Miramella demissa Mulder, 2023

Miramella demissa Mulder View in CoL sp. nov. ( Figs 1–6)

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Diagnosis. External appearance is generally similar to other species of the genus. The new species can be distinguished from the parapatric congeners, Miramella irena and M. albanica by the unique shape of the male copulatory stilets, color and shape of the tegmen, pattern of the pronotum, pattern and color of the hind femora and color of the hind tibia. Only submicropterous specimens of Miramella demissa sp. nov. are known. In both males and females, the tegmina do not overlap and barely touch each other.

Etymology. Demissa is the nominative single female declension of the Latin adjective demiss (-a, us,-um) meaning ‘low-lying/of low altitude/descended’ and refers to its occurrence at relatively low altitudes, uncommon for representatives in the genus.

Material examined. Holotype ♂, MONTENEGRO, municipality of Bar, Velja Gorana ; 41°59'29.0394"N, 19°15'41.0394"E; alt. ca. 90 m; 21.VII.2022; CJM 20220150, RMNH.INS.1622326, J. Mulder leg. GoogleMaps ; Paratypes: ♀, MONTENEGRO, municipality of Bar, Velja Gorana ; 41°59'29.0394"N, 19°15'41.0394"E, alt. ca. 90 m; 21.VII.2022; CJM 20220151, RMNH.INS.1622327 GoogleMaps ; 1♂, 2♀♀; MONTENEGRO, municipality of Ulcinj, Gornja Briska ; 42°07'07.68"N, 19°13'08.2554"E; alt. ca. 470 m; 21.VII.2022, CJM 20220118, CJM 20220046, CJM 20220119; J. Mulder leg. GoogleMaps

Description Male holotype. HABITUS . Body 23.8 mm. General appearance in concordance with congeneric species. Overall color greenish with black markings alive ( Fig. 1). HEAD. Extensive black markings dorsally. Flagellum of the antenna green with some orange-brown. THORAX. Pronotum length 5.2 mm, green with bold black markings on the dorso-lateral edge and along the midline. Paranota below black band mainly green ( Figs 4A, 4D). WINGS. Tegmina semi-transparent greenish with some black veins, distal end on the third abdominal tergite. Hind wings smaller, approximately 80% of the tegmen length ( Figs 4A, 4D). LEGS. Left hind femur length 11.9 mm. Hind femur length 4.4× as long as wide; knee almost completely shiny black besides the proximal border of the cover plate; almost complete black ring at 2/3 of the length, separated from the knee by a white ring of almost same size; just in front of the middle of the femur as seen from the coxa a black spot on the inner side (inner upper marginal area, inner medial area) and outer side (upper marginal area and the upper carinula); except black at the position of the femoral ring: outer lower marginal area whitish with a long black spot, medial area mainly yellowish green, lower outer faced carinula whitish ( Fig. 5A); left hind tibia length 9.9 mm., outer side of hind tibia shiny black, distally light colored, two rows of 10 tibial spines on both hind legs whitish on the base and distal half black; fore and middle legs orange-green ( Fig. 6A). ABDOMEN. Tergites green with two rows of black markings. Sternites light green. GENITALIA. Left and right dorsal valves of the aedeagus not connected, touching over almost their entire length, less sclerotized and partly transparent, upright, conical, weakly triangular at base in cross section, slightly bend in ventral direction, dorsal concavity especially visible as a groove near the tip; ventral valves well sclerotized, apical parts divergent, tips in an angle of 60 degrees ( Figs 2A, 3A).

Variations. in males ( n=2 for genitalia, n=5 for color or morphological characteristics): No obvious differences were seen between the males, except the paratype being overall a bit smaller (length 21.8 mm) and a deviant number of 12 spines in outer row of right tibia and in the specimen from Virpazar the tegmen does not reach the third tergite.

Description Female paratypes (n=3). HABITUS . Body length 26.3–26.9 mm; characters largely same as holotype, but larger. THORAX. Pronotum length 5.9–6.2 mm, green with bold black markings on the dorso-lateral edge and along the midline. LEGS. Left hind femora length 14.0– 15.3 mm, hind femora relatively longer than in male, length-width ratio 4,9; coloration same as in males; left hind tibia length 12.5–13.1 mm, outer side tibia black in proximal half; outer side of hind tibia shiny black, distally light colored, two rows of 10 tibial spines on both hind legs ( 11 in right outer row in one female); tibia whitish on the base and distal half black ( Fig 5D); fore and middle legs orange-green. GENITALIA. Only the ovipositor valves were shortly investigated. No apparent differences compared to M. albanica were noticed. There are no outstanding characteristic features.

Differential diagnosis. Only few European congeneric species exist, of which only M. albanica and M. irena are known to occur in this part of the Balkans. The new species is compared to these two species. The noticed differences are well beyond intraspecific variation. Miramella demissa sp. nov. can be distinguished from M. irena and M. albanica by the following characters:

GENITALIA. The male genitalia of Miramella irena differ considerably from both other species. The apical parts of the aedeagus are much bigger. The dorsal valves are straight and on the dorso-lateral side there is a large and deep longitudinal convexity or groove on almost the full length in dried condition. The dorsal edge of the dorsal valves proceeds far and straight downwards without bulbous base ( Figs 2C, 3C). Only M. demissa sp. nov. and M. albanica are further compared. In Miramella demissa sp. nov. the upper half of the ventral valves bend laterally outward in both males available. Between the slender tips there is an angle of about 60 degrees, while the ventral valves of M. albanica are more or less straight in dorsal view in all specimens checked ( n=15). In the specimen depicted by Galvagni (1986b) an angle of 35 degrees is seen. The dorsal valves are characterized by a pair of non-connected but closely placed conical tubes. Its cross section is more or less angular with round edges, becoming sharper during the drying process. In lateral aspect, the ventral side is convex to flat at the top and slightly convex in dried condition. Its dorsal side is slightly convex ending with a longitudinal groove near the top. In M. albanica , the dorsal valves are much shorter, thinner and are placed widely apart on a broad basis or slope, thus starting on a higher level than the ventral valves. In lateral view a clear concave or angular shape between the base and the slender upper part can been seen in the dorsal edge ( n=17). The bases of dorsal valves in M. albanica show a, sometimes dorsally protruding, wellsclerotized connective tissue. This concavity and connecting tissue were already depicted in the images drawn by Galvagni (1986a). This structure is missing in both males of M. demissa sp. nov. and in some cases the structure runs down in the shape of a crest on the sheet of the aedeagus ( Figs 2A, 3A).

HIND FEMORA. Coloration is different in several aspects (see Fig. 5), from which the most obvious characters are in the lower marginal area and adjacent lower carinula of the outer face and the coloration of the knee. In all males and females of M. demissa sp. nov. checked ( n=8) the outer upper and lower marginal area and upper and lower carinula are black at the location of the complete femoral ring. The lower outer marginal area is whitish with black in a variable extent. Reddish color is absent in all specimens checked ( n=8: 5♂♂, 3♀♀). In male M. albanica this lower marginal area is whitish, the lower carinula whitish with black at the location of the incomplete femoral band only at the dorsal part ( n=212 out of 214). In female M. albanica practically in 100% of the specimens checked, the lower marginal area as well as the lower carinula are pink-red ( n=214 out of 215), with black of the partial band sometimes entering the dorsal half of the lower carinula. In M. irena the lower marginal area is consistently red in both sexes and the lower (and upper) carinula are black. The most distal femoral black ring in M. demissa sp. nov. is almost complete (sometimes missing in inner lower marginal area), in M. albanica , this band is usually less broad longitudinally and only covers the dorsal part up to halfway on the lower carinula. An exemption of the latter state has been found twice ( 2 males from Bosnia-Herzegovina) among 429 specimens checked. In females this black band can be very faint, small or even largely missing ( n=215). In M. irena , due to the black lower carinula and red lower marginal area, black coloration is present dorsally and absent at the lower outer marginal areas, nevertheless it is often absent at the upper marginal area. A second, partial femoral band just in front of the middle of the femur as seen from the coxa is present in M. demissa sp. nov. on the inner face, running over the dorsal carina. It includes the outer upper marginal area and upper carinula, sometimes even on a small part of the medial area.

In dorsal view, next to the black knee, the femur shows two distinct black bands. This character is variable in M. irena , but often the black coloration is missing at the outer upper marginal area. In M. albanica , the second band is often absent or rudimentary on the outer side (especially in females) and on the inner side it is present dorsally down to and including the upper carinula. The femoral part of the knee in M. demissa sp. nov. is largely (80–90%) shiny black, only leaving out the outer edge of the external ventral genicular lobe, opposed to partly black in M. albanica , leaving out a large part (55–90%) of the external ventral genicular lobe and usually also the dorsal rim, often even the whole dorsal side ( n=429). Black coloration on the knee in M. irena is variable and often absent from parts of the dorsal side and significant parts of the ventral lobe.

HIND TIBIA. In M. demissa sp. nov. the outer side of the tibia is largely black colored in both sexes, becoming pink-whitish distally in all specimens checked ( n=8). In M. albanica , the tibia is light colored (pink-beige to orangish) in all checked specimens ( n=429). Sometimes there is a small dark part present dorsoproximally ( Fig. 6). In M. irena , this character shows more variation and the tibia are often largely black.

PRONOTUM. In all females of M. demissa sp. nov. checked in this respect ( n=3) a bold black midline is present on the pronotum. A midline is rare in female M. albanica (5 out of 212). All males of M. demissa sp. nov. show a bold black midline ( n=5). In M. albanica , 86% of the males ( n=96) showed this state, but it strongly depends on the population. In all 12 males evaluated from Hajla, Kosovo no midline was present ( Fig. 4). A pronotal midline is common in M. irena .

TEGMINA. The submicropterous tegmina of M. demissa sp. nov. are much more wing-like than the slender and scale-like (squamipterous) winglets of M. albanica and almost touch each other. The tegmina are very short in comparison with the longer-winged M. irena , in which considerable variation can also occur within and between populations. The visible part of the tegmina behind the pronotum is -2 to +8% longer than pronotum midline length ( n=6) compared to about -10% in M. albanica and 60– 110% in M. irena . The tegmina reach the hind border of the second or somewhere on the third tergite in M. demissa sp. nov. In M. albanica , they just reach up to the hind border of the first tergite or onto the second tergite and are slender and slightly orientated downwards. In M. irena , they reach until the fourth tergite or much further and touch or overlap each other ( Fig. 4). The tegmina of M. demissa sp. nov. are greenish and semi-transparent with a whitish venation, however some veins are black. In both other species the tegmina are brownish and not transparent. The inner border is light colored in all three species, but less conspicuous in M. demissa sp. nov.

Distribution. Miramella demissa sp. nov. was found at two localities, 15 km far from each other. Next to these localities several other records were found. I reckon those records to M. demissa too, based on high probability. Some were accompanied by images confirming the specific characters. After my findings, Slobodan Ivković kindly pointed me to an observation by Nikolay Skobelev of presumably the same taxon some weeks prior to mine with an image on iNaturalist (https://inaturalist.org/observations/ 123325958) on June 25, 2022 from Gluhi Do, municipality of Bar ( 42°12'58.32"N, 18°59'46.68"E), alt. ca. 620 m. On the Russian travelers website Forum Vinsky a Ukrainian man under the pseudonym aoz1961 placed a record with an image of this species (https://forum.awd.ru/viewtopic.php?f=1101&t=392608) from Poseljani, Cetinje municipality 42°18'28.4394"N, 19°03'5.76"E, alt. ca. 20 m, in 2020. Although this is not a scientific source, it provides a magnificent in situ image and one of the few distribution points. Also, a record from Virpazar ( 42°14'52.7994"N, 19°05'14.9994"E) in 2017 by Slobodan Ivcovic and Laslo Horvat at ca. 5 m asl. with an image on the website Orthoptera Species File ( Cigliano et al., 2022) under Miramella albanica should be attributed to the new species. In Ivković et al. (2020) this location is described as Virpazar ‘reedy shore of Orahovštica river’. In the same publication, a second locality from 2012 was given by Godinje ‘macchia brook at the village’ with altitude 30 m and location 42°13'15.5994"N, 19°06'43.1994"E. The Miramella albanica record of Pavićević et al. (2006) from ‘Velika beach and its rural hinterland of Štoj’ in Ulcinj municipality most likely belongs to the new species too. Much older records from Sutorman, Bar municipality, of Miramella alpina and M. albanica by respectively Burr (1906) and Us (1968) can safely be assigned to M. demissa sp. nov. too. The eight localities are visualized in a map in Figure 7. The known distribution area is still small. It consists of the flanks of Rumija Mountain between the sea and Skadar Lake. The species can be considered as endemic for Montenegro, but as the border is within some kilometers, the existence of nearby Albanian localities cannot be excluded.

Ecology. Species within the Miramella group usually inhabit mountains at higher regions up to arctic altitudes ( Chintauan-Marquier et al., 2013). Miramella irena occurs mainly in altitudes between 1.000 and 2.000 m, but is known up to 2400 m and even at sea level in Friuli, Italy. In addition, M. albanica occurs at altitudes of 1.000 –2.000 m asl in Montenegro. The records of M. demissa sp. nov. on the other hand are known from lowland, from sea level up to about 600 m asl and close to the coast. The type locality is characterized by exuberant lowland vegetation with grasses, herbs and bushes. Orthoptera species found at this locality were Arachnocephalus vestitus Costa, 1855, Tylopsis lilifolia (Fabricius, 1794) , Leptophyes laticauda (Frivaldszky, 1868) , Poecilimon jonicus (Fieber, 1853) , Tettigonia viridissima (Linnaeus, 1758) , Decticus albifrons (Fabricius, 1775) , Pezotettix giornae (Rossi, 1794) , Calliptamus italicus (Linnaeus, 1758) , Anacridium aegyptium (Linnaeus, 1764) , Oedipoda caerulescens (Linnaeus, 1758) and Aiolopus strepens (Latreille, 1804) .

At the second locality the animals lived in a relatively shadow-rich situation within the edge of a chestnut forest. Other Orthoptera species found around this locality were Vichetia oblongicollis (Brunner von Wattenwyl, 1882) , Eupholidoptera schmidti (Fieber, 1861) , Pholidoptera dalmatica (Krauss, 1879) , Pholidoptera femorata (Fieber, 1853) , Pachytrachis gracilis (Brunner von Wattenwyl, 1861) , Chorthippus dorsatus (Zetterstedt, 1821) and Pseudochorthippus parallelus (Zetterstedt, 1821) .