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Citation: Shi X, Zhao Y, Shih C, Ren D (2014) Two new species of Archaeohelorus (Hymenoptera, Proctotrupoidea, Heloridae) from the Middle Jurassic of China. ZooKeys 369: 49–59. doi: 10.3897/zookeys.369.6561
Two new fossil species, Archaeohelorus polyneurus sp. n. and A. tensus sp. n., assigned to the genus Archaeohelorus Shih, Feng & Ren, 2011 of Heloridae (Hymenoptera), are reported from the late Middle Jurassic, Jiulongshan Formation of Inner Mongolia, China. Based on the well-preserved forewings and hind wings of these specimens, the diagnosis of the Archaeohelorus is emended: forewing 2cu-a intersecting Cu and Rs+M at the same point or postfurcal, and hind wing may have tubular veins C, Sc+R, R, Rs, M+Cu, M and Cu distinct, or simplified venation. The new findings also elucidate the evolutionary trend of forewing and hind wing venation and body size for the Heloridae from the late Middle Jurassic to now.
Fossil wasps, Heloridae, Archaeohelorus, Middle Jurassic, China
Proctotrupoidea Latreille, 1802, including 11 extant families, is a significant group within Hymenoptera for their long evolutionary history, special morphology and diversity (
Up to date, fossil Heloridae contains 8 genera and 12 species, which have been summarized by
So far, 7 species of Heloridae from China have been reported, including Archaeohelorus hoi Shih, Feng & Ren, 2011; Gurvanhelorus beipiaoensis Shi, Shih & Ren, 2012; Protocyrtus validus Zhang & Zhang, 2001; Sinohelorus elegans Shi, Shih & Ren, 2012; Spherogaster coronata Zhang & Zhang, 2001; and Spherogaster saltatrix Shi, Shih & Ren, 2012.
Extant helorids contains only one genus, Helorus Latreille, 1802, with 12 known valid species mostly in the Holarctic Region. They are parasitoids of larvae of chrysopid lacewings (Neuroptera: Chrysopidae: Chrysopinae: Chrysopa species) (
Recently, we collected two well-preserved fossil specimens referable to Heloridae from the late Middle Jurassic Jiulongshan Formation at Daohugou Village, Ningcheng County, Inner Mongolia, China. Based on these new findings, the diagnosis of Archaeohelorus Shih, Feng & Ren, 2011 is emended and two new species, Archaeohelorus polyneurus sp. n. and Archaeohelorus tensus sp. n., are described. This is the second report of Heloridae in the late Middle Jurassic.
The age of the Daohugou fossil-bearing beds is interpreted to be late Middle Jurassic (ca 165 Ma;
All the materials have been collected near Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China; the late Middle Jurassic (Bathonian-Callovian boundary, 165 Ma). All fossil specimens are housed in the Key Lab of Insect Evolution & Environmental Changes, College of Life Sciences, Capital Normal University, Beijing, China.
The specimens were examined dry or under alcohol using a M165 C dissecting microscope (Leica) and are illustrated with the aid of a drawing tube attachment. The figures were drawn by Adobe Photoshop CS5 and CorelDraw 12.0. Morphological terminology and the system used here follow those of
Archaeohelorus hoi Shih, Feng & Ren, 2011
Forewing 2cu-a intersecting Cu and Rs+M at the same point or postfurcal. Hind wing may have tubular veins C and Sc+R separated at base, R developed, M+Cu robust and forking at the basal part, M and Cu distinct, or simplified venation.
Archaeohelorus hoi Shih, Feng & Ren, 2011, Archaeohelorus polyneurus sp. n. and Archaeohelorus tensus sp. n.
This genus was established by
http://zoobank.org/BFECD83C-516A-4F08-BBB3-72A5EB6DFF41
http://species-id.net/wiki/Archaeohelorus_polyneurus
Figures 1, 2The specific nameis from Greek word “polyneurus”, means “many veins”, referring to the complete venation of the hind wing preserved.
Holotype, CNU-HYM-NN2012052, dorsal view, gender unknown. A well-preserved body with almost complete forewings and hind wings and part of legs, but head missing. Paratype: CNU-HY-NN2008010, dorsal view, a well-preserved almost complete body with forewings and right hind wing and part of legs, previously mis-identified as a paratype of Archaeohelorus hoi.
Jiulongshan Formation, Middle Jurassic, Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China.
In forewing, 2cu-a intersecting Cu and Rs+M at the same point and cell r obtuse-angled triangular (vs. approximately right-angled triangular in Archaeohelorus hoi). Hind wing with Sc+R confluent with C and extended to costal margin, R developed. M+Cu distinct and forking at the basal part, M and Cu robust.
Upon further examination, we found that a paratype of Archaeohelorus hoi, CNU-HYM-NN2008010, has forewing 2cu-a intersecting Cu and Rs+M at the same point and cell r obtuse-angled triangular (vs. approximately right-angled triangular in Archaeohelorus hoi); and hind wing with tubular vein C parallel with Sc+R at base, M+Cu robust and forking at the basal section, M long, 1-Cu, 2-Cu distinct. These venational characters are consistent with the diagnostic characters of Archaeohelorus polyneurus sp. n., hence we transfer CNU-HYM-NN2008010 as a paratype of Archaeohelorus polyneurus sp. n.
A medium-sized body with both forewings and hind wings well-preserved, but, without head (Fig. 1). Forewing broad. Mesosoma suboval, nearly 1.35 times as long as wide; mesoscutum trapezoidal with notauli distinct and concave; tegula triangular; scutellum broad with two rows of pits; metanotum relatively wide with plenty of pits; propodeum transverse, 2.89 times as broad as long.
Holotype of Archaeohelorus polyneurus sp. n. CNU-HYM-NN-2012052. A Photo B line drawing C line drawing of forewing D line drawing of hind wing. Scale bars: 1 mm. (Online figure in color.)
Metasoma suboval with six segments; first metasomal segment transverse, second segment with several longitudinal ridges; third and fourth segments trapezoidal with shorter longitudinal ridges anteriorly; other ones smooth; the end of terminal segment not preserved.
Forewing broad and subtriangular. Pterostigma long and acute apically, not widened beyond mid-length. C robust and extending to apex of forewing. 2r-rs arising from basal one third of pterostigma, slightly oblique apicad, slightly longer than pterostigmal width. R robust, cell r obtuse-angled triangular and closed by R, M+Cu, 1-Rs and 1-M; Rs straight and intersecting with distal part of C. Cell 1+2r six-sided and surrounded by R, 1-Rs, 1-Rs+M, 2-Rs+M, 2-Rs, and 2r-rs. M+Cu straight and distinct; M and Cu distinct, M straight, Cu break after 1m-cu. 1-Rs as long as 1-M and slightly inclined toward wing base. Cell 1mcu small and subtriangular. M and Rs branching at 30% from 1m-cu of the length between 1m-cu and 2r-rs. 1cu-a and 2cu-a distinct and reaching A. 1cu-a in line with 1-M; and 2cu-a intersecting Cu and Rs+M at the same point.
Hind wing with tubular veins C parallel with Sc+R at base; R developed and Rs preserved; M+Cu robust and forking at the basal section, M and 1-Cu long and distinct, 2-Cu long.
Right foreleg with only coxa and femur partially preserved. Right midleg with partial trochanter, spindle-shaped femur and partial tibia preserved. Right hindleg with partial coxa, trochanter trapezoidal and small, robust spindle-shaped femur, long tibia swollen distally with spurs, tarsi with five segments, basitarsus longest and two claws fixing the end of pretarsus.
Measurements (in mm). Mesosoma length 3.36, width 2.48; metasoma > 3.38 long; lengths of the first to fifth metasomal segments are 0.34, 0.51, 0.72, 0.94, and 0.55; forewing length 6.52, width > 3.05.
Body medium-sized (Fig. 2). Head oval and antennae filiform and thick, scape swollen and bell-shaped, only first and second flagellomeres preserved. Mesosoma, forewing and metasoma same as holotype, the end of terminal segment not preserved. Hind wing with tubular vein C parallel with Sc+R at base, M+Cu robust and forking at the basal section, M long, 1-Cu, 2-Cu distinct. Left foreleg and right midleg with only coxa and femur partially preserved. Left midleg with femur robust and tibia long, left hindleg with femur robust and spindle-shaped, long tibia swollen distally. Right hindleg with femur robust and spindle-shaped, tibia partially preserved.
Paratype of Archaeohelorus polyneurus sp. n. CNU-HYM-NN-2008010. A Photo B line drawing C line drawing of forewing D lLine drawing of hind wing. Scale bars: 1 mm. (Online figure in color.)
Measurements (in mm). Body length > 14.6; head length 1.89, width 1.53; mesosoma length 3.58, width 2.52; metasoma length 3.68; forewing length > 6.54, width > 2.88.
This species can be assigned to Archaeohelorus Shih, Feng & Ren, 2011 by its metasoma with six segments and the first segment narrow and transverse. Forewing 1-Rs as long as 1-M. Cell 1mcu small and subtriangular, 1cu-a in line with 1-M; and 2cu-a intersecting Cu and Rs+M at the same point. Compared with Archaeohelorus hoi, Archaeohelorus polyneurus has preserved hind wing with Sc+R confluent with C and extended to costal margin, R developed. M+Cu distinct and forking at the basal part, M and Cu robust; in forewing, cell r obtuse-angled triangular (vs. approximately right-angled triangular in Archaeohelorus hoi). Besides, Archaeohelorus polyneurus has a much larger body size than Archaeohelorus hoi, length of forewing 6.52 mm or > 6.54 mm (vs. 3.17 mm in Archaeohelorus hoi).
http://zoobank.org/84E4B637-E6E6-4799-929E-1536DA5213A5
http://species-id.net/wiki/Archaeohelorus_tensus
Figure 3The specific name “tensus” means stretching and long, referring to the shape of the forewing.
Holotype, CNU-HYM-NN2012056p/c, part and counterpart, dorsal view, male. A well-preserved almost complete body with antenna, forewings and part of legs and partial hind wings.
Holotype of Archaeohelorus tensus sp. n. CNU-HYM-NN-2012056 p/c, part and counterpart. A Photo of part B photo of counterpart C line drawing of part D line drawing of forewing E terminal segments of counterpart (under alcohol). Scale bars: 1 mm. (Online figure in color.)
Jiulongshan Formation, Middle Jurassic, Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China.
Forewing 2cu-a postfurcal with intersection of Cu and Rs+M. First abscissa of Rs (1-Rs) and basal section of M (1-M) arched toward basal of wing. Hind wing with tubular veins C, Sc+R, and Rs preserved.
A smalladult specimen with a total body length of 4.4 mm. Head suboval and large relative to mesosoma in width. Eyes large, and located at the both sides of the head.
Antennae filiform, thick, with 6 segments preserved on left and 5 segments preserved on right. Scape swollen and bell-shaped; pedicel short and quadrate; subsequent flagellomeres not well preserved.
Mesosoma subhexagonal and broader than head; pronotum narrower than head, short, probably covered by mesonotum; mesoscutum distorted in the middle and trapezoidal with notauli not distinct but preserved; tegula large and subtriangular; scutellum round with pits anteriorly; propodeum narrow, transverse and areolated.
Metasoma suboval with seven segments; first metasomal segment transverse with several longitudinal ridges; second segment trapezoidal with less longitudinal ridges anteriorly; other ones smooth; the male terminalia triangular, partially covered by the previous segment.
Forewing broad and subtriangular. Pterostigma long and acute apically, not widened beyond mid-length. C robust and extending near the apex of forewing. 2r-rs arising from basal one third of pterostigma, slightly longer than the width of pterostigma, and slightly oblique apicad. R robust and cell r closed with C, R and 1-RS. Rs straight and intersecting with distal part of C. Cell 1+2r longer and narrower relatively and six-sided surrounded by R, 1-Rs, 1-Rs+M, 2-Rs+M, 2-Rs, and 2r-rs. M+Cu straight and distinct; M and Cu distinct, almost straight. 1-Rs as long as 1-M and slightly inclined toward wing base. Cell 1mcu small, subtriangular and relatively slender with 2-M+Cu 4.3 times as long as 1-M. M and Rs branching at 30% from 1m-cu of the length between 1m-cu and 2r-rs. 1cu-a and 2cu-a distinct and reaching A. 1cu-a in line with 1-M; and 2cu-a postfurcal with Rs+M distinctly.
Hind wing, with tubular veins C parallel with Sc+R at base, Rs short, intersecting distal part of C.
Left foreleg, midleg and hindleg and right foreleg with only coxa and femur partially preserved, hind femur much thicker, nearly three times as long as wide; right midleg with partial spindle-shaped femur and partial tibia preserved, right hindleg with partial coxa, trapezoidal trochanter, robust spindle-shaped femur and relatively thin tibia and some parts of tarsus preserved.
Measurements (in mm): Body length 4.67, head length1.39, width 0.68, mesosoma length 1.68, width 1.36; metasoma 2.23 long; lengths of metasomal segments are 0.20, 0.39, 0.48, 0.38, 0.25 and 0.32; forewing length > 2.96, width > 1.36.
This species is assigned to Archaeohelorus Shih, Feng & Ren, 2011 by its seven separated metasomal segments, the first metasomal segment transverse with several longitudinal ridges. Forewing 1-Rs as long as 1-M. Cell 1mcu small and subtriangular, 1cu-a in line with 1-M. Compared with Archaeohelorus hoi, Archaeohelorus tensus has preserved hind wing with tubular veins C, Sc+R, R, and Rs preserved; in forewing, 2cu-a distinctly postfurcal with intersection of Cu and Rs+M (vs. 2cu-a intersecting Cu and Rs+M at the same point in Archaeohelorus hoi); cell 1mcu distinctly slender, 4.3 times as long as wide (vs. 2.4 times in Archaeohelorus hoi); 1-Rs and 1-M arched toward basal of wing (vs. 1-Rs and 1-M straight in Archaeohelorus hoi). It also differs from Archaeohelorus polyneurus by its postfurcal 2cu-a, slender cell 1mcu.
In the vast and extensive Daohugou fossil insect collection (>200, 000 insect fossil specimens) at the Capital Normal University, only ten helorids are collected so far: two are Archaeohelorus polyneurus sp. n., one is Archaeohelorus tensus sp. n., and seven specimens are previously described Archaeohelorus hoi Shih, Feng & Ren, 2011. It is likely that helorids might have been a very small group in the Middle Jurassic.
As a relict family, helorids have survived from the Middle Jurassic to now. Among all the fossil specimens, only Archaeohelorus polyneurus has distinct and complicated hind wing venation, which were unknown before. Therefore, the new findings are important supplement to helorid record and suggest that the trend for the morphological evolution of the Heloridae is as follows:
In the Middle Jurassic, Archaeohelorus polyneurus had forewing cell r obtuse-angled triangle, and hind wing venation clear and more complex (C and Sc+R separated at base, M+Cu short and robust, forking at the basal part, M and Cu distinct) and a much larger body (forewing length > 6.52mm). On the other hand, Archaeohelorus hoi and Archaeohelorus tensus had forewing cell r approximately right-angled triangular, and hind with simplified venation (only C and Sc+R and/or R preserved) and smaller body (forewing length ≈ 3 mm). In the Late Jurassic to the Early Cretaceous, Spherogaster coronata Zhang & Zhang, 2001 had hind wing with only C, R and M present, costal area extremely narrow, and a much larger body size with forewing length of 12 mm. In extant helorids, forewing cell r approximately right-angled triangular, and hind wings have tubular veins C and Sc+R and nebulosus veins of M+Cu, M, Cu and A (Goulet and Huber 1993). The body size is small with forewing about 3.3 mm long (
We are sincerely grateful to Qiang Yang and Taiping Gao (College of Life Sciences, Capital Normal University) for their valuable comments and suggestion on the manuscript. This work was supported by the National Basic Research Program of China 973 Program Grant 2012CB821906; National Natural Science Foundation of China Grants, 31230065 and 41272006; Great Wall Scholar and KEY project of Beijing Municipal Commission of Education (grant KZ201310028033), Program for Changjiang Scholars and Innovative Research Team in University.