New species of Cicadocoris ( Hemiptera : Coleorrhyncha : Progonocimicidae

A new progonocimicid bug named Cicadocoris parvus sp. n. is described from the mid-Jurassic Haifanggou Formation at Daohugou, Ningcheng County, Inner Mongolia, northeastern China. It differs from other species by being distinctly smaller, veins dSc, R1 and Rs run parallel to one another and are nearly evenly spaced on left tegmen, widest length of apical half/basal half of left tegmen is 1.1 and that of right tegmen is 1.0. Thus, there are at least three species of Cicadocoris (Progonocimicidae) described from Daohugou. All these species are relatively abundant in the Haifanggou Formation and are remarkable representatives of the early Yanliao biota. ZooBank Article LSID: BE181216-421E-4594-9A48-A789D0B3B86F * Corresponding author; e-mail: dyhuang@nigpas.ac.cn INTRODUCTION Coleorrhyncha is a small suborder of Hemiptera comprised of three families: the extant Peloridiidae Breddin, 1897, the extinct Progonocimicidae Handlirsch, 1906 and Karabasiidae Popov, 1985 (Popov & Shcherbakov, 1991; Wang et al., 2009). Peloridiidae is a small family with currently a limited distribution in wet moss in temperate and sub-Antarctic rainforests in the Southern Hemisphere (e.g. Chile, Argentina, New Zealand, New Caledonia, southeastern Australia; Burckhardt, 2009; Burckhardt et al., 2011). The fossil progonocimicids are recorded from the Upper Permian to the Lower Cretaceous and have a remote Gondwana origin. They were widely distributed, occurring in Permian deposits in Australia (Tillyard, 1926), Triassic in Kyrgyzstan (Becker-Migdisova, 1958; Popov & Shcherbakov, 1991), Australia (Evans, 1956, 1961, 1963; Wootton, 1963) and Argentina (Martins-Neto, 2003), Jurassic in Kyrgyzstan (Popov, 1982, 1985), United Kingdom (Popov et al., 1994), China (Hong, 1983; Dong et al., 2012, 2013, 2014), Russia (Popov, 1985, 1988), Germany (Handlirsch, 1939; Popov & Wootton, 1977), Luxembourg (Szwedo, 2011), and Cretaceous in Lebanon (Szwedo et al., 2011), United Kingdom (Handlirsch, 1906; Klimaszewski & Popov, 1993), Argentina (Martins-Neto, 2003), Mongolia Eur. J. Entomol. 114: 355–364, 2017 doi: 10.14411/eje.2017.045


MATERIAL AND METHODS
The material studied includes a total of 40 individuals (21 males, 12 females and 7 undetermined); most of them are com- Type species.Cicadocoris kuliki Becker-Migdisova, 1958 (original designation).
Etymology.Derived from the Latin parvus meaning "small", or "little" to indicate the small size of this new species.
Diagnosis.Small size, differs from other species by its distinctly smaller body size (2.8-3.9 mm); veins dSc, R1 and Rs sub-parallel and nearly evenly spaced on left tegmen; left tegmen, widest length of apical half/basal half is1.1 and that of right tegmen is 1.0; 21 visible ridges along median section of female ovipositor; wing length/body length is 0.83 in male, 0.92 in female.
Tegmen (Figs 2, 3) length-width ratio 2.5, with widest part 3/4 along the length of wing.Anterior margin weakly arcuate, with apex at apex of vein M 3+4 .Precostal carina narrow and horizontal.Costal area narrow, not widening towards dSc, dSc straight.Stem of R slightly convex at point of dSc origin, dividing into R 1 and Rs halfway between the fork in M and costal margin, slightly beyond basal forking of M. Arculus short and sub transverse.M 1+2 dividing before r-m, M 3+4 longer than stem M. Stem CuA beyond arculus curved.CuA 2 transverse.Length of radial cell 0.36 times as long as tegmen, length of medial cell 0.34 times as long as tegmen.Right tegmen cu-a, m-cu, base of M 3+4 , M 1+2 and M 1 form distinct and continuous transverse arc, distance from dSc origin to that of R 1 1.1-1.5 times longer than that from the origin of R 1 to that of Rs, veins dSc, R 1 and Rs sub-parallel, M 1 arcuate and closer to Rs than M 2 , width of apical half 1.0 times that of basal half.Left tegmen without arc, origin of R 1 in the middle of that of dSc and Rs, veins dSc, R 1 and Rs parallel and evenly spaced, M 1 on left tegmen straight, equals Rs and M 2 , width of apical half 1.1 times that of basal half.Hindwing rounded apically, with apex at apex of vein M 1+2 .Vein Rs connected to M 1+2 by a sub transverse cross vein r-m a little distal of the origin of M 1+2 .Stem M forks into M 1+2 and M 3+4 little beyond middle of wing.Vein M 3+4 fused with vein CuA 1 .Stem of CuA bifurcates into veins CuA 1 and CuA 2 at same level as the stem of M branches.

DISCUSSION
This new species is assigned to Cicadocoris based on the following constitutive characters: small head; clavate antennae; rostrum thin, extends to base of hind coxae; trapezoidal pronotum; hind tibia with two large movable conical spurs; hind tarsi three-segmented; tegmen at claval apex wider than 1/3 of full length; costal margin arcuate; precostal carina narrow; M 3+4 longer than stem of M.
Cicadocoris parvus sp.n. shares several critical characters with C. sinensis (Hong 1983), such as costal area not widening toward dSc, stem of M shorter than M 3+4 , and the ratio of the widest length of apical half/basal half of the left tegmen larger than the right tegmen.In addition, the body length of this new species ranges from 2.8-3.9 mm and that of C. sinensis is 3.7-5.4mm, with an overlapping region.Thus the body sizes of these two species are not strictly different.However, the new species differs from C. sinensis in the following characters: (1) veins dSc, R 1 and Rs sub-parallel; (2) normal distance from dSc origin to R 1 origin 1.1-1.5 times longer than R 1 origin to Rs origin on right tegmen (1.5-2.5 times in C. sinensis); (3) the ratio of the widest length of apical half is 1.1 times that of the basal half on left tegmen and 1.0 times on right tegmen (1.2 times and 1.1 times for C. sinensis, respectively); ovi-positor has 24 visible ridges along its median section, 8-9 ridges /100 μm (13 visible ridges, 3-4 ridges /100 μm).
C. parvus closely resembles C. brunneus (Hong, 1983) in its venation.The body length of C. brunneus is 5.2-7.0 mm, circa twice as large as the new species.C. parvus sp.n., C. sinensis and C. brunneus always co-occur in the major fossil layers in the Daohugou beds, which excludes the possibility that the new species and C. brunneus are the same species.
C. parvus differs from the type species Cicadocoris kuliki Becker-Migdisova, 1958 in its body length 2.8-3.9mm(vs.2.8-5.8mm in C. kuliki); M 2 does not have a common stalk with M 1 (vs.M 2 has a common stalk with M 1 ); costal area does not widen towards dSc (vs.costal area much wider toward dSc).
Five species of Cicadocoris are so far described from the Daohugou beds.Two species namely C. varians and C. assimilis erected based on a single specimen (Dong et al., 2012(Dong et al., , 2013)).Among them, at least C. varians is probably a junior synonym of C. sinensis because of taphonomic deformation.Three of them (C.sinensis C. brunneus and C. parvus) are abundant in the Daohugou beds at Ningcheng, Inner Mogolia and localities near Haifeng (type section of the Haifanggou Fm.), Beipiao, Liaoning Province (Jiang et al., 2016).This is convincing evidence of the stratigraphic similarity of the Daohugou beds and the type locality of the Haifanggou Formation.Cicadocoris are distinct representatives of an early assemblage of the famous Yanliao biota that became extinct, probably due to a great tectonic movement and volcanic explosion (Huang, 2015;Jiang et al., 2016).

Fig. 1 .
Fig. 1.A -stratigraphic column for the Haifanggou Fm. near Daohugou, Ningcheng County, Inner Mongolia, NE China (red arrows indicate the fossil layers that yielded new species); B -map showing the location of Daohugou and Yujiagou (red triangles); C -map showing locations near Daohugou (red spots).

Fig. 4 .
Fig. 4. Cicadocoris parvus sp.n., males, from the Haifanggou Formation at Daohuguo.A -NIGP165307, general habitus; B -NIGP165307(A), general habitus and body structures; C -NIGP165308, general habitus with tegmina under rock; D -enlargement of C, showing the details of rostrum and legs; E -enlargement of C, showing details of hind leg; F -enlargement of G, showing the tiny teeth on the apices of the tibia and tarsi (indicated by white arrows); G -NIGP165309, general habitus; H -NIGP165310, general habitus; Ienlargement of H, showing details of the tiny teeth on the apices of tibia and tarsi; J -enlargement of H, showing the apical teeth on hind tibia (white arrows); K -enlargement of H, details of claws.B, D, E, F, G, H, I, J, K moistened with 70% ethanol.Scale bars represent 1 mm in A, B, C, G, H; 0.5 mm in D; 0.2 mm in E, F, I; 50 μm in J, K.

Fig. 5 .
Fig. 5. Photographs of males of Cicadocoris parvus sp.n., from the Haifanggou Formation at Daohugou.A -NIGP165311, general habitus; B -enlargement of A, showing details of mouthparts and antennal socket; C -enlargement of A, showing the tiny teeth on the apices of tibia and tarsi; D -NIGP165312, general habitus; E -enlargement of D, showing the details of hind legs and pygophore; F -NIGP165313, general habitus; G -enlargement of F, showing the details of antennae (indicated by black arrows) and antennal socket (indicated by white arrows); H -enlargement of d, showing the details of pygophore.A, B, C, E, F, G, H moistened with 70% ethanol.Scale bars represent 1 mm in A, D, F; 0.5 mm in B, E; 0.2 mm in G, H, C.

Fig. 6 .
Fig. 6.Photographs of females of Cicadocoris parvus sp.n., from the Haifanggou Formation at Daohugou.A -NIGP165314, general habitus; B -NIGP165314(A), general habitus; C -enlargement of A, showing the details of ovipositor; D -enlargement of E, showing the details of ovipositor; E -NIGP165315, general habitus; F -NIGP165315 (E), general habitus and body structures; G -enlargement of A, showing the details of hind leg; H -NIGP165316, general habitus; I -enlargement of H, showing the details of hind legs; J -enlargement of H, showing the tiny teeth on the apices of the tibia and tarsi.B, C, F, G, H, I , J moistened with 70% ethanol.Scale bars represent 1 mm in A, B, E, F, H; 0.2 mm in C, D, G, I, J.

Fig. 7 .
Fig. 7. Photographs of females of Cicadocoris parvus sp.n., from the Haifanggou Formation at Daohugou (A-E, G-J) and Yujiagou (F).A -NIGP165317, general habitus; B -NIGP165317(A), showing general habitus; C -NIGP165318, showing general habitus; D -NIGP165319, the details of ovipositor; E -NIGP165325, general habitus; F -NIGP165326, general habitus; G -enlargement of B, showing details of hind legs and ovipositor; H -enlargement of E, showing details of ovipositor; I -enlargement of D, showing details of ovipositor; J -enlargement of B, showing the tiny teeth on apices of tarsomere and claws.B, G, I, J moistened with 70% ethanol.Scale bars represent 1 mm in A, B, C, D, E, F; 0.5 mm in G; 0.2 mm in H, I, J.
Note: M -male; F -female; U -unknown sex; all the values are in mm.