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Myanmarinidae, a new family of basal Apocrita (Hymenoptera: Stephanoidea) from mid-Cretaceous Burmese amber
Accepted Manuscript Myanmarinidae, a new family of basal Apocrita (Hymenoptera: Stephanoidea) from mid-Cretaceous Burmese amber Qi Zhang, Alexandr P. Rasnitsyn, Bo Wang, Haichun Zhang PII:
S0195-6671(17)30366-X
DOI:
10.1016/j.cretres.2017.09.015
Reference:
YCRES 3705
To appear in:
Cretaceous Research
Received Date: 19 August 2017 Revised Date:
23 September 2017
Accepted Date: 24 September 2017
Please cite this article as: Zhang, Q., Rasnitsyn, A.P., Wang, B., Zhang, H., Myanmarinidae, a new family of basal Apocrita (Hymenoptera: Stephanoidea) from mid-Cretaceous Burmese amber, Cretaceous Research (2017), doi: 10.1016/j.cretres.2017.09.015. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
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ACCEPTED MANUSCRIPT Myanmarinidae, a new family of basal Apocrita (Hymenoptera:
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Stephanoidea) from mid-Cretaceous Burmese amber
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Qi Zhanga,b, Alexandr P. Rasnitsync,d,*, Bo Wanga,e, Haichun Zhanga
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a
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Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008,
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China
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b
University of Chinese Academy of Sciences, Beijing 100049, China
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c
A.A. Borissiak Palaeontological Institute, Russian Academy of Sciences, 117647
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Moscow, Russia
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d
Natural History Museum, Cromwell Road, London SW7 5BD, UK
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Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology,
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Chinese Academy of Sciences, 1, Beichen West Road, Beijing 100101, China
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State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of
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* Corresponding author. A.A. Borissiak Palaeontological Institute, Russian
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Academy of Sciences, 117647 Moscow, Russia.
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Email: [email protected]
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Abstract
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A new family of basal apocritan wasps is established based on three new
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species discovered from the mid-Cretaceous Burmese amber and tentatively allied
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to the superfamily Stephanoidea. The family Myanmarinidae Zhang and Rasnitsyn
ACCEPTED MANUSCRIPT fam. nov. contains one new genus Myanmarina Zhang and Rasnitsyn gen. nov.,
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including three new species: Myanmarina lisu Zhang and Rasnitsyn sp. nov.,
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Myanmarina kachin Zhang and Rasnitsyn sp. nov. and Myanmarina lahu Zhang
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and Rasnitsyn sp. nov. Comparisons are made between Myanmarinidae and its
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similar families. Some general remarks are also made about the new family in
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taxonomy and biology.
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Keywords: Hymenoptera, Myanmarinidae, Myanmarina, mid-Cretaceous, Burmese
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amber
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1. Introduction
Burmese amber is currently one of the richest sources of knowledge about the
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past of insects and some other segments of the non-marine biosphere (Rasnitsyn,
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1996; Zherikhin and Ross, 2000; Grimaldi et al., 2002; Ross et al., 2010; Rasnitsyn
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et al., 2016; Guo et al, 2017). Indeed, the Burmese amber arthropod assemblage
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accounts for 252 families which is the highest figure among the Cretaceous
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Lagerstätten (Rasnitsyn et al., 2016; the data by the end of 2013). Years later Ross
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(2017) lists 356 arthropod families and 481 insect species there, and Guo et al.
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(2017) cite 73 described species in Hymenoptera. Hymenoptera represents a
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significant part of the modern and, since the Jurassic, past world biodiversity: the
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above source counts 30 families, that is, 12 percent of the total arthropod diversity
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in the Burmese amber. Like other taxa, Hymenoptera displays a number of
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ACCEPTED MANUSCRIPT described (Aptenoperissidae, Bryopompilidae, Melittisphecidae, Othniodellithidae,
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Syspathoxyelidae, cf. Rasnitsyn et al., 2017; Rodriguez et al 2016; Poinar and
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Danforth, 2006; Engel et al., 2016a, 2016b; respectively) and undescribed (in
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preparation) endemic families within the Burmese amber assemblage. Description
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of one more such family is the objective of the present publication.
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2. Material and methods
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The present research is based on the amber collected from the amber mines
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situated in the Hukawng Valley of Kachin State, Myanmar (locality in Kania et al.,
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2015: fig. 1). The rock containing the Burmese amber was radiometrically dated at
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98.79 ± 0.62 Ma (Shi et al., 2012). The date corresponds to the early Cenomanian,
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that is, the earliest Late Cretaceous. However, the amber displays clear traces of
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re-deposition (Ross, 2015) and so can be older than enclosing rocks. That is why
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we prefer to refer to the amber age informally as mid-Cretaceous.
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The material was studied under Nikon SMZ-10 R stereoscopic microscope and
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Nikon Optiphot compound microscope with magnifications up to 800× at the State
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Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology
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and Palaeontology, Chinese Academy of Sciences, and using Leica M165C
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stereomicroscope with a Leica DFC 420 camera at the A.A. Borissiak
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Paleontological Institute, Russian Academy of Sciences, in Moscow. In most
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instances, incident and transmitted light were used simultaneously. Helicon Focus
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Pro ×64 was used to stack photos for better depth of field. The line drawings were
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prepared based on photographs using image-editing softwares (CorelDraw X7 and
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ACCEPTED MANUSCRIPT Adobe Photoshop CS6). The specimens are housed in the Nanjing Institute of
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Geology and Palaeontology, Chinese Academy of Sciences (NIGPAS).
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Morphological terminology and symbols to wing vein homology are standard,
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except that ampersand (&) linking two adjacent vein sections denotes those aligned
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and forming a seemingly entire vein section. Particularly, 1-RS&1-M means the
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first (basal) sections of RS and M joining to form a smooth, seemingly entire vein
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(basal vein in older nomenclature; cf. Fig. 1 C). All taxonomic acts established in
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the present work have been registered in ZooBank (see below), together with the
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electronic publication LSID: urn:lsid:zoobank.org:pub:FC26F5AC-E0FF-44CB-
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B621-C68973E0BEAF
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3. Systematic paleontology
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Order Hymenoptera
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Suborder Vespina
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Superfamily Stephanoidea
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Family Myanmarinidae Zhang and Rasnitsyn, fam. nov.
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(LCID: urn:lsid:zoobank.org:act:A6499035-F929-449A-8477-
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Type genus Myanmarina Zhang and Rasnitsyn, gen. nov.
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Diagnosis. Male (female unknown). Stature elongate, subcylindrical, as in
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Stephanidae. Body size small (1.5-3 mm). Head subglobular, without crown of
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teeth around fore ocellus, with big eyes, circular occipital carina and long genal
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and hypostome). Antenna 11- or 12-segmented, not geniculate, flagellar segments
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more or less uniform, possibly with elongate (multiporous plate) sensillae.
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Mandible short, wide, with 2 or, possibly, 3 subequal teeth, with cutting edge
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subparallel to mandible rotation axis, not distinctly oblique. Maxillary palps with 4
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visible segments, apical three ones narrow and elongate. Labial palps not observed,
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probably much reduced. Mesosoma long and narrow, subcylindrical or somewhat
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depressed (possibly a preservational distortion), its morphology little known except
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that the pronotum is short centrally in dorsal view and the mesoscutum long.
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Propleurae more or less elongate, usually forming a distinct neck. Propodeum
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elongate, straight in lateral view, with no downward bending towards metasomal
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articulation. Legs elongate, ordinary; fore tibia with single, thin, arcuate spur.
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Wings densely pubescent, with comparatively long marginal setae, with venation
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much reduced. Forewing with tubular veins only in anterobasal part, namely fused
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C+R up to some 0.6 wing length, M+Cu, strongly oblique basal vein (1-RS&1-M),
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cu-a, 1-Cu and 1A up to cu-a only (Cu may continue after cu-a as nebulous vein).
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All distal wing surface free of veins, supported with system of nested radiated
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folds like in Spathiopterygidae. Hind wing short and very narrow, reaching level of
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upper end of basal vein (that is, level of 1-RS base), with only C+R tubular and
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with 3 hamuli. Metasoma linear, with 8 external segments, 1st segment distinctly
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narrowed basal but details of articulation not clear, male claspers wide and often
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long, pointing obliquely downward.
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Genera included. Type genus only.
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Genus Myanmarina Zhang and Rasnitsyn, gen. nov.
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(LCID: urn:lsid:zoobank.org:act:BB9726BE-39FB-44A5-901A-
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A7791CFEEC21) Type species. Myanmarina lisu Zhang and Rasnitsyn, sp. nov.
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Etymology. From Myanmar, the country where all the three species of the new genus come. Gender feminine.
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Diagnosis. As for family because of monotypy.
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Species included. Three species.
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Myanmarina lisu Zhang and Rasnitsyn, sp. nov.
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(LSID: urn:lsid:zoobank.org:act:5AC41744-1151-4846-A7C6-
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Fig. 1
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FD58CBE42E3A)
Derivation of name. After the Lisu, an ethnic group in northern Myanmar
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where the fossil material was collected. Noun in apposition. Holotype. Male, NIGP166310 (Fig. 1A-D). Only few part of legs damaged,
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diagenetic deformation present, with some scattered debris around; small air
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bubbles distributed around the whole body including the antennae and legs make it
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difficult to discern the dorsal surface sculpture.
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Paratype. Male, NIGP166311 (Fig. 1E-G). One forewing damaged, the other bent; many scattered debris around. Dark body and lots of small bubbles obscuring
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ACCEPTED MANUSCRIPT particular details of the fossil. The type specimens were originally found as
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syninclusions and subsequently separated for study.
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Locality and Horizon. Hukawng Valley, Kachin State, Myanmar; lowermost Cenomanian, mid-Cretaceous. Description. NIGP166310. Head subglobose; ocelli small, well separated from
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compound eyes; compound eyes prominent, berry-like in appearance. Antenna 11-
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segmented, inserted moderately low on head, densely covered with fine setae;
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scape relatively straight; pedicel almost as long as scape; first flagellomere
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extremely long, thinner than pedicel basally and becoming gradually thicker;
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second about as long as scape and pedicel combined; third and fourth slightly
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longer than second and fifth, remaining ones shorter than fifth but still more than
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twice as long as wide, apicalmost flagellomere rounded. Maxillary palps almost as
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long as head height, with three apical segments long and thin, basal visible
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segment probably short. Mesosoma imbricate, long-column-shaped and wider than
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high; pronotum short, mesoscutum long with indistinct (possibly percurrent)
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notauli. Legs long with respect to body length and covered with minute setae; tibial
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spur formula 1-1-1; basitarsomeres almost as long as remainder of corresponding
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tarsomeres together; all femora shorter than their tibia, and hind femur slightly
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swollen. Length ratio of tarsomeres in all legs (similar in all congeners) ca. 10:5:3-
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3.5:1.5-1.7:2-2.5 (precise measurements often difficult because of imperfect
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preservation state and improper position of legs in amber). Forewing longer than
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metasoma, spatulate, with well-defined marginal fringe, fringe setae longest along
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hind margin (possibly also along wing apex which is obscured), moderate along
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ACCEPTED MANUSCRIPT veins on fore margin, growing shorter toward anterior preapical margin; membrane
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with dense and comparatively long setae all over surface distal of venation, with
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basal (1rm) and subbasal (1cua) cells naked; C+R reaching maximum wing width
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(0.64 wing length); M+Cu faded in basal third, branching in wing basal 0.2, 1-
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RS&1-M thickest vein, reaching C+R at 0.33 wing length, cu-a aligned with 1A,
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meeting Cu distinctly distal of M+Cu apex; free Cu long, nebulous, subparallel to
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wing hind margin, continuing into weak fold; five stronger folds running fan-like
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toward wing margin anterior of Cu. Hind wing very narrow, some 0.4 as long as
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forewing, with fringe of moderately long setae along all visible hind margin and
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apex and with 3 hamuli present. Metasoma narrow, about as long as head and
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mesosoma combined, and wider than high, with segments difficult to discern; first
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segment trapezoid, elongate, following apparently shorter; claspers wide and long,
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triangular in side view, otherwise genitalia not visible.
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NIGP166311, providing additional characters such as: mandible possibly tridentate, maxillary palps narrow and long; fore leg with coxa long ovate,
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trochanter small, tibia with inner row of setae; mid and hind coxa almost as long as
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femur.
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Measurements. NIGP166310. Total body length, 2.7 mm; head length, 0.3 mm,
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width, 0.4 mm; mesosomal length,1.0 mm; metasomal length, 1.4 mm; antennal
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length, 1.4 mm; forewing length, 1.9 mm, width, 0.7 mm; hindwing length 0.7 mm.
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NIGP166311. Total body length, 2.3 mm; head length, 0.3 mm, width, 0.3 mm;
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mesosomal length,0.7 mm; metasomal length, 1.3 mm; antennal length, 1.3 mm;
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forewing length, 1.38 mm, width, ca. 0.6-0,65 mm; hindwing length 0.55 mm.
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ACCEPTED MANUSCRIPT Remarks. The species differs from all congeners in having antennae distinctly
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thickened apically, hind coxae and genital claspers very long. It is noteworthy that
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high size variability is present in the species, with forewing length differing in
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almost 1.4 times between two males available. Fig.1
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Myanmarina kachin Zhang and Rasnitsyn, sp. nov.
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(LSID: urn:lsid:zoobank.org:act:148831C2-8910-4229-8E5E-4F36DC8F465D)
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Fig. 2.
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Derivation of name. After the Kachin, an ethnic group in northern Myanmar where the fossil material comes. Noun in apposition.
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Holotype. Male, NIGP166309. Nearly complete with diagenetic deformation,
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with antennae and hind legs partly damaged; amber inner layer surface light brown;
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the surface sculpture not discernible.
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Locality and Horizon. Hukawng Valley, Kachin State, Myanmar; lowermost
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Cenomanian, mid-Cretaceous. Description. Body dark brown to black. Head subglobose. Eyes not easily
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recognizable. Antenna inserted moderately low on head, slightly widened apically,
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11-segmented; scape slightly expanded ventrally, about twice the length of pedicel;
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pedicel small, slightly longer than wide; first flagellomere thin and extremely long,
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arched softly and becoming thicker apically; flagellomeres 2-5 similar in size and
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shape, about half as long as first and slightly thinner in both ends than middle part;
ACCEPTED MANUSCRIPT flagellomeres 6-8 shorter and broader, flagellomere 9 almost as long as 8 and
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slightly pointed at apex. Maxillary palps as visible as long as head high, with 4
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segments visible, apical three narrow and elongate, first visible segment much
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thicker and probably short. Mesosoma subcylindrical with reticulated surface
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(possibly because of cuticle degradation), narrow and more than 3 times as long as
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wide. Pronotum short and mesoscutum long; propodeum elongate, straight in
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lateral view, with no downward bending towards metasomal articulation. Legs
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long and thin on the whole, coxa long and moderately stout; trochanter thin;
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tarsomere length as in type species. Protibial spur simple and curved. Fore and mid
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legs with femur cylindrical and shorter than tibia; tibia expanded apically. Fore
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basitarsus as long as tibia; mid basitarsus longer than tibia. Hind legs with femur
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slightly swollen; tibia longer than femur, with single tibial spur. Forewing hyaline
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with moderately dense setation, with short fringe along hind margin (possibly also
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along wing apex which is obscured). Venation incompletely visible, with fused
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C+R long, about 3/5 length of wing; 1-Rs&1-M strongly oblique, very thick,
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connecting to C+R near its midlength; nested forking folds present distal of
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preserved veins, with hind fold apparently continuing into nebulous free Cu. Hind
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wing short and very narrow, with 3 hamuli present. Metasoma linear, longer than
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mesosoma, with segment boundaries difficult to identify precisely. Claspers wide
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and short, other details of genitalia invisible.
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Measurements. NIGP166309. Total body length, 3.2 mm; head length, 0.4 mm,
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width, 0.2 mm; mesosomal length,1.2 mm; metasomal length, 1.6 mm; antennal
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length, 1.5 mm; forewing length, 1.8 mm, width, 0.7 mm.
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ACCEPTED MANUSCRIPT Remarks. The species differs from all congeners by its larger size and the hind
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femur comparatively short and rather stout (instead of long and at most slightly
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swollen in both M. lisu and M. lahu). Fig.2
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Myanmarina lahu Zhang and Rasnitsyn, sp. nov.
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(LSID: urn:lsid:zoobank.org:act:ECC6AE11-5B6C-4306-9ECB-
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BE229F17BCE1) Fig. 3
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Derivation of name. After the Lahu, an ethnic group in northern Myanmar where the fossil material was discovered. Noun in apposition.
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Holotype. Male, NIGP166312 (Fig. 3A-D). Nearly complete fossil missing
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only one wing apex, deformed in many respects; amber color faint yellow and not
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well transparent, inner surface deep yellow colored; many scattered debris around;
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surface sculpture and many details of morphology of holotype are difficult to see
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and interprete correctly because of diagenetic deformations as well as deep dark
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color.
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Paratype. Male, NIGP166313 (Fig. 3E-F). Head and part of legs damaged, lots
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of scattered debris and some filaments around; amber not well transparent, dark
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body and lots of small bubbles obscuring particular details of the fossil.
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The type specimens were originally found as syninclusions and separated subsequently.
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ACCEPTED MANUSCRIPT Locality and Horizon. Hukawng Valley, Kachin State, Myanmar; lowermost
Cenomanian, mid-Cretaceous. Description. NIGP166312. Specimen dark brown to black. Head subglobose, longer than high. Eyes berry-like, moderately large, reaching mandibular base;
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ocelli indistinguishable. Antenna inserted low on head, below eye midheight, 12-
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segmented; scape ventrally slightly expanded, about twice the length of pedicel;
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flagellomeres densely covered with thin setae, with first three antennal segments
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pale compared to remainders; first one moderately long, thinner than pedicel
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basally, second to sixth as long as first, seventh to tenth slightly shorter than sixth,
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apical one rounded. Mandible narrowed apically. Maxillary palps some half as
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long as head high, with three apical segments long and thin. Mesosoma very long,
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wider than height. Pronotum extremely short centrally, hardly visible in dorsal
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view; propleurae elongate forming distinct neck; mesoscutum long; propodeum
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elongate and straight. Legs long with respect to body length and covered with
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minute setae; coxa long, trochanters small; fore and mid femora longer than
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respective tibia, hind femur slightly swollen and shorter than tibia; tarsomere
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length ratio as in type species. Forewing as preserved similar to that in other
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congeners. Hindwing short with membrane, without wing venation apart from C+R,
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and with three hamuli present. Metasoma long and narrow, with 8 external tergites,
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second longest, eighth shortest. Claspers wide and short, other parts of genitalia not
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visible.
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ACCEPTED MANUSCRIPT Specimen NIGP166313 not well visible, very small, head big, mandible
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narrowed apically, distinctly with two subequal teeth, hind coxa and femur long.
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Flagellomeres possibly with multiporous plate sensilla (Fig. 3F).
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Measurements. NIGP166312. Total body length, 2.9 mm; head length, 0.4 mm, width, 0.3 mm; mesosomal length,1.1 mm; metasomal length, 1.7 mm; antennal
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length, 1.8 mm; forewing length, 1.9 mm; hind wing length ca. 0.65 mm.
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NIGP166313. Head length, 0.3 mm; mesosomal length, 0.85 mm; antennal length, 1.7 mm.
Remarks. The species differs from all congeners in having the antenna 12-
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segmented, the first flagellomere hardly 1.5 times as long as apical one and the
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maxillary palps short (antenna 11-segmented with the first flagellomere several
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times as long the apical one, and maxillary palp long in both M. lisu and M. lahu).
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Fig.3
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4. Discussion
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The present fossils are imperfectly preserved, and therefore they are
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insufficiently known in morphology. However, the morphological characters
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known from these specimens are distinctive enough so as so warrant description of
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the fossils as a family of their own. Accommodation of the new family in the wasp
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tree is not straightforward as well. Its distinctive similarity to the
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Spathiopterygidae in wing morphology cannot be inherited because of too different
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organization of propodeum and metasoma. In Spathiopterygidae, the first
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metasomal segment is transformed into a petiole and attached low on the
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ACCEPTED MANUSCRIPT propodeum at the end of the backward slope of the latter. In combination with
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other features scrutinized by Engel et al. (2013), this is clearly indicative of
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Diaprioidea. In contrast, propodeum of the new family is straight in side view
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lacking a posterior slope, and hence a real constriction is absent between the
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propodeum and the metasoma. This character state is characteristic of Stephanidae
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and extinct families Ephialtitidae and Aptenoperissidae, that is, of the superfamily
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Stephanoidea (Rasnitsyn and Zhang, 2010; Rasnitsyn et al., 2017; Zhang et al.,
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submitted). That is why we attribute Myanmarinidae to Stephanoidea, at least until
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a sound contradictory information appears.
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Myanmarinidae can be easily differentiated from all three other families of Stephanoidea in their small size, 11-segmented antenna and particularly in their
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specific mode of reduction of the wing venation: particularly with two
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incompletely separated basalmost cells margined with thick veins within the basal
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third of wing's length, with no other closed cells, and with distinctly radially folded
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outer wing membrane. Unlike the new family, Ephialtitidae and male
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Aptenoperissidae (Zhang et al., submitted) always have more complete venation
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(female apterous), and Stephanidae have venation varying from complete to
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strongly reduced but never displaying deep differentiation between the wing base
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and the remainder wing blade.
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5. Concluding remarks The biology of the new family could be very tentatively inferred from the general appearance of the available fossils. Their long and narrow body and
ACCEPTED MANUSCRIPT comparatively big and movable head are suggestive of insects parasitizing hidden,
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probably xylobiotic preys. This agrees additionally with the proposed taxonomic
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position of Myanmarinidae, because Stephanidae are known to develop at expense
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of xylophagous beetles (Achterberg, 2002), and Ephialtitidae are hypothesized to
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behave similarly (Rasnitsyn, 1980). The distinct variation of the body size within
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the males of M. lisu sp. nov. is compatible with the hypothesis as well.
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Acknowledgements
This research was supported by the National Natural Science Foundation of
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China (41272013, 41572010, 41622201 and 41688103), the Chinese Academy of
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Sciences (XDPB05), and Youth Innovation Promotion Association of CAS (No.
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2011224).The financial support of the UCAS (UCAS[2015]37) Joint PhD Training
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Program to ZQ is acknowledged. We offer our sincere gratitude to Dr. Ekaterina A.
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Sidorchuk for providing technical guidance, and the anonymous reviewers for the
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very useful comments on the earlier version of the manuscript.
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Grimaldi, D.A., Engel, M.S., Nascimbene, P.C., 2002. Fossiliferous Cretaceous amber from Myanmar (Burma): its rediscovery, biotic diversity, and
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paleontological significance. American Museum Novitates 3361, 1-71.
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Guo, M., Xing, L., Wang, B., Zhang, W., Wang, Sh., Shi, A., Bai, M., 2017. A
356 357 358 359 360
Kania, I., Wang, B., Szwedo, J., 2015. Dicranoptycha Osten Sacken, 1860 (Diptera,
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catalogue of Burmite inclusions. Zoological Systematics 42(3), 249-379.
Limoniidae) from the earliest Upper Cretaceous Burmese amber. Cretaceous
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Research 52, 522-530.
Poinar, G.O.Jr., Danforth, B.N., 2006. A fossil bee from Early Cretaceous Burmese amber. Science, 314, p. 614.
Rasnitsyn, A.P., 1980. Origin and evolution of Hymenoptera. Trudy
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Paleontologicheskogo lnstituta Acadernii Nauk SSSR, 174. Nauka Press,
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Moscow, 192 p. (in Russian).
363 364
ACCEPTED MANUSCRIPT Rasnitsyn, A.P., 1996. Burmese amber at the Natural History Museum. Wrostek 23,
19-20. Rasnitsyn, A.P., Zhang, H.C., 2010. Early evolution of Apocrita (Insecta,
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Hymenoptera) as indicated by new findings in the Middle Jurassic of
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Daohugou, Northeast China. Acta Geologica Sinica (English Edition) 84, 834-
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873.
Rasnitsyn, A.P., Bashkuev, A.S., Kopylov, D.S., Lukashevich, E.D.,
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Ponomarenko, A.G., Popov, Yu.A., Rasnitsyn, D.A., Ryzhkova, O.V.,
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Sidorchuk, E.A., Sukatsheva, I.D., Vorontsov, D.D., 2016. Sequence and scale
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of changes in the terrestrial biota during the Cretaceous (based on materials
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from fossil resins). Cretaceous Research 61, 234-255. Rasnitsyn, A.P., Poinar, G.Jr., Brown, A.E., 2017. Bizzare wingless parasitic wasp
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from mid-Cretaceous Burmese amber (Hymenoptera, Ceraphronoidea,
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Aptenoperissidae fam. nov.). Cretaceous Research 69, 113-118.
379 380 381 382 383
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Rodriguez, J., Waichert, C., Dohlen C.D.von., Poinar, G.Jr., Pitts, J.P., 2016. Eocene and not Cretaceous origin of spider wasps: Fossil evidence from amber.
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Acta Palaeontologica Polonica 61(1), 89-96.
Ross, A.J. 2017. Burmese (Myanmar) amber taxa, on-line checklist v.2017.2. 73pp. http://www.nms.ac.uk/explore/stories/natural-world/burmese-amber/
Ross, A., 2015. Insects in Burmese amber. Entomologentagung 02-05.03. Frankfurt/M. Programm und Abstracts. Frankfurt/Main, p. 72.
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ACCEPTED MANUSCRIPT Ross, A., Mellish, C., York, P., Crighton, B., 2010. Burmese amber, in Penney, D.
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(Eds.), Biodiversity of fossils in amber from the major world deposits. Siri
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Scientific Press, Manchester, 208-235.
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Shi, G., Grimaldi, D.A., Harlow, G.E., Wang, J., Wang, J., Wang, M., Lei, W., Li, Q., Li, X., 2012. Age constraint on Burmese amber based on U-Pb dating of
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zircons. Cretaceous Research 37, 155-163.
Zhang, Q., Rasnitsyn, A.P., Wang, B., Zhang, H., submitted. New data about the
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enigmatic wasp from mid-Cretaceous Burmese amber (Hymenoptera,
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Stephanoidea, Aptenoperissidae). Cretaceous Research.
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Zherikhin, V.V., Ross, A.J., 2000. A review of the history, geology and age of Burmese amber (Burmite). Bulletin of the Natural History Museum, London
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(Geology) 56, 3-10.
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Figure captions
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Fig.1. Myanmarina lisu sp. nov. A-D. Holotype NIGP166310, E-G. Paratype
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NIGP166311. A, B, E, F. Photomicrographs and reconstructions of habitus in side
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views. C. Wing base with indication of vein nomenclature (standard). D. Dorsal
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view. G. Dorsolateral view.
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Fig.2. Myanmarina kachin sp. nov., holotype NIGP166311. Reconstruction (A)
and photomicrograph (B) of habitus in side view.
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Fig.3. Myanmarina lahu sp. nov. A-D. Holotype NIGP166312, E-F. Paratype
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NIGP166313. A, B. Oblique ventral view. C. Dorsal view. D. Forewing. E. Head.
ACCEPTED MANUSCRIPT F. Flagellar segments showing possible multiporous plate sensilla (slit-like
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foramina among circular ones).
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S0195-6671(17)30366-X
DOI:
10.1016/j.cretres.2017.09.015
Reference:
YCRES 3705
To appear in:
Cretaceous Research
Received Date: 19 August 2017 Revised Date:
23 September 2017
Accepted Date: 24 September 2017
Please cite this article as: Zhang, Q., Rasnitsyn, A.P., Wang, B., Zhang, H., Myanmarinidae, a new family of basal Apocrita (Hymenoptera: Stephanoidea) from mid-Cretaceous Burmese amber, Cretaceous Research (2017), doi: 10.1016/j.cretres.2017.09.015. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
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ACCEPTED MANUSCRIPT Myanmarinidae, a new family of basal Apocrita (Hymenoptera:
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Stephanoidea) from mid-Cretaceous Burmese amber
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Qi Zhanga,b, Alexandr P. Rasnitsync,d,*, Bo Wanga,e, Haichun Zhanga
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a
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Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008,
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China
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b
University of Chinese Academy of Sciences, Beijing 100049, China
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c
A.A. Borissiak Palaeontological Institute, Russian Academy of Sciences, 117647
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Moscow, Russia
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d
Natural History Museum, Cromwell Road, London SW7 5BD, UK
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e
Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology,
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Chinese Academy of Sciences, 1, Beichen West Road, Beijing 100101, China
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State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of
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* Corresponding author. A.A. Borissiak Palaeontological Institute, Russian
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Academy of Sciences, 117647 Moscow, Russia.
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Email: [email protected]
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Abstract
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A new family of basal apocritan wasps is established based on three new
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species discovered from the mid-Cretaceous Burmese amber and tentatively allied
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to the superfamily Stephanoidea. The family Myanmarinidae Zhang and Rasnitsyn
ACCEPTED MANUSCRIPT fam. nov. contains one new genus Myanmarina Zhang and Rasnitsyn gen. nov.,
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including three new species: Myanmarina lisu Zhang and Rasnitsyn sp. nov.,
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Myanmarina kachin Zhang and Rasnitsyn sp. nov. and Myanmarina lahu Zhang
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and Rasnitsyn sp. nov. Comparisons are made between Myanmarinidae and its
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similar families. Some general remarks are also made about the new family in
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taxonomy and biology.
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Keywords: Hymenoptera, Myanmarinidae, Myanmarina, mid-Cretaceous, Burmese
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amber
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1. Introduction
Burmese amber is currently one of the richest sources of knowledge about the
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past of insects and some other segments of the non-marine biosphere (Rasnitsyn,
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1996; Zherikhin and Ross, 2000; Grimaldi et al., 2002; Ross et al., 2010; Rasnitsyn
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et al., 2016; Guo et al, 2017). Indeed, the Burmese amber arthropod assemblage
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accounts for 252 families which is the highest figure among the Cretaceous
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Lagerstätten (Rasnitsyn et al., 2016; the data by the end of 2013). Years later Ross
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(2017) lists 356 arthropod families and 481 insect species there, and Guo et al.
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(2017) cite 73 described species in Hymenoptera. Hymenoptera represents a
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significant part of the modern and, since the Jurassic, past world biodiversity: the
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above source counts 30 families, that is, 12 percent of the total arthropod diversity
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in the Burmese amber. Like other taxa, Hymenoptera displays a number of
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ACCEPTED MANUSCRIPT described (Aptenoperissidae, Bryopompilidae, Melittisphecidae, Othniodellithidae,
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Syspathoxyelidae, cf. Rasnitsyn et al., 2017; Rodriguez et al 2016; Poinar and
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Danforth, 2006; Engel et al., 2016a, 2016b; respectively) and undescribed (in
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preparation) endemic families within the Burmese amber assemblage. Description
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of one more such family is the objective of the present publication.
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2. Material and methods
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The present research is based on the amber collected from the amber mines
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situated in the Hukawng Valley of Kachin State, Myanmar (locality in Kania et al.,
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2015: fig. 1). The rock containing the Burmese amber was radiometrically dated at
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98.79 ± 0.62 Ma (Shi et al., 2012). The date corresponds to the early Cenomanian,
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that is, the earliest Late Cretaceous. However, the amber displays clear traces of
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re-deposition (Ross, 2015) and so can be older than enclosing rocks. That is why
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we prefer to refer to the amber age informally as mid-Cretaceous.
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The material was studied under Nikon SMZ-10 R stereoscopic microscope and
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Nikon Optiphot compound microscope with magnifications up to 800× at the State
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Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology
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and Palaeontology, Chinese Academy of Sciences, and using Leica M165C
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stereomicroscope with a Leica DFC 420 camera at the A.A. Borissiak
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Paleontological Institute, Russian Academy of Sciences, in Moscow. In most
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instances, incident and transmitted light were used simultaneously. Helicon Focus
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Pro ×64 was used to stack photos for better depth of field. The line drawings were
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prepared based on photographs using image-editing softwares (CorelDraw X7 and
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ACCEPTED MANUSCRIPT Adobe Photoshop CS6). The specimens are housed in the Nanjing Institute of
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Geology and Palaeontology, Chinese Academy of Sciences (NIGPAS).
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Morphological terminology and symbols to wing vein homology are standard,
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except that ampersand (&) linking two adjacent vein sections denotes those aligned
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and forming a seemingly entire vein section. Particularly, 1-RS&1-M means the
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first (basal) sections of RS and M joining to form a smooth, seemingly entire vein
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(basal vein in older nomenclature; cf. Fig. 1 C). All taxonomic acts established in
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the present work have been registered in ZooBank (see below), together with the
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electronic publication LSID: urn:lsid:zoobank.org:pub:FC26F5AC-E0FF-44CB-
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B621-C68973E0BEAF
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3. Systematic paleontology
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Order Hymenoptera
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Suborder Vespina
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Superfamily Stephanoidea
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Family Myanmarinidae Zhang and Rasnitsyn, fam. nov.
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(LCID: urn:lsid:zoobank.org:act:A6499035-F929-449A-8477-
63D8BCCB0191)
Type genus Myanmarina Zhang and Rasnitsyn, gen. nov.
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Diagnosis. Male (female unknown). Stature elongate, subcylindrical, as in
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Stephanidae. Body size small (1.5-3 mm). Head subglobular, without crown of
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teeth around fore ocellus, with big eyes, circular occipital carina and long genal
ACCEPTED MANUSCRIPT bridge (the space where contralateral genae meet each other between postocciput
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and hypostome). Antenna 11- or 12-segmented, not geniculate, flagellar segments
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more or less uniform, possibly with elongate (multiporous plate) sensillae.
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Mandible short, wide, with 2 or, possibly, 3 subequal teeth, with cutting edge
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subparallel to mandible rotation axis, not distinctly oblique. Maxillary palps with 4
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visible segments, apical three ones narrow and elongate. Labial palps not observed,
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probably much reduced. Mesosoma long and narrow, subcylindrical or somewhat
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depressed (possibly a preservational distortion), its morphology little known except
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that the pronotum is short centrally in dorsal view and the mesoscutum long.
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Propleurae more or less elongate, usually forming a distinct neck. Propodeum
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elongate, straight in lateral view, with no downward bending towards metasomal
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articulation. Legs elongate, ordinary; fore tibia with single, thin, arcuate spur.
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Wings densely pubescent, with comparatively long marginal setae, with venation
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much reduced. Forewing with tubular veins only in anterobasal part, namely fused
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C+R up to some 0.6 wing length, M+Cu, strongly oblique basal vein (1-RS&1-M),
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cu-a, 1-Cu and 1A up to cu-a only (Cu may continue after cu-a as nebulous vein).
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All distal wing surface free of veins, supported with system of nested radiated
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folds like in Spathiopterygidae. Hind wing short and very narrow, reaching level of
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upper end of basal vein (that is, level of 1-RS base), with only C+R tubular and
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with 3 hamuli. Metasoma linear, with 8 external segments, 1st segment distinctly
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narrowed basal but details of articulation not clear, male claspers wide and often
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long, pointing obliquely downward.
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Genera included. Type genus only.
ACCEPTED MANUSCRIPT
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Genus Myanmarina Zhang and Rasnitsyn, gen. nov.
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(LCID: urn:lsid:zoobank.org:act:BB9726BE-39FB-44A5-901A-
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A7791CFEEC21) Type species. Myanmarina lisu Zhang and Rasnitsyn, sp. nov.
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Etymology. From Myanmar, the country where all the three species of the new genus come. Gender feminine.
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Diagnosis. As for family because of monotypy.
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Species included. Three species.
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Myanmarina lisu Zhang and Rasnitsyn, sp. nov.
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(LSID: urn:lsid:zoobank.org:act:5AC41744-1151-4846-A7C6-
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Fig. 1
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FD58CBE42E3A)
Derivation of name. After the Lisu, an ethnic group in northern Myanmar
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where the fossil material was collected. Noun in apposition. Holotype. Male, NIGP166310 (Fig. 1A-D). Only few part of legs damaged,
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diagenetic deformation present, with some scattered debris around; small air
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bubbles distributed around the whole body including the antennae and legs make it
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difficult to discern the dorsal surface sculpture.
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Paratype. Male, NIGP166311 (Fig. 1E-G). One forewing damaged, the other bent; many scattered debris around. Dark body and lots of small bubbles obscuring
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ACCEPTED MANUSCRIPT particular details of the fossil. The type specimens were originally found as
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syninclusions and subsequently separated for study.
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Locality and Horizon. Hukawng Valley, Kachin State, Myanmar; lowermost Cenomanian, mid-Cretaceous. Description. NIGP166310. Head subglobose; ocelli small, well separated from
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compound eyes; compound eyes prominent, berry-like in appearance. Antenna 11-
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segmented, inserted moderately low on head, densely covered with fine setae;
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scape relatively straight; pedicel almost as long as scape; first flagellomere
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extremely long, thinner than pedicel basally and becoming gradually thicker;
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second about as long as scape and pedicel combined; third and fourth slightly
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longer than second and fifth, remaining ones shorter than fifth but still more than
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twice as long as wide, apicalmost flagellomere rounded. Maxillary palps almost as
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long as head height, with three apical segments long and thin, basal visible
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segment probably short. Mesosoma imbricate, long-column-shaped and wider than
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high; pronotum short, mesoscutum long with indistinct (possibly percurrent)
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notauli. Legs long with respect to body length and covered with minute setae; tibial
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spur formula 1-1-1; basitarsomeres almost as long as remainder of corresponding
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tarsomeres together; all femora shorter than their tibia, and hind femur slightly
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swollen. Length ratio of tarsomeres in all legs (similar in all congeners) ca. 10:5:3-
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3.5:1.5-1.7:2-2.5 (precise measurements often difficult because of imperfect
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preservation state and improper position of legs in amber). Forewing longer than
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metasoma, spatulate, with well-defined marginal fringe, fringe setae longest along
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hind margin (possibly also along wing apex which is obscured), moderate along
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ACCEPTED MANUSCRIPT veins on fore margin, growing shorter toward anterior preapical margin; membrane
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with dense and comparatively long setae all over surface distal of venation, with
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basal (1rm) and subbasal (1cua) cells naked; C+R reaching maximum wing width
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(0.64 wing length); M+Cu faded in basal third, branching in wing basal 0.2, 1-
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RS&1-M thickest vein, reaching C+R at 0.33 wing length, cu-a aligned with 1A,
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meeting Cu distinctly distal of M+Cu apex; free Cu long, nebulous, subparallel to
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wing hind margin, continuing into weak fold; five stronger folds running fan-like
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toward wing margin anterior of Cu. Hind wing very narrow, some 0.4 as long as
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forewing, with fringe of moderately long setae along all visible hind margin and
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apex and with 3 hamuli present. Metasoma narrow, about as long as head and
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mesosoma combined, and wider than high, with segments difficult to discern; first
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segment trapezoid, elongate, following apparently shorter; claspers wide and long,
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triangular in side view, otherwise genitalia not visible.
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NIGP166311, providing additional characters such as: mandible possibly tridentate, maxillary palps narrow and long; fore leg with coxa long ovate,
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trochanter small, tibia with inner row of setae; mid and hind coxa almost as long as
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femur.
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Measurements. NIGP166310. Total body length, 2.7 mm; head length, 0.3 mm,
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width, 0.4 mm; mesosomal length,1.0 mm; metasomal length, 1.4 mm; antennal
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length, 1.4 mm; forewing length, 1.9 mm, width, 0.7 mm; hindwing length 0.7 mm.
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NIGP166311. Total body length, 2.3 mm; head length, 0.3 mm, width, 0.3 mm;
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mesosomal length,0.7 mm; metasomal length, 1.3 mm; antennal length, 1.3 mm;
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forewing length, 1.38 mm, width, ca. 0.6-0,65 mm; hindwing length 0.55 mm.
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ACCEPTED MANUSCRIPT Remarks. The species differs from all congeners in having antennae distinctly
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thickened apically, hind coxae and genital claspers very long. It is noteworthy that
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high size variability is present in the species, with forewing length differing in
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almost 1.4 times between two males available. Fig.1
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Myanmarina kachin Zhang and Rasnitsyn, sp. nov.
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(LSID: urn:lsid:zoobank.org:act:148831C2-8910-4229-8E5E-4F36DC8F465D)
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Fig. 2.
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Derivation of name. After the Kachin, an ethnic group in northern Myanmar where the fossil material comes. Noun in apposition.
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Holotype. Male, NIGP166309. Nearly complete with diagenetic deformation,
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with antennae and hind legs partly damaged; amber inner layer surface light brown;
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the surface sculpture not discernible.
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Locality and Horizon. Hukawng Valley, Kachin State, Myanmar; lowermost
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Cenomanian, mid-Cretaceous. Description. Body dark brown to black. Head subglobose. Eyes not easily
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recognizable. Antenna inserted moderately low on head, slightly widened apically,
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11-segmented; scape slightly expanded ventrally, about twice the length of pedicel;
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pedicel small, slightly longer than wide; first flagellomere thin and extremely long,
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arched softly and becoming thicker apically; flagellomeres 2-5 similar in size and
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shape, about half as long as first and slightly thinner in both ends than middle part;
ACCEPTED MANUSCRIPT flagellomeres 6-8 shorter and broader, flagellomere 9 almost as long as 8 and
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slightly pointed at apex. Maxillary palps as visible as long as head high, with 4
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segments visible, apical three narrow and elongate, first visible segment much
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thicker and probably short. Mesosoma subcylindrical with reticulated surface
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(possibly because of cuticle degradation), narrow and more than 3 times as long as
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wide. Pronotum short and mesoscutum long; propodeum elongate, straight in
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lateral view, with no downward bending towards metasomal articulation. Legs
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long and thin on the whole, coxa long and moderately stout; trochanter thin;
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tarsomere length as in type species. Protibial spur simple and curved. Fore and mid
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legs with femur cylindrical and shorter than tibia; tibia expanded apically. Fore
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basitarsus as long as tibia; mid basitarsus longer than tibia. Hind legs with femur
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slightly swollen; tibia longer than femur, with single tibial spur. Forewing hyaline
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with moderately dense setation, with short fringe along hind margin (possibly also
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along wing apex which is obscured). Venation incompletely visible, with fused
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C+R long, about 3/5 length of wing; 1-Rs&1-M strongly oblique, very thick,
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connecting to C+R near its midlength; nested forking folds present distal of
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preserved veins, with hind fold apparently continuing into nebulous free Cu. Hind
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wing short and very narrow, with 3 hamuli present. Metasoma linear, longer than
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mesosoma, with segment boundaries difficult to identify precisely. Claspers wide
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and short, other details of genitalia invisible.
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Measurements. NIGP166309. Total body length, 3.2 mm; head length, 0.4 mm,
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width, 0.2 mm; mesosomal length,1.2 mm; metasomal length, 1.6 mm; antennal
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length, 1.5 mm; forewing length, 1.8 mm, width, 0.7 mm.
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ACCEPTED MANUSCRIPT Remarks. The species differs from all congeners by its larger size and the hind
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femur comparatively short and rather stout (instead of long and at most slightly
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swollen in both M. lisu and M. lahu). Fig.2
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(LSID: urn:lsid:zoobank.org:act:ECC6AE11-5B6C-4306-9ECB-
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BE229F17BCE1) Fig. 3
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Derivation of name. After the Lahu, an ethnic group in northern Myanmar where the fossil material was discovered. Noun in apposition.
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Holotype. Male, NIGP166312 (Fig. 3A-D). Nearly complete fossil missing
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only one wing apex, deformed in many respects; amber color faint yellow and not
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well transparent, inner surface deep yellow colored; many scattered debris around;
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surface sculpture and many details of morphology of holotype are difficult to see
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and interprete correctly because of diagenetic deformations as well as deep dark
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color.
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Paratype. Male, NIGP166313 (Fig. 3E-F). Head and part of legs damaged, lots
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of scattered debris and some filaments around; amber not well transparent, dark
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body and lots of small bubbles obscuring particular details of the fossil.
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The type specimens were originally found as syninclusions and separated subsequently.
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Cenomanian, mid-Cretaceous. Description. NIGP166312. Specimen dark brown to black. Head subglobose, longer than high. Eyes berry-like, moderately large, reaching mandibular base;
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ocelli indistinguishable. Antenna inserted low on head, below eye midheight, 12-
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segmented; scape ventrally slightly expanded, about twice the length of pedicel;
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flagellomeres densely covered with thin setae, with first three antennal segments
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pale compared to remainders; first one moderately long, thinner than pedicel
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basally, second to sixth as long as first, seventh to tenth slightly shorter than sixth,
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apical one rounded. Mandible narrowed apically. Maxillary palps some half as
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long as head high, with three apical segments long and thin. Mesosoma very long,
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wider than height. Pronotum extremely short centrally, hardly visible in dorsal
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view; propleurae elongate forming distinct neck; mesoscutum long; propodeum
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elongate and straight. Legs long with respect to body length and covered with
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minute setae; coxa long, trochanters small; fore and mid femora longer than
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respective tibia, hind femur slightly swollen and shorter than tibia; tarsomere
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length ratio as in type species. Forewing as preserved similar to that in other
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congeners. Hindwing short with membrane, without wing venation apart from C+R,
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and with three hamuli present. Metasoma long and narrow, with 8 external tergites,
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second longest, eighth shortest. Claspers wide and short, other parts of genitalia not
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visible.
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ACCEPTED MANUSCRIPT Specimen NIGP166313 not well visible, very small, head big, mandible
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narrowed apically, distinctly with two subequal teeth, hind coxa and femur long.
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Flagellomeres possibly with multiporous plate sensilla (Fig. 3F).
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Measurements. NIGP166312. Total body length, 2.9 mm; head length, 0.4 mm, width, 0.3 mm; mesosomal length,1.1 mm; metasomal length, 1.7 mm; antennal
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length, 1.8 mm; forewing length, 1.9 mm; hind wing length ca. 0.65 mm.
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NIGP166313. Head length, 0.3 mm; mesosomal length, 0.85 mm; antennal length, 1.7 mm.
Remarks. The species differs from all congeners in having the antenna 12-
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segmented, the first flagellomere hardly 1.5 times as long as apical one and the
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maxillary palps short (antenna 11-segmented with the first flagellomere several
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times as long the apical one, and maxillary palp long in both M. lisu and M. lahu).
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Fig.3
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4. Discussion
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The present fossils are imperfectly preserved, and therefore they are
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insufficiently known in morphology. However, the morphological characters
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known from these specimens are distinctive enough so as so warrant description of
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the fossils as a family of their own. Accommodation of the new family in the wasp
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tree is not straightforward as well. Its distinctive similarity to the
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Spathiopterygidae in wing morphology cannot be inherited because of too different
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organization of propodeum and metasoma. In Spathiopterygidae, the first
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metasomal segment is transformed into a petiole and attached low on the
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other features scrutinized by Engel et al. (2013), this is clearly indicative of
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Diaprioidea. In contrast, propodeum of the new family is straight in side view
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lacking a posterior slope, and hence a real constriction is absent between the
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propodeum and the metasoma. This character state is characteristic of Stephanidae
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and extinct families Ephialtitidae and Aptenoperissidae, that is, of the superfamily
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Stephanoidea (Rasnitsyn and Zhang, 2010; Rasnitsyn et al., 2017; Zhang et al.,
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submitted). That is why we attribute Myanmarinidae to Stephanoidea, at least until
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a sound contradictory information appears.
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Myanmarinidae can be easily differentiated from all three other families of Stephanoidea in their small size, 11-segmented antenna and particularly in their
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specific mode of reduction of the wing venation: particularly with two
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incompletely separated basalmost cells margined with thick veins within the basal
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third of wing's length, with no other closed cells, and with distinctly radially folded
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outer wing membrane. Unlike the new family, Ephialtitidae and male
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Aptenoperissidae (Zhang et al., submitted) always have more complete venation
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(female apterous), and Stephanidae have venation varying from complete to
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strongly reduced but never displaying deep differentiation between the wing base
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and the remainder wing blade.
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5. Concluding remarks The biology of the new family could be very tentatively inferred from the general appearance of the available fossils. Their long and narrow body and
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probably xylobiotic preys. This agrees additionally with the proposed taxonomic
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position of Myanmarinidae, because Stephanidae are known to develop at expense
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of xylophagous beetles (Achterberg, 2002), and Ephialtitidae are hypothesized to
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behave similarly (Rasnitsyn, 1980). The distinct variation of the body size within
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the males of M. lisu sp. nov. is compatible with the hypothesis as well.
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Acknowledgements
This research was supported by the National Natural Science Foundation of
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China (41272013, 41572010, 41622201 and 41688103), the Chinese Academy of
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Sciences (XDPB05), and Youth Innovation Promotion Association of CAS (No.
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2011224).The financial support of the UCAS (UCAS[2015]37) Joint PhD Training
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Program to ZQ is acknowledged. We offer our sincere gratitude to Dr. Ekaterina A.
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Sidorchuk for providing technical guidance, and the anonymous reviewers for the
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very useful comments on the earlier version of the manuscript.
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Figure captions
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Fig.1. Myanmarina lisu sp. nov. A-D. Holotype NIGP166310, E-G. Paratype
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NIGP166311. A, B, E, F. Photomicrographs and reconstructions of habitus in side
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views. C. Wing base with indication of vein nomenclature (standard). D. Dorsal
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view. G. Dorsolateral view.
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Fig.2. Myanmarina kachin sp. nov., holotype NIGP166311. Reconstruction (A)
and photomicrograph (B) of habitus in side view.
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Fig.3. Myanmarina lahu sp. nov. A-D. Holotype NIGP166312, E-F. Paratype
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NIGP166313. A, B. Oblique ventral view. C. Dorsal view. D. Forewing. E. Head.
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foramina among circular ones).
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