Corresponding author: Dong Ren (
Academic editor: D. Shcherbakov
Two new species of the genus
Today, the
Currently, 34 fossil berothid species have been described from various localities (listed in Table 1; others are described but unnamed, only illustrated or represented by larvae). The family was previously only known tentatively from the Jurassic:
A list of known fossil
Species | Age | Locality | References | |
---|---|---|---|---|
1 | Early Cretaceous (Early Berriasian) | Durlston Bay, England (Lulworth Formation) |
|
|
2 | Early Cretaceous (Valanginian/Hauterivian) | Lebanese amber (Jezzine) |
|
|
3 | Early Cretaceous (Valanginian/Hauterivian) | Lebanese amber (Jezzine) | ||
4 | Early Cretaceous (Valanginian/Hauterivian) | Lebanese amber (Jezzine) | ||
5 | Early Cretaceous (Valanginian/Hauterivian) | Lebanese amber (Jezzine) |
|
|
6 | Early Cretaceous (Neocomian) | Lebanese amber (Houarij) |
|
|
7 | Early Cretaceous (Barremian/Aptian) | Lebanese amber (Hammana) |
|
|
8 | Early Cretaceous (Barremian) | Yixian Formation, China |
|
|
9 | Early Cretaceous (Late Aptian) | Crato Formation, Brazil |
|
|
10 | Early Cretaceous (Late Aptian) | Crato Formation, Brazil |
|
|
11 | Early Cretaceous (Early Aptian) | Spanish amber (El Sophao) |
|
|
12 | Early Cretaceous (Late Albian) | Archingeay, France |
|
|
13 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
14 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
15 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
16 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
17 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
18 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
19 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
20 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
21 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
22 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
23 | Early Cretaceous (Late Albian) | Burmese amber |
|
|
24 | Early Cretaceous (Late Albian) | Burmese amber | ||
25 | Late Cretaceous (Cenomanian) | Bezonnais, France |
|
|
26 | Late Cretaceous (Cenomanian) | Obeshchayushchiy, NE Siberia (Ola Formation) |
|
|
27 | Late Cretaceous (Turonian) | Raritan (New Jersey) amber |
|
|
28 | Late Cretaceous (Turonian) | Raritan (New Jersey) amber |
|
|
29 | Late Cretaceous (Turonian) | Raritan (New Jersey) amber |
|
|
30 | Late Cretaceous (Turonian) | Raritan (New Jersey) amber |
|
|
31 | Late Cretaceous (Turonian) | Raritan (New Jersey) amber |
|
|
32 | Late Cretaceous (Santonian) | Yantardakh, N Siberia |
|
|
33 | Late Cretaceous (Campanian) | Canadian amber |
|
|
34 | Late Cretaceous (Campanian) | Canadian amber |
|
|
35 | Late Cretaceous (Campanian) | Canadian amber | ||
36 | Early Eocene | Oise amber, France |
|
|
37 | Early Eocene | Hat Creek amber, British Columbia |
|
|
38 | Late Eocene | English amber |
|
|
39 | Late Eocene | Baltic amber |
|
|
40 | Late Eocene | Baltic amber |
|
|
41 | Late Eocene | Baltic amber | ||
42 | Late Eocene | Baltic amber | ||
43 | Late Eocene | Baltic amber |
|
|
44 | Late Eocene | Baltic amber | ||
45 | Late Eocene | Baltic amber | V.Makarkin, S.Wedmann, T.Weiterschan (ongoing research) | |
46 | Late Eocene | Rovno amber, Ukraine | E.Perkovsky, V.Makarkin (ongoing research) |
This study is based on three specimens collected from Daohugou Village (Shantou Township, Ningcheng County, Inner Mongolia, China) and housed in the Key Laboratory of Insect Evolution and Environmental Changes, College of Life Sciences, Capital Normal University, Beijing, China (CNUB; Dong Ren, curator). These insect-bearing beds are here considered as belonging to the Jiulongshan Formation and are dated Bathonian, Middle Jurassic (
Specimens were examined using a Leica MZ12.5 dissecting microscope; line drawings were prepared with CorelDraw 12 graphics software with the aid of Adobe Photoshop; photographed by a Nikon SMZ1000 stereomicroscope.
Venational terminology principally follows
Abbreviations used in the text and figures are as the follows: 1A–3A, first to third anal veins; CuA, CuP, anterior and posterior branches of the cubital vein (Cu); MA, MP, anterior and posterior branches of the medial vein (M); R1, anterior branch of the radial vein (R); Rs1, most proximal branch of the radial sector (Rs); Rs2, branch of the radial sector located distal to Rs1; Rs3, branch of the radial sector located distal to Rs2; Sc, subcostal vein.
Forewing: costal space strongly narrowed basally; humeral veinlet not recurrent and branched; Sc, R1 fused distally; Sc+R1 with 9-11 veinlets, mostly simple; all subcostal veinlets simple; M forked far distal to origin of Rs; CuA pectinate, with seven branches; few crossveins in radial space arranged mainly in 1-2 ‘inner’ gradate series.
Three species from the Middle Jurassic of China (Jiulongshan Formation):
The venation of these two new species is very similar to that of
Differs from both other species of
Body indistinctly preserved. Antennae moniliform, incomplete; preserved segments transverse (wider than long). Prothorax short. Mesonotum of usual neuropteran morphology. Legs covered with short hairs; fore-, mid-legs relatively short; hind-leg tibia long; fore-, hind-leg basitarsus longest segment of tarsus. Abdomen very poorly preserved.
Forewing with broad-rounded apex, 6.7 mm long, 3.0 mm wide. Costal space moderately broad, strongly dilated at proximal 1/5 of wing length, narrowed basally. Subcostal veinlets simple, regularly arranged, closely spaced. Sc distally fused with R1 far from wing apex; Sc+R1 with 9-11 simple veinlets. Subcostal space broad, with one basal crossveins located immediately after origin of Rs. R1 space narrower than subcostal space; three widely-spaced crossveins before fusion of Sc, R1, one after. Rs with 11 (right wing), 10 (left wing) parallel pectinate regularly-spaced branches; six proximal branches with 2-4 terminal forks, other branches once forked. Rs1 originating near origin of Rs. M appears fused basally for short distance; forked much distal to origin of Rs1. MA, MP almost parallel, distally with one, two quite long forked branches respectively. Cu divided into CuA, CuP proximal to origin of Rs. CuA pectinate, with 7 branches, some once forked. CuP once deeply forked. Anal veins very poorly preserved; 1A, 2A apparently once deeply forked each; 3A simple. Four gradate series of crossveins posterior to stem of Rs partly preserved (series 1-4 of
Hind wing poorly preserved, approximately 6.5 mm long, 2.6 mm wide. Costal space narrow, distally only slightly dilated. Subcostal veinlets simple, rather closely spaced. Sc distally fused with R1 far from wing apex; Sc+R1 with 13 simple veinlets. Subcostal space relatively narrow; no crossveins detected. R1 space broad, dilated basally; two crossveins before fusion of Sc, R1, one after. Rs originating far from wing base, with eight branches, each forked distally 1-3 times except Rs1 which deeply forked four times. Fork of M not detected. MA once forked distally; MP dichotomously branched distally. CuA long, almost parallel to hind margin, its branches poorly preserved. CuA space relatively broad. CuP fragmentary preserved, quite short. Anal veins not preserved. Crossveins posterior to stem of Rs not detected except one distal between MP, CuA (4m-cu).
Holotype CNU-NEU-NN2011002P (part), CNU-NEU-NN2011002C (counterpart), deposited in CNUB. A nearly complete specimen.
Daohugou Village, Shantou township, Ningcheng county, Inner Mongolia, China. Jiulongshan Formation, Middle Jurassic.
The species is named in honor of the distinguished Russian paleoentomologist Prof. Alexandr Pavlovich Rasnitsyn.
Differs from
Forewing with broad-rounded apex, about 6.0 mm long (as preserved, estimated complete length about 6.5 mm), 2.6 mm wide. Costal space moderately broad, most dilated at proximal 1/5 of wing length. Subcostal veinlets simple, regularly arranged, less closely spaced than in previous species. Sc distally fused with R1 far from wing apex; Sc+R1 with nine veinlets (eight simple, one forked). Subcostal space broad, with two basal crossveins. R1 space nearly as wide as subcostal space; six crossveins before fusion of Sc and R1, one after. Rs with nine pectinate, regularly spaced branches; four proximal-most branches with 2-4 terminal forks, other branches once forked. Rs1 originating at some distance from origin of Rs. M not fused basally; forked much distal to origin of Rs1. MA, MP almost parallel, distally with one (simple) , two (one simple) branches respectively. Cu divided into CuA, CuP proximal to origin of Rs. CuA pectinate, with 7 branches; proximal-most branch once forked. CuP once deeply forked. Anal veins incompletely preserved; 1A with single marginal fork; 2A with two marginal short branches; 3A very incomplete, with single fork preserved. Four gradate series of crossveins posterior to stem of Rs, all incomplete. First series consists of three crossveins: 1r-m (located at origin of Rs), 1m-cu, 1a1-a2 (longer than previous); Second series includes two crossveins: 2m-cu (connecting MP, CuA), 2icu (connecting CuA, anterior branch of CuP). Third (‘inner’) series with six crossveins (3rs-rs7, 3rs5-rs4 to 3rs2-rs1; two between Rs3, Rs2). Fourth (‘outer’) series with five crossveins (from 4rs2-rs1 to 4m-cu; two between Rs1, MA). Wing one color, fuscous. Veins mainly dark brown as preserved.
Holotype CNU-NEU-NN2011003, deposited in CNUB. A nearly complete forewing.
Daohugou Village, Shantou township, Ningcheng county, Inner Mongolia, China. Jiulongshan Formation, Middle Jurassic.
From the Latin
Hind wing approximately 6.5 mm long, 2.7 mm wide. Humeral lobe not extended; frenulum poorly-developed consisting of few bristles. Costal space narrow, dilated towards apex. Subcostal veinlets simple, more closely-spaced apically. Sc distally fused with R1 far from wing apex; Sc+R1 with seven long veinlets (one forked). Subcostal space relatively broad, with one basal crossveins. R1 space nearly as wide as subcostal space; four crossveins before fusion of Sc, R1. Rs with seven pectinate, regularly spaced branches; one branch deeply forked. Rs1 originating at some distance from origin of Rs. Proximal crossvein m-r long, connecting Rs1 near its origin with M. M forked distal to origin of Rs1. MA, MP almost parallel, distally with few branches. CuA long, slightly incurved, in general parallel to hind margin, with nine forkes branches originated at angle >45 degrees, one simple branch. CuP short, with two branched preserved. 1A–3A not preserved. Crossvein between CuA, 1A (or CuP). One crossvein between Rs, Rs6 in ‘inner’ gradate series (possibly anomalous). Six crossveins (from Rs4 to CuA) in ‘outer’ gradate series preserved. Wing one color, fuscous. Veins appear mainly dark brown.
Specimen CNU-NEU-NN2011004, deposited in CNUB. A nearly complete hind wing.
Daohugou Village, Shantou township, Ningcheng county, Inner Mongolia, China. Jiulongshan Formation, Middle Jurassic.
The venation of this hind wing is typical for
This specimen is tentatively assigned to
As the type species of the genus
The forewing venation of
The hind wing venation of “
Triassic berothid-like taxa have been treated as belonging to the family
We thank S. Bruce Archibald (Simon Fraser University, Burnaby, Canada) for correction of English. This research is supported by the National Natural Science Foundation of China (Nos. 40872022, 31071964), the Nature Science Foundation of Beijing (No. 5082002) and Scientific Research Key Program KZ200910028005, and PHR Project of Beijing Municipal Commission of Education.