Phylogenetic analysis
We conducted Bayesian Inference (BI) and Maximum Parsimony (MP) analyses using morphological data to place the fossil taxa and resolve the relationships within Sternorrhyncha. Therefore, we mainly included those morphological characters that were also discernible in the fossils that were selected. The data matrix used for the analysis consisted of 10 taxa (Fulgoromorpha taken as an outgroup, and 9 Sternorrhyncha ingroups, including extinct groups, see Supplementary information 1 Table S1) and 42 characters (see Supplementary information 1 Table S2). The characters were treated as non-additive and unordered. The list of characters and the nexus file containing the character matrix is available in Appendix (Tables S1 and S2).
The detailed results of phylogenetic analyses are presented in the Appendix. Both phylogenetic methods (MP and BI) were highly congruent in their resultant topologies (Supplementary information 1 Figs S1, S2a–c). According to the resulting phylogenies, the fossil described below forms a group of its own (Fig. 1), included in a clade of Psylliformes, related to Psyllodea and Aleyrodomorpha, but deserving of recognition as a different infraorder.
Phylogenetic position of Dingla shagria gen. sp. nov. on most parsimonius tree. Numbers at nodes represent posterior probabilities and bootstrap values. Image of planthopper Pyrops candelaria: Max Pixel Public Domain CC0 (modified); pincombeid Pincombea sp. redrawn from46; male scale insect: Pavel Kirillov CC-BY-SA2.0 (modified); Coccavus supercubitus redrawn from46; aphid Macrosiphum rosae: Karl 432 CC-BY-SA4.0 (modified); protopsyllidiid Poljanka hirsuta redrawn from47; liadopsyllid Liadopsylla apedetica redrawn from48; whitefly Aleyrodes proletella: Amada44 CC-BY-SA4.0 (modified); psyllid Trioza urticae photo by Jowita Drohojowska.
Systematic palaeontology
Order Hemiptera Linnaeus, 1758.
Suborder Sternorrhyncha Amyot et Audinet-Serville, 1843.
Clade Psylliformes sensu Schlee, 1969.
Dinglomorpha Szwedo & Drohojowska infraord. nov
Diagnosis
Fore wing with costal veins complex carinate (Pc carinate as in Psylliformes), ScP present as separate fold at base of common stem R + MP + CuA (unique character); common stem R + MP + CuA weakened at base (unique character); areola postica reduced (homoplasy with Aleyrodoidea); clavus present, with single claval vein A1. Hypandrium present as small plate (as in Psylliformes).
Dingloidea Szwedo & Drohojowska superfam. nov
Diagnosis
Fore wing membranous with modified venation—veins thickened, areola postica reduced; antennae 10-segmented; 3 ocelli present; stem MP present, connected with RP and CuA; abdomen widely fused with thorax; no wax glands on sternites.
Dinglidae Szwedo & Drohojowska fam. nov
urn:lsid:zoobank.org:act:D0A1C785-62D3-4E07-9A3B-FFAE3C13B704.
Type genus Dingla
Szwedo et Drohojowska gen. nov.; by present designation.
Diagnosis
Imago. Head with compound eyes narrower than thorax. Eyes entirely rounded, postocular tumosity present; lateral ocelli placed dorsolaterally, near anterior angle of compound eye in dorsal view, median ocellus present. Antennae 10-segmented, with bases in frons to compound eyes, rhinaria scarce (?). Pronotum in mid line longer than mesopraescutum. Fore wing with thickened costal margin, basal portion of stem R + MP + CuA weak, distal portion of stem R + MP + CuA convex, forked at about half of fore wing length, branch RA short; pterostigmal area thickened. Common stem MP + CuA short, branches RP, MP and CuA parallel on membrane. Rostrum reaching metacoxae. Metacoxa without meracanthus. Metadistitarsomere longer than metabasitarsomere, claws distinct, long and narrow, no distinct additional tarsal structures. Male anal tube long. Hypandrium in form of small plate, styli long, narrow and acutely hooked at apex.
Dingla Szwedo & Drohojowska gen. nov
LSID urn:lsid:zoobank.org:act:5053D386-4A13-445C-8036-9C69D885561F.
Type species Dingla shagria
Szwedo et Drohojowska sp. nov.; by present designation and monotypy.
Etymology
The generic name is derived from the adjective ‘dingla’ meaning ‘old’ in Jingpho language, which is spoken in Kachin state where the amber originates from. Gender: feminine.
Diagnosis
Vertex in mid line about as long as wide between compound eyes. Frons flat, widely triangularly incised at base. Antenna with 10th antennomere longer than penultimate one, widened, membranous apically, with terminal concavity. Pronotum about twice as wide as long. Mesopraescutum narrow, about as wide as pronotum; mesoscutum wide, with scutellar sutures not reaching anterior margin; mesoscutellum widely pentagonal. Fore wing with branch R forked anteriad of branch MP + CuA forking. Tip of clavus at level of MP + CuA forking. Hind wing with terminals RP and M subparallel and weakened in apical portion. Metafemur not thickened, metatibia without apical spines.
Dingla shagria Szwedo & Drohojowska sp. nov
LSID urn:lsid:zoobank.org:act:3EA05FB0-B783-4D7A-98EA-10B02F50B83D (Figures 2, 3).
Dingla shagria gen. sp. nov., holotype male, No. MAIG 5,979: body in dorsal view (a); body in ventral view (b); drawing of body in dorsal view (c); drawing of head in ventral view with clypeus (d); fore wing (e); apical antennomere (f); antennomeres 6th—10th (g); head in dorsal view (h); head in ventral view (i); head in lateral view (j); male genitalia in dorsal view (k); male genitalia in ventral view (l); male genitalia in lateral view (m); scale bars: 0.5 mm a, b, c, e; 0.1 mm f, g, k, l, m; 0.2 mm j, h, i; 0.25 mm d.
Dingla shagria gen. sp. nov., paratype male, No. MAIG 5,980: body in dorsal view (a); body in ventral view (b); head in ventral view with median ocellus (c); paratype male, No. NIGP172398, body in dorsal view (d); body in ventral view (e); fore tibia (f); paratype male, No. NIGP172399 body in dorsal view (g); body in ventral view (h); mid leg (i); hind leg (j); scale bars: 0.5 mm a, b, g, h, j; 0.4 mm d, e, f; 0.2 mm c; 0.25 mm i.
Etymology
The specific epithet is derived from the noun ‘shagri’ meaning ‘insect’ in Jingpho language spoken in the Kachin State, when the amber was collected.
Material
Holotype male. MAIG 5979, Paratype male, MAIG 5980, deposited in Museum of Amber Inclusions, Laboratory of Evolutionary Entomology and Museum of amber Inclusions, Department of Invertebrate Zoology and Parasitology, Faculty of Biology, University of Gdańsk, Gdańsk, Poland; paratype male NIGP172398, paratype male NIGP172399, deposited in Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China.
Locality and horizon
Kachin amber, Noije Bum hill, Hukawng Valley, Kachin State, northern Myanmar. Terminal Aptian/earliest Cenomanian.
Diagnosis
Pedicel (2nd antennomere) elongate, slightly thickened, 3rd antennomere longer than second and 4th; antennomeres 4th to 8th subequal in length. Protibia with row of thin setae in apicad half. Probasitarsomere about half as long as prodistitarsomere. Subgenital plate small, subquadrate, parameres long and narrow, parallel; about 3 times as long as wide at base, with hooked acute apex. Male anal tube tubular, slightly widening apicad, merely shorter than parameres.
Description
Male. Measurements (in mm): Total length 1.76 to 2.13; Body length total (including claspers) 1.76–2.13; Head including compound eyes width 0.37–0.52; head length along mid line 0.18–0.24; vertex width 0.2–0.26; Forewing length 1.32–1.79; forewing width 0.62–0.74; Claspers length 0.2–032; Antennomere 1st 0.04–0.08; antennomere 2nd 0.8–0.13; antennomere 3rd 0.08–0.16; antennomere 4th 0.06–0.12; antennomere 5th 0.06–0.09; antennomere 6th 0.06–0.1; antennomere 7th 0.06–0.09; antennomere 8th 0.0–0.09; antennomere 9th 0.06–0.09; antennomere 10th 0.0.8–0.01; Profemur + protrochanter cumulative length 0.26–0.46; protibia length 0.29–0.34; probasitarsomere length 0.06–0.09; prodistitarsomere length 0.08–0.13; mesofemur + mesotrochanter cumulative length 0.3–0.4; mesotibia length 0.36–0.4; mesobasitarsomere length 0.05–0.1; mesodistitarsomere length 0.13–015; metafemur + metatrochanter cumulative length 0.39–0.56; metatibia length 0.5–0.68; metabasitarsomere length 0.1–0.15; metadistitarsomere length 0.1–0.18.
Vertex about half as long as width of head with compound eyes; slightly narrower than wide at base; disc of vertex slightly concave; sutura coronalis absent. Scapus cyllindrical, longer than wide, pedicel slightly longer than scapus, barrel-shaped, wider than 3rd antennomere. Antennomere 3rd longer than 2nd antennomere (pedicel) antennomeres 5th to 9th subequal in length; antennomere 9th with subapical rhinarium; antennomere 10th (apical) longer then penultimate one, spoon-like widened apically, with rhinarium placed subapically. Median and lateral ocelli visible from above. Compound eyes large, not divided, with distinct, non-differentiated ommatidia; postocular protuberances narrow. Frons convex, with distinct triangular, concave median portion; median ocellus at margin with vertex; postclypeus and apical portion of loral plates distinctly incised to frons; postclypeus about twice as long as wide; anteclypeus tapering ventrad; lora semicircular, long, with upper angles slightly below upper margin of postclypeus, lower angles not exceeding half of anteclypeus length. Rostrum with apex reaching metacoxae; scapus short, wide, placed in distinct anterolateral concavity.
Pronotum large, as long lateral as in midline; about 2.6 times as wide as long in mid line; disc of pronotum convex; anterior margin convex, slightly protruding between compound eyes; posterior margins converging posteriad; posterior margin slightly concave. Mesopraescutum with anterior margin covered by pronotum, with anterior margin convex, lateral margins expanded posterolaterad, with posterior margin convex posteriomediad, slightly concave posterolaterad. Mesoscutum distinctly wider than long in mid line; anterior margin merely concave medially, lateral margins distinctly diverging posteriad, posterolateral angles acute, distinct, posterior margin W-shaped, with distinct median concavity; disc of mesoscutum convex with indistinct longitudinal concavities (apodemes? sutures?). Mesoscutellum narrow, with anterior margin acutely convex, lateral margins subparallel, posterior margin straight, disc of mesoscutellum concave, with posteromedian furrow. Metascutum and metascutellum not visible.
Fore wing about 2.5 times as long as wide; narrower at base, widening posteriad, rounded in apical margin; widest at ¾ of its length. Costal margin thickened, veins thick, distinctly elevated; basal portion of stem R + MP + CuA weak, distal portion of stem R + MP + CuA convex, forked at about half of forewing length, branch RA short; pterostigmal area thickened; common stem MP + CuA short, branches RP, MP and CuA parallel on membrane; areola postica absent; clavus present, with apex exceeding half of forewing, with single claval vein A1.
Hind wing about 0.8 times as long as forewing, with costal margin with two groups of regularly dispersed setae, basal group with seven longer and stiff setae and median group with 10 shorter, stout setae; terminals RP and M subparallel and weakened in apical portion.
Profemur and mesofemur subequal in length; protibia slightly shorter than mesotibia; pro- and metadistitarsomeres slightly longer than pro- and mesobasitarsomeres. Metacoxa without meracanthus; metafemur longer than pro- and mesofemur; metatibia distinctly longer than pro- and mesotibia; metadistitarsomere distinctly longer than metabasitarsomere; tarsal claws long, narrow, without arolium or empodium.
Abdomen with segments III to VIII almost homonomic in length, widely connected to thorax, subgenital portion narrowing. Subgenital plate small, subquadrate, parameres long and narrow, parallel; about 3 times as long as wide at base, with hooked acute apex. Male anal tube tubular, slightly widening apicad, merely shorter than parameres.
Source: Ecology - nature.com
