Research Article |
Corresponding author: Sigitas Podenas ( sigitas.podenas@gamtc.lt ) Academic editor: Gunnar Kvifte
© 2022 Sigitas Podenas, Sun-Jae Park, Hye-Woo Byun, Virginija Podeniene.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Podenas S, Park S-J, Byun H-W, Podeniene V (2022) Hexatoma crane flies (Diptera, Limoniidae) of Korea. ZooKeys 1105: 165-208. https://doi.org/10.3897/zookeys.1105.82495
|
Hexatoma Latreille, 1809 is a large genus of short-palped crane flies with a worldwide distribution. Accounting for more than 60 percent of global species, 362 extant species occur in Asia. Prior to our study, Hexatoma crane flies on the Korean Peninsula (both the Democratic People’s Republic of Korea and the Republic of Korea) had been studied for 90 years starting in 1930, but only three species had been recorded, with one of them being a misidentification. This study adds six species to the fauna of the Korean Peninsula, all of which belong to the subgenus H. (Eriocera) Macquart, 1838. General information on genus and subgenus morphological characters is presented in this paper, with a redescription of each species based on Korean specimens, as well as illustrations of both sexes, the elevation range, the period of activity, habitat information, general distribution, and a distribution map for the Korean Peninsula. Three species H. (E.) ilwola Podenas, sp. nov., H. (E.) pianigra Podenas, sp. nov. and H. (E.) serenensis Podenas, sp. nov. are described as new to science. Hexatoma (E.) lygropis (Alexander, 1920) is deleted from the Korean species list as a misidentification. This publication is a continuation of our previous work on short-palped crane flies (Limoniidae) from Korea.
East Palaearctic, Limnophilinae, North Korea, South Korea, Taiwan, taxonomy
Investigations into Korean short-palped crane flies, family Limoniidae (Diptera), began more than a century ago with the first specimens collected as early as 1915 (
The original description of the genus Hexatoma (
The genus has a worldwide distribution and includes 596 extant species (
Larvae of all known Palaearctic Hexatoma species are aquatic, most of them developing in rivers with sandy or gravel bottoms, some Nearctic species developing in bogs (
The second publication on Korean crane flies (
Commensing our studies, we expected a high diversity of Hexatoma crane flies in Korea due to the abundance of suitable habitats, specifically rivers with sandy or gravel bottoms.
Since 2012, crane flies have been collected annually in different localities, at different times and using different methods throughout the country. Despite original and subsequent descriptions of East Palearctic species over a long period of time, some of these species were known only from the original descriptions and no illustrations were available. The aim of our study was to document, redescribe, illustrate, and prepare keys for all Korean crane fly species identified to date. In this article, we provide photographs of important taxonomical details, such as antennae, wings and male and female terminalia. We also include distribution maps of the Korean species, as well as a key for all the species of the Korean Peninsula. This publication is a continuation of our previous work on short-palped crane flies (Limoniidae) from Korea. We plan similar treatments of the subfamily Chioneinae and the family Pediciidae which potentially could yield many more species for the Peninsula.
Crane flies available for this study (Table
Adult crane flies were collected in various ways, including by insect nets, with Malaise traps, LED light traps, black light traps, Mosquito Magnet traps (Pro Model, Woodstream Corp., Lititz, PA), New Jersey traps and at light sources. Some specimens were preserved dry in envelopes in the field and were later mounted at the laboratory on their side on a paper point, with legs generally surrounding the insect pin. Other specimens were preserved in 96% ethanol (ETOH). Some specimens were slide mounted in Euparal; the genitalia of males and ovipositors of females were cleared overnight in approximately 10% potassium hydroxide (KOH) and preserved in microvials filled with glycerol on the same pin as the dry insect, or on a separate pin if the crane fly was preserved in ETOH.
Locality | Year | Coordinates (N*, E*) | Collector | Method | Collection |
---|---|---|---|---|---|
S. Korea, Suigen, Chosen | 1930 | 37°16.00'N, 127°01.00'E | I. Tabashi | Net |
|
N. Korea, Ompo (Onbo, Hamgyeongbuk-do,Gyeongsung-gun) | 1937 1938 1939 | 41°30.81'N, 129°34.69'E | A. M. Yankovsky | Net |
|
N. Korea, Seren Mts. (Hamgyeongbuk-do, Gyeongsung-gun) | 1938 | 41°41.24'N, 129°18.55'E | A. M. Yankovsky | Net |
|
N. Korea, Kankyo Nando, Puksu Pyaksan (Yanggang-do, Pungseo-gun, Mt. Buksubaeksan) | 1939 | 40°41.99'N, 127°42.96'E | A. M. Yankovsky | Net |
|
N. Korea, Chonsani (Yanggang-do, Daehongdan-gun) | 1940 | 41°59.62'N, 128°45.15'E | A. M. Yankovsky | Net |
|
S. Korea, #12, Hwy. #20, 8 mi. SW of Kangnung (Gangwon-do, Gangneung, Seongsan-myeon, Eoheul-ri) | 1954 | 37°42.00'N, 128°47.00'E | G. W. Byers | Net |
|
S. Korea, #25, #26 Central National Forest, 18 mi. NE Seoul (Gyeonggi-do, Namyangju-ai, Sudong-myeon, Naebang-ri) | 1954 | 37°44.89'N, 127°17.62'E | G. W. Byers | Net |
SMEK, |
N. Korea, Prov. South Pyongan, Pyongyan, Hotel garden | 1971 | 39°00.63'N, 125°45.10'E | S. Horvatovich, J. Papp | Net |
|
S. Korea, Gyeonggi-do, Paju-si, Aengmubong | 1973 | 37°45.46'N, 126°55.65'E | Y. Kim | Net |
|
S. Korea, Gyeonggi-do, Pocheon-si, Soheul-eup, Gwangneung Forest | 1973 | 37°45.05'N, 127°09.70'E | O. Lee | Net |
|
S. Korea, Seoul, Mt. Suraksan | 1974 | 37°41.79'N, 127°04.93'E | – | – |
|
S. Korea, Gyeongsangbuk-do, Yeongju-si, Punggi-eup, Samga-ri, Mt. Sobaeksan | 2000 2001 | 36°55.28'N, 128°30.33'E | – | – |
|
S. Korea, Gyeongsangbuk-do, Bonghwa-gun, Mt. Seondalsan | 2000 2001 | 37°02.38'N, 128°42.55'E | – | – |
|
S. Korea, Gyeongsangbuk-do, Yeongyang-gun, Ilwol-myeon, Yonghwa-ri, Mt. Ilwolsan, Yonghwasa Temple | 2001 | 36°48.71'N, 129°07.55'E | – | – |
|
S. Korea, Jeollabuk-do, Namwon, Sannae-myeon, Buun-ri, Namwonsi Sannaemyeon Baemsagol | 2009 | 35°21.21'N, 127°34.95'E | S. W. Jung | Net |
|
S. Korea, Gangwon-do, Jeongseon-gun, Imgye-myeon, Dojeon-ri | 2011 | 37°32.15'N, 128°54.17'E | H.-W. Byun et al. | Malaise trap |
|
S. Korea, Gangwon-do, Pyeonchang-gun, Jinbu-myeon, Dongsan-ri, Odaesan NP | 2012 | 37°44.26'N, 128°35.50'E | S. Podenas | Net |
|
S. Korea, Jeollanam-do, Gurve, Masan-myeon, Hwangjeon-ri | 2013 | 35°14.62'N, 127°29.38'E | S. Podenas, H.-W. Byun | Net |
|
S. Korea, Gangwon-do, Chuncheon, Dongsan-myeon, Bongmyeong-ri, KNU experimental Forest | 2014 | 37°46.74'N, 127°48.94'E | S. Podenas | Net |
|
S. Korea, Gyeonggi-do, Gapyeong-gun, Buk-myeon, Jeokmok-ri | 2014 | 37°58.61'N, 127°26.59'E | D.-G. Kim, M.-D. Baek, H.-D. Gang, Ch. Uy | Net |
|
S. Korea, Gyeonggi-do, Gapyeong-gun, Buk-myeon, Jeokmok-ri, Garim-gyo (Br.) | 2015 | 37°58.55'N, 127°26.49'E | Y. J. Bae | Malaise trap |
|
S. Korea, Jeollanam-do, Gurye-gun, Toji-myeon, Naeseo-ri, Piagol valley | 2015 | 35°16.31'N, 127°34.29'E | S. Podenas | Net |
|
2016 | 35°16.40'N, 127°34.15'E | ||||
2019 | 35°15.50'N, 127°34.93'E | ||||
35°15.95'N, 127°34.85'E | |||||
35°16.03'N, 127°34.66'E | |||||
S. Korea, Gyeongsangbuk-do, Gyeongju-si, Yangbuk-myeon, Janghang-ri | 2016 | 35°45.74'N, 129°21.84'E | S. Podenas, H. M. Baek | Net |
|
S. Korea, Gyeonggi-do, Yangpyeong, Cheongun-myeon, Dowon-ri | 2017 | 37°32.70'N, 127°47.69'E | S. Podenas | At light |
|
S. Korea, Gyeonggi-do, Paju-si, Gunnae-myeon, Jeongja-ri, Warrior Base Training Area | 2017 | 37°55.07'N, 126°44.50'E | T.A. Klein, H.-C. Kim | NJ trap |
|
Information on the examined material is given as it is on the labels, except coordinates, altitudes and measurement units which are given according to journal requirements. Also given are any additional labels kept with the specimen or additional notes on the same label, such as “metatype” written by Dr. Ch. P. Alexander, who originally described the species. For specimens collected by S. Podenas and his colleagues, the collecting date on the label is followed by a number in brackets. Different localities where insects were collected on the same date were given separate numbers and all information from those localities, whether in the field notes, databases, photographs, or other locality information, were marked with the specific number. Specimens are arranged according to the collecting date.
Prior to these studies all East Palaearctic and most Oriental species of Hexatoma (Eriocera) were studied and photographed. Special attention was payed to species recorded from neighbouring countries, like China, Japan, and Russia. Only four East Palaearctic species were not accessible to the authors of this publication and other entomologists who kindly helped with illustrations or photographs. These species are H. (E.) caesia (Savchenko, 1979), H. (E.) cleopatroides (Men, 2015) (
Crane flies were observed using an Olympus SZX10 dissecting microscope. Photographs were taken with a Canon EOS R5 digital camera through a Canon MP–E 65 mm macro lens and through Mitutoyo M Plan Apo 10× and 20× lenses mounted on the same camera.
The terminology of adult morphological features generally follows that of
The general distribution of species is given according to
Hexatoma
Nematocera
Anisomera
Peronecera
Trimacromera
Hexatoma nigra Latreille, 1809 (southern Europe).
Medium-sized to large crane flies with body length 6.5–32.0 mm and wing length 7.5–21.0 mm. Body coloration varies from yellow or orange to brown and black, some species have very distinct coloration.
Head. Rounded posteriorly without neck–like extension. Vertex wide with distinct tubercle. Length of antenna varies from short, hardly reaching wing base, if bent backwards, to very long, when it exceeds body length up to 4×. Antennae sexually dimorphic. Males usually have longer antennae than females, but that is because of elongated basal segments of the male flagellum. Antenna has reduced number of segments, less than typical 14–16-segmented antenna of most short–palped crane flies, often male antenna 6- or 7-segmented, that of female 8–11-segmented. Verticils missing or indistinct, but male flagellum often with two longitudinal rows of short erect spines medially.
Thorax. Some species with very setose thorax, setae could be long, dense, and erect. Some species with more dense and longer pubescence in males than in females. Prothorax very narrow but wide. Mesonotal prescutum usually without, sometimes with, small indistinct tubercular pits, pseudosutural fovea small. Prescutum and presutural scutum with three or four longitudinal stripes. Pleuron usually without stripes, could be bare or setose, depending on species. Meron usually big, thus middle and posterior coxae widely separated. Wing long and narrow, patternless or with very distinct pattern, sometimes completely dark, even black, but often with light “window” in the middle, stigma present or missing. Macrotrichiae missing on wing cells. Arculus present, humeral vein close to arculus. Vein Sc long, reaching wing margin far beyond branching point of Rs, sc-r slightly before tip of Sc. Radial sector with two or three branches reaching wing margin. R1 short, nearly transverse, or slightly elongate, R3 and R4 diverging. Cell r3 with long stem. Cell m1 present or missing; two, three or four branches of M reaching wing margin. Discal cell present or missing. Position of cross-vein m-cu differs according to species. Vein CuP usually slightly arched at distal part, anal vein long, slightly sinuous or arched, reaching wing margin close to the level of Rs base. Anal angle distinct, widely rounded. Wing cells without macrotrichiae. Wing squama setoseless. All legs with tibial spurs, usually fore leg with single spur, middle and posterior legs with two spurs each. Claw simple or with single subbasal spine.
Abdomen. Tergites with paired transverse sutures. Male terminalia approximately as wide as the rest of the abdominal segments, slightly elongate. Epandrium (ninth tergite) wider than longer, posterior margin simple without additional structures. Each gonocoxite elongate, two pairs of terminal gonostyli, the shape of which are only slightly variable among different species. Aedeagus simple, short, and straight. Ovipositor usually with long and narrow cerci and hypovalvae, distal part of cercus slightly raised upwards, acute. Some species with shortened ovipositor bearing fleshy cerci and hypovalvae.
596 species belong to the genus Hexatoma worldwide, they are divided into six subgenera:
H. (Eriocera) Macquart, 1838 (556 extant and three fossil species),
H. (Cladolipes) Loew, 1865 (three species, one of them with two subspecies),
H. (Coreozelia) Enderlein, 1936 (one Western Palearctic species),
H. (Euhexatoma) Alexander, 1936 (one Oriental species),
H. (Hexatoma) Latreille, 1809 (23 species, one of them with two subspecies),
H. (Parahexatoma) Alexander, 1951 (12 species, Afrotropics only) (
Six fossil species are described from the Eocene, three of them in H. (Eriocera), three not assigned to subgenera (
1 | Radial sector with three branches (Figs |
2 |
– | Radial sector with two branches (Fig. |
Hexatoma (Cladolipes) Loew, 1865 |
2 | Discal cell present (Figs |
3 |
– | Discal cell missing, two branches of M reaching wing margin (Figs |
5 |
3 | Supernumerary cross-veins missing in cells r3, r4 and r5 (Figs |
4 |
– | Supernumerary cross-veins in cells r3, r4 and r5 (Fig. |
Hexatoma (Euhexatoma) Alexander, 1936 |
4 | Vein Sc reaching wing margin beyond Rs branching point, R2 beyond fork of R3 and R4 (Figs |
Hexatoma (Eriocera) Macquart, 1838 |
– | Vein Sc reaching wing margin at Rs branching point, R2 at fork of R3 and R4 (Fig. |
Hexatoma (Coreozelia) Enderlein, 1936 |
5 | Ovipositor short with fleshy valves | Hexatoma (Hexatoma) Latreille, 1809 |
– | Ovipositor with long and slender valves | Hexatoma (Parahexatoma) Alexander, 1951 |
Hexatoma (Eriocera)
Macquart, 1838: 78, pl. 10, fig. 2;
Caloptera
Eriocera
Evanioptera
Pterocosmus
Allarithmia
Oligomera
Physecrania
Arrhenica
Penthoptera
Androclosma
Globericera
Coreozelia
Hexatoma macquarti (Enderlein, 1912) (= Eriocera nigra Macquart, 1838, = Hexatoma macquarti (Enderlein, 1912)) (Brazil).
Most characters as for the genus. Medium-sized to large crane flies with body length 6.5–32.0 mm and wing length 7.5–21.0 mm. Most species dark colored, but some could be orange-yellow (e.g., H. masakii Alexander, 1934).
Head. Rounded, vertex with distinct tubercle. Antennae sexually dimorphic. Male antenna longer than that of female, sometimes few times longer than body, 6- or 7-segmented, female antenna 8–11-segmented. Verticils missing or indistinct, but male flagellum often with two longitudinal rows of short erect spines.
Wing. Radial sector with three branches, discal cell always present, three or four branches of M reaching wing margin.
Terminalia. Male terminalia slightly elongate, not wider than preceding abdominal segments. Epandrium transverse, posterior margin slightly concave. Gonocoxite elongate with two pairs of terminal gonostyli. Outer gonostylus long, narrow with spine-shaped apex. Inner gonostylus long, fleshy, and setose. Aedeagus simple, usually short, and straight (Figs
Subgenus H. (Eriocera) includes 556 extant species (seven of them with two subspecies each). It has a worldwide distribution with the highest diversity in the Oriental region, 286 species (four of them with two subspecies each), the Neotropics, 143 species, and the Eastern Palearctic, 65 species. Thirty-three species (one of them with two subspecies) are recorded from Nearctic, 29 species (one with two subspecies) from Afrotropics, five species from Australasia, and four species from West Palaearctic (
Hexatoma (Eriocera) gifuensis Alexander, 1933
Hexatoma (Eriocera) ilwola Podenas, sp. nov.
Hexatoma (Eriocera) masakii Alexander, 1934
Hexatoma (Eriocera) pernigrina Alexander, 1938
Hexatoma (Eriocera) pianigra Podenas, sp. nov.
Hexatoma (Eriocera) serenensis Podenas, sp. nov.
Hexatoma (Eriocera) stackelbergi Alexander, 1933
Hexatoma (Eriocera) ussuriensis Alexander, 1934
Hexatoma wings 1 H. (Cladolipes) simplex (Loew, 1865) 2 H. (Coreozelia) cimicoides (Scopoli, 1763) 3 H. (Eriocera) gifuensis Alexander, 1933 4 H. (Euhexatoma) triphragma Alexander, 1936, holotype 5 H. (Hexatoma) khasiensis Alexander, 1962, holotype 6 H. (Parahexatoma) angustatra Alexander, 1963, holotype. Scale bars: 1.0 mm. (1 redrawn after
1 | Entire body, including legs and wings, coal black (Figs |
2 |
– | Body patterned with brown, gray, or orange (Figs |
3 |
2 | Wing cell m1 missing (Fig. |
Hexatoma (Eriocera) pernigrina Alexander, 1938 |
– | Wing cell m1 present (Fig. |
Hexatoma (Eriocera) pianigra Podenas, sp. nov. |
3 | Male | 4 |
– | Female | 9 |
4 | Wing cell m1 present (Figs |
5 |
– | Wing cell m1 missing (Figs |
6 |
5 | Antenna 3× as long as the rest of the body (Fig. |
Hexatoma (Eriocera) serenensis Podenas, sp. nov. |
– | Antenna distinctly shorter than the rest of the body (Fig. |
Hexatoma (Eriocera) ilwola Podenas, sp. nov. |
6 | Antenna at least twice as long as entire body, usually more than that (Figs |
7 |
– | Antenna not reaching wing root if bent backwards | Hexatoma (Eriocera) stackelbergi Alexander, 1933 |
7 | Abdomen orange yellow (Fig. |
Hexatoma (Eriocera) masakii Alexander, 1934 |
– | Abdomen brown or dark brown (Figs |
8 |
8 | Paramere with dorsal branch parallel-sided, lower branch wide, plate-shaped, anterior apodeme of aedeagus with wide lateral lobes (Figs |
Hexatoma (Eriocera) gifuensis Alexander, 1933 |
– | Paramere with dorsal branch wedge-shaped, lower branch elongate, anterior apodeme of aedeagus without lateral plates (Figs |
Hexatoma (Eriocera) ussuriensis Alexander, 1934 |
9 | Abdomen orange yellow. Costal wing area darkened (Figs |
Hexatoma (Eriocera) masakii Alexander, 1934 |
– | Abdomen brown or dark brown (Figs |
10 |
10 | Wing cell m1 missing (Figs |
11 |
– | Wing cell m1 present (Figs |
13 |
11 | Wing stigma distinct, dark brown (Figs |
12 |
– | Wing stigma very small, nearly missing (Fig. |
Hexatoma (Eriocera) stackelbergi Alexander, 1933 |
12 | Wing stigma elongate, oval, radial sector arched at base (Fig. |
Hexatoma (Eriocera) ussuriensis Alexander, 1934 |
– | Wing stigma approximately as long as wide, radial sector angulate at base (Fig. |
Hexatoma (Eriocera) gifuensis Alexander, 1933 |
13 | Thorax brown (Fig. |
Hexatoma (Eriocera) ilwola Podenas, sp. nov. |
– | Thorax gray (Fig. |
Hexatoma (Eriocera) serenensis Podenas, sp. nov. |
Hexatoma (Eriocera) gifuensis
Holotype
, male (wing and genitalia slide mounted), Japan, Gifu, 6 June 1931, Kariya leg. (
(Fig.
Body dark brown. Male body length 9.5 mm, wing length 14.3 mm. Female body length 12.5–13.5 mm, wing length 12.0–12.2 mm.
Head. Dark brown, postero–laterally yellowish. Vertical tubercle large, dark brown, yellowish laterally. Eyes widely separated, distance between them at the base of the antennae nearly the same as length of both basal antennomeres. Male antenna 47.3 mm long, ~ 3× as long as the entire body (Fig.
Thorax. Cervical sclerites dark brown. Pronotum short but wide, brown. Prescutum and presutural scutum grayish brown with three longitudinal dark brown stripes. Medial stripe separated anteriorly by narrow grayish line, which is missing posteriorly. Tubercular pits missing, pseudosutural fovea brown. Dorsopleural membrane dark brown, yellowish anteriorly. Postsutural scutum with each lobe brown with concave elongate dark brown spot in middle, area between lobes brown. Scutellum dark brown, lighter along posterior margin. Mediotergite entirely brown. Pleuron uniformly dark brown. Episternum bare, setoseless. Meron comparatively small, second and third pairs of legs close together. Wing (Fig.
Abdomen
. Tergites dark brown, narrowly yellow laterally, with two pairs of transverse indistinct sutures. Sternites dark brown along middle, yellow laterally. Male terminalia (Fig.
Ca. 20 m altitude.
Second half of July.
Unknown. Attracted to light.
Honshu and Shikoku islands of Japan. Recorded in the Korean Peninsula for the first time.
(Fig.
Large brownish gray species with body length 19.0–31.8 mm. Rostrum brown. Head and thorax with short and scarce pubescence. Male antenna reaching to approximately middle of abdomen if bent backwards. Prescutum and presutural scutum with four distinct dark brown stripes. Wing translucent with distinct stigma. Cell m1 present. Halter with dark knob. Femur yellow with narrowly blackened distal part. Abdominal sternites yellowish. Epandrium of male genitalia with wide V-shaped emargination. Gonostyli approximately equal in length. Posterior margin of inner gonostylus rounded, apical part slightly arched. Paramere V-shaped. Aedeagus simple, short, straight. Ovipositor with nearly straight cercus. Hypovalva long, distal part widened and setose, apex distinctly narrows into setiforme structure.
Species is named after type locality, Ilwol mountain.
Body coloration brownish gray. Body length of male 19.0 mm, female 23.0–31.8 mm, wing length of male 20.8 mm, female 16.3–20.6 mm.
Head
(Fig.
Thorax. Cervical sclerites brown, dusted with gray. Pronotum much wider than long, gray with narrowly yellowish anterior margin. Prescutum light bluish gray, presutural scutum bluish gray laterally, brownish gray posteriorly. Prescutum and presutural scutum with four distinct dark brown stripes (Fig.
Abdomen. Tergites dark brown, dusted with gray, narrowly orange along lateral margin, posterior margin narrowly orange starting from fourth tergite. All tergites with two pairs of transverse sutures and covered with very short yellowish setae. Sternites dark brown basally, obscure yellow laterally and posteriorly, dusted with gray. Male terminalia (Fig.
Hexatoma (Eriocera) ilwola Podenas, sp. nov. 12 holotype, male, dorsal view 13 head and thorax, dorsal view, paratype, female 14 male antenna, holotype 15 female antenna, paratype 16 female wing, paratype 17 male genitalia, dorsal view, holotype 18 aedeagal complex, dorsal view 19 ovipositor, lateral view, paratype. Scale bars: 1.0 mm (12–17, 19); 0.1 mm (18).
From slightly above 200 m to slightly above 500 m.
Whole of August.
Sandy and rocky margins of medium–sized mountainous rivers covered with deciduous forest and scarce grassy vegetation (Fig.
Korean Peninsula.
Hexatoma ilwola sp. nov. is most similar to H. aequinigra Alexander, 1934b, which is described and known only from the southern part of the Far East of Russia. Hexatoma aequinigra was described from the female, which is distinctly bigger than that of H. ilwola sp. nov. Hexatoma aequinigra has dense and long pubescence on head and thorax, while it is short and scarce in H. ilwola sp. nov. Hexatoma aequinigra has dark brown basal antennomeres, which are paler in H. ilwola sp. nov. Hexatoma aequinigra has pale yellow halter with dark brown knob, while the halter of H. ilwola sp. nov. is grayish brown with dark brown knob. Abdominal sternites of H. aequinigra are dark brown, but widely yellowish in H. ilwola sp. nov. Another similar species is H. sachalinensis, which is also known from the Far East of Russia, but it has a brownish black rostrum, dark brown femora, and brownish black tibiae. The rostrum of H. ilwola sp. nov. is brown, the legs yellow to brownish yellow. Unfortunately, the male of H. aequinigra is unknown and the male terminalia of H. sachalinensis have not been illustrated, thus comparison of the structure of male terminalia is not possible at the moment.
Hexatoma (Eriocera) masakii
Holotype
, male (pinned, antennae, legs and wing slide mounted), South Korea, Suigen, Chosen, 14 August 1930, I. Tabashi leg. (
Body
.Thorax brown, head, and abdomen orange-yellow (Fig.
Head. Orange-yellow, narrowly grayish along posterior eye margin, sparsely covered with short erect brown setae. Vertex with distinct uniformly pale orange-yellow tubercle. Eyes widely separated in male, distance between them at base of antennae nearly the same as length of both basal antennomeres. Male antenna 8-segmented, 16.0–19.5 mm long, ~ 2× as long as wing (Fig.
Thorax. Cervical sclerites and pronotum dark brown. Prescutum blackened. Presutural scutum and prescutum semi-polished dark brown, covered with scattered short brown setae and very sparse brownish pruinosity, longitudinal stripes missing. Tubercular pits missing, pseudosutural fovea distinct. Dorsopleural membrane yellow frontally. Postsutural scutum with each lobe blackish, area between lobes polished dark brown. Scutellum brown, sparsely dusted with brownish pruinosity. Mediotergite brown, darkened posteriorly. Pleuron brown, very sparsely dusted with gray. Episternum bare, setoseless, ventral margin of katepisternum blackish. Meron well developed, second and third pairs of legs staying apart. Wing (Figs
Abdomen. Abdominal segments orange yellow. Tergite laterally narrowly blackened, with paired transverse suture at ~ 1/3 of length. Sternite with lateral margin narrowly blackened and with longitudinal spot in the middle. Eight sternite without black spot in the middle. Lateral and ventral abdominal lines interrupted at posterior margins of segments. Whole ninth segment compact, making genital ring, yellow dorsally, pale brown ventrally. Male genitalia (Figs
Slightly above 100 m.
Middle of August.
Unknown.
(Fig.
Hexatoma (Eriocera) pernigrina
Holotype
, male (pinned), North Korea, Ompo, alt. 140 m, 14 June 1937, A. Y. Yankovsky leg. (
North Korea, 6 males, 2 females (pinned), Ompo, alt. 90 m, 9 June 1937, A. Y. Yankovsky leg. (
Body
coloration opaque black (Fig.
Head. Opaque black dorsally, dull black ventrally, sparsely covered with erect black setae. Vertex with small tubercle. Eyes widely separated in both sexes, distance between them at base of antennae nearly the same as length of scape. Antenna black at base, turning dark brown towards apex, 7-segmented in male (Fig.
Thorax. Cervical sclerites and pronotum black. Prescutum and presutural scutum opaque black with four semi–polished stripes, areas between stripes covered with dense short setae. Tubercular pits missing, pseudosutural fovea black, semi-polished. Postsutural scutum with each lobe black covered with grayish pruinosity, area between lobes polished-black. Scutellum dull black with narrow transverse wrinkles. Mediotergite black, laterally covered with grayish pruinosity. Pleuron black, sparsely dusted with gray. Wing (Fig.
Abdomen. Abdominal segments black with narrowly grayish posterior margin. Posterior segments dusted with gray, more densely on sternites. Tergites with paired transverse sutures frontally. Male terminalia (Figs
From less than 50 m to more than 1200 m.
From beginning of May through to middle of September.
Margins of mountainous small- and medium-sized streams densely covered with deciduous forests. Species is attracted to light.
(Fig.
(Fig.
Large black species (Fig.
The species is named after the type locality, the Pia River, and for the black color of the body (= nigra).
Body coloration black, semi-polished. Body length of male 14.0 mm, female 26.5–31.5 mm, wing length of male 16.3 mm, female 17.8–21.0 mm.
Head. Black, densely covered with brownish gray pruinosity and scattered short erect black setae. Eyes marginated by narrow whitish gray. Vertical tubercle large, rounded, with indistinct medial groove, concolorous with the rest of the head. Eyes widely separated in both sexes, distance between them at the base of antennae equal to length of scape. Male antenna 7-segmented, 5.2 mm long, reaching to approximately base of halter if bent backwards. Scape large, 2× as long as wide, 4× as long as pedicel, dark brown to blackish, sparsely dusted with brownish. Pedicel wider than long, black. Flagellum entirely black, densely covered with semi–erect black setae. Basal flagellomere longer than both basal antennomeres taken together and slightly longer than second flagellomere, third flagellomere longest, apical segment very small, button-shaped. Female antenna (Fig.
Thorax. Cervical sclerites dark brown dorsally, brown laterally, densely dusted with gray. Pronotum dark brown, postero–lateral angle polished rusty brown. Prescutum and presutural scutum densely dusted with orange–brownish gray, with three distinct stripes, medial stripe laterally semi-polished dark brown, divided along middle with densely dusted area, lateral stripe polished black. Tubercular pits missing, pseudosutural fovea black, semi-polished. Postsutural scutum with each lobe black, sparsely dusted. Area between lobes densely covered with pruinosity. Scutellum dark brown, densely dusted, covered with sparse short erect setae. Mediotergite dark brown densely dusted with grayish brown. Pleuron dark brown, dusted with grayish brown. Wing (Fig.
Abdomen. Male abdomen black, semi-polished, dusted with brownish pruinosity, covered with erect sparse whitish setae, longer on sternites, shorter on tergites. Posterior margins of tergites and sternites narrowly grayish. Tergites with two pairs of transverse sutures. Female abdomen dark brown, coloration of sternites slightly varies individually from brown to dark brown, in some females basal sternites pale brown, in some seventh sternite pale brown to yellowish brown. Male terminalia (Figs
300–500 m.
From beginning of June through to mid–August.
Mountainous medium–sized rivers with sandy or fine gravel covered margins surrounded by dense mixed forests (Fig.
South Korea.
There are a few black Eriocera species with cell m1 recorded from territories close to the Korean Peninsula, but some of them have unknown males. Among those with males described, the male terminalia are usually unstudied and separation of them is mostly based on external features such as coloration or comparative length of separate structures. Hexatoma aequinigra Alexander, 1934b is known only from the female, the size and general appearance of which is similar to that of H. pianigra sp. nov., but the species can be easily separated by leg coloration, the femur of H. pianigra sp. nov. is black with a narrowly brownish base, while that of H. aequinigra is yellow with only the tip blackened. Hexatoma atripes Alexander, 1934b is also described from the female only, the measurements of which are also close to H. pianigra sp. nov., but the halter has a yellow stem and blackened knob, while the halter of H. pianigra sp. nov. is entirely black. The male of H. issikii (Alexander, 1928) is somewhat larger than H. pianigra sp. nov., it has a yellow mesonotum, a pleuron with a broad stripe and a yellow halter with only the knob blackened. All these structures are completely black in H. pianigra sp. nov. Hexatoma lygropis (Alexander, 1920) is a somewhat larger species with a velvety black body, H. pianigra sp. nov. is semi-polished with a sparse cover of pruinosity. Hexatoma nigrotrochanterata (Alexander, 1932) is similar in size to H. pianigra sp. nov., but both species can be easily separated based on leg coloration. The femur of H. pianigra sp. nov. is black, while that of H. nigrotrochanterata is yellow with only the apical part blackened. Hexatoma pieliana Alexander, 1940 is described from the female with the male unknown, but it can be easily separated from H. pianigra sp. nov., because it has yellow legs and orange yellow abdominal sternites. Males of H. imperator Alexander, 1953b, H. jozana (
(Fig.
Large crane fly with body length 16.0–23.5 mm. Body dark brown, densely dusted with gray. Male antenna ~ 3× as long as the whole body. Prescutum and presutural scutum with four stripes. Wing clear with dark stigma. Cell m1 present. Abdominal tergites dark brown, lateral margins brownish orange. Male gonocoxite nearly 3× as long as width at base. Inner gonostylus nearly parallel-sided, distal part smoothly narrows to blunt apex. Aedeagus simple, short, straight. Paramere with long narrow branches widely separate at base. Ovipositor brownish orange.
Species is named after type locality, the Seren mountains.
Body coloration dark brown densely dusted with gray pruinosity. Body length of male 16.0–16.5 mm, female 23.5 mm, wing length of male 19.0–20.5 mm, female 20.0 mm.
Head. Gray due to dense pruinosity, densely covered with long yellowish erect setae in male, pubescence less dense and distinctly shorter in female. Vertical tubercle large, rounded, dark brown, densely covered with long yellowish erect setae. Area between bases of antennae and tubercle brownish yellow. Eyes widely separated in both sexes, distance between them at base of antennae equals length of scape and pedicel taken together. Male antenna 7-segmented, 50.0–50.3 mm long, ~ 3× as long as entire body (Fig.
Thorax. Cervical sclerites dark brown, densely dusted with gray dorsally. Pronotum gray, lateral angle brownish orange-yellow. Prescutum and presutural scutum brownish gray, pale gray laterally with four distinct dark brown stripes (Fig.
Abdomen. Tergites dark brown with dense cover of gray pruinosity, dusting less intense along middle and posterior margin, covered with long erect yellowish setae. Basal tergite with brownish orange frontal margin, remaining tergites narrowly gray along posterior margin. Lateral margins brownish orange. Second tergite with two distinct pairs of transverse sutures, remaining tergites with second pair less distinct. Sternites dark brown dusted with gray, narrowly orange laterally, each with paired transverse suture at base. Male terminalia (Fig.
Hexatoma (Eriocera) serenensis Podenas, sp. nov. 40 aedeagal complex, dorsal view, paratype 41 holotype, male, dorsal view 42 head and thorax, dorsal view, holotype, male 43 male wing, paratype 44 male genitalia, dorsal view, paratype 45 female, lateral view, paratype 46 ovipositor, lateral view, paratype. Scale bars: 0.1 mm (40); 1.0 mm (41–46).
Slightly above 600 m.
Mid-July.
Unknown.
North Korea, Seren Mountains.
Hexatoma serenensis sp. nov. is most similar to H. superba Savchenko, 1976, which is described and known only from Kunashir Island, Russia (Table
Comparison of Hexatoma aequinigra, H. superba, and H. serenensis sp. nov.
Character | H. aequinigra | H. superba | H. serenensis sp. nov. |
---|---|---|---|
Pubescence of female head | dense and long | scarce and medium-long | medium-long |
Vertical tubercle | reddish on either side of midline of vertical tubercle | same color as rest of the head, brownish gray | dark brown, rest of the head gray |
Male antenna | – | slightly more than twice as long as body | 3.5× as long as body |
Pubescence of thorax | dense and long | scarce and medium-long | dense and long |
Area separating medial prescutal stripes | slightly wider than the stripes | distinctly narrower than stripes | as wide as stripes |
Lateral margin of prescutum | suffused with pale brown | with additional dark spot | uniformly pale gray |
Femur | yellow, tip narrowly blackened | basal 1/3 brownish yellow | basal 1/2 brownish yellow |
Cell m1 | as long as its stem | as long as 1/2 of its stem | as long as its stem |
Abdominal tergites | female dark brown with yellowish brown lateral margin | male uniformly dark brown | male dark brown with brownish orange lateral margin |
Female body length (mm) | 33 | 26–27 | 23.5 |
Macrotrichiae on distal wing veins | abundant | missing | missing |
Hexatoma (Eriocera) stackelbergi
Paratype
, male (antenna, fore leg, and both wings slide-mounted), Russia, E. Siberia, Ussuri, Tigrowaja, Suchan distr., 43°15.00'N, 133°00.00'E, [alt. 250 m], 11 June1927, Stackelberg leg. (
(Fig.
Body
coloration brownish gray (Fig.
Head. Gray, Brownish gray anteriorly, pale gray posteriorly, covered with short whitish erect setae. Vertical tubercle large, divided by medial grove longitudinally. Eyes widely separated, distance between them at base of antennae approximately equals to length of scape. Basal segments of antenna brown. Scape elongate, 1.6× longer than wide, 2.7× as long as pedicel, darker at base, dusted with gray. Pedicel short, subglobular. Basal flagellomere 1.6× longer than scape, nearly cylindrical, second flagellomere 0.6× as long as basal. Flagellomeres covered with short semi–erect whitish pubescence. Rostrum very short, dark brown, palpus black, labella pale brown.
Thorax. Cervical sclerites pale brown, dusted with gray. Pronotum much wider than long, yellowish brown with gray dusting. Prescutum and presutural scutum gray with three darker brown longitudinal stripes and covered with sparse short whitish setae. Medial stripe separated longitudinally by narrow darker line. Tubercular pits missing, pseudosutural fovea small, pale brown. Postsutural scutum with each lobe gray with indistinct darker brownish spots anteriorly, at middle and posteriorly. Scutellum brownish or yellowish gray with longer yellowish setae along posterior margin. Mediotergite brownish gray, more brownish posteriorly. Pleuron gray with brownish spots where gray pruinosity scarcer. Laterotergite with dense long yellowish setae posteriorly. Wing (Fig.
Abdomen. Tergites brown, dusted with gray, with one pair of transverse sutures, covered with short yellowish setae. Sternites yellowish brown. Male hypopygium large, black, epandrium wider than long, polished black (
Above 1800 m in Korea, ca. 250 m in Russia.
Late August in Korea, middle of June in the Far East of Russia.
Unknown in Korea. Larvae of this species develop in the bottom gravel of large and medium size rivers. Last instar larvae and pupae can be found in the riparian zone, usually in gravel, sand or under stones in Mongolia (
Far East of Russia and Mongolia. Recorded on the Korean Peninsula for the first time.
Hexatoma stackelbergi was known only from three type specimens, all males, listed in
Hexatoma (Eriocera) ussuriensis
Paratypes: Russia, male (antenna, fore leg, wing, and genitalia slide–mounted), E. Siberia, Ussuri, Bikin, river Bikin, 8 July 1927, Martynov leg. (
(Fig.
Body
coloration gray to dark brown (Fig.
Head. Brownish gray, narrowly pale gray along eye margin, covered with whitish erect setae. Vertical tubercle very large, rounded, brown, dusted with grayish yellow postero-dorsally, without polished black summit mentioned in original description (
Thorax. Cervical sclerites brown, dusted with gray. Pronotum much wider than long, dark brown with gray dusting. Prescutum and presutural scutum gray with four distinct dark brown longitudinal stripes and covered with comparatively sparse long whitish setae. Area separating medial stripes slightly narrower than the stripes themselves. Tubercular pits missing, pseudosutural fovea small, polished–brown. Postsutural scutum with each lobe gray with dark brown central area, which also dusted with gray. Area between lobes dark brown anteriorly, pale posteriorly. Scutellum gray with pale fronto-lateral angle. Mediotergite brown, dusted with gray. Pleuron gray, posterior margin of anepimeron with dense long yellowish setae. Wing (Fig.
Abdomen. Two basal tergites brown, remaining dark brown with narrowly brownish yellow lateral margins. Three basal sternites grayish brown, remaining getting darker towards apex, lateral margins brownish to grayish yellow. Abdominal segments covered with long whitish setae, that are denser laterally. Male terminalia (Figs
Ca. 20 m in Korea. 30 m to 650 m in Russia, Japan, and Mongolia (
Beginning of August in Korea. June–July in Russia, Japan, and Mongolia (
Unknown in Korea. Shores of different types of running waters, from shores of large- and medium-sized rivers on plains to medium-sized and small rivers in the foothills in boreal forest, mixed forest, forest-steppe, and steppe landscape zones in Russia and Mongolia (
Eastern part of Russia, Mongolia and Hokkaido Island, Japan. Recorded on the Korean Peninsula for the first time.
Paratype
, male (pinned, wing slide–mounted), China, Formosa [Taiwan], Koshun, 25 April – 25 May 1918, J. Sonan, K. Miyake, M. Yoshino leg. (
China, 2 males, 1 female (pinned), Formosa [Taiwan], Koshun, 25 April – 25 May 1918, J. Sonan, K. Miyake, M. Yoshino leg. (
The first record of this species from Korea is that of
Crane flies belonging to the genus Hexatoma Latreille, 1809 (Diptera, Limoniidae) are very diverse in the Eastern Palaearctic, the fauna of which includes 72 species (
Hexatoma (Eriocera) lygropis (Alexander, 1920) 54 male, dorsal view 55 erroneous image of H. lygropis on which were based all Korean records 56 male wing 57 male genitalia, dorsal view 58 male genitalia with ninth segment removed, dorsal view. Scale bars: 1.0 mm (54, 56); 0.5 mm (57, 58). (55 after
Sampling localities of Korean Hexatoma (Eriocera) 59 H. (E.) gifuensis Alexander, 1933 60 H. (E.) ilwola sp. nov. 61 H. (E.) masakii Alexander, 1934 62 H. (E.) pernigrina Alexander, 1938 63 H. (E.) pianigra sp. nov. 64 H. (E.) serenensis sp. nov. 65 H. (E.) stackelbergi Alexander, 1933 66 H. (E.) ussuriensis Alexander, 1934.
Our warmest thanks to all Korean friends and colleagues who helped us during our visits to South Korea and all those who helped to collect crane flies. We are very grateful to Professor Y.J. Bae for the specimens from Korea University, Seoul, the Republic of Korea; to J.C. Thomas for help with Korean specimens from the University of Kansas, U. S. A.; to Dr F. Shockley and Dr T. Dikow (
This work was supported by a grant from the National Institute of Biological Resources (