StemCell Therapy

Systematic paleontology

Aneuretopsychidae Rasnitsyn and Kozlov, 1990.

Burmopsyche gen. nov.

LSID (Life Science Identifier). urn:lsid:zoobank.org:act:15E5EB7B-004B-4AA7-A807-F2EE5C095EF0.

Etymology. The generic name is derived from Burma, the original English name of Myanmar, and -psyche, a typical suffix of generic names in Aneuretopsychidae. Gender: feminine.

Type species. Burmopsyche bella sp. nov.

Other species included. Burmopsyche xiai sp. nov.

Diagnosis. Head triangular in dorsal view, with ocelli raised on ocellar dome, with lateral and medial ocelli at edges of dome. Maxillary palps three-segmented. Antennae moniliform, slender, shorter than proboscis. Wings membranous, without spots or patterns of infuscation aside from pigmented pterostigma between termination of Sc and R1. Forewing ovate to subtriangular, relatively broad at base, with more or less distinct tornus and narrow but extended clavus; Sc reaching middle of wing length or beyond, with strong humeral vein and short distal branch to costal margin; R1 long and simple, braced to Rs by three crossveins; Rs separating from R1 proximally, in basal one-fifth of wing; Rs four-branched with two forks, of which Rs1+2 fork distinctly shorter than that of Rs3+4; stem of M elongate and straight, forking nearly at wing midlength, distal to Rs fork; M three-branched; connection between M and CuA obtuse Y-shaped, with prominent, strong upper arm (M5) and lower arm (base of CuA) much shorter, crossvein-like; CuA long, reaching at least three-fourths of wing length or more, straight in basal half, then broken lineshaped, with three crossveins to M; CuP long, braced to CuA by two strong crossveins, which form a characteristic hexagonal cell; three anal veins long and subparallel to CuP and to each other; A1 and A2 connected by strong, oblique crossvein forming pseudo-loop (apomorphy of family); A3 with long oblique crossvein to wing margin. Hind wing subtriangular, with wide folding vannus and well-developed jugum; hind wing R1 bearing a distinctive row of setal hooks along its medial third (potentially to link with posterior portion of forewing during flight); venation of remigium more or less similar to forewing. Legs slender, with setation of femora and especially tibiae arranged in transverse linear series, those of tibiae and tarsi longer and stiffer on inner surfaces; distitarsi slightly expanded with ventral surfaces flattened to faintly concave and covered in microtrichia; pretarsal claws large, broadly arched, and widely separated by broad and blunt unguitractor. Female cercus dimerous (two cercomeres).

Remarks. The new genus undoubtedly belongs to Aneuretopsychidae based on the opisthognathous siphonate mouthparts, the presence of pseudo-loop in the anal area of the forewing, and the well-developed jugum of the hind wing. It differs from other genera of Aneuretopsychidae by the relatively small body size [except for Aneuretopsyche minima; Rasnitsyn and Kozlov, 1990 (2)], three-segmented maxillary palps, and three-branched M in both fore- and hind wings.

In addition to the two species described here, there are two additional undetermined specimens embedded in the same piece of amber, at least one of which may represent a third species of the genus (see note S1 and fig. S1).

Burmopsyche bella sp. nov. (Figs. 1, A to J, and 2, A, C, D, and fig. S2)

(A) Burmopsyche bella gen. et sp. nov.; holotype, NIGP166158a (left); paratype, NIGP166158b (right). (B and C) Burmopsyche bella gen. et sp. nov.; holotype, NIGP166158a. (B) Habitus in lateral view. (C) Head and proboscis, enlarged from the red template in (B). (D to F) Burmopsyche bella gen. et sp. nov.; paratype, NIGP166158b. (D) Habitus in lateral view. (E) Head and proboscis, enlarged from the red template in (D). (F) Genitalia, enlarged from the green template in (D). (G to J) Burmopsyche bella gen. et sp. nov.; paratype, NIGP166159. (G) Habitus in dorsal view. (H) Line drawing of (G). (I) Proboscis. (J) Genitalia, enlarged from the green template in (G). (K) Reconstruction of mouthparts of Aneuretopsychidae (note the opisthognathous orientation of mouthparts). (L to P) Burmopsyche xiai sp. nov.; holotype, NIGP171685. (L) Genitalia, enlarged from the green template in (O). (M) Habitus in right lateral view. (N) Proboscis, enlarged from the red template in (M). (O) Habitus in left lateral view. (P) Proboscis, enlarged from the red template in (O). An, antennae; Ce1, cercus segment 1; Ce2, cercus segment 2; Fc, food channel; Ga, galea; Hy, hypopharynx; Mxp, maxillary palp. Scale bars, 2 mm (A, B, D, M, and O), 1 mm (G and H), 0.5 mm (C, E, and P), and 0.2 mm (F, I, J, L, and N). Photo credit: B.W., Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences.

(A) Burmopsyche bella gen. et sp. nov.; holotype, NIGP166158a. (B) Burmopsyche xiai sp. nov.; holotype, NIGP171685. (C) Burmopsyche bella gen. et sp. nov.; paratype, NIGP166158b; forewing. (D) Burmopsyche bella gen. et sp. nov.; paratype, NIGP166159; forewing. Scale bars, 1 mm.

LSID. urn:lsid:zoobank.org:act: 204B591A-2B9F-40F8-88C0-67D8C5FB5BFC.

Etymology. The specific epithet is from the Latin bellus, meaning beautiful.

Materials. NIGP166158, two adults preserved in an amber piece: holotype, NIGP166158a, a well-preserved incomplete specimen of unknown sex (distal part of abdomen is missing, legs partly preserved); paratype, NIGP166158b, a nearly complete female (forewings apices missing) embedded in the same piece of amber. Paratype, NIGP166159, a nearly complete female of somewhat smaller size, with right wings partially destroyed.

Locality and horizon. Noije Bum Village, Tanaing Town, northern Myanmar; lowermost Cenomanian, Upper Cretaceous.

Diagnosis. Body length 7 to 8 mm. Forewing length 8 to 10 mm; fork of Rs1+2 slightly shorter than stem; fork of Rs3+4 about 1.3 as long as its stem, longer than fork of Rs1+2; M1 simple, M2+3 with distinct fork; CuA long, in basal half straight, in distal half broken linelike, hexagonal cell as long as distal portion of CuA.

Description. Head spheroidal in lateral view, length 1.02 mm, width 0.77 mm, approximately half of greatest width of thorax, with prolonged siphonate mouthparts. Three prominent ocelli present on vertex (not visible in holotype, but well preserved in paratype NIGP166159); compound eyes large and rounded, occupying most of the head. Antennae connected to head in frontal region below ocelli; scapus poorly visible, elongate; pedicellus relatively large, apically dilated; flagellum moniliform, with approximately 49 to 50 flagellomeres (42 in paratype NIGP166159), up to three times wider than long; each flagellomere slightly flattened, with three setal whorls (proximal, medial, and distal), with medial whorl composed of the most prominent setae and positioned slightly closed to proximal whorl. Entire length of antenna (in holotype) 4.10 mm. Proboscis thin, flexible, consisting of three lobes, apparently lacking any terminal absorptive structures such as pseudolabellae. Hypopharynx greatly slender, about 3.80 mm long, needle-like at tip; galeal sheaths of proboscis as long as hypopharynx. Other head and mouthpart structures, including clypeus, labrum, and maxillary palpi not visible as preserved.

Thorax length 3.12 mm, width 1.79 mm, pronotum very small, mesonotum and metanotum subequal in size. Legs entirely covered with annulate setae (shown in paratype NIGP166158b); coxae and femora not shortened; tibiae slightly longer than femora; tarsi five-segmented, basitarsus longest, nearly as long as remaining tarsomeres combined; pretarsi with two curved claws and unpaired arolium. Abdomen (based mainly on the paratype NIGP166158b) stout, slightly shorter than wings, preabdominal segments obscured, postabdominal (VI to XI) segments tapering apically. Cerci dimerous, distal cercomere filamentary (in paratype NIGP166158b, dorsally curved as preserved; in NIGP166159, straight), with dense, nonisometric setae.

Forewing oblong-subtriangular, with distinct tornal angle at about 0.7 wing length with blunt apex. Forewing length about 10.5 mm, maximum width 4.15 mm, length/width ratio 2.5:1. Sc long, straight, reaching beyond middle of wing, with strong humeral vein at base and short distal branch to costal margin, in apical portion connected to R1 by short crossvein sc-r1. R1 long, simple, in distal third gently curved toward wing apex. Pterostigma well developed, extensive, at base extending toward RS1+2 along r1rs1+2 crossvein, in distal part bounded by R1. Rs separating from R1 in basal one-fifth of wing, four-branched, forking somewhat before middle of wing; fork of Rs1+2 slightly shorter than stem; fork of Rs3+4 about 1.3 as long as its stem, longer than fork of Rs1+2; stem of Rs3+4 with distinct curve at first crossvein to M. Base of medial vein braced to base of R1 by short crossvein r-m. M forking at center of wing distal to Rs fork, having three branches meeting wing margin, M1 simple, M2+3 with distinct fork. CuA long, ending nearly at three-fourths of wing length just beyond tornus, in basal half straight, in distal half broken linelike, braced to CuP by two long crossveins, of which distal one is strongly oblique; hexagonal cell as long as distal portion of CuA. CuP long, ending somewhat before or beyond level of Sc apex, apical portion of CuP after distal cua-cup crossvein sharply curved toward wing margin. Three anal veins long and subparallel to CuP and to each other, A2 reaching middle of wing length, at level of M fork; A1 and A2 connected by strongly oblique crossvein, which is relatively short and located slightly distal to cubito-medial Y-vein; A2 markedly curved at this point. Space posterior to third anal vein wide, crossed by oblique crossvein from A3 to wing margin. Crossveins numerous, concentrated mainly in distal half of wing, arranged in three more or less distinct rows; crossveins mostly forwardly inclined or upright.

Hind wing delicate, somewhat smaller than forewing, subtriangular in shape, with distinct apical angle and smooth tornus. Hind wing length about 8.45 mm, width of remigium about 3.6 mm. Bases of all main veins close to wing base. Costal space narrow, Sc long and simple, with short apical crossvein to R1. Pterostigma not developed. Vein R1 with distinctive row of long, anteriorly directed hooked setae. Branching pattern of Rs and M and arrangement of crossveins similar to forewing. Spaces between cubital and anal veins expanded, crossed by long crossveins: three in intercubital space, one in space CuP-A1, and one in space A1-A2 (in addition to two short basal crossveins a1-a2). Veins CuP and A1 fused for short distance before separating close to wing base. Vannus folded twice, vannal (claval) fold apparently located behind A1; vein A3 and jugal (anal) fold not discernible, as well as jugal veins, if present.

Burmopsyche xiai sp. nov. (Figs. 1, L to P, and 2B)

LSID. urn:lsid:zoobank.org:act: C925539E-A099-47F8-A9AA-4537153887DC.

Etymology. The specific name honors Mr. Fangyuan Xia, director of the Lingpoge Amber Museum in Shanghai.

Holotype. NIGP171685, a nearly complete female with distal parts of antennae missing.

Locality and horizon. Noije Bum Village, Tanaing Town, northern Myanmar; lowermost Cenomanian, Upper Cretaceous.

Diagnosis. Body length about 12 mm. Forewing length about 9 mm; fork of Rs1+2 about as long as its stem; fork of Rs3+4 about 2.5 as long as stem; M1+2 with deep fork, M3 simple and more or less smooth at crossvein junctions; CuA long, ending nearly at 0.8 of wing length just beyond tornus, hexagonal cell relatively short, half as long as distal portion of CuA.

Description. Body length 12.1 mm, thorax length 3.0 mm, abdomen length 7.2 mm. Head length 1.9 mm, seriously distorted; with prolonged siphonate mouthparts. Most of the head and body structures obscured. Preserved part of antenna, 1.63 mm. Maxillary palpi short, about 0.53 mm long, three-segmented. Proboscis about 3.08 mm long.

Thorax length 3.07 mm, width 0.62 mm. Legs with short, dense setae (visible only in hind tibiae and tarsi), hind legs distinctly longer than fore and mid legs. Length: profemur 1.31 mm, protibia 1.92 mm, protarsus 1.48 mm; mesofemur 1.54 mm, mesotibia 1.97 mm, mesotarsus 1.64 mm; metafemur 2.28 mm, metatibia 2.08 mm, metatarsus 1.81 mm; tarsus pentamerous, basitarsus longest, but shorter than other four tarsomeres combined; pretarsus obscured. Abdomen slender, tapering apically, length 7.16 mm, with two cercomeres.

Forewing ovate, with rounded apex, claval area and posterior (inner) margin not visible (overlapped by hind wing), tornal angle unclear. Forewing length 8.95 mm, maximum width 4.3 mm, length/width ratio 2:1. Sc long, straight, reaching beyond middle of wing, with strong humeral vein at base possibly having a short vein to costal margin at about middle of wing length; Sc near apex connected to R1 by short crossvein sc-r1. R1 long, simple, gently curved toward apex of wing. Pterostigma not preserved. Rs separating from R1 in basal one-fifth of wing length, forking somewhat before middle of wing; fork of Rs1+2, about as long as its stem; fork of Rs3+4, about 2.5 as long as stem, longer than fork of Rs1+2; stem of Rs3+4 with distinct curve at first crossvein to M. Medial vein fork near center of wing somewhat distal to Rs fork, having three branches meeting wing margin, M1+2 with deep fork, M3 simple and more or less smooth at crossvein junctions. Base of CuA slightly inclined backward, forming an obtuse Y-vein with long M-arm (M5). CuA long, ending nearly at 0.8 of wing length just beyond tornus, in basal half straight, in distal half broken lineshaped, braced to CuP by two relatively short crossveins; hexagonal cell relatively short, half as long as distal portion of CuA. CuP long, ending at least at the level of Sc apex or beyond; apical part of CuP not visible but must be quite long and without sharp curve to wing margin. A1 long and subparallel to CuP, its distal portion not visible; other anal veins visible only at their bases. A1 and A2 connected by long, strongly oblique crossvein possibly located somewhat before cubito-medial Y-vein; portion of A2 distal to this crossvein not visible. Space posterior to A3 narrow, crossed by short oblique crossvein from A3 to wing margin. Crossveins numerous, concentrated mainly in distal half of wing, indistinctly arranged in three rows, mostly upright or inclined backward.

Hind wing subtriangular in shape, with obtuse apex and smooth tornus; length about 7.6 mm, width of remigium 3.35 mm. Both hind wings slightly bent at costal margin toward wing plane, such that Sc not visible. Pterostigma not developed. Branching pattern of remigium more or less similar to forewing, except for more expanded intercubital space crossed by three long crossveins instead of two as in forewing. Vannus folded, details of venation not discernible.

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Mouthpart homologies and life habits of Mesozoic long-proboscid scorpionflies - Science Advances

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