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Systematic Paleontology

Order Permopsocida Tillyard, 1926 sensu et stat. nov.

Included families

Permian to Liassic (with some doubt) Psocidiidae Tillyard, 1926, Permian Permopsocidae Tillyard, 1926, and Jurassic to earliest Upper Cretaceous (with a problematic Permian taxon) Archipsyllidae Handlirsch, 1906, incl. the new archipsyllid genus Psocorrhyncha.

Emended diagnosis

(Figs 1 and 2, Figs S2–6). Head somewhat flattened and depressed; clypeus not strongly swollen; mandibles elongate, with a strong molar plate and a long incisor; four maxillary palpomeres; three labial palpomeres; paraglossae long and sclerotized, appearing as half tubes; paraclypeal lobes present; median part of anteclypeus membraneous; gena divided into two parts by a furrow; ocell-ocular distance < inter-ocellular distance; tarsi four-segmented; fore- and hind wings of similar size, shape, and venation; subcosta posterior ScP present; radius posterior RP two-branched; median vein M normally four-branched (five-branched in one genus); areola postica present; two anal veins present; pterostigmata between costa C and radius anterior RA, of identical shape in all wings; RA forming a pronounced posterior curve below pterostigmata; radius R with a pronounced angle at level of base of M; M + CuA basally fused with R, separating from radius far from wing base; long crossvein cua-cup present between cubitus posterior CuP and cubitus anterior CuA; abdomen with strong basal constriction; cerci absent; female ovipositor well-developed and sclerotized.

Family Archipsyllidae Handlirsch, 1906.

Psocorrhyncha burmitica gen. et sp. nov

Type species of genus

Psocorrhyncha burmitica sp. nov.


Male holotype NIGP161473 and male paratype NIGP161474 at Nanjing Institute of Geology and Paleontology (NGIP, Academia Sinica, China); female allotype SMNS Bu-157 and female paratype SMNS Bu-135 at State Museum for Natural History in Stuttgart (SMNS, Germany).

Type locality

Hukawng Valley, Kachin State, Myanmar (Burma). The exact outcrop among the various amber mines in this valley is unknown, because the specimens were acquired from traders.

Type horizon

Burmese amber (Burmite)11,12, Earliest Upper Cretaceous, earliest Cenomanian, absolute age 98.79 ± 0.62 million years ago (mya) established by U-Pb dating of zircons from the rind of the unprocessed amber13. Nuclear magnetic resonance spectra and the presence of araucaroid wood fibers in amber samples indicate an araucarian (possibly Agathis) tree as source for the resin14.


The generic name refers to the resemblance of this taxon with the Psocodea and its affinities with the Hemiptera (old name Rhynchota). The gender of the name is feminine. The specific epithet refers to the country of origin.


Forewing ScP short, ending on C at level of base of M + CuA and re-emerging distally as a faint phantom-vein ending on R (the fusion of forewing ScP with C is a character present in the other Archipsyllidae as putative synapomorphy, but it is re-emerging as a distinct vein in these genera, instead of being phantom-like); hind wing ScP fused with R.


Psocorrhyncha burmitica is the youngest fossil record of Archipsyllidae. A redescription of the enigmatic Permian psocidiid species Dichentomum tinctum Tillyard, 1926, and a discussion of all other taxa previously attributed to Permopsocida is provided online in the Supporting Information (S1 Text).


The description is based mainly on holotype NIGP161473, completed by information from the three other fossils.

Body 2.4 mm long between apex of abdomen and base of antennae, and glabrous; head with rostrum 0.9 mm long; head capsule 0.4 mm long; occiput abruptly bent; compound eyes well developed, 0.28 mm wide and well separated; dorsal part of head between compound eyes divided in two parts by weak furrow: a posterior part (looking like a corypha of Fulgoromorpha15,16), divided into two pronounced lobes each bearing a smooth but pronounced lobe, separated by a median sulcus; and a vertical anterior part (looking like a metopa of Fulgoromorpha15) anterior of compound eyes, bearing two well-separated lateral ocelli, each being closer to eye than to other ocellus; anterior ocellus positioned far from lateral ocelli, on a line separating dorsal part of head from frons (Fig. 2g, Fig. S3e); frons narrow, as long as narrow sclerotized postclypeus, which is separated from anteclypeus by a furrow; anteclypeus short, 0.4 times shorter than labrum, composed by two lateral parts (paraclypea), rounded elongate, more sclerotized and higher than membranous median part (Fig. 2b,g, Fig. S3b); mouthparts hypognathous but clearly movable relative to head capsule (as documented by forming different angles with head capsule in different specimens) (Fig. 1a, Fig. S3a, Fig. S4a,b); labrum elongate, 0.28 mm long, three times as long as wide, apically spatulate and rounded, flat and thin, with small apical setae; mandibles elongate, 0.29 mm long and 0.09 mm wide at base (paratype specimen NIGP161474), three times as long as wide at base, with a broad base and distal two-thirds narrow; molar plates well developed bearing three distinct teeth on left mandible and only two on right mandible; incisor far from molar plate, with a strong apical tooth and two smaller basal teeth (Fig. 2f, Fig. S2a,h); anterior condyle of mandible connected with latero-basal angle of paraclypeus (Fig. S2a,e); posterior condyle connected to distal margin of gena; gena large and broadly quadrangular with transverse furrow dividing it obliquely, anterior part distinctly concave, bearing condyle of mandible; posterior part more convex than anterior part (Fig. 2e,g), apparently bearing a small sensilla along its posterior margin below compound eye (paratype specimen NIGP161474); subgena between anterior part of gena and mandible; postgena between gena and maxilla (Fig. S2b); maxillary palps long with four palpomeres (Fig. 1g, Fig. 2a,b,g, Fig. S4b), apical palpomere long, 0.18 mm long, subapical palpomere 0.07 mm long, shorter than apical palpomere and with an apical bevel cut, basal palpomere short, 0.18 mm long, second palpomere as long as apical one, 0.17 mm long; cardo and stipes well separated, articulation of maxilla visible17; lacinia long, as long as galea, spoon-like, i.e., broadened in its distal part but apically narrowed and without subapical tooth, detached from stipes and deeply inserted into head (Fig. 2c,d,g, Fig. S2c,d); galea broader than lacinia, with distal half broadened, apex bearing short setae, distally ending close to apex of mandible, apically serving as guide for mandibles due to ‘T-profile’ cross-section (Fig. S2c,d,g,); three labial palpomeres (Fig. 2c,d,g), with basal palpomere shortest, 0.05 mm long, second palpomere 0.1 mm long, third palpomere 0.09 mm long; labium with elongate prementum and half-tube-shaped paraglossae as guide for laciniae; antennae inserted well below compound eyes, well separated, with a subquadrate scape 0.11 mm long and 0.10 mm wide, pedicel as long as scape but narrower (Fig. S2g, Fig. S3e, Fig. S4b); 14 elongate flagellomeres, finely annulated, with individual lengths decreasing progressively toward apex; first, second, and third flagellomeres bearing an apical, elliptical flat sensilla (Fig. S5a,b), and first flagellomere bearing also a basal one; membraneous zone between flagellomeres simple, without mechanism for rupturing antennae (as in Psocodea18); no sclerotized ring at base of first flagellomere in cavity of pedicel; scape inserted on head capsule by a dicondylic articulation (acute lateral antennifer and weaker, median articulation point on head capsule, see Fig. S2g); no cephalic trichobothria.

Prothorax developed as narrow neck bearing an anterior sclerotized ring with small indentations and posterior part desclerotized (Fig. 2a); mesothorax and metathorax higher than prothorax, separated by subvertical pleural furrow; mesothoracic scutum deeply concave; wings inserted high on meso- and metathorax; tegula present at forewing base.

Legs long and thin; profemur 0.5 mm long, protibia 0.7 mm long, protarsus 0.4 mm long; mesofemur 0.5 mm long, mesotibia 0.7 mm long, mesotarsus 0.4 mm long; metafemur not enlarged, 1.3 mm long, 0.1 mm wide, metatibia 0.9 mm long, 0.03 mm wide, metatarsus 0.6 mm long; tibiae with two strong apical spurs and a row of spines; 4-segmented tarsi (Fig. S4e,g); tarsomeres bearing a row of spines, tarsomeres without plantulae; strong apical pretarsal claws without basal tooth, a fleshy and broad arolium present between pretarsal claws (Fig. S4f).

Forewing and hind wing elongate, of nearly same size and shape; forewing 2.6 mm long, 0.7 mm wide; ScP ending on costal margin C 0.5 mm from wing base, and re-emerging 0.3 mm distally to reach radius R as a phantom-vein (Fig. S6c); area between R and C broad, 0.17 mm wide; R, M, and CuA fused into a common stem at wing base, making a weak posterior curve for 0.52 mm; then M + CuA and R separating, with R and basal stem R + M + CuA forming a pronounced angle at this point (Fig. 1b,c); RP and RA separating 0.15 mm distal of base of M + CuA; convex RA with pronounced posterior curve surrounding darkly pigmented pterostigma, 0.42 mm long and 0.14 mm wide, pterostigma basally delimited by a vein (Fig. S6b); a crossvein perpendicular to RA and to RP exactly below middle of pterostigma; concave RP with only one distal fork, 1.3 mm from its base; M and CuA separating immediately distal of point of re-emergence of M + CuA, or CuA emerging directly on stem R + M + CuA just basal of base of M (depending on specimen); neutral stem of M long, 0.85 mm long before first fork; anterior branch of M with a deep fork distally and branches ending near wing apex (but in paratype specimen NIGP161474, this vein is simple in one wing while it is forked in the second); posterior branch of M with a more open fork and shorter branches ending on posterior wing margin; convex CuA short before crossvein cua-cup terminates on it, cua-cup aligned with distal part of CuA; distal part of CuA long, 0.5 mm long before areola postica; areola postica long and narrow, parallel to posterior wing margin, with CuA1 curved and CuA2 short; cua-cup weaker than CuA and M, 0.40 mm long between base of CuP and CuA (Fig. 1b,c); concave CuP weakly curved and simple; two convex simple anal veins basally curved. Forewing articulation partly visible in specimen NIGP161473: humeral plate (HP) and basisubcostale (BSc) united but well separated from basiradiale (BR) and second axillary sclerite (2Ax) by two deep furrows that extend transversely from wing base and tegula (Fig. S6a).

Hind wing 2.3 mm long, 0.71 mm wide; nearly identical to forewing, with following differences: wing narrower, with narrower pterostigma; ScP longer than in forewing, ending on R 0.52 mm from wing base (Fig. 1d); area between R and costal margin C much narrower than in forewing, 0.11 mm wide; cua-cup weak, ending on M + CuA; stem of M + CuA relatively long distal of its separation from radius, 0.14 mm long; areola postica very faint with CuA1 phantom-like.

A strong constriction between thorax and abdomen present due to small first abdominal segment, bearing small lateral lobes (Fig. 1a, Fig. S3a,c); sternum I not visible. Abdomen ca. 1.3 times as long as thorax plus head; abdominal terga short and of nearly same length; cerci absent.

Male appendages symmetrical (Fig. S5c), with a large, sclerotized spoon-like hypandrium; a short epiproct partly hidden by a fecal pellet (composed of pollen) extended from anus, and two, long subvertical paraprocts, 0.23 mm long, with a subbasal hook, a trichobothrial field on external surface of epiproct; aedeagus large, 0.25 mm long, broadly triangular, with three small, lateral spines; endosoma extruded exhibiting ductus ejaculatorius and gonopore II; hypandrium (sternite IX) long, spoon-like, 0.37 mm long; some sperm is visible in the abdomen.

Female ovipositor curved upwards (Fig. S3d,f), with ventral valvulae (gonapophyses VIII) with ventral margin bearing small denticles and a dorso-apical part bearing a raking structure; dorsal valvulae (gonapophyses IX) triangular, narrow, and elongate, ending with a small upward denticle, and less sclerotized than ventral valvulae; gonoplacs broad and weakly sclerotized, with an apical lobe; gonocoxites VIII large, broadly quadrangular in an anterior position; gonocoxite IX triangular and small at base of gonoplacs; epiproct and paraprocts of same length, shorter than gonoplacs, pointed at apices; tergum X longer than tergum IX; laterotergite VIII with a distal membraneous zone; subgenital plate with two broad arms; sternum IX reduced; tergum IX + X narrow; trichobothrial field on a gibbosity of epiproct.

Phylogenetic analysis

We conducted a cladistic analysis using morphological data to correctly place crucial fossil taxa and resolve the relationships within Acercaria (Hypoperlidae, Psocodea, Permopsocida, Thripida, and Hemiptera). Therefore, mainly those morphological characters that are also discernible in the fossils have been selected. The data matrix used for the analysis consists of 16 taxa (four outgroup taxa in Polyneoptera and Holometabola, and 12 of the ingroup, see Table S2) and 62 characters (see Table S3). The characters were treated as non-additive and unordered. The matrix was constructed with WinClada ver. 1.00.08 (see Table S4) and analysed with the parsimony software package TNT19. Using New Technology search method with default parameters resulted in a single topology, presented in Fig. S12, and the resulting acercarian phylogeny in Fig. 3. Its length is 100 steps, CI = 0.730, and RI = 0.833. The Bremer support of subclades are indicated in Fig. S12. This tree is slightly better resolved than the strict consensus tree of the two most parsimonious trees resulting from Traditional search method with default options. It supports a monophyletic Acercaria with Hypoperlidae as sister group of all other Acercaria; Permopsocida resolves as sister group of Thripida + Hemiptera (Condylognatha), and Psocodea as sister group of Permopsocida + Condylognatha. The new fossil genus and species Psocorrhyncha burmitica is recovered within the monophyletic Permopsocida as sister group of Archipsylla.

The results of our phylogenetic analysis agree with most other recent studies3,5 in the relationships among the extant acercarian orders. However, there is one important difference to the most recent, extensive phylogenomic analysis of insects by the 1Kite project2, which proposed a paraphyletic Acercaria with Psocodea as sister group of Holometabola. The authors of the 1Kite project remarked, ‘convincing morphological features and fossil intermediates supporting a monophyly of Acercaria are lacking’. Contrarily to the op cite analysis, Acercaria monoplyly is well recovered and supported by a large set of morphological autapomorphies, even if some of these characters are unknown in some fossil groups like Permopsocida or absent in early stem group representatives like Hypoperlidae1,10. These characters include the following: postclypeus large and with large cibarial dilator muscles; asymmetrical mandibles; laciniae transformed into stylet-like, slender rods, detached not directly connected to stipes and retractile, withdrawn deep into head capsule (a complex and strong character!); labial palps reduced (max. three palpomeres) or lost; cibarial pump (with similar sclerites and muscles especially in Psocodea and Thysanoptera); presence of an areola postica at least in forewings (character subject to reversions); neutral crossvein cua-cup between concave CuP and convex CuA, weaker than CuA; a common stem R + M + CuA at wing base; 1st abdominal sternum strongly reduced or absent; cerci completely reduced (one-segmented in Hypoperlidae); abdominal ganglia concentrated in a single ganglionic mass; max. four malpighian tubules; biflagellate spermatozoa; and acrosome of spermatozoa without perforatorium (last three characters not observable in fossils). We therefore assume that the 1Kite result concerning the phylogenetic position of Psocodea could be due to a systematic error (e.g. long branch attraction) or methodological artefact.

Remark. The reduction of the number of tarsomeres to max. four is no longer an acercarian apomorphy as there are five in Hypoperlidae.

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