Research Article |
Corresponding author: Manuel Brazidec ( manuel.brazidec@gmail.com ) Academic editor: Ricardo Pérez-de la Fuente
© 2023 Manuel Brazidec, Frédéric Legendre, Vincent Perrichot.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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The Lancepyrinae are an extinct subfamily of Bethylidae known exclusively from Cretaceous amber deposits of Lebanon, Spain, Taimyr and Myanmar. In this study, we describe and illustrate four new genera and five new species of lancepyrine wasps from the Albian of Hkamti and late Albian-early Cenomanian of Kachin (Myanmar): Azepyris delamarrei gen. et sp. nov., Burmapyris ohmkuhnlei sp. nov., Gwesped groehni gen. et sp. nov., Paralanceis chotardi gen. et sp. nov. and Yunbayin rossei gen. et sp. nov. These taxa not only highlight the taxonomic diversity of the Lancepyrinae during the mid-Cretaceous but they also reveal the morphological disparity of the subfamily. To establish the phylogenetic relationships of these fossils and to check the monophyly of the Lancepyrinae, we add them to a pre-existing morphological matrix and perform a cladistic analysis. We retrieve the subfamily as poorly supported yet monophyletic, with the newly described taxa deeply nested in it. A key to the genera of Lancepyrinae is provided. Finally, we erect the subfamily Cretabythinae subfam. nov. for the genera Cretabythus Evans, 1973, Holopsenelliscus Engel, 2019 and Megalopsenella
Chrysidoidea, Bethylidae, fossil record, taxonomy, Kachin amber, Hkamti amber
With no less than 2900 extant species, whose biology is assumed to be parasitic on lepidopteran and coleopteran larvae, the Bethylidae are the largest family within the Chrysidoidea (
In this paper, we report and describe various new genera and species of Lancepyrinae from mid-Cretaceous Burmese amber, and we explore the monophyly of the subfamily. We also present an illustrated key to the genera of Lancepyrinae.
Three of the amber pieces studied herein originated from the deposits of Noije Bum, in the Hukawng Valley of Kachin State, northern Myanmar (see map in
Specimen IGR.BU-062 was acquired by one of us (V.P.). Specimens IGR.BU-060 and IGR.BU-063 were donated by Dr Christoph Öhm-Kühnle (Herrenberg, Germany) and specimen IGR.BU-061 by Corentin Jouault (MNHN, Paris, France) to the Geology Department and Museum of the University of Rennes, France (IGR). This material is housed in the amber collection of the IGR. Specimen GPIH.5058 (CCGG nº11341) was loaned for study by Carsten Gröhn (Glinde, Germany) and is deposited in the amber collection of the Leibniz Institute for the Analysis of Biodiversity Change, Hamburg, Germany (GPIH, Carsten Gröhn coll. CCGG). Specimen
The amber pieces have been trimmed and polished to facilitate the observation of the specimens, using thin silicon carbide papers on a grinder polisher (Buehler MetaServ 3000). Observations and photographs were conducted with a Leica DMC4500 camera attached to a Leica M205C stereomicroscope. All images are digitally stacked photomicrographic composites of several focal planes, which were obtained using Helicon Focus 6.7. Adobe Illustrator CC2019 and Photoshop CC2019 software were used to compose the figures and ImageJ 1.53 for measurements (
The description of the characters follows the nomenclature of
In order to test the monophyly of the Lancepyrinae, all genera considered as belonging to this subfamily—either in this contribution or in previous works (i.e.,
Data matrix used in the phylogenetic analyses (– = inapplicable; ? = missing).
Taxa | Characters | ||||||
1–10 | 11–20 | 21–30 | 31–40 | 41–50 | 51–60 | 61–69 | |
Clystopsenella | 2010001101 | 0010001000 | 0010000100 | 0010001000 | 1110000100 | 0001001010 | 001101100 |
Archaepyris | ??1?0??00? | 1????000?0 | 01???????? | ??10?????? | 1110010100 | 1101101??? | 00?1?0??? |
Azepyris | 1010111200 | 001??00000 | 01001-0020 | 0111001000 | 1110010100 | 1101101?01 | 0001?110? |
Burmapyris | 1010001200 | 101??00000 | 01001-0020 | 0111000000 | 1110010100 | 1101101?01 | 0001?110? |
Cretepyris | 0010011220 | 1????0000? | ???0??00?? | ??1?????00 | 1110010100 | 11(01)1101?01 | ??11?1?0? |
Gwesped | 1000111201 | ?01??00000 | 0??01-0020 | 0112001000 | 1110010100 | 1101101?01 | 0001?110? |
Lancepyris | 2010111210 | 0010100000 | 01001-0000 | 0111001000 | 1110010100 | 1101001000 | 1011?110? |
Liztor | 1010011210 | 101??0000? | 01001-0?20 | 0?1100??00 | 1110010100 | 111-1?1?01 | ?011?1??? |
Protopyris | 2010011200 | 101??00000 | 01001-0020 | 0111000000 | 1110010100 | 0101101?01 | 0001?110? |
Paralanceis | 1010101201 | 101??00000 | 01001-0010 | 011100?000 | 1110010100 | 0101001?01 | 0001?110? |
Yunbayin | 2010111200 | ?01??00000 | 01001-0020 | 0?1100??00 | 1110010100 | 1101101?01 | 0001?110? |
Zophepyris | 1010011210 | ?????0001? | 01001-0??0 | 0?1?001?00 | 1110010100 | 111-001?01 | ?001?110? |
Pycnomesitius | 2010001100 | 0000100010 | 0010001001 | 0111001010 | 1110010100 | 111-1-1000 | 101110111 |
Sulcomesitius | 2010101120 | 0000100000 | 0010001001 | 2111001010 | 1110010100 | 111-1-1001 | 101110111 |
Apenesia | (02)010100111 | 01121(01)0000 | (01)100(01)110(01)0 | (02)1(01)0(01)(01)1(01)0(01) | 0110010100 | 111-1-1001 | 10(01)001100 |
Dissomphalus | (12)0101001(01)(01) | 01101(01)000(01) | (01)000(01)110(01)0 | (02)1(01)0(01)(01)1(01)0(01) | 0110010100 | 111-1-1001 | 10(01)001100 |
Pristocera | (12)0101001(12)1 | 01001(01)0000 | (01)010(01)110(01)0 | 21(01)0(01)(01)1(01)0(01) | 0110010100 | 111-1-1001 | (01)0(01)001100 |
Pseudisobrachium | (12)01010011(01) | 01(01)01(01)000(01) | (01)000(01)110(02)0 | (02)1(01)01(01)1(01)0(01) | 0110010100 | 111-1-1001 | 10(01)0(01)1100 |
Bethylus | 1200111110 | 1001200000 | 0101001100 | 1111100110 | 1100010110 | 11001-1000 | 111111101 |
Eupsenella | 0210111110 | 0000100000 | 0101001100 | 0111000110 | 1100010110 | 0000011000 | 111111101 |
Lytopsenella | 2210011110 | 0000100000 | 0101011100 | 0111000110 | 1110010110 | 0000011000 | 101111101 |
Sierola | 2210111110 | 1000100000 | 0101011100 | 0111010110 | 1110010110 | 0100011000 | 111111101 |
Bakeriella | 2010100110 | 0010100000 | 0000011011 | 0111001110 | 1110010110 | 111-1-1000 | 101011001 |
Epyris | 2010100110 | 0000100000 | 0100011020 | 0111001110 | 1110010110 | 111-1-1000 | 101011001 |
Holepyris | 1010100110 | 0000100000 | 0100011010 | 0111101010 | 1110010110 | 111-1-1000 | 101011001 |
Elektroepyris | 2200101121 | 0011100000 | 0011011020 | 21120?1?00 | 1110010100 | 111-1-0000 | ?011?1?0? |
Allobethylus | 1111101101 | 0100201000 | 0000011020 | 0112001100 | 1110010100 | 111-1-1001 | 00111110? |
Cephalonomia | 2101111101 | 010(12)(23)01110 | (01)0001-1110 | 01(01)2101(01)0(01) | 00001-(01)0-0 | 111-1-1101 | 001101101 |
Sclerodermus | 2011101101 | 0110101110 | (01)000011020 | 21(01)1001(01)1(01) | 00001-00-0 | 111-1-1100 | 001101101 |
Gynopteron | 2010100200 | 0102100000 | 0100011110 | 011100??10 | 1110010100 | 111-1-1?00 | ??00?0?0? |
Cretabythus | 201?111100 | 0110?0000? | 01?00??000 | 0110000000 | 1110010101 | 0001001?01 | ?011?110? |
Holopsenelliscus | 101?111110 | ?11000001? | 0??01-0100 | 011100?000 | 1110010100 | 0001001?01 | 1011?110? |
Holopsenella
Family Bethylidae Haliday, 1839
Cretabythus Evans, 1973.
Small to mid-sized wasps (body length 2.5−7 mm); body not particularly pubescent; head prognathous; frons flat; compound eyes developed; antenna with 13 antennomeres; maxillary palpus with six palpomeres; median clypeal lobe short, not projecting; occipital carina present; dorsal pronotal area wider than long; propleuron more or less developed; prosternum concealed, nearly obscured by procoxae and propleura; anteromesoscutum with notaulus and parapsidal signum present; mesoscutellum posteriorly rounded; metanotum developed, separating mesoscutellum from metapectal-propodeal complex; metapectal-propodeal complex not elongate, without posterior spines or projections; both sexes macropterous; tegula present; fore wing with C, Sc+R, M+Cu, A, Rs+M, Rs, R1, basal segments of M and Cu tubular; [C], [R], [1Cu], [1R1], [1M] and [2R1] cells closed; [2Cu] closed in Cretabythus; hind wing only with C vein present; femora incrassate; tarsal claws slightly arched; metasoma without particular modifications.
Cretabythus Evans, 1973, Holopsenelliscus Engel, 2019, Megalopsenella
Lower Cenomanian to Santonian, in the deposits of northern Myanmar and Russia (Taimyr).
As no taxonomic treatment has been provided for the leftover genera of the obsolete Holopsenellinae, i.e., Cretabythus Evans, 1973, Megalopsenella
Lancepyris Azevedo and Azar, 2012.
Small-sized wasps (body length mainly around 2−3 mm, max 5.4 mm); body black to dark castaneous, not particularly pubescent; antenna with 13 antennomeres, rarely 12; ocelli present; dorsal pronotal area elongate, narrowing anteriorly; posterior margin of dorsal pronotal area concave; mesoscuto-mesoscutellar groove present; metanotum developed medially; metapectal-propodeal complex not posteriorly produced into spines; both sexes macropterous; tegula present; fore wing with C, Sc+R, M+Cu and A tubular; Rs+M vein tubular and straight, rarely reduced to spectral; 2r-rs&Rs long, sometimes reaching anterior margin; m-cu vein sometimes present, closing [1M] cell; pterostigma large; tarsal claws slightly arched; second metasomal tergite about as long as third.
Azepyris delamarrei sp. nov.
The genus name is a combination of Az-, honouring both Dr. Celso O. Azevedo and Dr. Dany Azar who first named the Lancepyrinae, and -epyris, a suffix traditionally used to name bethylids. Gender masculine.
Female. Body depressed, elongate; head longer than wide; eye elliptical, longer than high, located anteriorly on head; clypeus with median lobe projecting forward and lateral lobe not developed; mandible with three teeth, apical tooth longest; 11 flagellomeres; anterior ocellus not crossing supra-ocular line (Fig.
Azepyris delamarrei gen. et sp. nov., holotype female number
Following the key to the genera of Lancepyrinae of
The specific epithet is a patronym honouring Yann Delamarre, a student and the senior author’s fellow from the palaeontology program at the University of Rennes. The specific epithet is to be treated as a noun in the genitive case.
Holotype
Hkamti site, Hkamti district, Sagaing Region, Myanmar; early Albian, ca. 110 Ma, Early Cretaceous.
As for genus.
Body rather depressed, elongate, poorly pubescent (length 5.35 mm). — Head prognathous, longer than wide; LH: 1.04 mm, WH: ca. 0.70 mm, HE: 0.60 mm, VOL: 0.20 mm; frons flat, punctate; compound eye elliptical, longer than high, not covering head length, located on anterior half of head, closer to mandible than to occipital carina; clypeus with median lobe rather projecting forward, lateral lobe poorly developed; mandible long, with three teeth, apical tooth longest; antenna filiform; scape 2.17 times as long as pedicel (length 0.26 mm); flagellomeres 1−10 cylindrical, all longer than wide (length 0.09-0.10 mm); flagellomere 11 longest, tapering at apex: ocelli forming short triangle, anterior ocellus not crossing supra-ocular line; occipital carina present, complete, forming weak arch. — Mesosoma flattened, with dorsum smooth (length 1.62 mm); propleuron slightly visible in dorsal view, ‘neck-shaped’; dorsal pronotal area 1.56 times as long as anteromesoscutum (length 0.50 mm), narrow, anterior flange developed, lateral margin slightly incurved, posterior margin concave; anteromesocutum wider than dorsal pronotal area, posterior margin straight; notaulus deeply impressed, not reaching posterior margin of anteromesoscutum, convergent; parapsidal signum poorly marked; mesoscuto-mesoscutellar suture with reniform sulcus connecting lateral foveae; mesoscutellum subquadrate, posterior margin slightly convex; metanotum rather developed, overlapping mesoscutellum posteriorly sensu
Burmapyris azevedoi Jouault, Perrichot and Nel, 2021
The specific epithet is a patronym honouring Dr Christoph Öhm-Kühnle, who generously donated the specimen for study. The specific epithet is to be treated as a noun in the genitive case.
Holotype IGR.BU-060, a complete female; housed in the amber collection of the Geology Department and Museum of the University of Rennes, France (IGR).
Noije Bum, Hukawng Valley, Kachin State, northern Myanmar; late Albian-early Cenomanian, ca. 99 Ma, mid-Cretaceous.
Female. Head elongate, ovoid (Fig.
Burmapyris ohmkuhnlei sp. nov., holotype female number IGR.BU-060. A habitus in dorsal view; B mesosoma in dorsal view; C fore wing in dorsal view; D head and antenna in dorsal view; E habitus in lateral view; F head and anterior mesosoma in lateral view. Scale bars: 1 mm (A, E); 0.5 mm (B, C, D, F).
Body weakly depressed and poorly pubescent (length 3.80 mm). — Head prognathous, elongate, ovoid; LH: 0.83 mm, WH: 0.66 mm; WF: 0.52 mm, HE: 0.23 mm; OOL: 0.31 mm; WOT: 0.10 mm; DAO: 0.05 mm; VOL: 0.43 mm; compound eye elliptical, longer than high, located anteriorly on head; clypeus with median lobe triangular, lateral lobe not developed; space between toruli depressed; antenna filiform, reaching mesoscutellum posteriorly; scape 1.73 times as long as pedicel (length 0.19 mm), dorso-ventrally flattened; 11 flagellomeres cylindrical, distinctly longer than wide except flagellomere 1 (flagellomere 1 length 0.09 mm; flagellomeres 2−10 length 0.10−0.12 mm), slightly longer than wide, shorter than pedicel; flagellomere 11 longest, tapering at apex (length 0.16 mm); mandible decussate, with four to five (?) teeth, first and second apical teeth longest, remaining shorter; ocellar triangle small, anterior ocellus posterior to supra-ocular line; occipital carina present, forming small arch posterior to ocelli. — Mesosoma with dorsum smooth (length 1.46 mm); propleuron only slightly visible dorsally; dorsal pronotal area narrower anteriorly, lateral margin straight, posterior margin widely concave; anteromesoscutum shorter than dorsal pronotal area, notaulus impressed on posterior two thirds of anteromesoscutum, convergent, absent anteriorly, parapsidal signum present; mesoscuto-mesocutellar sulcus reniform, narrow; mesoscutellum posteriorly overlapped by long metanotum; metapectal-propodeal complex rectangular, without postero-lateral spine, median carina present, continuing on propodeal declivity. Fore wing hyaline, at least reaching third metasomal segment (LFW: 2.46 mm); C, Sc+R, M+Cu, 1A tubular; 1Rs&1M only slightly angled at junction with Rs+M; 1Rs shorter than 1M; 1M and cu-a aligned; Rs+M tubular, reaching apex of pterostigma; pterostigma slightly rounded; short stub of post-stigmal abscissa of R1 present; 2r-rs&Rs arising on distal half of pterostigma, long but not closing [2R1] cell; [2R1] cell large. Legs slender, only profemur swollen; tibial spur formula 1-2-2; tarsal claws slightly curved; arolium small. — Metasoma smooth, fusiform and elongate (length 1.51 mm); petiole short; six exposed tergites, covering sternites laterally; sting exserted.
Following the key to the genera of Lancepyrinae of
Gwesped groehni sp. nov.
The genus name is taken from the word for ‘wasp’ in the regional Breton language (Brittany, northwestern France). Gender feminine.
Female. Body flattened (Fig.
Gwesped groehni gen. et sp. nov., holotype female number GPIH.5068. A habitus in dorsal view; B habitus in ventral view (arrow = sting); C head in dorsal view; D head in ventral view; E fore wing in dorsal view; F line drawing of fore wing. Scale bars: 1 mm (A, B, E, F); 0.5 mm (C, D). Note: The apparent structure at the apex of the metasoma is the result of a bubble formed in the amber, combined with some artifacts produced during the stacking process.
Following the key to the genera of Lancepyrinae of
The specific epithet is a patronym honouring Carsten Gröhn, who generously made the specimen available for study. The specific epithet is to be treated as a noun in the genitive case.
Holotype GPIH.5068, a complete female; housed in the Carsten Gröhn amber collection (under CCGG no. 11341) of the Leibniz Institute for the Analysis of Biodiversity Change, Hamburg, Germany (GPIH).
Noije Bum, Hukawng Valley, Kachin State, northern Myanmar; late Albian-early Cenomanian, ca. 99 Ma, mid-Cretaceous.
As for genus.
Body depressed and poorly pubescent (length 3.82 mm). — Head prognathous, quadrate, about as long as wide, with metallic reflections; LH: 0.65 mm, WH: 0.56 mm; WF: 0.39 mm, HE: ca. 0.19 mm; frons flat, slightly punctate; compound eye rounded, located anteriorly on head; clypeus with median lobe only slightly projecting forward, lateral lobe poorly developed; antenna short, barely reaching pronotum posteriorly; scape 2.2 times as long as pedicel (length 0.20 mm); pedicel longer than flagellomeres 1−9; 10 flagellomeres, all compact, as long as wide; flagellomere tapering at apex, flagellomere 10 slightly longer than flagellomeres 1−9 (0.05 mm vs. 0.11 mm); occipital carina present. — Mesosoma narrow with dorsum smooth (length 1.56 mm); propleuron elongate, ‘neck-shaped’, visible dorsally; dorsal pronotal area narrower anteriorly, lateral margin slightly incurved, posterior margin straight; anteromesoscutum short, notaulus and parapsidal signum present; mesoscuto-mesoscutellar sulcus conspicuous, widely connecting lateral grooves; mesoscutellum subquadrate; metanotum developed, overlapping mesoscutellum posteriorly; metapectal-propodeal complex with metapostnotal median carina, lateral margin slightly outcurved, posterior corner with small lateral dentiform projection. Fore wing hyaline, posteriorly reaching fourth metasomal segment (LFW = 2.19 mm); C, Sc+R, M+Cu, 1A tubular; 1Rs and 1M aligned; cu-a slightly post-furcal to 1M; Rs+M tubular, reaching apex of pterostigma; pterostigma rounded; 2r-rs arising from distal half of pterostigma; 2r-rs&Rs tubular, reaching wing margin; [2R1] cell large; post-stigmal abscissa of R1 tubular for half-length of [2R1] cell. Legs slender; tibial spur configuration 1-2-2; tarsal claws slightly curved; arolium small. — Metasoma fusiform, smooth (length 1.61 mm); petiole relatively long, conspicuous; six tergites visible, convex; short sting exserted.
Paralanceis chotardi sp. nov.
The genus name is a combination of Para- and -lanceis, contraction of Lancepyris, for the similarities between the latter and the new proposed genus. Gender masculine.
Female. Body stout (Fig.
Following the key to the genera of Lancepyrinae of
The specific epithet is a patronym honouring Matthieu Chotard, a former student and the senior author’s fellow from the palaeontology program at the University of Rennes, and now working on the morphology of extinct and extant birds. The specific epithet is to be treated as a noun in the genitive case.
Holotype IGR.BU-061, a complete female; housed in the Geology Department and Museum of the University of Rennes, France (IGR).
Hkamti site, Hkamti district, Sagaing Region, Myanmar; early Albian, ca. 110 Ma, Early Cretaceous.
As for genus.
Body stout and poorly pubescent (length about 3.5 mm). — Head prognathous, ovoid; LH: 0.94 mm, WH: 0.74 mm; WF: 0.54 mm, HE: 0.28 mm; compound eye slightly longer than high, located anteriorly on head; clypeus with median lobe triangular and projecting forward, lateral lobe visible but much shorter than median lobe; toruli separated by flat surface; antenna filiform, short, barely reaching mesoscutellum posteriorly; scape 2.8 times as long as pedicel; 11 flagellomeres, slightly longer than wide; flagellomere 11 longest, tapering at apex; mandibles decussate at apex, with three teeth; occipital carina present. — Mesosoma with dorsum smooth (length 1.80 mm); propleuron elongate and visible dorsally; dorsal pronotal area narrower anteriorly, lateral margin straight, posterior margin slightly concave; anteromesoscutum short, notaulus and parapsidal signum hardly distinguishable; mesoscuto-mesoscutellar sulcus wide, connecting lateral grooves; mesoscutellum posteriorly overlapped by metanotum; metapectal-propodeal complex smooth dorsally, rectangular, without postero-lateral spine. Fore wing hyaline, micro-pubescent (LFW: 2.40 mm); C, Sc+R, M+Cu, 1A tubular; 1Rs and 1M aligned; cu-a slightly and 1M aligned; [1M] cell elongate, fully enclosed by tubular Rs+M, m-cu and 1Cu; stub of M tubular distally to [1M] cell; stub of 2Cu visible nebulous to spectral; pterostigma elongate; 2r-rs&Rs arising from distal half of pterostigma; 2r-rs&Rs tubular, reaching wing margin; post-stigmal abscissa of R1 long, meeting 2r-rs&Rs distally; [2R1] cell closed, lanceolate. Legs slender; tarsal claws slightly incurved. — Metasoma fusiform, smooth; petiole short; six tergites visible, partially covering sternites laterally; tergite 2 longest; sting exserted.
Yunbayin rossei sp. nov.
Yun Bayin is one of the 37 Nat spirits, a deity of the Burmese pantheon. Gender feminine.
Female. Head and mesosoma depressed (Fig.
Yunbayin rossei gen. et sp. nov., holotype female IGR.BU-062. A habitus in lateral view; B head and mesosoma in dorsal view; C fore wing in dorsal view; D line drawing of fore wing; paratype IGR.BU-063; E habitus in lateral view; F head and mesosoma in lateral view. Scale bars: 1 mm (A, E); 0.5 mm (B, C, D, F).
Following the key to the genera of Lancepyrinae of
The specific epithet is a patronym honouring Simon Rosse-Guillevic, a former student and the senior author’s fellow from the palaeontology program at the University of Rennes. The specific epithet is to be treated as a noun in the genitive case.
Holotype IGR.BU-062, a complete female; paratype IGR.BU-063, a complete female; both housed in the Geology Department and Museum of the University of Rennes, France (IGR).
Holotype from Hkamti site, Hkamti district, Sagaing Region, Myanmar; early Albian, ca. 110 Ma, Early Cretaceous. Paratype from Noije Bum, Hukawng Valley, Kachin State, northern Myanmar; late Albian-early Cenomanian, ca. 99 Ma, mid-Cretaceous.
As for genus
Body weakly depressed, elongate, poorly pubescent (length 2.43 mm). — Head prognathous, subquadrate and longer than high; LH: 0.42 mm, WH: 0.37 mm, WF: 0.25 mm, HE: 0.21 mm, VOL: 0.12 mm; frons flat; compound eye elliptical, longer than high, not covering head length, located on anterior half of head, closer to mandible than to occipital carina; antenna filiform, shorter than head + mesosoma combined; scape 2.8 times as long as pedicel (length 0.11 mm); pedicel longer than flagellomeres 1−10, thicker (length 0.04 mm); flagellomeres elongate, longer than wide; flagellomeres 1−10 gradually thickening, flagellomere 10 almost as long as wide (length 0.03 mm); flagellomere 11 longest (length 0.06 mm), tapering at apex; at least four maxillary palpomeres; occipital carina present. — Mesosoma flattened, with dorsum smooth (length 0.76 mm; height 0.21 mm); propleuron elongate and visible in dorsal view; prosternum short between procoxae; dorsal pronotal area narrower than anteromesoscutum, with lateral margin incurved, posterior margin slightly concave; anteromesoscutum shorter than dorsal pronotal area, with notaulus present and convergent posteriorly, parapsidal signum present; scutellum posteriorly overlapped by metanotum; metapectal-propodeal complex carinate, rectangular, without posterior spine. Fore wing hyaline, reaching at least third metasomal segment (LFW: 1.39 mm); C, Sc+R, M+Cu, 1A tubular; 1Rs&1M only slightly angled at junction with Rs+M; 1Rs shorter than 1M; cu-a slightly post-furcal to 1M; Rs+M tubular, reaching apex of pterostigma; pterostigma slightly rounded and elongate; short stub of post-stigmal abscissa of R1 present; 2r-rs&Rs arising on distal half of pterostigma, long but not closing [2R1] cell; [2R1] cell narrow, lanceolate. Hind wing without apparent venation; at least three hamuli. Proleg with protrochanter originating from apex of procoxa; femora slightly enlarged; tibial spur formula 1-2-2; tarsal claws only slightly curved; arolium present, as long as claws. — Metasoma cylindrical, not flattened (length 1.25 mm; height 0.39 mm); petiole long and conspicuous; seven visible segments; metasomal tergites decreasing in length; ovipositor not exserted. — Paratype. Similar to holotype but longer. Body length 2.92 mm; LH: 0.56 mm; fore wing with Rs+M hardly distinguishable distally but clearly tubular, reaching apex of pterostigma (LFW: 1.46 mm); proleg with internal musculature exquisitely preserved; mesosoma length 1.08 mm, height 0.27 mm; metasoma length 1.28 mm, height 0.31 mm; ovipositor exserted, long (length 0.34 mm).
Although the two specimens of the type series originate from deposits considered distinct in age, they do not show much variation. Therefore, we assign them to the same species, pending further material. To our knowledge, it is the first instance of a shared species between Hkamti and Noije Bum (Kachin) amber.
1a | Fore wing with Rs+M conspicuous and tubular (Fig. |
2 |
1b | Fore wing with Rs+M absent, spectral or weak (Fig. |
8 |
2a | Fore wing with [2R1] cell closed by 2r-rs&Rs and R1 (Fig. |
3 |
2b | Fore wing with [2R1] cell open apically (Fig. |
4 |
3a | Fore wing with [1M] cell fully enclosed by tubular veins (Fig. |
Paralanceis gen. nov. |
3b | Fore wing with [1M] cell open (Fig. |
Lancepyris Azevedo and Azar, 2012 |
4a | Fore wing with short pterostigma and Rs+M vein short, not enclosing [1M] cell or reaching apex of pterostigma (Fig. |
Archaepyris Evans, 1973 |
4b | Fore wing with elongate pterostigma and Rs+M long, enclosing [1M] cell or reaching apex of pterostigma (Fig. |
5 |
5a | Fore wing with [1M] cell fully enclosed by tubular veins (Fig. |
Protopyris Jouault and Nel, 2021 |
5b | Fore wing with [1M] cell open (Fig. |
6 |
6a | Mesosoma more flattened than metasoma, lateral margin of pronotum widely incurved (Fig. |
Yunbayin gen. nov. |
6b | Mesosoma equally or less flattened than metasoma, lateral margin of pronotum straight (Fig. |
7 |
7a | Metapectal-propodeal complex with small dentiform projections, fore wing with 2r-rs&Rs fully tubular (Fig. |
Gwesped gen. nov. |
7b | Metapectal-propodeal without such projections, fore wing with 2r-rs&Rs absent distally (Fig. |
Burmapyris Jouault, Perrichot and Nel 2021 |
8a | Fore wing with cu-a post-furcal to 1M (Fig. |
9 |
8b | Fore wing with cu-a aligned or antefurcal to 1M (Fig. |
10 |
9a | Fore wing with Rs+M weak but distinct, 1Rs&1M angled, pterostigma rounded and 2r-rs&Rs not fully tubular, metapectal-propodeal complex with small dentiform projections (Fig. |
Azepyris gen. nov. |
9b | Fore wing with Rs+M absent, 1Rs&1M straight, the pterostigma elongate and 2r-rs&Rs fully tubular, metapectal-propodeal complex without such projections (Fig. |
Zophepyris Engel et al. 2016 |
10a | Fore wing with 2r-rs&Rs arising on distal half of pterostigma (Fig. |
Cretepyris Ortega-Blanco and Engel, 2013 |
10b | Fore wing with 2r-rs&Rs arising on basal half of pterostigma (Fig. |
Liztor Ortega-Blanco and Engel, 2013 |
Pictures of metapectal-propodeal complex and schematical fore wing line drawing for characters of couplet 7. A top: dentiform projection of metapectal-propodeal complex, bottom: 2r-rs&Rs fully tubular (couplet 7a); B top: metapectal-propodeal complex without projection, bottom: 2r-rs&Rs absent distally (couplet 7b).
Schematical fore wing line drawing and pictures of metapectal-propodeal complex for characters of couplet 9. A top: Rs+M distinct, 1Rs&1M angled, pterostigma rounded, 2r-rs&Rs absent distally, bottom: dentiform projection of metapectal-propodeal complex (couplet 9a); B top: Rs+M absent, 1Rs&1M straight, pterostigma straight, 2r-rs&Rs fully tubular, bottom: metapectal-propodeal complex without projection (couplet 9b).
The search with equal weighting produced 15 most-parsimonious trees of length L = 184 steps, consistency index CI = 0.386 and retention index RI = 0.663 for 17 983 090 arrangements tried. Among those 15 trees, topological differences concerned both inter-subfamilial and intra-subfamilial relationships in the Pristocerinae, Lancepyrinae and Epyrinae. About two thirds of the trees (9/15) retrieved the Cretabythinae as the first diverging lineage and Lancepyrinae as the sister of the remaining Bethylidae. The Bethylinae were the first-diverging subfamily in the crown-group and the clade [Protopristocerinae + Epyrinae] was found to be sister to the [Mesitiinae + (Elektroepyrinae + (Scleroderminae + Pristocerinae))] clade. In the other trees (6/15), Cretabythinae were found as the first diverging lineage but the remaining subfamilies successively diverged as follows: Pristocerinae, Scleroderminae, Elektroepyrinae, Mesitiinae, Protopristocerinae, Epyrinae, Bethylinae and Lancepyrinae. A strict consensus was computed and we obtained one cladogram (Fig.
Strict consensus of 15 trees obtained under equal-weighting analyses. Character changes are indicated on each branch by circles, with character number and character state (as stated in Table
The search with implied-weighting produced 9 most parsimonious trees of length L = 188 steps, consistency index CI = 0.378 and retention index RI = 0.651 for 2 866 340 arrangements tried. Among those nine trees, topological differences only concerned intra-subfamilial relationships in the Pristocerinae and Lancepyrinae. A strict consensus was computed and we obtained one cladogram (Fig.
Strict consensus of 9 trees obtained under implied-weighting analyses. Character changes are indicated on each branch by circles, with character number and character state (as stated in Table
In both consensus trees, all seven non-monotypic subfamilies are retrieved as monophyletic, some being rather well-supported with bootstrap values ≥ 65 (Mesitiinae, Bethylinae, Scleroderminae, Pristocerinae and Epyrinae under implied-weighting). Equally, the Lancepyrinae were found to be monophyletic but less strongly supported. The Lancepyrinae are supported by two characters that are common to the equal- and implied-weighting trees but neither is uniquely apomorphic for the subfamily: the scape evenly wide (6:1, also in Bethylinae and Cephalonomia Westwood, 1833) and the frontal line absent (25:1, also in Holopsenelliscus, Cephalonomia and Pristocerinae). The equal-weighting analyses found four additional characters supporting the monophyly of the Lancepyrinae, none being uniquely apomorphic either: the pedicel longer than wide (8:2, also in Gynopteron Falières and Nel, 2019 and Pristocera Klug, 1808), the posterior ocelli close to vertex crest (22:1, also in Bethylinae, Gynopteron, Holepyris Kieffer, 1904, Epyris Westwood, 1832 and Apenesia Westwood, 1874), the dorsal pronotal area narrow anteriorly (27:0, also in Clystopsenella and Holopsenelliscus) and the presence of the Rs+M vein of the fore wing (53:0, also in Clystopsenella, Cretabythinae and Bethylinae).
With hundreds of new insect species being reported each year from mid-Cretaceous Burmese amber (
If taxonomic boundaries between each genus are clear, this study enlightens the wide range of morphologies within the Lancepyrinae. The example of the wing venation is particularly illustrative, ranging from the simplest, with Rs+M absent and two closed cells (Cretepyris), to the most complete known in Lancepyrinae, with five closed cells (Paralanceis gen. nov.). Given these morphological differences, one could wonder whether they truly represent a monophyletic group, especially in the case of species with reduced or absent Rs+M vein (this vein was a diagnostic feature of the subfamily at the moment of its description). The cladistic analysis performed here confirms the positions of Azepyris gen. nov., Liztor, Zophepyris and Cretepyris, which have reduced or absent Rs+M, within the Lancepyrinae. Nonetheless, the genus Cretepyris needs to be revised since the three type specimens of C. martini (plus another putative male, which is not part of the type series) show some variations (
The four newly described genera complicate the inference of the relationships among the lancepyrine genera. While the general trend proposed for the evolution of Hymenoptera implies a simplification of the wing venation over time (
Despite similar analytical parameters, our cladograms differ from that of
Indeed, these morphology-based hypotheses do not fully align with molecular-based hypotheses (
With five new species described herein, the subfamily Lancepyrinae is now composed of 12 species in 11 genera from several major deposits: Lebanese amber, Spanish amber, Myanmar amber and Taimyr amber. We confirmed the monophyly of this subfamily, while highlighting the diversity and the morphological disparity of the Lancepyrinae, and consequently of the Bethylidae, as early as the mid-Cretaceous. The relationships within the Lancepyrinae remain to be deciphered, as well as those between each subfamily, which should be achieved by combining molecular and morphological characters.
The authors have declared that no competing interests exist.
We are most grateful to Dr Christoph Öhm-Kühnle (Herrenberg, Germany) and Corentin Jouault (MNHN, Paris, France) for donating three of the specimens studied herein; and to Carsten Gröhn (Glinde, Germany) and Patrick Müller (Zweibrücken, Germany) for access to two other specimens with subsequent repositories in institutional collections. We sincerely thank Dr. Celso Azevedo (Espírito Santo, Brazil) for his precious comments on the first version of the draft and Dr. Wesley Colombo (Espírito Santo, Brazil) for his help on the construction of the matrix. We thank Prof. Denis J. Brothers and two anonymous reviewers for their constructive comments on the manuscript, as well as the subject editor, Dr. Ricardo Pérez-de la Fuente, for his guidance during the submission process. Finally, we thank the Willi Hennig Society for making available the TNT software for free. This work is part of Manuel Brazidec’s Ph.D. project CHRYSIS: “The role of greenhouses on the diversification and evolution of chrysidoid wasps”.