Order Hemiptera Linnaeus, 1758 Suborder Fulgoromorpha Evans, 1946 Superfamily Fulgoroidea Latreille, 1807 Family Inoderbidae Shcherbakov et Emeljanov, 2021
Revised diagnosis. Rather small planthoppers, with habitus variable
(derbid-like and tropiduchid-like). Lateral carinae of vertex and frons
foliaceous, vertex and frons without median carina; fastigium may be
developed; clypeus protruded, without median carina but with two lateral
carinae. Pronotum with disc elevated, without median carina; postocular
carinae may be developed; mesonotum in shape of diamond, with weakened or
without distinct median and lateral carinae, mesoscutellum not distinctly
separated. Tegmen with clear submarginal pterostigma, costal area wide with
several transverse veinlets, not continued with apical cell series; clavus
closed; stem MP forked much later than stem CuA with a few terminal branches
(three), stem MP area reduced; stem CuA forked very early (at same level of
fork of ScP
Body and appendages covered with filamentous wax, dorsum with numerous
wax plates (in females only?) producing copious wax strands, especially long
on posterior abdomen, wings not folded over abdomen in repose but held
laterally, head narrow, frons with small fastigium at midline, antennae with
scape and pedicel much elongate, tegmen slightly longer than body, narrowed
to base, transparent, basal cell subtrapezoidal and much elongated, costal
area with fewer than 10 transverse veinlets, CuA Body and appendages not covered with filamentous wax, wings folded
tectiform over abdomen in repose, head wide, frons without fastigium,
antennae with scape and pedicel slightly elongate, tegmen much longer than
body, very large and broad, translucent, basal cell relatively short,
triangular, costal area with more than 10 transverse veinlets, CuA
Subfamily Ingensalinae Luo, Song et Szwedo subfam. nov. urn:lsid:zoobank.org:act:09A85EE6-624C-4F80-B9D2-DE34027E131A
Diagnosis. Body and appendages not covered with filamentous wax. Wings hold
tectiform over abdomen in repose. Head wide, vertex and frons wider than
long, frons short, only visible in ventral view, without fastigium; antennal
pedicel barrel-like, with sensory plaque organs. Tegmen much longer than
body, very large and broad, about twice as long as wide, translucent,
shallowly tectiform, clavus with blunt apex, tornus well developed, basal
cell relatively short, triangular; costal area with more than 10 transverse
veinlets, CuA forked at same level of ScP
Genus urn:lsid:zoobank.org:act:C813BB00-005E-406C-BCCF-079BF1BA223D
The generic name is derived from the combination of two words
from the Latin language:
Type species only.
Head with compound eyes about
Holotype of
Age and occurrence. Mid-Cretaceous (early Cenomanian); amber from Kachin State, northern Myanmar.
Detailed photographs of the head, pronotum, and mesonotum of
(Figs. 1–4) urn:lsid:zoobank.org:act:A5922B96-2532-4454-973E-BBCFE7534C1C
The specific name is dedicated to Fangyuan Xia, Director of the Lingpoge Amber Museum in Shanghai, for his contribution to the study of this amber specimen.
Detailed photographs and line drawing of tegmen and hind wing of
Holotype. Specimen no. BA19006, deposited in the Lingpoge Amber
Museum in Shanghai. To avoid any confusion and misunderstanding, all authors
declare that the fossil reported in this study was collected before 2016 and was
not involved in armed conflict and ethnic strife in Myanmar. This specimen
is deposited in a public collection permanently that is always accessible to
scientists by contacting the corresponding author, in full compliance with
the International Code of Zoological Nomenclature and the instructions of
the International Palaeoentomological Society (Ride et
al., 1999; Szwedo et al., 2020). Burmese amber, cabochon,
Locality and horizon. Kachin amber, near Tanai Village in the Hukawng Valley of northern Myanmar, lower Cenomanian (mid-Cretaceous).
Vertex about 4 times wider than long at mid line, trigons
absent, frons about 3 times wider than long at mid line. Pronotum almost
as wide as mesonotum, mesonotum in mid line 3 times as long as pronotum
in mid line. Tegmen basal cell about twice as long as wide, with 12 transverse veinlets in costal area, ScP single, RA with two terminals, RP
with four terminals, MP with three terminals, CuA with six terminals; tegmen
with one prenodal
Adult (male). Tegmen uniformly brownish with darker brown
“pseudostigma” and larger, semicircular, transparent, light area posteriad
of “pseudostigma”. Total length ca. 7.2 mm (wings
Pronotum 0.42 mm long in mid line and 1.70 mm wide, saddle-shaped, with disc elevated, postocular carinae distinct, no median carina, anterior margin exceeding level half of compound eyes length, arcuate, posterior margin shallowly arcuate (Fig. 2a, b). Mesonotum 1.22 mm long and 1.73 mm wide, slightly convex; with three indistinct, incomplete carinae; no distinct mesoscutellum (Fig. 2a, e).
Tegulae relatively large (0.40 mm long and 0.25 mm wide), carinate (Fig. 2f).
Tegmen (Fig. 3a–f) very large, translucent, with clear dark and pale
pterostigma, shallowly tectiform (Fig. 3c). Tegmen broadest near half of its
length, about twice as long as wide (6.55 mm long, 3.10 mm wide at broadest
point). Costal area developed. Anterior angle widely arcuate, costal margin
arcuate, anteroapical angle widely arcuate, apex rounded, posteroapical
angle widely arcuate, shifted slightly basad than anteroapical angle, tornus
well developed, straight, not extending beyond line of posterior claval
margin Costal margin slightly thickened, appendix narrow, shifting to
transversely corrugated appendix from apex of costal area to claval apex;
clavus closed with apex blunt, at about
Hind wing (Fig. 3g, h) membranous, transparent, shorter than tegmen, at
least 4.74 mm long and 1.96 mm wide. Costal margin sinuate, anteroapical
angle broadly rounded, posteroapical angle also rounded, like apical margin.
Stem ScP
Proleg (Fig. 4a, b, d, f): procoxa elongate, profemur laterally compressed, carinate, with margins covered with numerous very short setae, at least 1.17 mm long and 0.27 mm wide; protibia narrow and long, about as long as profemur, subquadrate in cross section, with margins carinate, and covered with numerous very short setae along margins, 1.14 mm long and 0.15 mm wide; protarsomeres partly covered, I 0.17 mm long, widened towards apex, II 0.15 mm long, indistinct, III 0.22 mm long, cylindrical; without distinct claws and arolium. Mesoleg (Fig. 4c–f): mesofemur similar to profemur, flattened, carinate, with margins covered with numerous very short setae, 1.41 mm long and 0.22 mm wide; mesotibia slightly longer than protibia, narrow and long, subquadrate in cross section, with margins carinate and covered with numerous very short setae, 1.27 mm long and 0.18 mm wide; mesotarsomeres partly covered; without distinct claws and arolium (Fig. 4f). Metaleg (Fig. 4g–j): metafemur compressed laterally, shorter than metatibia, covered with numerous very short setae, more robust than metatibia; metatibia narrow and long, carinate, and covered with numerous very short setae, thinnest near the mid part, then widened towards apex, with at least five apical teeth, lacking lateral armature, 1.71 mm long and 0.09 mm wide at thinnest part; basimetatarsomere slightly longer than combined length of mid and apical metatarsomeres, cylindrical, dorsally deeply excavated and widened apicad, with a few (5?) apical teeth, 0.72 mm long and 0.08 mm wide near the mid part; midmetatarsomere shorter, about half of basimetatarsomere length, excavated dorsally, widened apicad with a few (5?) apical teeth, 0.52 mm long and 0.07 mm wide near the mid part; apical metatarsomere narrow, gradually widened apicad, 0.24 mm long and 0.03 mm wide; arolium present, claws not visible.
Detailed photographs of legs and abdomen of
Abdomen with nine segments, wide, flattened, 2.27 mm long including terminal, and 1.83 mm wide at broadest part, pregenital segments gradually contracted; posterior margin of last pregenital segment strongly arched (Fig. 4k). Male pygofer tubular, slightly longer than wide in ventral view (0.53 mm long and 0.49 mm wide), genital styles fused medially, male anal tube subquadrate and long, exceeding apices of genital styles in length (Fig. 4i). Female unknown.