Print
The first Elcanidae (Orthoptera, Elcanoidea) from the Daohugou fossil bed of northeastern China
expand article infoHe Tian, Jun-Jie Gu§, Xiang Chu Yin|, Dong Ren#
‡ Capital Normal University, Beijing, China
§ Sichuan Agricultural University, Chengdu, China
| Northwest Institute of Plateau Biology, Chinese Academy of Sciences, Xining, China
¶ Hebei University, Baoding, China
# Capital of Normal University, Beijing, China
Open Access

Abstract

A new species of Elcanidae (Orthoptera, Elcanoidea), Parelcana pulchmacula sp. nov., is described based on four new specimens from the Middle Jurassic Jiulongshan Formation of northeastern China. The new species differs from all other known Archelcaninae species by its combination of wing-venation characters. This new finding improves our knowledge of variation on wing venation in elcanid insects and constitutes the first record of Elcanidae from the Daohugou fossil bed (Yanliao Biota) of northeastern China.

Keywords

Archelcaninae, Inner Mongolia, Middle Jurassic, Parelcana, systematic paleontology, Yanliao Biota

Introduction

The extinct family Elcanidae is a cryptic group of Orthoptera insects due to their complex anatomical features. The caeliferan-like wing venation, characteristic of this family, groups them close to Caelifera in cladistic analyses (Béthoux and Nel 2002). However, Elcanidae also shares a long, filiform antennae and exerted ovipositors with the suborder Ensifera. The presence of these contrasting anatomical features makes the systematic position of Elcanidae unclear.

The Elcanidae existed from the Upper Triassic to the Cretaceous in Eurasia and America (Handlirsch 1906; Sharov 1968; Martins-Neto 1991; Gorochov et al. 2006; Poinar et al. 2007; Peñalver and Grimaldi 2010; Fang et al. 2015; Fang et al. 2018a, Fang et al. 2018b; Heads et al. 2018; Tian et al. 2019). So far 50 species in 16 genera have been described from compression fossils and ambers. These species have been divided into two subfamilies, the Archelcaninae and the Elcaninae, based on taxonomic characters (Gorochov et al. 2006). Elcanids evolved a unique character among orthopterans, i.e., the presence of various spurs on the distal part of the metatibia. These structures might have been associated with an improved capability to swim (Tian et al. 2019).

The Jurassic elcanids are well known from the UK (Handlirsch 1906; Whalley 1985), Germany (Handlirsch 1906; Ansorge 1996), Kazakhstan (Sharov 1968), and Kyrgyzstan (Sharov 1968). In China, two specimens have been reported, one from the Guangxi Province (from Early Jurassic), and one from the Hebei Province (from Middle Jurassic). Both were attributed to Elcana reticulata (Handlirsch, 1939), based on existing highly fragmented forewing sample specimens (Handlirsch 1939; Hong 1983; Lin 1986). Other Orthoptera are commonly discovered from the Daohugou fossil bed, at the widely known and profuse fossil assemblages of Yanliao Biota, northeastern China. Numerous species discovered from this fossil bed have been described, including ensiferans and caeliferans (Ren and Meng 2006; Ren et al. 2010, 2019; Gu et al. 2009, 2011, 2012, 2016; Wang et al. 2017). In this report we describe a new species of Elcanidae, Parelcana pulchmacula sp. nov., from the Daohugou fossil bed. The new species is erected based on four isolated but well-preserved forewings, providing new insights into the complex wing-venation patterns of elcanids.

Geological setting

The specimens described here were collected in the Daohugou Bed, located along the boundaries of the provinces of Hebei, Liaoning and Inner Mongolia (Fig. 1). The Daohugou Bed has been previously assigned to the Middle Jurassic Jiulongshan Formation (Ren et al. 2002, 2010, 2019). The Mesozoic section of the Daohugou Bed is mainly composed of tuffaceous conglomerates, tuffaceous siltstones, tuffaceous mudstones, tuffaceous shales, and volcanic breccias. Isotopic radiometric dating of rock samples from the Daohugou area enabled assessment of the age of the Jiulongshan Formation at circa 168–164 million years (Chen et al. 2004; He et al. 2004; Liu et al. 2006; Yang and Li 2008; Chang et al. 2014). This indicates that the age of the Jiulongshan Formation falls within the Bathonian–Callovian boundary interval (Xu et al. 2016).

Figure 1. 

Location map for Parelcana pulchmacula sp. nov.

Materials and methods

The wing specimens were examined with a Nikon SMZ 25 microscope, and photographed with a Nikon DS-Ri 2 digital camera system. Line drawings were prepared using Adobe Illustrator CC 17.0.0 and Adobe Photoshop CC 14.0 graphics software. The measurements were taken using Adobe Illustrator. The lengths of wings were measured from the apex to the visible base of the wing; the widths of wings refer to the maximum width of the wing. The specimens are deposited in the Key Lab of Insect Evolution & Environmental Changes, Capital Normal University (CNU), Beijing, China.

Wing-venation analyses followed the interpretation proposed by Béthoux and Nel (2001, 2002). Corresponding abbreviations used in taxonomical descriptions are as follows: CP, posterior costa; ScA, ScP, anterior and posterior subcosta, respectively; RA, RP, anterior and posterior radius, respectively; M, media; MA, MP, anterior, posterior media, respectively; CuA, CuP, anterior, posterior cubitus, respectively; CuPaα, the anterior branch of first posterior cubitus; CuPaβ, the posterior branch of first posterior cubitus; CuPb, the second posterior cubitus; AA1, first branch of anterior anal vein.

Systematic paleontology

Class Insecta Linnaeus, 1758

Order Orthoptera Olivier, 1789

Suborder Ensifera Chopard, 1920

Superfamily Elcanoidea Handlirsch, 1906

Family Elcanidae Handlirsch, 1906

Subfamily Archelcaninae Gorochov, Jarzembowski & Coram, 2006

Parelcana Handlirsch, 1906

Type species

Parelcana tenuis Handlirsch, 1906.

Composition

Parelcana tenuis Handlirsch, 1906 (Jurassic, Dobbertin, Germany), P. anglicana Handlirsch, 1939 (Jurassic, Binton, UK), probably P. dubia Handlirsch, 1939 (Jurassic, Gloucester, UK) and Parelcana pulchmacula sp. nov. (Handlirsch 1939; Cigliano et al. 2019).

Revised diagnosis

ScP with numerous branches ending at the anterior margin; M with 3 branches before RP fused with MA1; short CuA; CuPaβ, CuPb, and AA1 detached from each other.

Comments

All species of genus Parelcana are based on forewing structure. The forewing of Parelcana differs from other genera in Archelcaninae by the presence of 3 branches of M before RP fuses with MA1, and presence a very short and nearly vertical CuA.

Parelcana pulchmacula sp. nov.

Fig. 2

Diagnosis

ScP with 6–8 branches ending in anterior margin; CuA very short and fusion with CuPaα before ScA ends in anterior margin, CuA+CuPaα long and S-shaped; occurrence of two big and round dark spots in distal half of wing and one small spot covering the area of CuPa.

Etymology

From the latin ‘Pulch-’ for beautiful and ‘macula’ for patches, referring to the beautiful spots and coloration of the forewing.

Type materials

Holotype , CNU-ORT-NN2016041; Paratypes, CNU-ORT-NN2016035; CNU-ORT-NN2016036; CNU-ORT-NN2016042.

Locality and age

Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China; Jiulongshan Formation, Middle Jurassic.

General description

Forewing 18.4–20.9 mm long and 4.3–5.0 mm wide (maximum width recorded). Costal area long and narrow; CP nearly straight, ending in anterior margin after the forking of M+CuA, generating numerous distinct oblique branches ending in the anterior margin; ScA slightly curved, ending in the anterior margin before 1/3 of total wing length; ScP reaching anterior margin at nearly half-length of wing and generating 6–8 oblique branches ending in the anterior margin; stem R+M+CuA forking into R and M+CuA after the divergence point of CuPa; stem R long and distinctly strong, branched into RA and RP near the mid-length of wing; area between ScP and R narrow; RA slightly curved towards posterior wing margin before its first branch, reaching anterior margin close to apex with 16–18 oblique branches; RP with 10–12 comb-like branches reaching wing margin, most of them reaching posterior margin, with several distal terminals dichotomizing and reaching anterior margin; area between RA and RP relatively wide; M forking into MA and MP near to the end of ScA; MA branching into MA1 and MA2 close to the end of ScP; MA1 with 2 branches, with the first fused with RP; MP simple, originates after ScA ends at anterior margin; CuA extremely short, 0.16 to 0.20 mm long, originates before CP ends at anterior margin; CuA almost vertical against the posterior margin; free CuPa short, 0.23 to 0.38 mm long, directed to anterior wing margin, forking into CuPaα and CuPaβ before (Fig. 2A–D) or at the level (Fig. 2G, H) of the bifurcation point of M+CuA; free part of CuPaα approximately three times longer than CuPa, then fused with CuA; CuA+CuPaα simple, long and S-shaped, reaching posterior margin at 2/3 of wing length; CuPaβ simple, similar to CuPaα in shape; CuPb simple; areas between CuPaβCuPb and CuPbAA1 narrow; CuPaβ, CuPb, and AA1 detached each other; AA1 strong and straight; area between branches of RP and M covered with simple and straight crossveins. Dark colorations cover the areas between ScP–R and RA–anterior margin, and also along several rows of the crossveins between branches of RP and M; occurrence of two big and round spots in distal half of wing, one located between the branches of RP, one located at the boundary of RP branches and MA1; one small round spot covers the area of CuPa.

Figure 2. 

Photo and line drawing of Parelcana pulchmacula sp. nov. A, B holotype, CNU-ORT-NN2016041 C, D CNU-ORT-NN2016035 E, F CNU-ORT-NN2016042 G, H CNU-ORT-NN2016036. The inclined and dotted lines in the middle of the wings of D and F represent the cracks in the specimen. The dotted line on the wing venation represents the imaginary line of the wing. Scale bar: 2 mm.

Specimen description

CNU-ORT-NN2016041 (Fig. 2A, B). Holotype, forewing nearly complete with only basal and posterior margin partially missing, 18.4 mm long and 4.3 mm wide (the maximum width, the same below). CP with 3 oblique branches preserved; ScA with 2 branches ending in anterior margin; ScP with 7 branches ending in anterior margin; RA with 17 oblique branches; RP with 10 pectinate branches reaching wing margin; RP fused with anterior branch of MA1 slightly after the ramification point of MA1; CuPaβ reaches the posterior wing margin distally to the end of ScP.

CNU-ORT-NN2016035 (Fig. 2C, D). Paratype, forewing lost anal region and split into two pieces at about mid-length by an oblique crevice, preserved 20.7 mm long and 4.8 mm wide. CP with 3 oblique branches preserved; ScA with 1 branch connected with CP; ScP with 1 branch connected with ScA and 7 branches ending in anterior margin; RA with 16 oblique branches; RP with 12 pectinate branches reaching wing margin; shortly after origination of posterior branch of MA1, RP fused with anterior branch of MA1.

CNU-ORT-NN2016042 (Fig. 2E, F). Paratype, forewing lost basal and anal regions, and an oblique crevice split it into two pieces at about mid-length, remaining part 19.1 mm long and 4.5 mm wide. ScA with 1 branch connected with CP; ScP with 8 branches ending in anterior margin; RA with 18 oblique branches; RP with 12 pectinate branches reaching wing margin; RP fused with anterior branch of MA1 at same level as origination of posterior branch of MA1.

CNU-ORT-NN2016036 (Fig. 2G, H). Paratype, forewing lost anal region, 20.9 mm long and 5 mm wide. CP with 8 oblique branches; ScA with 1 branch connected with CP; ScP with 6 branches ending in anterior margin; RA with 18 oblique branches; RP with 10 pectinate branches reaching wing margin; RP fused with anterior branch of MA1 after the ramification point of MA1; CuPaβ reaches the posterior wing margin basally to the end of ScP.

Discussion

This new species can be assigned to Archelcaninae by its relatively wide area between RA and RP, and free distal part of CuPaβ, CuPb and AA1. Its simple ScA, the presence of 3 branches of M before RP fuses with MA1 and a very short CuA support assignment to the genus Parelcana. Parelcana pulchmacula sp. nov. shares with P. tenuis a short and vertical CuA, but differs from P. tenuis in its larger size, greater number of branches of ScP and RP, a long and S-shaped CuA+CuPaα, a wider area between CuPb and anals, and the coloration pattern of the forewing. The new species is notably different from P. anglicana in its greater number of branches of ScP, free and vertical CuA, wider area between CuA+CuPaα and anal region, and fusion pattern of CuA and CuPaα. Parelcana dubia was erected based on a fragmentary forewing. It differs from P. pulchmacula sp. nov. in having a wider area between RA and RP and the branching pattern of RP. Parelcana pulchmacula sp. nov. is also different from the other two known Chinese Jurassic elcanids. It is much larger than the specimen from the Early Jurassic with an estimated wing length of approximately 9.5 mm. The other specimen from the Middle Jurassic of Hebei was originally assigned to Elcana reticulata based on an isolated forewing with only the distal half. Most of the diagnostic characters were missing, making comparisons with the new species difficult. Based on the line drawing patterns described for the wing (see Hong 1983, fig. 28), it might be an elcanid, but its generic assignment is questionable.

Variation in forewing size and wing-venation pattern is common in fossil orthopterans and their relatives from the Palaeozoic to the Mesozoic (Prokop and Ren 2007; Gu et al. 2010, 2011, 2014; Béthoux et al. 2012), even between the left and right forewings of the same individual (Gu et al. 2009). For elcanids insects, the documentation on variation of wing venation is scarce since most of the species are described based on limited and often poorly preserved samples. We observed some variation of forewing venation within specimens of P. pulchmacula sp. nov.. The number of branches of ScP, RA and RP was not consistent (Fig. 2). Moreover, CuPaβ reaches the posterior wing margin distally to the end of ScP in CNU-ORT-NN2016041(Fig. 2A, B), but basally to the end of ScP in CNU-ORT-NN2016036 (Fig. 2G, H).

Based on the known data of wing venation of fossil orthopterans (Gu et al. 2010, 2011, 2014; Béthoux et al. 2012), the differences found in four P. pulchmacula sp. nov. specimens should be considered as variations within a species. The location of the fusion point of RP and MA1 is an important character usually used as a distinctive generic character of Elcaninae (Gorochov et al. 2006). In P. pulchmacula sp. nov., three of these fusion points show a clear pattern of M with 3 branches before RP is fused with MA1 (Fig. 3A–C). However in specimen CNU-ORT-NN2016042 (Fig. 3D), 2MA1 branches off at the level of RP when reaching MA1. Although this kind of difference always occurs between different species or genus, the integral similarity of wing venation between the four specimens of P. pulchmacula sp. nov. indicates that these specimens belong to the same species.

Figure 3. 

Details of the middle part of the forewings of Parelcana pulchmacula sp. nov., showing the variable location of the fusion point of RP and MA1 in the forewing. A CNU-ORT-NN2016035 B CNU-ORT-NN2016036 C holotype, CNU-ORT-NN2016041 D CNU-ORT-NN2016042. Scale bar: 0.5 mm.

Acknowledgments

We sincerely appreciate the critical and valuable comments from the editor and anonymous reviewers. We thank Dr Jose Marcelino from the Centre for Ecology, Evolution and Environmental Changes, University of the Azores, Portugal for improving our manuscript. This research is supported by grants from the National Natural Science Foundation of China (No. 41872020, 41688103, 31730087), the Program for Changjiang Scholars and Innovative Research Team at University (IRT-17R75) and the Project of High-level Teachers in Beijing Municipal Universities (No. IDHT20180518).

References

  • Ansorge J (1996) Insekten aus dem oberen Lias von Grimmen (Vorpommern, Norddeutschland). Neue Paläontologische Abhandlungen 2: 1–132.
  • Béthoux O, Nel A (2002) Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa 96(1): 1–88. https://doi.org/10.11646/zootaxa.96.1.1
  • Béthoux O, Gu JJ, Yue YL, Ren D (2012) Miamia maimai n. sp., a new Pennsylvanian stem-orthopteran insect, and a case study on the application of cladotypic nomenclature. Fossil Record 15(2): 103–113. https://doi.org/10.1002/mmng.201200008
  • Chang SC, Zhang HC, Hemming SR, Mesko GT, Fang Y (2014) 40Ar/39Ar age constraints on the Haifanggou and Lanqi formations: when did the first flowers bloom. Geological Society London Special Publications 378(1): 277–284. https://doi.org/10.1144/SP378.1
  • Chen W, Ji Q, Liu DY, Zhang Y, Song B, Liu XY (2004) Isotope geochronology of the fossil-bearing beds in the Daohugou area, Ningcheng, Inner Mongolia. Geological Bulltetin of China 23(12): 1165–1169. [in Chinese with English abstract]
  • Fang Y, Wang B, Zhang HC, Wang H, Jarzembowski EA, Zheng DR, Zhang Q, Li S, Liu Q (2015) New Cretaceous Elcanidae from China and Myanmar (Insecta, Orthoptera). Cretaceous Research 52: 323–328. https://doi.org/10.1016/j.cretres.2014.05.004
  • Fang Y, Heads SW, Wang H, Zhang HC, Wang B (2018a) The first Archelcaninae (Orthoptera, Elcanidae) from the Cretaceous Jehol Biota of Liaoning, China. Cretaceous Research 86: 129–134. https://doi.org/10.1016/j.cretres.2018.02.008
  • Fang Y, Muscente AD, Heads SW, Wang B, Xiao S (2018b) The earliest Elcanidae (Insecta, Orthoptera) from the Upper Triassic of North America. Journal of Paleontology 92(6): 1028–1034. https://doi.org/10.1017/jpa.2018.20
  • Gorochov AV, Jarzembowski EA, Coram RA (2006) Grasshoppers and crickets (Insecta: Orthoptera) from the Lower Cretaceous of southern England. Cretaceous Research 27(5): 641–662. https://doi.org/10.1016/j.cretres.2006.03.007
  • Gu JJ, Qiao GX, Ren D (2010) Revision and New Taxa of Fossil Prophalangopsidae (Orthoptera: Ensifera). Journal of Orthoptera Research 19(1): 41–56. https://doi.org/10.1665/034.019.0110
  • Gu JJ, Béthoux O, Ren D (2011) A exceptionally-preserved new species of Barchaboilus (Orthoptera: Prophalangopsidae) from the Middle Jurassic of Daohugou, China. Zootaxa 2909: 64–68. https://doi.org/10.11646/zootaxa.2909.1.7
  • Gu JJ, Qiao GX, Ren D (2012) The first discovery of Cyrtophyllitinae (Orthoptera, Haglidae) from the Middle Jurassic and its morphological implications. Alcheringa: An Australasian Journal of Palaeontology 36(1): 27–34. https://doi.org/10.1080/03115518.2011.576535
  • Gu JJ, Yue YL, Wen WC, Zong LY, Ren D (2014) A review of researches on Palaeozoic insects in China. Acta Enomologica Sinica 57(1): 123–132. [in Chinese with English abstract]
  • Handlirsch A (1906–08) Die fossilen Insekten und die Phylogenie der rezenten Formen. Ein Handbuch für Paläontologen und Zoologen, 1430 pp. [Engelman, Leipzig: pp. 1–640 was published in 1906, pp. 641–1430 was published in 1908]
  • Handlirsch A (1939) Neue Untersuchungen Über die fossilen Insekten. II. Teil. -Ann. Naturhist. Mus. Wien 49, 1–240.
  • He HY, Wang XL, Zhou ZH, Zhu RX, Jin F, Wang F, Ding X, Boven A (2004) 40Ar/39Ar dating of ignimbrite from Inner Mongolia, northeastern China, indicates a post-Middle Jurassic age for the overlying Daohugou Bed. Geophyical Research Letters 31: 1–4. https://doi.org/10.1029/2004GL020792
  • Heads SW, Thomas MJ, Heads YN (2018) A new genus and species of Elcanidae (Insecta: Orthoptera) from Cretaceous Burmese amber. Zootaxa 4527(4): 575–580.
  • Hong YC (1983) Taxonomic description. In Middle Jurassic Fossil Insects in North China. Rong L-b ed, Geological Publishing House, Beijing, 42–48. [in Chinese]
  • Liu YX, Liu YQ, Zhong H (2006) LA-ICPMS zircon U-Pb dating in the Jurassic Daohugou beds and correlative strata in Ningcheng of Inner Mongolia. Acta Geologica Sinica (English Edition) 80: 733–742. https://doi.org/10.1111/j.1755-6724.2006.tb00296.x
  • Lin QB (1986) Taxonomic description. In Early Mesozoic Fossil Insects from South China. Nanjing Institute of Geology and Palaeontology ed, Science Press, Beijing, 53–54. [in Chinese]
  • Martins-Neto RG (1991) Sistemática dos Ensifera insecta, (Orthopteroida) da Formação Santana (Cretáceo inferior do nordeste do Brasil). Acta Geologica Leopoldensia (32): 110–113.
  • Ren D, Gao KQ, Guo ZG, Ji SA, Tan JJ, Song Z (2002) Stratigraphic division of the Jurassic in the Daohugou area, Ningcheng, Inner Mongolia. Geological Bulltetin of China 21(8–9), 584–591. [in Chinese with English abstract]
  • Ren D, Meng XM (2006) New Jurassic Protaboilins from China (Or thoptera, Prophalangopsidae, Protaboilinae). Acta Zootaxonomica Sinica 31: 513–519. [in Chinese with English abstract]
  • Ren D, Shih CK, Gao TP, Yao YZ, Zhao YY (2010) Silent Stories–Insect Fossil Treasures from Dinosaur Era of the Northeastern China. Science Press, Beijing, 322 pp.
  • Ren D, Shih CK, Gao TP, Wang YJ, Yao YZ (2019) Rhythms of Insect Evolution–Evidence from the Jurassic and Cretaceous in Northern China. Wiley Blackwell, NJ, 710 pp. https://doi.org/10.1002/9781119427957
  • Peñalver E, Grimaldi DA (2010) Latest occurrences of the Mesozoic family Elcanidae (Insecta: Orthoptera), in Cretaceous amber from Myanmar and Spain. Annales de la Société entomologique de France (N.S. ) 46(1–2): 88–99. https://doi.org/10.1080/00379271.2010.10697641
  • Poinar G, Gorochov AV, Buckley R (2007) Longioculus burmensis, n. gen., n. sp. (Orthoptera, Elcanidae) in Burmese amber. Proceedings of the Entomological Society of Washington (109): 649–655.
  • Prokop J, Ren D (2007) New significant fossil insects from the Upper Carboniferous of Ningxia in northern China (Palaeodictyoptera, Archaeorthoptera). European Journal of Entomology 104: 267–275. https://doi.org/10.14411/eje.2007.041
  • Sharov AG (1968) Phylogeny of the Orthopteroidea. Proceedings of the Paleontological Institute, Russian Academy of Sciences, 217 pp. [in Russian]
  • Tian H, Gu JJ, Huang F, Zhang H, Ren D (2019) A new species of Elcaninae (Orthoptera, Elcanidae) from the Lower Cretaceous Yixian Formation at Liutiaogou, Inner Mongolia, NE China, and its morphological implications. Cretaceous Research 99: 275–280. https://doi.org/10.1016/j.cretres.2019.03.010
  • Yang W, Li SG (2008) Geochronology and geochemistry of the Mesozoic volcanic rocks in Western Liaoning: implications for lithospheric thinning of the North China Craton. Lithos 102: 88–117. https://doi.org/10.1016/j.lithos.2007.09.018
  • Xu X, Zhou ZH, Corwin S, Wang Y, Ren D (2016) An Updated Review of the Middle-Late Jurassic Yanliao Biota: Chronology, Taphonomy, Paleontology and Paleoecology. Acta Geologica Sinica [English edition] 90(6): 2229–2243. https://doi.org/10.1111/1755-6724.13033
  • Wang H, Fang Y, Zhang QQ, Lei XJ, Wang B, Jarzembowski EA, Zhang HC (2017) New material of Sigmaboilus (Insecta, Orthoptera, Prophalangopsidae) from the Jurassic Daohugou Beds, Inner Mongolia, China. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 107: 177–183. https://doi.org/10.1017/S1755691017000172
  • Whalley PES (1985) The systematics and palaeogeography of the Lower Jurassic insects of Dorset, England. Bulletin British Museum Natural History, (Geol. ) 39(3): 107–189.