Revision of the genus Mniophila STEPHENS , 1831 ( Coleoptera : Chrysomelidae ) With 7 figures and 3 maps

A revision of the genus Mniophila STEPHENS, 1831 is provided. Redescriptions of the genus and species, differential diagnoses, and distribution are given. New data on biology are presented. Three new species: Mniophila caucasica sp. n. from Caucasus (Russia, Abkhazia, Georgia, Azerbaijan), Mniophila transcaucasica sp. n. from Armenia and Georgia, and Mniophila taurica sp. n. from Crimea are described. The species level status of Mniophila bosnica APFELBECK, 1914 is confirmed. The status of Mniophila muscorum turcica L. MEDVEDEV, 1970 is elevated to a species level. The generic position of Mniophila exulans SAMUELSON, 1973 is discussed.


Discussion:
The species of the genus Mniophila inhabit primary mountain territories and foothills of Europe, Caucasus and Asia Minor and occur in deciduous and mixed forests of the nemoral origin one of indicators of which is Fagus.Ecological and morphological features of the genus Mniophila determine its distribution to a significant extent.First, a cryptic way of life with all life stages and trophic relations connected to moss in mesophilic or mesohygrophilic biotopes.Second, a rather small body size (1.1-1.6 mm), and reduced hind wings limit their dispersal ability.These features apparently determined the recent distribution of the genus Mniophila.Another important factor was the distribution of plant formations favourable to the distribution of this genus.Some paleobotanical data can be used to correlate the modern range of Mniophila.Recent mountain and foothill nemoral forests in Europe, Caucasus and Asia Minor are significantly fractionated.The forests in Western and Central Europe are more homogenous and continuous than those in Eastern Europe, Caucasus, and Asia Minor that are often restricted to large territories (SHELYAG-SOSONKO 1980;DIDUKH 1992).This is partly connected with a landscape because mountain territories usually are nemoral forest refugia.Thus, the range of Mniophila is continuous in Western and Central Europe while significantly split up in its eastern and southeastern parts, such as, Crimea, Caucasus, and Asia Minor.Straits, large rivers, and valleys separate mountain territories with nemoral forests in these regions.Obviously, in the past when the range of nemoral forests was more continuous and embraced larger territories in Europe, Caucasus, and Asia Minor (SHELYAG-SOSONKO et. al. 1987;KLEOPOV 1990;DIDUKH 1992) the conditions were more favourable for range expansion of Mniophila.When territories with nemoral forests were limited, barriers arose between them and interchange of flora and fauna diminished, and they became valuable refugia.So, exchange between these territories was significantly limited.The disjunctions of ranges of the species of Mniophila apparently are caused by these reasons.Recently, for the above-mentioned reasons, the exchange between populations of Crimea and Balkans, or Caucasus, was probably, impossible.These are some of the reasons why these populations are recognized as good species.Another criterion is morphological.The population's morphological variability was studied and allowed me to reveal some stable characters, such as body shape, head structure, antennae, prothorax, legs, and male genitalia.These characters are stable at the level of population.The character states of these characters can be used to distinguish species from each other.It is worthwhile to emphasize that stable complex of characters confirms the chosen species concept in Mniophila.In other words, e.g.Mniophila taurica sp.n. differs from Mniophila caucasica sp.n. in the same features as from Mniophila bosnica.Thus, the problem of a choice between subspecific or specific status in the present case, in my opinion, should be solved in favour of specific status for the following reasons.First, as mentioned above geographical distribution with restricted or impossible exchange between populations in most cases.Second, apparently sympatric distribution of some species (it can be supposed for Mniophila bosnica and Mniophila muscorum; Mniophila transcaucasica sp.n. and Mniophila caucasica sp.n.) and absence of intermediate forms.Third, there is a stable complex of morphological characters with equal differences between the species.Additionally, it can be mentioned that the degree of morphological differences between these species, in my opinion, is sufficient for recognizing them as good species and have the same level of difference as in many other genera of Alticini.The morphological uniformity of species of Mniophila does not provide a basis for establishing subgenera or species groups.This uniformity did not allow the production of a workable key for all species.There is only the affinity of M. taurica sp.n., M. caucasica sp.n., M. transcaucasica sp.n., and M. turcica on one side and M. muscorum and M. bosnica on the other side.The most reliable character for determination is the structure of aedeagus.The distribution of the species can also be used.For the help in identification keys to European and Caucasian species of genus are given.The genus Mniophila included a single species for a long time.M. bosnica was regarded only as a variation or subspecies until the paper of GRUEV (1979).Mniophila exulans SAMUELSON, 1973 was then described from Fiji (SAMUELSON 1973).The new species was attributed to the genus Mniophila with some reservations.This author mentioned some characters that are not shared with the genus Mniophila: "because it has upper frons and vertex strongly delimited instead of weakly and antennal segment 8 of normal size instead of reduced" (SAMUELSON 1973: 61).He also mentioned the genera Taizonia CHEN, 1934, Amphimeloides JACOBY, 1887 and Kamala MAULIK, 1926 that can be compared or related to this species.In my opinion attributing M. exulans to Mniophila has no sufficient bases.As mentioned above morphological characters possibly can be used against this generic assignmenet.The spermatheca figured by SAMUELSON (1973: fig. 27j) can also be applied to this argument.The range of distribution of M. exulans causes objections as well.As mentioned above the species of genus Mniophila are distributed in mountains and foothills of Europe and Asia Minor and connected with nemoral forests of mountain systems of Alpine folding.Primarily European genera Aeschrocnemis WEISE, 1888, Apteropeda CHEVROLAT, 1839, Minota KUTSCHERA, 1859, and Orestia GERMAR, 1845 have similar distributions.In the present time Nepalese species of Orestia, and Himalayan, Chinese, and Japanese species of Minota were described but their generic position requires confirmation.Nonetheless, such disjunctions of ranges can be explained from the genesis of Palaearctic flora and fauna point of view.In the same time the origin and composition of Fijian flora and fauna have another history.Therefore, it is unlikely that a genus of the European fauna (with limited dispersal ability) has its own representative in the fauna of Oceania.My opinion has also been confirmed by A. SAMUELSON (pers. comm.) that Mniophila exulans represents a separate and undescribed genus.The generic position of Mniophila is unclear.The group Mniophilites was established in the alticine classification developed by CHAPUIS (1875).The rank of this group can be regarded as subtribe.After Chapuis this group was mentioned in the catalogues of HORN (1889 -Mniophilae) and LENG (1920 -Mniophilini).Undoubtedly, Mniophila differs in many characters from most Alticini.Such features like living in moss, very small size and globose shape of a body, reduced hind wings along with other characters usually rarely present together among other alticines.These are specialized features for a cryptic way of life.Similar characters are presented in some other alticine genera like Mniophilosoma WOLLASTON, 1854, Apteropeda, Minota, Clavicornaltica SCHERER, 1974, Kiskeya KONSTANTINOV et CHAMORRO-LACAYO, 2006, and some others genera.Comparison between these genera reveals that adaptations for a cryptic way of life and corresponding morphological characters apparently originated independently but in similar ways and resulted in their similarity.Significant morphological specialization raises difficulties in revealing of relations of Mniophila and other genera with the present state of knowledge.At the same time establishing of a separate high-level taxon (tribe or subtribe) for each genus will not clear up the situation but will result in fractionation of the classification.The only genus that can be compared with Mniophila is Mniophilosoma.The features shared by both genera are size and globose shape of a body and its proportions, shape of antennae and labrum, structure of thorax, male and female genitalia.Mniophila differs from Mniophilosoma in the structure of a head, particularly shallow and curved ocular sulci, long and narrow frontal ridge, frontal calli larger, pronotal lateral sides rounded, eighth antennomere smaller than seventh and ninth, elytra shorter.The question about the relationship of these genera cannot be resolved unambiguously because of probability of convergent similarity.

Biological remarks:
The genus Mniophila belongs to "minotoid" morpho-ecological group (NADEIN 2005).The other genera belonging to this group are Mniophilosoma, Minota, Apteropeda, Orestia, Clavicornaltica, Kiskeya, some species of Psylliodes LATREILLE, 1825.This morpho-ecological group is characterized by the following characters: small or very small size, 1-3 mm; colour dark, usually black, sometimes with metallic lustre; body compact, rounded, subspherical or elliptic-cylindrical, very convex; head drawn into prothorax, almost invisible from above; antennae and legs short, often swollen, legs fit into depressions on ventral side of body, and antennae into deep grooves lateral to frontal ridge.The winglessness is typical for the "minotoid" form as well as for many beetles that inhabit mountains.The trophic relations of Mniophila have been insufficiently studied.The host plants of Mniophila were recorded for the first time by KALTENBACH (1874).This author recorded Digitalis, Plantago (Plantaginaceae) and Teucrium (Lamiaceae) as host plants of leaf-mining larva.In that time the host plants of adults were not recorded and habitation in moss was the only indirect record for their trophic relations.After KALTENBACH (1874) these host plants were mentioned in the works of KASZAB (1962), MOHR (1966), MEDVEDEV & ROGINSKAYA (1988), GRUEV & TOMOV (1986), JOLIVET &HAWKESWOOD (1995), andDOGUET (1994), etc. Actually such trophic selection raises some doubts.It is a fact that adults of Mniophila can only be collected in moss.It means that beetles never transfer to other plants.This could be reliable confirmation that their host plants are mosses.This would confirm the opinion of COX (1997) that the larvae are not leaf miner of dicots and all stages of their life cycle are spent in moss.The observations on the first instar larva in Great Britain presented by this author revealed that it is most likely an external feeder on mosses.According to COX (1997)  The copulation of beetle in Abkhazia was recorded from 23 June to 8 July.The larvae, apparently, finish their development and are ready to pupate at the end of summer or beginning of autumn while imagoes emerge in spring.This supposition is based on collecting of adults in the first half of May in Crimea.COX (1997) mentioned that adults in Great Britain occur in moss throughout the year.Adults of Mniophila usually occur in moss at the trunk base and prominent roots of trees, fallen trees, stones and rocks of various sizes, soil, logs, and tree branches.Beetles prefer the mosses usually with a middle length of stems and not very dense growing on a stone and at the base of tree (usually deciduous, sometimes conifers).As a rule the beetles can be found in fresh, not dry or wet moss.Usually adults occur on moss and sometimes inside of moss' bed.The most preferable substrates are oriented horizontally or sloping.More rarely beetles occur on completely vertical substrate.Beetles that occur in mosses on trees prefer large stands of Fagus or Carpinus with thick base and with projecting trunks.Trees or stones favourable for Mniophila are usually situate in more or less shaded conditions.But sometimes adults occur in well lighted biotopes like stony slopes at glades.Beetles usually do not occur at high altitude above ground, the maximal recorded height was 1.5 meters above ground.Activity of adults has usually been recorded during the day but their activity in evening was recorded also.Adults moving up the trunk of Fagus was recorded in evening.At the altitude of 1300 m beetles were collected in the evening while the air temperature was 12-13 °C; at the altitude ca.2000 m the temperature recorded was 7-10 °C at the same time.Beetles occur singularly or in small groups up to 2-5 individuals on a unit of substrate.Such groups occur sporadically and do not form mass accumulations.Usually beetles are not rare in favourable biotopes.Correspondence between the kind of substrate and the altitude is recorded.At an altitude of 200-500 m beetles usually were in the moss at trees base.Then with gradual increase in altitude a preferable substrate is moss on stones.In Abkhazia at the altitude of ca.1300 m the beetles were collected exclusively in the moss on stony slope under Rhododendron bushes hanging down.Observation on the behavior of the species of Mniophila demonstrates that individuals inhabiting moss on stones usually occur on the moss motionlessly or crawl from site to site.The individuals inhabiting moss on trees bases and trunks usually occur in deeper moss.When threatened adults exhibit an escape response (thanatosis) where they fall downward.Observations show that beetles usually jump reluctantly and at small distance.Probably, inhabiting in hidden conditions and small size promoted development of another tactics of behavior when jumping activity has secondary value.Key to the European species of Mniophila STEPHENS, 1831 1.
Aedeagus ventrally with apical third short, straight margins and nearly straight apex with denticle to elongated, narrow, with narrow apex without denticle (Fig. 2); eyes convex, body usually slightly elongated (Fig. 1F); male first metatarsomere short and narrow (Fig. 1K), colouration black usually with greenish metallic lustre; pronotum long with base narrowly rounded (Fig. 1G

Redescription:
Body dark to black, shining, with greenish or rarely bronzy luster.Antennae and legs brown.Body shape rounded with weakly elongated elytral apices to distinctly elliptical.Head large; vertex wide; eyes gently convex to usually clearly convex.Antennae with thick segments.Head surface covered with well developed, large shagrination.Pronotum long with clearly rounded A-E -body outline: A-C -female, D, E -male; F -head; G -pronotum outline; H -hind femur; I -hind tibia of male; J -fore tibia of male; K -hind tarsus of male; L -fore tarsus of male; M -antenna.Scale bar: A-E -1 mm, F-M -0.5 mm.
base.Pronotal surface covered with shagrination; punctation large, sparce, shallow, weakly visible.Elytra with greatly variable punctation.Usually punctation well developed, small, dense, striae confused; secondary punctation well developed, often as large as strial.Rarer strial punctation regular and secondary punctation smaller or almost indistinct.Legs thickened; tibiae straight or weakly curved; hind femora wide.First protarsomere of male usually short and moderately narrow.Aedeagus (Fig. 2) ventrally with apical third short with straight margins and nearly straight apex with denticle to elongated, narrow, with narrow apex without denticle.From lateral view apical third clearly gradually narrowed to apex, sometimes wider.Body length -1.20-1.68mm, width -0.89-1.21mm.

Differential diagnosis:
Differs from M. bosnica in structure of aedeagus (Fig. 2): apical third in posterior view elongate with straight margins instead of short with round margins, apex straight or nearly straight with a denticle instead of apex rounded without denticle (Fig. 3J, K); body more elongate (Fig. 1A-E); eyes convex (Fig. 1F); pronotum longer (Fig. 1G); first metatarsomere thinner and shorter (Fig. 1K), first protarsomere of male shorter (Fig. 1L), colouration of the body darker, dark brown to black, usually with weak green luster; elytral punctation usually denser, punctation often confused, secondary punctation well developed, usually as large as punctures in striae.From M. taurica sp.n. differs in structure of aedeagus: apex usually elongate, with narrowing straight margins in posterior view or apical third with straight margins instead of sides toward apex parallel-sided, apical 1/4 with straight margins (Fig. 5K), denticle always present and well developed instead of denticle short or poorly developed; shagrination of the head more developed; eyes more convex; legs thicker; hind femur broader (Figs 1H, 5D); punctation of pronotum usually more developed; differs also in the shape of pronotum.

Discussion:
Mniophila wroblewskii WAŃKOWICZ was described as a separate species and later was regarded as a variation of Mniophila muscorum by WEISE (1883).This variation was also recorded from Caucasus and Crimea (WEISE 1906;HEIKERTINGER & CSIKI 1939;DOGUET 1994).Study of the material from Caucasus and Crimea revealed no Mniophila muscorum in these regions but three species new to science.The name "wroblewskii" was based on European specimens of Mniophila and treated as variation of Mniophila muscorum.Therefore, this name cannot be used for one of the new species from Caucasus or Crimea.Study of the variability of Mniophila muscorum reveals neither forms and variations nor subspecies.The main basis for description of the new forms or variations was the elytral punctation character.The state of this character is significantly variable and there is no correlation with geographic distribution.The character of punctation varies greatly at the level of population.There are individuals with more or less confused or regular punctation in the same population as well as with reduced or developed punctation.Therefore, I cannot agree with MOHR (1966) and STREJČEK (1993) on the subspecies status of Mniophila wroblewskii.The same concerns also Mniophila muscorum fa.seriatopunctata described by ROUBAL (1932) and Mniophila muscorum fa.fortepunctata described by HORION (1939).These were described as definite forms and cannot be recognized as a species or as a subspecies (ICZN 2000, Arts. 45.5, 45.6, 45.6.4).The descriptions of these forms were based on intraspecific variability of elytral punctation and they are not localized geographically.

Redescription:
Body brown with bronzy lustre or without, sometimes dark-brown, margins of pronotum and elytra sometimes lighter; legs brown.Body distinctly rounded, wide, elytral apices not elongated or weakly elongated.Head large, vertex wide, eyes flattened, rarely weakly convex.Surface of a head with well developed shagrination.Antennae (especially in males) clearly thickened.Pronotum short with very wide and weakly rounded base.Pronotal surface covered with well developed, large shagrination; punctation large, shallow, usually poorly visible.Elytra with punctation arranged in regular striae, sometimes partly confused, secondary punctation smaller than strial, not dense, rarely striae more confused and secondary punctation large.Legs distinctly thickened, wide (especially in males).First protarsomere of male large, long, and wide.Tibiae straight or weakly curved.Hind femora wide.Aedeagus (Fig. 3J, K) ventrally with apical 1/3 with clear, often rounded margins and rounded apex without distinct denticle, medial third usually narrower than apical, from lateral view apical third wide, sharply narrowed to apex.Body length -1.28-1.61mm, width -0.95-1.19mm.

Differential diagnosis:
Differs from M. muscorum: structure of aedeagus (Fig. 3J, K) with apex short in posterior view rounded, apex laterally rounded, apically without denticle instead of apex elongated or short with straight margins and well developed denticle (Fig. 2); body more rounded (Fig. A-C); eyes flattened; prothorax shorter (Fig. 3E); first protarsomere of male thicker and longer (Fig. 3G); first metatarsomere of male longer (Fig. 3F); colouration of the body somewhat lighter, brown to dark brownish with bronzy lustre; elytral punctures usually sparser, usually arranged in regular striae, secondary punctation smaller, usually weakly developed.From M. turcica differs: in structure of aedeagus: apex short at view from behind with rounded sided, apex rounded, without denticle instead of aedeagus ventrally with apical third gradually narrowed to apex; apex almost straight with large denticle; punctated pronotum, tibiae usually thinner and less curved, segments of antennae wider.From M. taurica sp.n. differs: structure of aedeagus with apex short in posterior view, laterally rounded, apex rounded, without denticle instead of aedeagus ventrally toward apex parallel-sided, wide, apical 1/4 with straight, narrowing sides and with distinct and straight apex with rather short and wide denticle or with poorly developed one; coloration of the body (without green reflection); legs and antennae thicker, head larger and of another structure, shagrination of the head more developed, punctation of elytra usually more regular and larger; differs also in the shape of pronotum.

Discussion:
Mniophila bosnica was described as separate species by APFELBECK (1914).Later this species was regarded as variation of Mniophila muscorum (HEIKERTINGER 1930).A subspecies level status was applied by MEDVEDEV (1970) without any comments.The species level status of Mniophila bosnica was restored by GRUEV (1979) based on morphological differences in the structure of aedeagus, the present study confirms this.Mniophila bosnica is closely related to Mniophila muscorum but differs in the aedeagal structure and other characters mentioned above.The data on the distribution of Mniophila bosnica were limited by records from Bosnia and Montenegro (GRUEV 1979;GRUEV & DÖBERL 1997;WARCHAŁOWSKI 2000).A detailed examination of the material revealed that this species is more widely distributed.The distribution presented here is based on the material examined in the course of this study.It is highly possible that real range of distribution of this species is larger.Mniophila turcica L. MEDVEDEV, 1970 stat.n. (Figs 4,10) Mniophila muscorum turcica L. MEDVEDEV, 1970: 317, figs 1a, b. -GRUEV & DÖBERL 1997: 241. -ASLAN et al. 1999: 397.Type locality: Turkey: Rize.

Redescription:
Body dark brown with weak bronzy luster; legs and antennae brown.Body almost rounded with weakly elongated elytral apices.Head large, long, eyes moderately convex; frons long, frontal ridge strongly convex; antennal grooves rather deep.Head surface covered with shagrination.Antennae not thick, segments short.Pronotum moderately long with wide and widely rounded base.Its surface covered with well developed shagrination, impunctate.Elytra smooth, punctation weakly developed, punctures large, poorly visible, shallow; in apical third invisible.Punctures arranged in partly confused striae; secondary punctation not developed.Notch between metathoracic cavities concave.Legs rather thick, clearly curved.First protarsomere of male moderately wide, not very long; metafemora wide.Aedeagus ventrally with apical third gradually narrowed to apex; apex almost straight with large denticle; basal 2/3 gradually widened toward apical third; from lateral view apical 2/3 almost straight, slightly curved apically.Body length -1.2-1.5 mm, width -0.98 mm.
notch between metathoracic cavities concave; first metatarsomere longer and narrower (Fig. 4C); dorsal punctation poorly developed.From M. caucasica sp.n. differs: in structure of aedeagus, apex almost straight instead of aedeagal apex rather wide, straight, apical 1/4 in posterior view more or less sharply narrowed to apex; apical segments of antennae shorter; tibiae thicker and more curved; first protarsomere (Fig. 4D) and that of mesotarsomere of males shorter and wider; dorsal punctation poorly developed and shagrination well developed.From M. bosnica differs: in structure of aedeagus, ventrally with apical third gradually narrowed to apex, the latter almost straight with large denticle instead of aedeagal apex with rounded sides and apex, the latter without denticle; pronotum impunctate; tibiae usually more curved (Fig. 4E, F), antennal segments thinner (Fig. 4G).

Discussion:
This form has been described as a subspecies by MEDVEDEV (1970).Study of the type material reveals that this form is well differentiated from other species of the genus and should be recognized at the level of species.This form differs from M. muscorum and other species in a set of characters and their states are at a level significant enough to regard this form in a specific rank.
The above description of this species is based on two paratypes.The quite confused elytral punctation is mentioned in the original description.The two paratypes examined have elytral punctation is arranged in partly confused striae.

Etymology:
The specific epithet refers to geographical distribution of the new species that is endemic to Crimea.

Description:
Body black, shining, often with greenish luster; legs and antennae yellow-reddish to light brown.Body almost rounded, with weakly elongated elytral apices.Head small, vertex moderately wide; eyes convex.Head surface covered with poorly developed, smoothed shagrination, sometimes vertex almost smooth.Antennae thin.Pronotum comparatively long, with distinctly rounded base, its surface covered with fine shagrination, punctation small, weakly visible among surface's microsculpture.Elytra with small, dense punctation, striae usually confused; secondary punctation usually well developed, size as large as in striae or nearly so; rarely striae regular.Legs thin; first protarsomere of male almost not widened or weakly widened; tibiae straight or slightly curved, metafemora narrow.Aedeagus (Fig. 5K-M) ventrally toward apex parallel-sided, wide, apical 1/4 with straight, narrowing sides and with distinct and straight apex with rather short, wide denticle or denticle poorly developed; from lateral view apical half gradually narrowed to apex.Body length -1.34-1.59mm, width -0.95-1.21mm.

Differential diagnosis:
From M. caucasica sp.n. differs: in structure of aedeagus (Fig. 5K-M) with poorly developed denticle or with wider and shorter one, from lateral view narrower; flattened eyes (Fig. 5C); apical antennal segments shorter (Fig. 5E); shape of pronotum with more rounded and more elongated base (Fig. 5F); head more elongated with vertex narrower, hind femora narrower (Fig. 5D).From M. transcaucasica sp.n. differs: in structure of aedeagus, wide, ventrally toward apex parallelsided, apical 1/4 with straight, narrowing sides and with distinct and straight apex with rather short, wide denticle or poorly developed instead of aedeagus ventrally with apical third gradually narrowed to apex, the latter obtuse with well developed denticle; head longer, shagrination of head less developed, frontal calli more developed; pronotum longer with more convex base; tibiae less curved and usually straight (Fig. 5G, H); hind femora narrower; notch between metathoracic cavities concave; first protarsomere of male narrower.From M. turcica differs: in structure of aedeagus, wide, ventrally toward apex parallel-sided, apical 1/4 with straight, narrowing laterally, apex distinct and straight with rather short, wide denticle or poorly developed compared to aedeagus ventrally with apical third gradually narrowed to apex, the latter almost straight with large denticle; tibiae nearly straight and usually less curved; shagrination of head less developed; hind femora narrower; pronotum punctate; tibiae thicker; eyes more flattened.
Key to the Caucasian species of Mniophila STEPHENS, 1831 1.
From M. turcica differs: in structure of aedeagus, ventrally with apical 1/4 more or less sharply narrowed to apex, the latter rather wide, straight instead of aedeagus ventrally with apical third gradually narrowed to apex, the latter almost straight; head shorter with deep antennal grooves; apical antennal segments longer; tibiae thinner and less curved; usually well developed punctation of pronotum and usually poorer developed shagrination.From M. taurica sp.n. differs: in structure of aedeagus, ventrally with apical 1/4 more or less sharply narrowed to apex, the latter rather wide, straight, denticle well developed instead of aedeagus ventrally toward apex parallelsided, wide, apical 1/4 with straight, narrowing sides and with distinct and straight apex with rather short, wide denticle or poorly developed; head shorter and wider, eyes more convex, apical antennal segments longer; hind femora thicker; differs also in the shape of pronotum with less rounded base.

Etymology:
The specific epithet refers to geographical distribution of the new species that is distributed in Transcaucasus.

Description:
Body brown to dark-brown with weak bronze luster or without one; legs reddish-brownish.Body nearly elongated to clearly rounded.Head large, short; vertex covered with well developed, large shagrination; frontal calli almost not convex; supracallinal sulci poorly visible; frontal ridge short, weakly convex; eyes flattened.Antennae with segments short and thick.Pronotum short, widely transversal, with very widely rounded base, its surface usually covered with well developed, large shagrination; punctures large, sparse, poorly visible.Elytra impunctate to well developed, dense punctation, punctures not large; striae partly confused, secondary punctation well developed.Notch between metathoracic cavities straight (Fig. 7C).Legs not very thick, tibiae moderately curved; metafemora wide; first protarsomere of male wide and thick.Aedeagus (Fig. 7J, K) ventrally with apical third gradually narrowed to apex; apex obtuse with well developed denticle; from lateral view nearly gradually curved from basal third, weakly and gradually narrowed toward apex.

Differential diagnosis:
From M. caucasica sp.n. differs: in structure of aedeagus (Fig. 7J, K), ventrally with apical third gradually narrowed to apex; apex obtuse instead of aedeagus ventrally with apical 1/4 more or less sharply narrowed to apex, the latter rather wide, straight; hind tibiae distinctly curved (Fig. 7F), first tarsomeres of male shorter and thicker (Fig. 7H, I); eyes more flattened; ocular sulci and frontal calli less developed (Fig. 7E); apical antennal segments shorter and thicker (Fig. 7D), differs in pronotum shape (Fig. 7B); notch between metathoracic cavities straight (Fig. 7C); last metatarsomere thinner (Fig. 7H).From M. taurica sp.n. differs: in structure of aedeagus, ventrally with apical third gradually narrowed to apex; apex obtuse with well developed denticle instead of aedeagus ventrally toward apex parallel-sided, wide, apical 1/4 with straight, narrowing sides and with distinct and straight apex with rather short, wide denticle or poorly developed; head shorter, frontal calli less developed, shagrination of head more developed; pronotum shorter with less elongated base; hind femora thicker; tibiae thicker and more curved; notch between metathoracic cavities straight; first protarsomere of male wider.From M. turcica differs: in structure of aedeagus, apex obtuse instead of apex almost straight; head shorter, frontal ridge less convex, antennal grooves shallower; notch between metathoracic cavities straight; eyes more flattened; tibiae thinner; punctation of pronotum well developed; first metatarsomere shorter and wider.