Descriptions of new species of Issikiomartyria (Lepidoptera, Micropterigidae) and a new genus Melinopteryx gen. n. with two new species from Japan

Micropterigidae is considered to be the sister group of all other extant Lepidoptera. In Japan, 17 species of five genera have been recorded including three endemic genera, Issikiomartyria Hashimoto, 2006, Kurokopteryx Hashimoto, 2006 and Neomicropteryx Issiki, 1931, all of which are associated with the liverwort genus Conocephalum Hill. We discovered four new species of Issikiomartyria from snowy regions in Northeastern Japan, and two new species of a new genus Melinopteryx gen. n. from the subalpine zone of the Akaishi Mountain Range. All these new taxa, I. hyperborea sp. n., I. leptobelos sp. n., I. catapasta sp. n., I. trochos sp. n., M. coruscans sp. n. and M. bilobata sp. n. are also associated with Conocephalum liverworts. Furthermore, females of I. akemiae Hashimoto, 2006 and I. plicata Hashimoto, 2006 are described here for the first time. Our extensive surveys revealed that the fine-scale endemism of Issikiomartyria restricted to the fragmented area facing the Japan Sea. Keys to Issikiomartyria species based on the adult morphology are provided.

All species of Japanese endemic genera, comprising at least 14 species, feed exclusively on Conocephalum Hill.liverworts and also seem to occupy similar ecological niches (Hashimoto 2006, Imada et al. 2011).Remarkably, the liverwort-feeding micropterigids in Japan have experienced diversification without changing hostplants, during or after the Miocene (Imada et al. 2011), which represent one of the largest radiations known to have taken place on a single host-plant genus.These Conocephalum-feeders are widely distributed across the major islands of Japan (i.e.Honshu, Shikoku, Kyushu) yet the species do not co-occur with one another (Imada et al. 2011).
Our thorough field surveys for molecular phylogenetic analysis have revealed that the micropterigid fauna in the Northeastern Japan is unexpectedly diverse.We reveal a new genus that inhabits an elevational zone of ca.1500-1800 m in the Akaishi Mountain Range, which is sister to Issikiomartyria in the molecular phylogenetic analysis of Imada et al. (2011).In addition, we newly discovered that four new species of Issikiomartyria were distributed across the area ranging from the northern end of Honshu to the northern part of Yamagata Prefecture, which has not sufficiently been investigated.Issikiomartyria is a genus consisting of five species, and each species tends to be found from geographically fragmented areas facing the Japan Sea (Issiki 1953;Hashimoto 2006).Furthermore, we describe females of I. akemiae Hashimoto, 2006 andI. plicata Hashimoto, 2006, for the first time.
In this study, males and females of four Issikiomartyria species new to science and females of two known species are described.Also, a new genus is established based on two species new to science.An updated account of the distribution of micropterigids in the northeastern Japan based on the detailed additional sampling records is provided.

Methods
Adults and larvae of micropterigid moths were collected from temperate forests in Japan, and larvae were reared in plastic cases with their host-plants.A total of 226 adult pinned specimens were used for this study.For genital dissections, the whole abdomen was removed and macerated for 30 min in 10 % KOH.Residual scales and tissues were then washed in distilled water to remove KOH, immersed in 50 % ethanol and dehydrated in an ethanol series.Genitalia were then stained with 5 % chlorazol black E for 10 min, and dehydrated in a series of 70−100 % ethanol.After washing, specimens were mounted and stored in 70 % glycerol.For some specimens, to examine the wing veins, wings were removed and scales were removed by brushing in 70 % EtOH.Observation and measurements were made under an Olympus BX53F microscope at 10-40× with the aid of a micrometer scale.
All the specimens examined in this study are deposited in the following collections: National Museum of Nature and Science (NMNS), Graduate School of Human and Environmental Studies, Kyoto University (KUHE).
Terminology follows Gibbs and Kristensen (2011) and Davis and Landry (2012)

Diagnosis.
Aedeagus with three pairs of dorsal fins, a pair of lateral triangular fins, and a ventral longitudinal fin.Genital chamber with a large genital sclerite with four paddle-shaped accessory sclerites at posterior end.
Description.The generic description is based on M. coruscans sp.n. and M. bilobata sp.n.
Head capsule densely covered by microtrichia; genal area glossy and naked; most of clypeus, frons, and vertex covered with brownish yellow piliform scales.Ocelli present.Antenna moniliform, approximately as long as forewing in male, longer than in female; densely covered with fuscous piliform scales on scape and pedicel; scape the largest segment, three times longer than most basal flagellum; pedicel small, as long as most basal flagellum.SOI (Kristensen and Nielsen 1979)  Foretibial epiphysis absent.Antero-lateral processes of pronotum present, weakly sclerotized.
Wing venation as shown in Fig. 3A, B. Fore-and hindwings obtuse at apex, forewing with brown to purple luster, cilia shining grayish brown.Forewing with Sc forked, R1 unforked; R3 stalked with R4+5.Hindwing with a main stem of R absent; most anterior vein of hindwing forked near terminal end (Sc1 and Sc2+R1).Abdomen grayish brown, covered with piliform and lamellar scales, scattered with dark orange piliform scales on venter and genital segments in male.Sternum V gland present; orifice of gland a narrow slit.
Male abdomen and genitalia.Sternum VIII membranous.Segment IX a complete ring, well sclerotized, with a posterior expansion dorsally.Valva triangular, broad-ly membranous at proximo-dorsal surface, with a proximo-ventral ridge; anterior portion fused with median plate; median plate large, roughly fan-shaped.Phallobase strongly curved, without ventral longitudinal ridge.Aedeagus stout at caudal end, with three pairs of fins dorso-medially; a pair of lateral triangular fins extending horizontally; a pair of ventral fin extending vertically; dorsal apex of aedeagus acute and ventral one slightly forked, longer than dorsal one; gonopore opening horizontally; vesica with large as the intergeneric COI divergence among other Japanese micropterigid genera (for example, 7.8 ± 0.1% between Neomicropteryx and Kurokopteryx), indicated by Imada et al. (2011).Melinopteryx has a 2-segmented labial palp, while 1-segmented in Issikiomartyria, except for I. bisegmentata Hashimoto, 2006 which is 2-segmented.Melinopteryx is characteristic in its aedeagus with three pairs of dorsal fins, a pair of lateral triangular fins, a ventral longitudinal fin or protrusion extending vertically in the male.In female, a large sclerite with more than four accessory sclerites in genital chamber is also unique in Melinopteryx.Fore-and hindwing without a radial cell and large bulbous corpus bursae in female are shared with Issikiomartyria.
Etymology.The genus name is a compound noun derived from the Greek words transliterated into Latin, "melinos" (honey-color) and "pteryx" (wing), referring to the adult wing color of the species of this genus.The gender is feminine.serrate minute projections.Tergum X broader than long, with a pair of long ventral plates (venter X plates) extending antero-ventrally at base of terminal processes.
Female abdomen and genitalia.Segment IX forming a complete ring, strongly sclerotized, with a dorso-lateral concavity, without lateral protrusion.Segment X composed by lateral sclerites and one or two dorsal sclerotized plates; lateral sclerites simple, broader than long, having digitate projections with an apical seta at terminal inner margin.Corpus bursae large, globular, membranous, with signa composed of three or four sclerites near caudal end.Genital chamber with a large sclerite (genital sclerite) and a few tiny sclerites; genital sclerite deeply furcated posteriorly into four paddle-shaped accessory sclerites.
Remarks.The genetic distance between Melinopteryx coruscans sp.n. (labelled as 'Issikiomartyria' sp. in Imada et al. 2011) and its sister clade Issikiomartyria spp.(6.3 ± 0.9%, based on the Kimura 2-parameter model) was almost as Diagnosis.Aedeagus with a pair of ventral longitudinal fins, extending more than half of aedeagal length; female corpus bursae with two forms of signa consisting of two semicircular sclerite and a long rectangular sclerite.
Description.Head dark brown, naked and glossy on both sides, sparsely covered with brownish yellow piliform scales with dark yellow scales on vertex.Antenna about same length of forewing in male, about 4/5 in female; with 67 (60-74) flagellomeres in males (n=7).Labial palp 2-segmented.Thorax grayish brown, sparsely covered with purple and brownish gold scales on prothorax with blue metallic scales, with dark yellow piliform scales on tegula.Forewing with brownish purple luster tinged with coppery, densely covered with golden luster over basal half of dorsum; cilia grayish brown, pale yellow on apex; ventral surface glossy grayish purple.Forewing length 5.1 mm (4.8-6.0,n=8) and 5.1 mm (n=1) in male and female, respectively.Hindwing glossy brownish purple scattered with piliform scales on basal half; cilia grayish brown; ventral surface same as forewing.Abdomen sparsely covered with grayish brown piliform scales.
Male abdomen and genitalia (Fig. 4).Mid-dorsal length of segment IX ring about 1/6 of ventral length.Valva with obtuse apex, with a very small proximo-ventral ridge; inner ventral margin broad without concavity.Aedeagus with a ventral longitudinal fin extending vertically more than half of aedeagal length; three pairs of dorsal fins present; a pair of lateral triangular fins extending horizontally.Tergum X with longitudinal convex at medial part, with a pair of triangular lobes disto-dorsally.
Variations.Geographic variation is recognizable between individuals in the populations at Irisawai and Shirabiso-touge.In the populations of Irisawai, the proximal portion of the male tergum X is stouter and much more developed than that of Shirabiso-touge; wing color tinged with strongly purplish scales at Irisawai population, whereas wing color of the individuals at Shirabiso-touge tend to be more tinged with coppery and golden scales.
Remarks.Melinopteryx coruscans sp.n. is distinguished from M. bilobata sp.n. based on the following character states: aedeagus with a ventral longitudinal fin extending more than half of aedeagal length; female segment X with a rectangular plate of dorsal sclerites; corpus bursae with two different forms of signa consisting of a pair of semicircular sclerites and a ribbon-shaped sclerite.This species corresponds to " 'Issikiomartyria' sp." in Imada et al. (2011).
Etymology.The specific name is a participle in the nominative singular from the Latin word "coruscans", which stands for flashing.

Distribution. The Western mountain range of the Akaishi Mountain Range of Japan (Honshu: Nagano Pref.).
Bionomics.There is a single generation per year; however, there may be one generation per two years in some populations at high elevation, where larvae exhibit two significantly different size during the same period of time.The habitat is the peak or valley of sub-alpine forests at approximately 1100-1820 m of the Akaishi Mountain Diagnosis.Aedeagus with a short ventral fin in male genitalia; female segment X with two reduced lobes of dorsal sclerite.
Description.Head dark brown, naked and glossy on both sides, sparsely covered with brownish yellow piliform scales with dark yellow scales on vertex.Antenna slightly longer than forewing in male; with 67 (64-73) flagellomeres in males (n=8).Labial palp 2-segmented.Thorax grayish brown, sparsely covered with purple and brownish gold scales on prothorax with blue metallic scales, with dark yellow piliform scales on tegula.Legs covered with glossy fuscous scales.Forewing with brownish purple luster tinged with coppery, densely covered with golden luster over basal half of dorsum; cilia grayish brown, pale yellow on apex; ventral surface glossy grayish purple.Forewing length 4.9 mm (4.6-5.0,n=10) in male.Hindwing glossy brownish purple scattered with piliform scales on basal half; cilia grayish brown; ventral surface same as forewing.Abdomen sparsely covered with grayish brown piliform scales.
Male abdomen and genitalia (Fig. 6).Mid-dorsal length of segment IX ring about 1/5 of ventral length.Valva with a small proximo-ventral ridge; inner ventral margin broad without concavity; apical end obtuse.Aedeagus with a ventral protrusion at base; with three pairs of short dorsal fins; with a pair of small, lateral triangular fins extending horizontally.Tergum X with squarish medial part, with a pair of spines disto-dorsally.
Female abdomen and genitalia (Fig. 7).Segment IX ring strongly sclerotized, shallowly concave dorso-laterally; mid-dorsal length about 1/2 of mid-ventral length, without lateral protrusion.Segment X consisting of a pair of lateral sclerites and two dorsal sclerotized plates; dorsal plates small, well sclerotized, being behind dorsal side of segment IX. Corpus bursae membranous, bulbous, possessing four semicircular signa.Genital chamber armed with a large sclerite with four plate-shaped accessory sclerites.
Remarks.Melinopteryx bilobata sp.n. is distinguished from M. coruscans sp.n. based on the following characteristics: aedeagus with a ventral protrusion at base; female segment X with a pair of small dorsal sclerites.
Etymology.The specific name is a compound adjective in the nominative singular from the Latin words, "bi-" (two) and "lobatus" (having diminutive lobes), referring to a pair of small dorsal sclerites of the female genitalia (Fig. 7).
Distribution.The Eastern mountain range of the Akaishi Mountain Range of Japan (Honshu: Shizuoka Pref.).
Head capsule densely covered by microtrichia, apart from genal area where glossy and naked; most of clypeus, frons, and vertex covered with brownish yellow piliform scales.Ocelli present.Antenna moniliform, approximately as long as forewing in male, longer than in female; scape the largest segment, twice longer than most basal flagellum; pedicel bulbous, larger than most basal flagellum; basal one or two flagellomeres cylindrical.SOI about 0.4.MIOI about 0.5.Interocellar sulcus almost complete.Postinterocellar sulcus distinct.Epicranial sulcus distinct between occipital foramen and postinterocellar sulcus, being as a short distance anterior to interocellar sulcus.Temporal sulcus as a darker line.Occipital sulcus almost complete, but slightly indistinct on dorso-lateral corner.Occipus fan-shaped.Mandibular teeth greatly reduced.Labial palp 1-or 2-segmented.Maxillary palp 5-segmented.Proximal prelabium obscure.Foretibial epiphysis absent.Antero-lateral processes of pronotum present, strongly sclerotized.Fore-and hindwings obtuse at apex, forewing with brown to purple luster, without any distinct maculation.Forewing with R1 unforked; R3 stalked with R4+5.Hindwing with a main stem of R absent; most anterior vein of hindwing forked near terminal end (Sc1 and Sc2 + R1).Sternum V gland present; orifice of gland a narrow slit.
Male abdomen and genitalia.Sternum VIII membranous.Segment IX a complete ring, well sclerotized, with a posterior expansion dorsally; posterior margin gradually expanded from dorsum to venter.Valva triangular, broadly membranous at inner surface, with a proximo-ventral ridge whose anterior portion fused with median plate; median plate large, roughly fan-shaped.Phallobase strongly curved, with or without longitudinal ventral ridge(s) on midline.Aedeagus with acute apex, ventrally forked slightly at caudal end; with two pairs of basal fins dorso-medially and a pair of lateral triangular fins; gonopore opening horizontally; vesica with serrate minute projections.Tergum X, broader than long, with a pair of long ventral plates (venter X plates) extending antero-ventrally at base of terminal processes.
Female abdomen and genitalia.Segment IX forming a complete ring, strongly sclerotized; anterior margin gradually expanded anteriorly from dorsum to venter; mid-dorsal length generally shorter than 2/5 of mid-ven-tral length; laterally protruded in some species.Segment X consisting of a pair of lateral sclerites and a dorsal sclerotized plate; lateral sclerites simple, broader than long, with digitate projections having an apical seta at terminal inner margin.Corpus bursae large, globular, membraneous, with signa composed of four sclerites.Ductus spermathecae arising from a hexagonal or round concavity.Genital chamber with small sclerite(s).
Comparative Remarks.The following characters are regarded as synapomorphies of Issikiomartyia: aedeagus with two pairs of hornlike dorsal projections and without any protrusion vertically in male: sclerite in female genital chamber greatly reduced.Diagnosis.Aedeagus with a pair of lateral triangular fins arising from ventral margin, extending horizontally.Female segment IX with a strong concavity extending from lateral to ventral sides.
Description.Head dark brown, naked and glossy on both sides, sparsely covered with yellow piliform scales with dark yellow scales on vertex.Antenna slightly longer than forewing in male; densely covered with fuscous piliform scales on scape and pedicel.Labial palp 1-segmented.Forewing length 3.8 mm (n=1) and 3.9 mm (n=1) in male and female.
Male abdomen and genitalia (Fig. 8).Mid-dorsal length of segment IX ring about 1/4 of ventral length.Valva sharply tapered apically, with a tiny proximo-ventral ridge whose anterior portion fused with median plate.Aedeagus with a pair of short distal fins and a pair of longer, proximal fins extending vertically, arising from dorsal side; a pair of lateral triangular fins ventrally-oriented, extending horizontally.Tergum X with a small medial part; shorter than half of valva; with a pair of spines disto-dorsally.
Female abdomen and genitalia (Fig. 9).Segment IX ring strongly sclerotized, concave both at lateral and ventral sides; mid-dorsal length about 2/5 of ventral length.Dorsal plate between segment X sclerites large, well sclerotized, enlarged dorso-caudally.Corpus bursae large, globular, membranous,  with signa composed of four tridenta-form sclerites near proximal end.Ductus spermathecae arising from a round concavity.Genital chamber with a small, triangular sclerite.
Remarks.Issikiomartyria hyperborea sp.n. is distinguishable from the known Issikiomartyria species by the following characters.In the male, aedeagus with a pair of latero-basal fins arising from ventral side.In the female, segment IX with a deep concavity extending from lateral to ventral sides; dorsal sclerite of segment X convex vertically in the middle.
Etymology.The specific name is an adjective in the nominative singular derived from the Greek word transliterated into Latin, "hyperboreus", indicating the mythical people of Greek mythology who lived "Beyond the North Wind".
Distribution.This species has only been found from Tsugaru peninsula of Japan (Fig. 14:1; Honshu: Aomori Pref).sclerite simple, broader than long; dorsal sclerite rounded at apex.Corpus bursae large, globular, membranous, with signa composed of four sclerites near proximal end.Ductus spermathecae from a round concavity.Genital chamber with a small, umbrella-shaped sclerite.
Remarks.Issikiomartyria leptobelos sp.n. is unique in valva with the digitiform apex in the male.This species is most similar to I. hyperborea sp.n., but can be distinguished by the segment IX without a concavity at lateral and ventral sides in the female.
Etymology.The specific name is a compound noun in apposition derived from the Greek words transliterated into Latin, "leptos" (fine) and "belos" (divine, arrow), referring to the digitiform valva of this species.
Distribution.This species has only been found from Hachimori-cho (Honshu: Akita Pref).
Bionomics.Larvae feed on the thalli of Conocephalum conicum.The locality is a forest path along a stream in the cool-temperate forests at 100-340 m.Type locality.Japan, Akita Pref: Hachimori (Honshu).
Male abdomen and genitalia (Fig. 10).Mid-dorsal length of segment IX ring about 1/4 of ventral length.Valva with digitiform, elongated apex; with a tiny proximo-ventral ridge.Aedeagus straight, with two pairs of dorsal fins: a pair of shorter, distal fins and a pair of longer proximal fins extending vertically; with a pair of lateral triangular fins extending upwards.Tergum X with squarish medial part, longer than 3/4 of valva; with a pair of protrusions disto-dorsally.
Female abdomen and genitalia (Fig. 11).Segment IX forming a complete ring, strongly sclerotized; strongly concave dorso-laterally.Segment X consisting of a pair of lateral sclerites and a dorsal sclerotized plate: lateral Diagnosis.Middle portion of tergum X in male undeveloped, approximately half of lateral portions.Female genital chamber with numerous sclerites at proximo-dorsally.
Description.Head dark brown, naked and glossy on both sides, sparsely covered with yellow piliform scales with dark yellow scales on vertex.Antenna longer than forewing in male, with 60 flagellomeres in female (n=1).Labial palp 1-segmented.Forewing length 4.3 mm (4.1-4.6,n=9) in male.
Male abdomen and genitalia (Fig. 12).Mid-dorsal length of segment IX ring about 1/4 of ventral length.Valva with elongated apex; with a very small proximo-ventral ridge.Aedeagus with two pairs of dorsal ridges: a pair of shorter distal fins and a pair of longer proximal fins extending vertically; with a pair of lateral triangular fins extending upwards.Tergum X with squarish medial part, with a pair of triangular lobes disto-dorsally.Female abdomen and genitalia (Fig. 13).Segment IX forming a complete ring, strongly sclerotized; with a pair of dorso-lateral protrusions near base.Segment X consisting of a pair of lateral sclerites and a dorsal sclerotized plate; dorsal sclerite longer than width, projected upward at caudal end.Corpus bursae large, globular, membranous, with long narrow neck region anteriorly; with signa composed of four saggitate sclerites near proximal end.Ductus spermathecae arising from a round concavity.Genital chamber with numerous tiny sclerites dorsally and a large, fan-shaped sclerite ventrally.
Remarks.Issikiomartyria catapasta sp.n. is most similar to I. trochos sp.n. in that lateral parts of tergum X extending dorsally, but can be distinguished by the following traits: two basal pairs of dorsal and lateral aedeagal fins closer to each other; numerous tiny sclerites scattered dorsally in female genital chamber; corpus bursae without tiny sclerites.
Etymology.The specific name is a noun in the genitive singular derived from a Latin word, "catapastus" (patchwork), referring to the numerous tiny sclerites in the female genital chamber of this species (Fig. 13).Diagnosis.Male tergum X with middle portion as large as lateral portions, lateral portions extending dorsally.Female corpus bursae with tiny sclerites on membrane surface near caudal end.
Male abdomen and genitalia (Fig. 14).Mid-dorsal length of segment IX ring about 1/4 of ventral length.Valva with a proximo-ventral ridge; inner ventral margin broad without concavity.Aedeagus with two pairs of dorsal ridges: a pair Etymology.The specific name is a noun in apposition from the Greek word, "trochos" (wheel, disk), referring to the unusually extended form of tergum X of this species.
Distribution.This species has been found from the northern part of the main island of Japan (Honshu: Yamagata Pref.).
Bionomics.Larvae feed on the thalli of Conocephalum conicum.The habitat is a forest path along mountain streams of cool-temperate forests at 340-595 m, where Fagus crenata and Quercus crispula dominate.

Description (based on female).
Head dark brown, naked and glossy on both side, sparsely covered with yellow pili-of shorter distal fins and a pair of longer proximal fins oriented about proximal quarter of aedeagal length, extending upwards; with a pair of lateral triangular fins extending horizontally.Tergum X with developed medial part, as large as lateral portions; lateral portions expanded dorsally.
Female abdomen and genitalia (Fig. 15).Segment IX forming a complete ring, strongly sclerotized; with a pair of dorso-lateral protrusions near base.Segment X consisting of a pair of lateral sclerites and a dorsal sclerotized plate; dorsal plate almost rectangular, slightly projected in middle.Corpus bursae large, globular, membranous, with long narrow neck region anteriorly; membrane surface with tiny sclerites scattered near caudal end; with signa composed of four saggitate sclerites near proximal end.Ductus spermathecae arising from a round concavity.Genital chamber smooth without sclerites, and a large fanshaped sclerite ventrally.
Remarks.Issikiomartyria trochos sp.n. is most similar to I. catapasta sp.n. in that lateral parts of tergum X extending dorsally, but can be distinguished by the following traits: dorso-and latero-basal aedeagal fins separated from each other; female genital chamber without small sclerites; corpus bursae with tiny sclerites on membrane surface near caudal end.form scales with dark yellow scales on vertex.Antenna longer than forewing in male.Labial palp 1-segmented.
Female abdomen and genitalia (Fig. 16).Segment IX forming a complete ring, strongly sclerotized, without lateral concavity.Segment X consisting of a pair of lateral sclerites and a dorsal sclerotized plate; lateral sclerites simple, as broad as long, with digitate projections having an apical seta at terminal inner margin.Corpus bursae large, membranous, with long narrow neck region anteriorly; with signa composed of four tridenta-form sclerites around neck region.Ductus spermathecae arising from a round concavity, forked in middle.Genital chamber with numerous tiny sclerites dorsally and a large fan-shaped sclerite ventrally.Description (based on female).Head dark brown, naked and glossy on both side, sparsely covered with yellow piliform scales with dark yellow scales on vertex.Antenna longer than forewing in male.Labial palp 1-segmented.
Remarks.I. akemiae can be distinguished by female genitalia having the ductus spermathecae forked in the middle.
Bionomics.Larvae feed on the thalli of Conocephalum conicum.Female abdomen and genitalia (Fig. 17).Segment IX forming a complete ring, strongly sclerotized; concave dorso-laterally.Segment X consisting of lateral sclerites and a dorsal sclerotized plate; lateral sclerites simple, as broad as long, with digitate projections having an apical seta at terminal inner margin.Corpus bursae large, membranous, with long narrow neck region anteriorly; with signa composed of four fan-shaped sclerites at distal of neck region; distal end densely sclerotized.Ductus sper-mathecae arising from a round concavity.Genital chamber with a sclerite ventrally.

Discussion on the distribution of Issikiomartyria
Our extensive field surveys have revealed that the distribution of Issikiomartyria is extended from northernmost to the central region of Honshu.Issikiomartyria species tend to be found from the geographically fragmented area facing the Japan Sea but not from the Pacific Ocean sides, although we conducted a census with considerable efforts throughout northeastern Honshu.The host-plant species is not likely to be the limiting factor of their distribution, because Conocephalum liverwort is widespread in the mainland of Japan (Imada Y and Kato M, pers. obs.).The Japan Sea side of the Japanese archipelago corresponds to the largely snow-covered area (Suzuki 1962) and harbors unique plant and animal species and it is called "Japan Sea elements" (Fukuoka 1966).The factors contributing the distribution of Issikiomartyria are unclear, they may be associated with heavy-snow conditions in winter.
To our knowledge, Issikiomartyria offers a largest example of regional diversification of insects in the northeastern Japan.Especially, it should be noted that I. hyperborea sp.n. may be the only insect species endemic to the Tsugaru peninsula so far known, which can be morphologically well-differenciated from the rest of Issikiomartyria spp.Several groups of animals represent genetic variations among the geographic populations in the northeastern Japan: terrestrial animals (Suzuki et al. 2004); Carabus ground beetles (Sota et al. 2001), freshwater fish (Yamamoto et al. 2004), and amphibious salamanders, Onychodactylus nipponoborealis (Poyarkov et al. 2012) and Hynobius lichenatus Boulenger, 1883 (Yoshikawa et al. 2008, Aoki et al. 2013).In addition, pronounced morphological differenciation is detected in a sinistral land snail group, Euhadra grata (Gude, 1900), occurring in the northeastern region.E. grata group consists of four morphologically distinct subspecies (Nishitani 1996, Kawana 2007), each of which range is allopatric one another.However, the geographic area where the genetic differentiations detected in these lineages are not correlated with the species ranges of Issikiomartyria.
Furthermore, we have discovered two species of Melinopteryx gen.n., M. coruscans sp.n. and M. bilobata sp.n., from highlands of the Akaishi Mountain Range.Melinopteryx gen.n. is sister to Issikiomartyria in a molecular phylogenetic analysis (Imada et al. 2011), and the species of both genera favor snowy regions of the Chubu and Tohoku regions of Japan, whereas some of the geographic populations are in proximity with either Neomicropteryx matsumurana or Kurokopteryx dolichocerata, which tend to occur in lower mountains.The species range of M. coruscans approximates two Japan endemic microptergid species: at the Akaishi Mountain Range, N. matsumurana (Fig. 19:48) and K. dolichocerata (Fig. 19:27,28) respectively inhabit northern and southern regions of Bunkui touge, a mountain ridge along the Median Tectonic Line, and M. coruscans has only been found in between narrow area higher than 1500 m.Likewise, M. bilobata sp.n. has been found from the southeastern area of the Akaishi Mountain Range (Fig. 19:3), where some populations of K. dolichocerata colonizes in adjacent but lower area (Fig. 19:33,34).Hence, this study supports prior findings that each Conocephalum-feeding micropterigid species belonging to the Japanese endemic micropterigid genera (Issikiomartyria, Kurokopteryx, Neomicropteryx) does not co-occur with one another (Hashimoto 2006, Imada et al. 2011).
The Japanese patterns of allopatry is even more extreme than that found in New Zealand Sabatinca yet contrasts markedly with a pattern of sympatry found in New Caledonia (Gibbs 1983;Gibbs and Lees 2014).Our study reinforces the potential research interest of Micropterigidae in differentially reflecting geological and ecological diversification processes at different spatial and temporal scales.

Figure 19 .
Figure 19.Locality records of micropterigid moths in the Northeastern Japan.Locality codes correspond to those in the text.Shaded are the areas higher than 1000 m elevation.Sampling records from Hashimoto (2006) are indicated as outlined symbols.
. Author's names are abbreviated: YI and MK stand for Yume Imada and Makoto Kato, respectively.