Contribution to the knowledge of the oribatid mite genus Leobodes (Acari, Oribatida, Nippobodidae)

Three new species of oribatid mites of the genus Leobodes (Oribatida, Nippobodidae) - L. becki n. sp., L. schalleri n. sp. and L. schawalleri n. sp. - are described from Nepal. Revised generic diagnosis and an identification key to 10 known species of Leobodes are provided.


Introduction
The oribatid mite genus Leobodes (Acari, Oribatida, Nippobodidae) was described by Aoki (1965a), with Leobodes mirabilis Aoki, 1965 as type species. At present, Leobodes comprises seven species, which are all distributed in the Oriental region (Subías 2004(Subías , online version 2021. During taxonomic identification of oribatid mites which were collected from the Jochen Martens Expeditions to Nepal in 1988 (Results of the Himalaya Expeditions of J. Martens from 1969-2004 and housed in the Institut für Organismische und Molekulare Evolutionsbiologie (Mainz, Germany), we found three new species, belonging to Leobodes. The main goal of our paper is to describe and illustrate these new species. In addition, the revised generic diagnosis and an identification key to known species of the genus are presented.

Material and methods
Specimens -Samples containing oribatid mites were collected by the following method: leaf litter and upper layer of soil were collected by hand, sieved by means of a "Käfersieb", larger arthropods were sorted out by hand on a large white plastic sheet and the remainder of sieved substrate filled into a number of small, easily transportable Berlese funnels, each containing a vial filled about half with 75% alcohol; all samples were extracted in open field conditions during three days/nights; no special warming was applied except for sunshine radiation. To protect the funnels over night against rain, they were placed in a tent. Final sorting of the samples as to different arthropod groups was done in the Mainz lab of the Martens group. See the Material examined section for detailed location data for each new species. Specimens (preserved in 70% solution of ethanol with a drop of glycerol) are deposited in two institutions: the Senckenberg Museum of Natural History, Görlitz, Germany (SMNH); and the Tyumen State University Museum of Zoology, Tyumen, Russia (TSUMZ).
Observation and documentation -Specimens were mounted in lactic acid on temporary cavity slides for measurement and illustration. Body length was measured in lateral view, from the tip of the rostrum to the posterior edge of the notogaster. Notogastral width refers to the maximum width of the notogaster in dorsal view. Lengths of body setae were measured in lateral aspect. All body measurements are presented in micrometers. Formulas for leg setation are given in parentheses according to the sequence trochanter-femur-genu-tibia-tarsus (famulus included). Formulas for leg solenidia are given in square brackets according to the sequence genu-tibia-tarsus. Drawings were made with a camera lucida using a Leica transmission light microscope "Leica DM 2500". Images were obtained with an AxioCam ICc3 camera using a Carl Zeiss transmission light microscope "Axio Lab.A1". SEM micrographs were made with the aid of a JEOL-JSM-6510LV SEM microscope.
Terminology -Morphological terminology partially used in this paper follows that of Grandjean: see Travé and Vachon (1975) for references, Norton (1977) for leg setal nomenclature, and Norton and Behan-Pelletier (2009) for overview.
Notogaster -Anterior part of notogaster with two pairs of connected condyles (medial pair large, lateral pair small). Anterior margin inconspicuous, fused with prodorsal hollow, sometimes depressed or with unpair median tubercle. Ten or 13 (if da, dm, dp developed) pairs of setiform, thickened or phylliform notogastral setae present; seta c inserted on ventrobasal part of medial condyle.
Anogenital region -Four pairs of genital, one pair of aggenital, two pairs of anal, and three pairs of adanal setae. Adanal lyrifissure located close and lateral to anal aperture.
Integument -Body color brown. Body surface with dense granulate microsculpturing. Prodorsal processes close to prodorsal aperture sparsely tuberculate (diameter of tubercle up to 10). Region between bothridium and acetabula I-III densely tuberculate (diameter of tubercle up to 4). Region between bothridium and pedotecta I with some longitudinal folds.
Etymology -The specific name is dedicated to Prof. Dr. Ludwig Beck (State Museum of Natural History, Karlsruhe, Germany), the well-known acarologist.
Remarks -In main morphological traits (prodorsal processes have prodorsal aperture; notogastral seta c setiform, with mediodistal part bent and directed posteriad, other notogastral setae thickened; distance between medial prodorsal condyles equal to lm-lm) Leobodes becki n. sp. is morphologically most similar to Leobodes praeconcavus Chen andWang, 2007 andL. yinae Aoki, 2000. However, the new species differs from both by the presence of interlamellar    seta of medium length, distinctly shorter than rostral and lamellar setae (versus long, not shorter than rostral and lamellar setae) directed posterolaterad (versus erect) in dorsal view. Also, L. praeconcavus has smooth (versus barbed) notogastral setae (except c) of medium length, lm reaching insertion lp (versus short, lm not reaching insertion lp); L. yinae has bifurcate (versus Table 1 Leg setation and solenidia of adult Leobodes becki n. sp., L. schalleri n. sp. and L. schawalleri n. sp.

Leg Tr
Fe

s, (pv)
Note: Roman letters refer to normal setae (except ɛ = famulus); Greek letters refer to solenidia. Single quotation mark (′) marks setae on the anterior and double quotation mark (″) setae on the posterior side of a given leg segment; parentheses refer to a pair of setae.
Integument -Body color brown. Body surface with dense granulate microsculpturing. Prodorsum close to insertions of lamellar setae and partially tutorium and pedotectum I with slight short ribs and tubercles (diameter of tubercle up to 10). Region between bothridium and acetabula I-III densely tuberculate (diameter of tubercle up to 4). Dorsal part of notogaster with sparse, small foveolae (diameter of foveola up to 4). Anogenital region (except ventrolateral parts and region between genital and anal apertures) with dense, larger foveolae (diameter of foveola up to 10).
Type deposition -The holotype and one paratype are deposited in the collection of the SMNH; 13 paratypes are deposited in the collection of the TSUMZ.
Etymology -This new species is dedicated to Friedrich Schaller (1920-2018), a famous German-Austrian zoologist. Among his numerous fields of research he was dedicated to ecology and behaviour of Collembola and other soil-bound invertebrates. For example, he discovered the indirect sperm transfer of collembolans.
Remarks -Leobodes schalleri n. sp. differs from all representatives of the genus by the presence of broadly phylliform (versus setiform or slightly phylliform only in basal part) notogastral setae c.
Integument -Body color brown. Body surface with dense granulate microsculpturing. Tutorium and pedotectum I foveate. Region between bothridium and acetabula I-III densely tuberculate (diameter of tubercle up to 4). Notogaster with sparse foveolae (diameter of foveola up to 10). Anogenital region (except ventrolateral parts and region between genital and anal apertures) with more dense foveolae (diameter of foveola up to 10).