactions of the Lepidopterological Society of Japan 58 (3): 341–363.
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