Revision of the Orbamia Herbulot, 1966 group of genera with description of two new genera, ten new species, and two new subspecies (Lepidoptera, Geometridae, Ennominae, Cassymini)

Abstract The genus Orbamia Herbulot, 1966 is revised. Two new genera are described: Rabomia Hausmann & Tujuba, gen. nov. (type species: Ectropis ? subaurata Warren, 1899), and Morabia Hausmann & Tujuba, gen. nov. (type species: Morabia politzari Hausmann & Tujuba, sp. nov.). Ten new species and two new subspecies are described: Rabomia obscurior Hausmann & Tujuba, sp. nov., from western Africa, Morabia politzari Hausmann & Tujuba, sp. nov., from Kenya, Morabia brunnea Hausmann & Tujuba, sp. nov., from Zambia, Orbamia marginata Hausmann & Tujuba, sp. nov., from Tanzania, Orbamia clarissima Hausmann & Tujuba, sp. nov., from Kenya, Orbamia clarior Hausmann & Tujuba, sp. nov., from Kenya, Orbamia obliqua Hausmann & Tujuba, sp. nov., from Zambia, Orbamia obliqua parva Hausmann & Tujuba, subsp. nov., from South Africa, Orbamia abiyi Hausmann & Tujuba, sp. nov., from Zambia, Tanzania, Ethiopia, Orbamia emanai Hausmann & Tujuba, sp. nov., from Ethiopia, Orbamia emanai lenzi Hausmann & Tujuba, subsp. nov., from Zambia and Malawi, and Orbamia balensis Hausmann & Tujuba, sp. nov. from Ethiopia. The taxon Lepiodes ocellata Warren, 1897 is raised from synonymy of O. octomaculata (Wallengren, 1872) to species rank (Zambia, Tanzania, Rwanda). The taxonomical analysis is based on both morphological and genetic cytochrome oxidase I (COI) data. Adults and male and female genitalia of all species are illustrated.


Introduction
After 260 years of intensive work, taxonomists worldwide have together achieved the formal descriptions of approximately 160,000 lepidopteran species (Van Nieukerken et al. 2011), of which some 24,000 are geometrid moths (cf. Scoble 1999;Scoble and Hausmann 2007;Van Nieukerken et al. 2011). Many more are likely to be described and some recent revisions of many genera revealed high percentages of undescribed species (e.g., Brehm 2015Brehm , 2018Hausmann 2003;Hausmann et al. 2016). This probably covers less than half the number of actual extant geometrid species on earth, an estimate which can be inferred from apparently undescribed species in natural history museums and in molecular databases like the Barcode of Life Data Systems (BOLD: Ratnasingham and Hebert 2007). We conclude that conventional taxonomy with an actual description rate of 80-100 species per year works too slowly for addressing the biodiversity of our Earth (taxonomic impediment: de Carvalho et al. 2007;Wheeler 2008). In these times where we are facing a serious extinction rate we cannot afford to wait for another 250-600 years for the taxonomic assessment of our biodiversity. In the recent literature, several pleas for an accelerated taxonomy have been made (Riedel et al. 2013a;Forum Herbulot 2014) and some taxonomists already published exemplary revisions with shortened descriptions (Riedel et al. 2013b;Meierotto et al. 2019).
In this taxonomic revision we follow such a model of accelerated taxonomy (cf. Riedel et al. 2013aRiedel et al. , 2013bForum Herbulot 2014;Meierotto et al. 2019), which should lead in future to an automated, easy and rapid transfer of genetic data, images and metadata directly from BOLD into manuscripts and which will allow for continuous updates. Similarly, all nomenclatorial information (valid names, synonyms, original descriptions with authorship and year, type localities, type specimens and their deposition) may in future be transferred in an automated way either from BOLD, from the Global Lepidoptera checklist (Banki et al. 2019) and/or from the Geometridae Mundi database once it will be completed by the Forum Herbulot initiative (cf. Löbel and Hausmann 2019). Similar to the approach of Meierotto et al. (2019) we believe that DNA barcodes in most cases are an excellent tool for species diagnosis but that all descriptions should be linked with a brief description of characters in words, supplemented with photographs.
In future, such revisions may also be organised in a flexible way, i.e., with the possibility to subsequently publish updated versions (keeping the previous versions visible) with more data, newly added species, and revised taxonomic concepts. In the framework of the ongoing research project "GBOL III -Dark Taxa" (SNSB -Bavarian State Collection of Zoology, Munich) similar workflows are planned to be tested and established.
The genus Orbamia Herbulot, 1966 is restricted to the Ethiopian region, where it was so far represented by five species (Scoble 1999;Hausmann 2006): Orbamia octomaculata (Wallengren 1872), Orbamia pauperata Herbulot, 1966, Orbamia renimacula (Prout, 1926), Orbamia subaurata (Warren, 1899) and Orbamia becki Hausmann, 2006. Although the external appearance of Orbamia is somewhat reminiscent of that of the tribe Boarmiini, Hausmann (2006) placed Orbamia in the tribe Cassymini, due to the long process extending from the base of the dorsal margin of the male genital valvae, similar to the equivalent present in Zamarada. Molecular evidence for this tribal assignment was given by Murillo-Ramos et al. (2019) and Brehm et al. (2019). Species of Orbamia are recognised by the following characters: wings with conspicuous, contrasting discal spots on all wings, antennae bipectinate in males and filiform in females, dorsal process in male genitalia strongly curved.

Materials and methods
In the present paper, the material housed in the Zoologische Staatssammlung München (ZSM), Munich, Germany, collected from 22 different African and Arabian countries, has been studied. Two relevant type specimens available in the ZSM and two from the Natural History Museum in London (NHMUK) were examined. Altogether, 298 specimens of the genera Orbamia, Rabomia gen. nov., and Morabia gen. nov. have been examined.
Comparative morphological methods and COI sequence divergences were used to delimit the taxa and to estimate their taxonomic status. We studied morphological characters of adults (including genitalia and wing venation). The abdomens and genitalia were prepared using the method of Hardwick (1950). The analysis is furthermore based on 58 genitalia slides and 72 DNA barcodes.
For DNA analyses, one or two legs were removed from dried specimens and stored in an individual tube, in absolute ethanol. DNA extraction, amplification and sequencing of the "barcode" region of the mitochondrial cytochrome c oxidase I (COI) gene region (658 base pairs) were carried out in the Canadian Centre for DNA Barcoding, Ontario, Canada, using standard high through-put protocols (Ivanova et al. 2006). Sequence divergences within and between species were calculated using the Kimura 2-parameter model (Kimura 1980), using the analytical tools provided by BOLD Systems v4 platform (Ratnasingham and Hebert 2007; http://www.boldsystems.org/). Intra-specific and inter-specific genetic distances are reported as maximum and minimum distances, respectively. The Barcode Index Number (BIN) of each species is reported which was obtained from the BOLD Systems v4 database. BINs represent a species-level taxonomic registry of the animal kingdom based on the analysis of nucleotide variation patterns in the barcode region of the cytochrome c oxidase I (COI) gene (Ratnasingham and Hebert 2013). This genetic information facilitates the species delimitation and constitutes the basis of future phylogenetic works at the genus level and below (Brehm 2015(Brehm , 2018. Label data and photographs of types and other barcoded specimens are accessible on BOLD, dataset DS-ORBAMIA (https://dx.doi.org/10.5883/DS-ORBAMIA). All new names are registered in ZooBank. Geo-references were taken from specimen labels.
Genetic data and phylogeny. Multigene analyses of Geometridae revealed evidence for assignation of the genus Orbamia to Cassymini and for sister group relationship with the African genus Pycnostega Brehm et al. 2019). The maximum likelihood analysis of COI barcode data underpins the monophyly of the genus Orbamia as conceived and circumscribed here (cf. Etymology. The name refers to the uninterrupted black line at the hindwing margin (Lat. margo/marginis = edge, border).
Differential features (COI sequences, photographs of adults and their genitalia see https://dx.doi.org/10.5883/DS-ORBAMIA): Adult: Forewing length: 11.5-12.5 mm. Upper side of wings: Ground colour dirty grey with brown suffusion. Underside: Ground colour whitish beige with some dark suffusion, terminal fascia conspicuous, uninterrupted on all wings. Male genitalia: Uncus long, triangular, valva slightly bent, dorsal process with small spinule at tip, cornutus short and stout (1.6 mm). Female genitalia unknown. Etymology. The name refers to the oblique position of the transverse lines of the forewing (Lat. obliquus, -a, -um = oblique).
Differential features (COI sequences, photographs of adults and their genitalia see https://dx.doi.org/10.5883/DS-ORBAMIA): Adult: Forewing length: 11-12 mm. Upperside of wings: Ground colour pale grey with brown pattern, transverse lines oblique. Underside: Ground colour beige with slight yellowish tinge, and with strong dark suffusion, terminal area with pattern reduced to a dark apical spot and a dark shadow on the hindwing apex. Male genitalia: Uncus short, stout, hook-shaped, saccus projection shallow, valva straight, narrow at tip, dorsal process with a stout hook at tip, Note. Synonym of O. octomaculata according to Scoble (1999) but the type series of ocellata belongs to two different species. Herewith we designate the red-ring-labelled male specimen from Zambia, Mpeta, as lectotype to fix the identity of the name and to stabilise nomenclature. The taxon is herewith upgraded from synonymy to species rank (stat. nov.), based on the below mentioned differences in DNA barcodes and genitalia. Pattern of upper-and underside of wings of the lectotype exactly matches that of the examined material from Tanzania, partly collected in closely adjacent localities to the type locality (e.g., Ruvuma and Iringa provinces Differential features (COI sequences, photographs of adults and their genitalia see https://dx.doi.org/10.5883/DS-ORBAMIA): Adult: Forewing length: 8.5-12 mm. Upperside of wings: Ground colour pale grey with slight brown suffusion, pattern brown with slight orange tinge. Underside: Ground colour whitish, orange between veins, apical spots on forewing conspicuous, sharply bordered, on hindwing terminal fascia usually diffuse, rarely restricted to apex. Male genitalia: Uncus narrow, digitiform, valva straight, broad, dorsal process with conspicuous, stout hook at tip, cornutus narrow and very long (2.8-3.0 mm). Female genitalia: Apophyses stout, apophyses anteriores comparatively long (2/3 length of apophyses posteriores), lamella ante-and post-vaginalis fused, oval, comparatively broad (length 0.7 mm, width 0.5 mm), signum weakly sclerotised, small, transverse ridge straight (0.2 mm). Etymology. Named after Emana Getu, a senior professor of Entomology, at Addis Ababa University for his immense contributions to the field of entomology.
Remarks. Morphological differences to the previous species small, but the genetic divergence correlates with darker wing colour, and a few characters in male genitalia (longer cornutus). Distribution areas of both species overlapping.  Upperside of wings: Ground colour much paler than in the nominotypical subspecies, very pale grey, with slight brown suffusion, mainly in the terminal area, pattern grey brown. Underside: Ground colour whitish beige, orange on veins, apical spots on forewing conspicuous, sharply bordered, on hindwing terminal fascia usually diffuse over more or less the whole termen. Male genitalia: Uncus narrow, digitiform, valva straight, broad, dorsal process with conspicuous, stout hook at tip, cornutus narrow and long (2.7 mm). Female genitalia: Lamella ante-and post-vaginalis fused, oval (length 0.75-0.85 mm), signum weakly sclerotised, small, transverse ridge straight (0.17-0.2 mm).
Remarks. Phylogenetically the most isolated species within this genus, based on large differences in morphology and genetics, the latter, however, just based on a short barcode fragment of the holotype. Etymology. The name refers to the type locality in the Bale mountains. Differential features (COI sequences, photographs of adults and their genitalia see https://dx.doi.org/10.5883/DS-ORBAMIA): Adult: Forewing length: 11-13 mm. Upperside of wings: Ground colour comparatively dark, with much dark brown suffusion, pattern not well contrasted. Underside: Ground colour beige, with pale orange tinge on veins, apical spots on forewing conspicuous, sharply bordered, on hindwing slightly darker at apex only, remnants of dotted terminal fascia on all wings. Male genitalia: Uncus stout, very long, saccus broad, valva straight, ´very broad at base, narrow at tip, dorsal process stoutly sclerotised, with conspicuous, stout hook at tip, cornutus narrow and short (1.1 mm). Female genitalia: Lamella ante-and post-vaginalis fused, heart-shaped, short (length 0.6 mm), signum small, transverse ridge straight (0.2 mm).
Genetic data and phylogeny. The maximum likelihood analysis of COI barcode data supports the monophyly of the genus Rabomia gen. nov. and sister group relationship with (Pycnostega+Dorsifulcrum) (cf. genus level need to be considered with caution, when they are inferred from COI data. More research is needed to investigate the potential (re-) assignment of Dorsifulcrum to the Cassymini after having been excluded from that tribe in Brehm et al. (2019).

Rabomia subaurata
Etymology. The name refers to the darker colouration of wings (Lat. obscurior = darker). Differential features (COI sequences, photographs of adults and their genitalia see https://dx.doi.org/10.5883/DS-ORBAMIA): Adult: Forewing length: 12-14 mm. Upperside of wings: Ground colour with darker suffusion than in R. subaurata, medial fascia dark and conspicuous on all wings, sometimes anastomosing with postmedial line on forewing. Underside: Ground colour yellow, discal spots larger than in R. subaurata, terminal fascia much broader, at centre of forewing termen interrupted by a small yellow spot or even uninterrupted. Male genitalia: Valva long and narrow, sacculus edged at tip, only shortly projecting, at the base of aedeagus a bundle of twelve comparatively short (0.15-0.2 mm) microcornuti. Female genitalia: Star-shaped signum large, diameter 0.8-1.0 mm.  Differential features (COI sequences, photographs of adults and their genitalia see https://dx.doi.org/10.5883/DS-ORBAMIA). Adult: Male antennae ciliate-fasciculate, female antennae filiform. Palpi of both sexes broad, bushy scaled, length 1.0-1.5 times diameter of eye. Hind tibia of both sexes with two pairs of unequal spurs. Upperside of wings with discal spots vestigial, medial line zigzagging, terminal line conspicuous, zigzagging on all wings. Underside beige, with a few yellowish scales and a sharp black spot in forewing apex. Male genitalia: Uncus very short, rounded saccus shallowly projecting, dorsal process of valva strongly setose, valva long and narrow, curved, strongly setose, mainly at centre, aedeagus with long and stout cornutus. Female genitalia: Apophyses long and fine. Lamellae ante-and post-vaginalis membranous, ductus bursae straight, anteriorly membranous, posteriorly dilated and towards antrum strongly sclerotised, corpus bursae membranous, pyriform, signum absent.

Morabia
Genetic data and phylogeny. The maximum likelihood analysis of COI barcode data suggests the monophyly of the genus Morabia gen. nov. and an isolated position from (Rabomia gen. nov. (Pycnostega+Dorsifulcrum)) and from genus Orbamia (cf. Table 1, Fig. 85). However, phylogenies at genus level need to be considered with caution, when they are inferred from COI data. Etymology. The name refers to Dr. Heinz Politzar (1938 for his great merits in collecting and studying African Lepidoptera (see Hacker and Hausmann 2010).