Four new species of oak gallwasps of the genus
The cynipid gallwasp fauna of the Eastern Palaearctic and Oriental region is poorly known: only 35 valid species are recognised from the Eastern Palaearctic, mostly from Japan and the Russian Far East, and only a few oak gallwasp species have yet been described or mentioned as “cynipid gallwasp” from the Oriental region [
The family Fagaceae in Eastern Asia, including Taiwan, is represented by five taxa which serve as hosts for oak gallwasps (Cynipini):
In the Holarctic, 23 valid species are known from the genus
We describe four new
The material examined was reared from galls on different
We follow the current terminology of morphological structures [
Images of wasp anatomy were produced with a digital Nikon Coolpix 4500 camera attached to a Leica DMLB compound microscope, followed by processing in CombineZP (Alan Hadley; a software package that makes possible the construction of an image with increased depth of field) and Adobe Photoshop 6.0.
The type material is deposited in the following institutions: NMNS, National Museum of Natural Science, Taichung, Taiwan (curator Mei-Ling Chan); PDL, Pest Diagnostic Laboratory (the former Systematic Parasitoid Laboratory, SPL), Tanakajd, Hungary (curator G. Melika); NCHU, collection of the Department of Entomology, National Chung Hsing University, Taichung, Taiwan (curator Chang-Ti Tang); USNM, U.S. National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (curator Matthew Buffington).
TAIWAN, Taitung County, Daren Forest Station, Daren Township, ex
The species named after the peculiar pentagonal shape of the gall.
The four described
Head anteriorly and laterally dark brown to chestnut brown; posteriorly, especially postocciput, postgena and postgenal bridge dark brown to black; labial and maxilly palps yellow; antenna yellow to dirty yellow, especially flagellomeres much darker dorsally; mesosoma chestnut brown to brown, pronotum laterally always lighter; mesoscutellum dark brown to black; metapleuron and central propodeal area always slightly darker than rest of mesosoma; legs uniformly yellow; metasoma dark brown, usually dorsally darker than laterally.
Head alutaceous, with sparse white setae, more dense on lower face, 2.1x broader than long from above; 1.2x broader than high in anterior view and slightly broader than mesosoma. Gena alutaceous, not broadened behind eye, 2.0x narrower than cross-diameter of eye, invisible in anterior view. Malar space alutaceous, with striae radiating from clypeus and reaching eye; height of eye 3.7x longer than length of malar space. Compound eyes slightly converging ventrally. POL 1.4x longer than OOL; OOL 1.5x longer than diameter of lateral ocellus, 1.6x longer than LOL; ocelli ovate, all three equal in length. Transfacial distance 1.1x longer than height of eye and 1.5x longer than height of lower face (distance between antennal rim and ventral margin of clypeus); diameter of antennal torulus 1.5x longer than distance between them, distance between torulus and eye margin only slightly greater than diameter of torulus. Lower face alutaceous, with striae radiating from clypeus and extending nearly to lower level of antennal rim, with elevated median area. Clypeus rectangular, flat, slightly broader than high, alutaceous, with deep anterior tentorial pits, distinct epistomal sulcus, and clypeo-pleurostomal line; ventrally rounded and not incised medially. Frons alutaceous, with deep smooth and shiny impression below median ocellus; vertex and occiput alutaceous with some delicate short irregular striae; interocellar area delicately coriaceous, slightly elevated. Postgena smooth, shiny, postocciput around occipital foramen impressed, smooth, shiny; posterior tentorial pits large, deep, elongate; postgenal bridge at least 2.0x higher than broad, lower part narrowed to a stripe; occipital foramen very slightly higher or equal to height of postgenal bridge, around 1.3x shorter than height of oral foramen. Antenna with 12 distinct flagellomeres (or 13, indistinct suture between F13 and F12 visible in some individuals), longer than head + mesosoma; pedicel 1.7 times as long as broad, F1 1.2x shorter than length of scape + pedicel, 1.15x longer than F2, F2 only slightly longer than F3, F3 = F4, F5–F11 shorter than F4 and nearly equal in length; fused F12 + F13 1.8x longer than F11; placodeal sensilla on F1–F12, in numerous rows.
Mesosoma longer than high in lateral view. Pronotum smooth, shiny; with few white setae and with irregular delicate wrinkles laterally, emarginate along lateral edge, followed by deep longitudinal invagination. Anterior rim of pronotum narrow, emarginate; propleuron alutaceous, with few setae, strongly concave in mediocentral part. Mesoscutum smooth, shiny; nearly as long as broad (width measured across basis of tegulae); notauli complete, deeply impressed for full length; median mesoscutal line absent; anterior parallel and parapsidal lines not impressed, hardly traceable; parascutal carina broad, anteriorly reaching notauli. Transscutal articulation deep, distinct. Mesoscutellum only slightly longer than broad, rectangular, with parallel sides; shorter than mesoscutum, uniformly coriaceous, overhanging metanotum; scutellar foveae transverse ovate, with shiny bottom and few wrinkles on it, divided by central elevated area which only slightly narrower than width of fovea (in some individuals foveae indistinctly separated by central elevated area and form a transverse impression along anterior margin of mesoscutellum). Mesopleuron, including speculum, smooth, shiny, with few white setae and irregular delicate wrinkles which forming a transverse line bent in mid height and going across entire width of mesopleuron; mesopleural triangle shiny, with few white setae and some delicate irregular short wrinkles. Metapleural sulcus reaching mesopleuron slightly above half height; preaxilla delicately coriaceous, shiny; dorsal and lateral axillar areas shiny, smooth, with few setae; axillar carina broad, with longitudinal striae; subaxillular bar narrow, smooth, shiny, in most posterior end slightly higher than height of metanotal trough. Metascutellum uniformly coriaceous, slightly higher than height of smooth, shiny ventral impressed area; metanotal trough smooth, shiny, without setae. Lateral propodeal carinae distinct, lyre-shaped, bent outwards at mid height, central propodeal area shiny, with numerous delicate irregular wrinkles; lateral propodeal area coriaceous, with dense white setae; nucha very short, with delicate longitudinal sulci dorsolaterally and laterally.
Forewing as long or slightly longer than body, with distinct brown veins, margin with long dense cilia; radial cell 3.75x longer than broad, R1 and Rs reaching wing margin; areolet large, triangular, well-delimited by distinct veins; Rs + M well-pigmented, reaching basalis in the lower third.
Tarsal claws simple, without basal lobe. Metasoma slightly shorter than head + mesosoma, slightly longer than high in lateral view, smooth, shiny, without setae; only 2nd metasomal tergites laterally with very few white setae; 2nd tergite extending to 1/3 length of metasoma; prominent part of ventral spine of hypopygium short, nearly 2.0x longer than broad in ventral view, with long white setae, extending beyond apex of spine but never forming a tuft. Body length 2.3–2.6 mm (
The colour pattern of the body the same as in female but usually much darker to black. Similar to female but compound eyes bigger and thus frons and lower face narrower than in female; also ocelli much bigger. Antenna longer than body, with 13 flagellomeres, proximal half of F1 broader than basal half, curved and excavated. Body length 2.1–2.4 mm (
The gall closely resembles the asexual galls of three Western Palaearctic
The sexual generation galls
Taiwan, Taitung County (Figure
TAIWAN, Taitung County, Dawu Township, ex
The species named after the triangular-pyramid shape of the gall.
Most closely resembles
Head uniformly dark brown to chestnut brown; with slightly lighter clypeus and lower face; labial and maxilly palps yellow; antenna brown, especially flagellomeres much darker dorsally; mesosoma chestnut brown to brown, tegula always lighter; mesoscutellum dark brown to black; legs uniformly yellow, except dark brown hind tibiae and tarsi; metasoma dark brown, usually dorsally darker than laterally.
Head alutaceous, with sparse white setae, more dense on lower face, 1.9x broader than long from above; 1.1x broader than high in anterior view and slightly broader than mesosoma. Gena alutaceous, not broadened behind eye, 2.6x narrower than cross-diameter of eye, invisible in anterior view. Malar space alutaceous, with striae radiating from clypeus and reaching eye; height of eye 3.5x longer as length of malar space; compound eyes slightly converging ventrally. POL 1.8x longer than OOL; OOL 1.1x longer than diameter of lateral ocellus, 1.3x longer than LOL; ocelli ovate, all three equal in length. Transfacial distance as long as height of eye and 1.4x longer than height of lower face (distance between antennal rim and ventral margin of clypeus); diameter of antennal torulus 1.4x longer than distance between them, distance between torulus and eye margin equal to diameter of torulus. Lower face with numerous delicate striae radiating from clypeus and extending to lower level of antennal rim and into area between eye and torulus; median area slightly elevated. Clypeus rectangular, flat, slightly higher than broad, alutaceous, with deep anterior tentorial pits, distinct epistomal sulcus and clypeo-pleurostomal line; ventrally rounded and not incised medially. Frons alutaceous; vertex and occiput alutaceous with some delicate short irregular striae; interocellar area delicately coriaceous, elevated. Postgena smooth, shiny, postocciput around occipital foramen impressed, smooth, shiny with parallel interrupted delicate numerous striae; posterior tentorial pits large, deep, elongate; lower half of postgenal bridge narrowed to a stripe; occipital foramen slightly higher or as high as height of postgenal bridge, around 1.3x shorter than height of oral foramen. Antenna with 12 distinct flagellomeres (or 13, indistinct suture between F12 and F13 visible in some individuals), longer than head + mesosoma; pedicel 1.3x longer than broad; scape + pedicel 1.2x longer than F1; F1 1.3x longer than F2, F2 only slightly longer than F3, F3 = F4, F5–F11 shorter than F4 and nearly equal in length; fused F12 + F13 1.9x longer than F11; placodeal sensilla on F1–F12, in numerous rows.
Mesosoma longer than high in lateral view. Pronotum smooth, shiny; with few white setae and with irregular delicate wrinkles laterally, emarginate along lateral edge, followed by deep longitudinal invagination. Anterior rim of pronotum narrow, emarginate; propleuron alutaceous, with few setae, strongly concave in mediocentral part. Mesoscutum smooth, shiny; nearly as long as broad (width measured across basis of tegulae); notauli complete, deeply impressed for full length; median mesoscutal line absent; anterior parallel and parapsidal lines not impressed, hardly traceable; parascutal carina broad, anteriorly reaching notauli. Transscutal articulation deep, distinct. Mesoscutellum only slightly longer than broad, rectangular, with parallel sides; shorter than mesoscutum, uniformly coriaceous, overhanging metanotum; scutellar foveae transverse ovate, with shiny bottom and few wrinkles on it, divided by central narrow elevated area (in some individuals foveae indistinctly separated by central elevated area and form a transverse impression along anterior margin of mesoscutellum). Mesopleuron smooth, shiny, with few white setae and irregular delicate wrinkles which forming a transverse line bent in the middle and going across entire width of mesopleuron, as well as in dorsoposterior one quarter of mesopleuron; mesopleural triangle shiny, with few white setae and some delicate irregular short wrinkles. Metapleural sulcus reaching mesopleuron slightly above half height; preaxilla delicately coriaceous, shiny; dorsal and lateral axillar areas shiny, smooth, with few setae; axillar carina broad, with longitudinal striae; subaxillular bar narrow, smooth, shiny, in most posterior end slightly higher than height of metanotal trough. Metascutellum uniformly coriaceous, slightly higher than height of smooth, shiny ventral impressed area; metanotal trough smooth, shiny, without setae. Lateral propodeal carinae distinct, lyre-shaped, bent outwards at mid height, central propodeal area smooth, shiny, with numerous delicate irregular wrinkles; lateral propodeal area coriaceous, with dense white setae; nucha very short, with delicate longitudinal sulci dorsolaterally and laterally.
Forewing as long or slightly longer than body, with distinct brown veins, margin with long dense cilia; radial cell 4.05x longer than broad, R1 and Rs reaching wing margin; areolet large, triangular, well-delimited by distinct veins; Rs + M well-pigmented, reaching basalis in the lower third.
Tarsal claws simple, without basal lobe. Metasoma nearly as long as length of head + mesosoma, longer than high in lateral view, smooth, shiny, without setae; only 2nd metasomal tergites laterally with very few white setae; 2nd tergite extending to 1/3 length of metasoma; prominent part of ventral spine of hypopygium short, nearly 2.0x longer than broad in ventral view, with long white setae, extending beyond apex of spine but never forming a tuft. Body length 2.1–2.4 mm (
The colour pattern of the body the same as in female but usually much darker to black. Similar to female but compound eyes bigger and thus frons and lower face narrower than in female; also ocelli much bigger. Antenna longer than body, with 13 flagellomeres, proximal half of F1 broader than basal half, curved and excavated; scape, pedicel and F1 yellow, much lighter than all subsequent flagellomeres; pedicel extremely long, nearly as long as length of scape. Body length 1.8–2.1 mm (
The gall closely resembles that of herein described
Only the sexual generation is known, inducing galls on
Taiwan, Taitung County (Figure
TAIWAN, Taitung County, Daren Forest Station, Daren Township, ex
The species named after the host plant,
Most closely related to
Head and antennae brown to light brown; posteriorly, especially postocciput, postgena and postgenal bridge usually darker; labial and maxilly palps yellow to dirty yellow; compound eyes black; mesoscutum, mesopleuron, metapleuron and propodeum dark brown, while pronotum, mesoscutellum always lighter; legs uniformly yellow, except darker tarsal claws and hind tibiae; metasoma brown, dorsally usually darker.
Head alutaceous, with sparse white setae, more dense on lower face, shiny; 2.35x broader than long from above; 1.3x broader than high in anterior view and slightly broader than mesosoma. Gena alutaceous, shiny, not or very slightly broadened behind eye, 2.0x narrower than cross-diameter of eye, invisible in anterior view. Malar space alutaceous, with striae radiating from clypeus and reaching eye; height of eye 3.5x longer than length of malar space. Compound eyes converging ventrally. POL 1.35x longer than OOL; OOL 1.7x longer than diameter of lateral ocellus, 1.8x longer than LOL; ocelli ovate, all three equal in length. Transfacial distance 1.2x longer than height of eye and 1.4x longer than height of lower face (distance between antennal rim and ventral margin of clypeus); diameter of antennal torulus nearly equal to distance between them, distance between torulus and eye margin 1.4x longer than diameter of torulus. Lower face alutaceous, with numerous delicate striae radiating from clypeus and extending to lower level of antennal rim, with elevated smooth, shiny median area. Clypeus rectangular, flat, slightly broader than high, smooth, shiny, with small anterior tentorial pits, indistinct epistomal sulcus and clypeo-pleurostomal line; ventrally rounded, narrowly emarginate and not incised medially. Frons, vertex and occiput smooth, shiny; interocellar area delicately coriaceous, slightly elevated. Postgena smooth, shiny, postocciput around occipital foramen impressed, smooth, shiny; posterior tentorial pits small, rounded; postgenal bridge short, triangular-like, continuously narrowed down to emarginate hypostomal carina; occipital foramen nearly 1.5x higher than height of postgenal bridge, around 1.4x shorter than height of oral foramen. Antenna with 12 or 13 flagellomeres (in some individuals suture between F12 and F13 indistinct), longer than head + mesosoma; pedicel 1.4x longer than broad, F1 slightly shorter than scape + pedicel, 1.15x longer than F2, F2 1.3x longer than F3, F3 slightly longer than F4, F5–F11 shorter than F4 and nearly equal in length; fused F12 + F13 2.0x longer than F11 (if suture between F12 and F13 well-developed, than F13 slightly longer than F12); placodeal sensilla on F1–F13, in numerous rows.
Mesosoma only very slightly longer or nearly as long as high in lateral view, not compressed dorsoventrally. Pronotum smooth, shiny; with few white setae and with irregular delicate wrinkles laterally, emarginate along lateral edge, followed by deep longitudinal invagination. Anterior rim of pronotum narrow, emarginate; propleuron alutaceous, with few setae, strongly concave in mediocentral part. Mesoscutum smooth, shiny; nearly as long as broad (width measured across basis of tegulae); notauli complete, deeply impressed for full length; median mesoscutal line absent; anterior parallel impressed, distinct, extending to 1/3 of mesoscutum length; parapsidal lines not impressed, hardly traceable; parascutal carina broad, anteriorly reaching notauli. Transscutal articulation deep, distinct. Mesoscutellum only slightly longer than broad, rectangular, with parallel sides; shorter than mesoscutum, uniformly coriaceous, overhanging metanotum; scutellar foveae transversely ovate, with shiny bottom and few wrinkles on it, divided by central narrow elevated area. Mesopleuron and speculum uniformly smooth and shiny, without transversely orientated striae at mid height; mesopleural triangle shiny, with few white setae and some delicate irregular short wrinkles. Metapleural sulcus reaching mesopleuron slightly above half height; preaxilla delicately coriaceous, shiny; dorsal and lateral axillar areas shiny, smooth, with few setae; axillar carina broad, with longitudinal striae; subaxillular bar narrow, smooth, shiny, in most posterior end slightly higher than height of metanotal trough. Metascutellum uniformly coriaceous, slightly higher than height of smooth, shiny ventral impressed area; metanotal trough smooth, shiny, without setae. Lateral propodeal carinae distinct, lyre-shaped, bent outwards at mid height, central propodeal area smooth, shiny, with numerous delicate irregular wrinkles; lateral propodeal area coriaceous, with dense white setae; nucha very short, with delicate longitudinal sulci dorsolaterally and laterally.
Forewing as long or slightly longer than body, with distinct brown veins, margin with long dense cilia; radial cell 4.0x longer than broad, R1 and Rs reaching wing margin; areolet large, triangular, well-delimited by distinct veins; Rs + M well-pigmented, reaching basalis in the lower third.
Tarsal claws simple, without basal lobe. Metasoma slightly shorter than head + mesosoma, slightly higher than long in lateral view, smooth, shiny, without setae; only 2nd metasomal tergites laterally with very few white setae; 2nd tergite extending to 1/3 length of metasoma; prominent part of ventral spine of hypopygium short, nearly 2.0x or less longer than broad in ventral view, with long white setae, extending beyond apex of spine but never forming a tuft. Body length 2.7–3.1 mm (
Lower face brown, while rest of head dark brown to black; mesosoma, dark brown to black, except always light brown pronotum (in some specimens head and mesosoma entirely brown); legs yellow; metasoma yellowish brown, dorsally and dorsolaterally much darker. Similar to female but compound eyes bigger and thus frons and lower face narrower than in female; also ocelli much bigger. Antenna longer than body, with 13 flagellomeres, proximal 1/3 of F1 broader than basal half, curved and excavated. Body length 2.1–2.4 mm (
Galls develop from leaf buds or on leaves. The body of the gall located on a long, thin stalk, 10–20 mm long, growing out from the bud or forming within the leaf petiole. The body of the gall rounded, spherical, 7.0–8.3 mm in diameter. The gall is green with the surface smooth, shiny, mottled with some whitish irregular spots. The parenchyma of the gall is green, juicy, with a centrally located larval chamber, 3.0–3.5 mm in length and diameter usually half of the length. Unilocular.
Only the sexual generation is known from galls on
Taiwan, Taitung and Pingtung Counties (Figure
Current distribution of
TAIWAN, Taoyuan County, Fushing Township, ex
The species named after the shape of the gall which is reminiscent of mammalian testicles.
See diagnosis to
The colour pattern as in the previous species,
Mesosoma only very slightly longer or nearly as long as high in lateral view, not compressed dorsoventrally. Pronotum smooth, shiny; with few white setae and with irregular delicate wrinkles laterally, emarginate along lateral edge, followed by deep longitudinal invagination. Anterior rim of pronotum narrow, emarginate; propleuron alutaceous, with few setae, strongly concave in mediocentral part. Mesoscutum smooth, shiny; nearly as long as broad (width measured across basis of tegulae); notauli complete, deeply impressed for full length; median mesoscutal line absent; anterior parallel and parapsidal lines not impressed, hardly traceable; parascutal carina broad, anteriorly reaching notauli. Transscutal articulation deep, distinct. Mesoscutellum slightly longer than broad, rectangular, with parallel sides; shorter than mesoscutum, uniformly coriaceous, overhanging metanotum; scutellar foveae transverse ovate, with shiny bottom and few wrinkles on it, divided by central narrow sharp carina. Mesopleuron and speculum uniformly smooth and shiny, without transversely orientated striae at mid height; mesopleural triangle shiny, with few white setae and some delicate irregular short wrinkles. Metapleural sulcus reaching mesopleuron slightly above half height; preaxilla delicately coriaceous, shiny; dorsal and lateral axillar areas shiny, smooth, with few setae; axillar carina broad, with longitudinal striae; subaxillular bar narrow, smooth, shiny, in most posterior end slightly higher than height of metanotal trough. Metascutellum uniformly coriaceous, slightly higher than height of smooth, shiny ventral impressed area; metanotal trough smooth, shiny, without setae. Lateral propodeal carinae distinct, lyre-shaped, bent outwards at mid height, central propodeal area smooth, shiny, with numerous delicate irregular wrinkles; lateral propodeal area coriaceous, with dense white setae; nucha very short, with delicate longitudinal sulci dorsolaterally and laterally.
Forewing as long or slightly longer than body, with distinct brown veins, margin with long dense cilia; radial cell 4.0x longer than broad, R1 and Rs reaching wing margin; areolet large, triangular, well-delimited by distinct veins; Rs + M well-pigmented, reaching basalis in the lower third.
Tarsal claws simple, without basal lobe. Metasoma slightly shorter than head + mesosoma, slightly longer than high in lateral view, smooth, shiny, without setae; only 2nd metasomal tergites laterally with very few white setae; 2nd tergite extending to 1/3 length of metasoma; prominent part of ventral spine of hypopygium short, nearly 2.0x longer than broad in ventral view, with long white setae, extending beyond apex of spine but never forming a tuft. Body length 3.1–3.3 mm (
The colour pattern of the body the same as in female but usually much darker to black. The same as female but compound eyes bigger and thus frons and lower face narrower than in female; also ocelli much bigger. Antenna longer than body, with 13 flagellomeres, proximal half of F1 broader than basal half, curved and excavated. Body length 3.1 mm (
The general appearance and the location of the gall closely resembles that of
Only the sexual generation is known from galls on
Taiwan, Taoyuan County (Figure
Currently, 23 species are known to be valid in the Holarctic genus
The genus
Recent analyses have indicated that even the Western Palaearctic
Key to the
(1) | – Female | (2) |
– Male | (5) | |
(2) | – Mesosoma distinctly longer than high in lateral view, compressed dorsoventrally; mesopleuron with distinct transversely orientated numerous delicate striae in mid height, forming distinct bent across entire length of mesopleuron (Figures | (3) |
– Mesosoma nearly as long as high in lateral view, not compressed dorsoventrally; mesopleuron and speculum uniformly smooth and shiny, without transversely orientated striae at mid height (Figures | (4) | |
(3) | – POL 1.4x longer than OOL (Figure | |
– POL 1.8x longer than OOL (Figure | ||
(4) | – Height of eye 3.5x longer than length of malar space; POL longer than OOL; OOL 1.7x longer than diameter of lateral ocellus, 1.8x longer than LOL (Figures | |
– Height of eye 2.25x longer than length of malar space; POL shorter than OOL; OOL 3.0x longer than diameter of lateral ocellus, 2.7x longer than LOL (Figures | ||
(5) | – POL at least 3.0x longer than OOL; ocelli large, at least 3.0x larger than length of OOL (Figures | (6) |
– POL slightly longer than OOL; ocelli smaller, length nearly equal to length of OOL (Figures | (7) | |
(6) | – Height of compound eye 8.1x higher than length of malar space and 1.2x higher than length of transfacial distance (Figure | |
– Height of compound eye 8.7x higher than length of malar space and 1.4x higher than length of transfacial distance (Figure | ||
(7) | – OOL 1.6x longer than length of lateral ocellus; interocellar area alutaceous, like frons (Figure | |