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New Palaeontinidae (Insecta: Hemiptera) from the Lower Cretaceous of southern England
Accepted Manuscript New Palaeontinidae (Insecta: Hemiptera) from the Lower Cretaceous of southern England Yuling Li, Edmund Jarzembowski, Jun Chen, Bo Wang PII:
S0195-6671(18)30341-0
DOI:
https://doi.org/10.1016/j.cretres.2018.11.019
Reference:
YCRES 4022
To appear in:
Cretaceous Research
Received Date: 14 August 2018 Revised Date:
15 October 2018
Accepted Date: 21 November 2018
Please cite this article as: Li, Y., Jarzembowski, E., Chen, J., Wang, B., New Palaeontinidae (Insecta: Hemiptera) from the Lower Cretaceous of southern England, Cretaceous Research, https:// doi.org/10.1016/j.cretres.2018.11.019. 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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New Palaeontinidae (Insecta: Hemiptera) from the Lower Cretaceous of
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southern England
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Yuling Lia,b, Edmund Jarzembowskic, Jun Chena,d, Bo Wanga,e
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a
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Geology and Palaeontology and Center for Excellence in Life and Paleoenvironment,
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Chinese Academy of Sciences, 39 East Beijing Road, Nanjing 210008, China.
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b
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276000, China.
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e
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Minerals, Shandong University of Science and Technology, Qingdao, Shandong
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266590, China.
University of the Chinese Academy of Sciences, Beijing 100049, China.
Department of Earth Sciences, Natural History Museum, London SW7 5BD, UK.
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Institute of Geology and Paleontology, Linyi University, Shuangling Road, Linyi
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Shandong Provincial Key Laboratory of Depositional Mineralization & Sedimentary
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State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of
Corresponding author: [email protected]
Abstract
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Two new species of butterfly bug, Valdicossus mikewebsteri sp. nov. and
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Ilerdocossus prowsei sp. nov. are described from the Lower Cretaceous of southern
ACCEPTED MANUSCRIPT England. Valdicossus mikewebsteri is the first complete hindwing of a palaeontinid
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from the UK. It is different from the type species in having a hindwing with a
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different colour pattern, vein Sc+R straight, and branch RA2 slightly curved
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posteriorly then recurved; the length of vein RP+MP1 is shorter than that of the type
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species and vein A1 is short. Ilerdocossus prowsei sp. nov. is described based on a
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well-preserved forewing from the Lower Weald Clay (upper Hauterivian) of England;
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it is different from other species of the genus in possessing a narrower forewing and
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vein CuA bifurcating beyond the nodal line; the membrane is pitted distal of the nodal
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line.
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Key words: Valdicossus, Ilerdocossus, Palaeontinidae, Wealden, England
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1. Introduction
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Palaeontinidae (Insect: Hemiptera) is an extinct family of hemipterous insects from the Triassic (Carnian) to Cretaceous (Aptian) (Szwedo, 2018),, and their fossils
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have been discovered in Europe, Central Asia, Russia, China, South Korea, Africa and
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South America (Shcherbakov and Popov, 2002; Menon et al., 2005; Chen et al., 2016;
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Nam et al., 2017). The first palaeontinid was described from the Middle Jurassic of
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England and mistaken for a lepidopteran (Butler, 1873; Tillyard, 1921). So far,
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Cretaceous palaeontinids, belonging to more than 30 species in 12 genera, have been
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discovered in Spain, Russia, China, England and Brazil (Handlirsch, 1906; Martynov,
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1926; Gómez-Pallerola, 1984; Whalley and Jarzembowski, 1985; Shcherbakov, 1988;
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Ren, 1995, 1998; Ueda, 1996, 2008; Martins-Neto, 1998; Menon and Heads, 2005;
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Menon et al., 2005; Wang et al., 2008a).
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The early Cretaceous palaeontinids from Europe are mainly reported from Spain and England. The Lithographic Limestone of Montsech, Lérida province, Spain is a
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facies development of the Caliza con Caraćeas Formation (Garrido-Megias and Rios
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Aragües, 1972) and was deposited in a lagoonal-lacustrine environment which
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became stagnant at times (Schairer and Janicke, 1970). The Spanish palaeontinids
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from the Lithographic Limestone are as follows: Pachepsyche vidali Meunier, 1902;
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Montsecocossus sererai Gómez-Pallerola, 1984; Ilerdocossus villaltai
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Gómez-Pallerola, 1984; Wonnacottella pulcherrima Whalley and Jarzembowski, 1985
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(Meunier, 1902; Gómez-Pallerola, 1984; Whalley and Jarzembowski, 1985). The
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Wealden Supergroup of southern England has yielded a unique and abundant Early
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Cretaceous insect fauna (Jarzembowski, 1995), but palaeontinids are uncommon. The
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first Wealden palaeontinid, a forewing, was reported from the Lower Weald Clay
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Formation which was probably attributable to the genus Ilerdocossus
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Gómez-Pallerola, 1984 (Jarzembowski, 1984). A second Wealden palaeontinid,
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Valdicossus chesteri, was described on the basis of a well-preserved hindwing which
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was the first hindwing of Palaeontinidae from the UK (Wang et al., 2008a). Here we
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describe two new species from two different localities: Valdicossus mikewebsteri sp.
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nov. is from a new palaeontinid locality, Smokejacks brickworks in southern England,
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and is the first complete hindwing of Palaeontinidae from the UK. Ilerdocossus
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prowsei sp. nov. is based on a well-preserved forewing from the former Clock House
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Brickworks, also in southern England.
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2. Material and methods
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The material was found by Mike Webster and one of us (EAJ) in the county of Surrey, southern England (Fig.1), and is deposited in the Natural History Museum
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(London) and Booth Museum of Natural History (Brighton) respectively. The
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specimens were prepared by EAJ with a Burgess vibrotool under a Soviet MBS-1
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binocular microscope.
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The wing venation nomenclature of Palaeontinidae used in this paper is based on the interpretations by Wang B. et al. (2009) and Nel et al. (2012). The line drawings
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were readjusted on photographs using image-editing software (Coreldraw 2017 and
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Adobe Photoshop CS). In the drawings, dashed lines denote the nodal line in the
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forewing, and grey solid lines indicate faint and extrapolated veins.
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3. Systematic palaeontology
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Order: Hemiptera Linnaeus, 1758
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Infraorder: Cicadomorpha Evans, 1946
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Superfamily: Palaeontinoidea Handlirsch, 1906
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Family: Palaeontinidae Handlirsch, 1906
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Genus Valdicossus Wang, Zhang, and Jarzembowski, 2008
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Type species. Valdicossus chesteri Wang, Zhang, and Jarzembowski, 2008; by original designation and monotypy.
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Type horizon and locality. Lower Weald Clay (lower Hauterivian), Cooden, East Sussex, southern England.
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Species included. Two species, Valdicossus chesteri Wang, Zhang and
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Jarzembowski, 2008 and V. mikewebsteri sp. nov, both from the Lower Cretaceous of
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England.
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Genus Valdicossus Wang, Zhang, and Jarzembowski, 2008
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Valdicossus mikewebsteri sp. nov.
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Fig. 2A-C
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Etymology. After Mike Webster, fossil finder.
Holotype. NHMUK II S4046a, b, part and counterpart; hind wing. Deposited in
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The Natural History Museum, London; Upper Weald Clay Formation, below BGS bed
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5c, Lower Cretaceous (lower Barremian), Smokejacks brickworks, Surrey, England.
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Diagnosis. Costal area wide and more than half length of wing. RA1 and RA2 arising from RA1+2 at same level of indentation; branch RA2 slightly curved
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posteriorly then recurved. Vein MP3+4 unbranched and slightly curved; vein CuP
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slightly curved anteriorly; vein Pcu curved posteriorly; A1 short; membrane with
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spotty colour pattern.
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Description. Hindwing oval, with length 16.8 mm long; width 12 mm wide.
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Costal margin with distinct indentation. Anterior margin curved in distal part. Costal
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area wide and more than half length of the wing, length/width ratio 2.9, maximal
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width in basal 1/4th. Vein R fused with vein Sc basally and branching into veins
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Sc+RA and RP. Branch Sc separating from vein Sc+RA and ending in costal margin
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just basad of wing indentation. RA1 and RA2 arising from RA1+2 opposite indentation,
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RA1 ending in costal margin just distad of indentation. Branch RA2 slightly curved
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posteriorly then recurved, ending on margin. Vein RP branching from stem R at 0.2 of
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wing length, then fusing with vein MP1 for a long distance basad of indentation and
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finally bifurcating a little basad of wing indentation. Vein RP+MP1 length 2.4 mm,
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veins MP1, MP2 and MP3+4 arising from stem MP at wing base; branch MP1 almost
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straight; branch MP2 slightly curved anteriorly; vein MP3+4 unbranched and slightly
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curved; stem CuA forking into veins CuA1 and CuA2 basad of indentation; branches
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CuA1 and CuA2 long, branch CuA2 slightly curved. Vein CuP slightly curved
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anteriorly. Vein Pcu curved posteriorly. Vein A1 short. Wing membrane infuscate with
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three transverse light fasciae.
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Genus Ilerdocossus Gómez-Pallerola, 1984
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Type species. Ilerdocossus villaltai Gómez-Pallerola, 1984; by original
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designation.
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Species included. Nine species, I. villaltai and I. pulcherrima (Whalley and
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Jarzembowski, 1985), I. hui (Ren et al., 1998), I. beipiaoensis (Ren et al., 1998) and I.
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exiguus (Ren et al., 1998), I. fengningensis (Ren et al., 1998), I. pingquanensis (Ren
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et al., 1998), and I. ningchengensis (Wang et al., 2008b), and new species I. prowsei
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sp. nov..
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Genus Ilerdocossus Gómez-Pallerola, 1984
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Ilerdocossus prowsei sp. nov.
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Figs. 3A-C
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Etymology. After Alan Prowse, fossil collector. Holotype. BMB 014927, 018805; forewing; Lower Weald Clay Formation
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beneath BGS bed 3/3a, Lower Cretaceous (upper Hauterivian); Clockhouse
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brickworks, near Capel, Surrey, England, UK.
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Diagnosis. Forewing narrow, large size; vein Cu divided into CuA and CuP at
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slightly curved; discal cell about one-third of wing length; stem MP dividing into
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MP1+2 and MP3+4 before nodal line; vein MP1+2 bifurcating distad of indentation; vein
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MP3+4 bifurcating basad of indentation; branch CuA1 strongly curved anteriorly,
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branch CuA2 curved posteriorly then recurved; clavus absent. Membrane pitted distad
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of nodal line.
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Description. Forewing triangular, with length 59 mm, width 25 mm. Costal
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margin slightly curved anteriorly. Nodal indentation distinct, at about basal 0.42 of
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wing length. Vein Sc arising basally, fused with stem R+MP at basal 0.17 of wing
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length. Stem R+MP curved, connected to branch CuA by crossvein r+mp-cua at same
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point where stem Cu branches into veins CuA and CuP. Crossvein r+mp-cua long
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and oblique. Branches RA, RP and M departing from stem R+MP at same point.
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Branch RA slightly curved anteriorly. Branch RP connected to branch MP1 by
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crossvein r-mp1 distad of indentation. Stem M dividing into MP1+2 and MP3+4 before
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nodal line. Branch MP1+2 branching into veins MP1 and MP2 distad of indentation.
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Branch MP3+4 branching into MP3 and MP4 at basal 0.36 of wing length. Branch MP1
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partly directed anteriorly, geniculate at junction with crossvein r-mp1. Branch MP3
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almost straight then curved anteriorly near margin. Crossvein mp4-cua long, oblique
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and slightly curved. Stem Cu branching into veins CuA and CuP at basal 0.09 of wing
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length. Branch CuA slightly curved posteriorly between crossveins r+mp-cua and
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mp4-cua, branching into veins CuA1 and CuA2 after crossvein mp4-cua, basad of nodal
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branch CuA2 curved posteriorly then recurved. Discal cell length 19 mm, width 8.5
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mm at nodal line, about one-third of wing length; antenodal region trapezoidal,
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postnodal region less than half of antenodal region in length. Nodal line traceable as a
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crease across RA and RP to MP1+2, M3+4 and then separating discal cell into two parts;
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it continues crossing CuA just before its initial division, clavus absent. Membrane
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pitted distal of nodal line.
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4. Discussion
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Valdicossus Wang et al., 2008 was erected based on a well-preserved but
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incomplete hindwing from the Lower Cretaceous Wealden Supergroup of southern
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England (Wang et al., 2008a). Valdicossus mikewebsteri sp. nov. was placed in the
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genus Valdicossus based on the following features: hindwing oval, anterior margin
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curved in distal part; costal area wide; vein R fused with vein Sc basally, and
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branching into veins Sc+RA and RP; vein Sc terminating in costal margin just basad
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of wing indentation; branch RP fused with vein MP1 for a long distance basad of level
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of indentation; veins MP1, MP2 and MP3+4 arising from stem MP at wing base; vein
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MP3+4 unbranched. The new species and the type species are from different locations
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and horizons; it differs from the latter in the hindwing having a different colour
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pattern; vein Sc+R straight; branch RA2 slightly curved posteriorly then recurved;
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vein RP bifurcating from vein RP+MP1 basad of vein Sc separating from vein Sc+RA,
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short. They both have the following characters: anterior margin curved in the distal
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part; vein R fused with vein Sc basally and branching into veins Sc+RA and RP; veins
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MP1, MP2 and MP3+4 arising from stem MP at the wing base; vein MP3+4 is
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unbranched and vein RP fused with vein MP1 for a long distance basad of the
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indentation.
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Ilerdocossus Gómez-Pallerola, 1984 was erected based on a forewing from the
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Lower Cretaceous of Lérida, Spain. Menon et al. (2005) considered both
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Wonnacottella and Liaocossus to be the junior synonyms of Ilerdocossus. Wang B. et
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al. (2008b) also considered three genera (Ilerdocossus, Wonnacottella, and Liaocossus)
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as belonging to the same genusand our material also provides further support this
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view.
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We placed Ilerdocossus prowsei sp. nov. in the genus Ilerdocossus based on the following features: forewing triangular, with narrow costal area; reduced clavus;
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nodal indentation distinct; RA, RP and MP separate at the same point; discal cell long;
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antenodal region trapezoid. It mostly resembles Wonnacottella pulcherrimma
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(Fig.3D)(Whalley and Jarzembowski, 1985) from Spain: they both have a narrow
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forewing with a similar venation, and vein MP1+2 bifurcating distad of the indentation;
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vein MP3+4 bifurcates basad of the indentation, branch CuA2 is curved posteriorly
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then recurved; and the membrane is pitted distal of the nodal line. But the former
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differs from the latter in having a much larger and more narrow forewing, vein Sc
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into CuA and CuP at the same point where vein CuA meets crossvein r+mp-cua; stem
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M divides into MP1+2 and MP3+4 before the nodal line, vein CuA bifurcates beyond
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the line, cross vein r-mp1 is long. It is also distinctly different from other species of
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Ilerdocossus in having a forewing with membrane pitted distal of the nodal line, and
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vein CuA bifurcating beyond the nodal line.
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5. Conclusions
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Based on two well-preserved fossils from England, the new species Valdicossus mikewebsteri sp. nov. and Ilerdocossus prowsei sp. nov. are proposed. Our new
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material further confirms that Ilerdocossus was probably widely distributed in Eurasia
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and was diverse in the Early Cretaceous whereas Valdicossus appears to be a Wealden
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endemic.
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Acknowledgements
This research was supported by the National Natural Science Foundation of
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China (41572010, 41622201, 41688103, 41502007), the Strategic Priority Research
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Program (B) of the Chinese Academy of Sciences (XDB26000000), and the State Key
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Laboratory of Palaeobiology and Stratigraphy (Nanjing Institute of Geology and
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Palaeontology, CAS) (183101 and 183105). This is a Leverhulme Emeritus
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Fellowship contribution for EAJ. We thank Terry Keenan (London) for help with
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photography.
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Figure legends：
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Fig.1. Map showing the location of insect fossil localities. S: Smokejacks brickworks;
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C: Clockhouse brickworks.
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Fig.2. A-C, Valdicossus mikewebsteri sp. nov. A, Holotype NHMUK II S4046a, part;
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B, counterpart of holotype NHMUK II S4046b. C, line drawing of holotype. D,
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Valdicossus chesteri Wang et al., 2008, hindwing redrawn from Wang et al. (2008a).
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Scale bars represent 5 mm.
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Fig.3. A-C, Ilerdocossus prowsei sp. nov. A, Holotype BMB 014927 (CHa5), part; B,
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counterpart of holotype BMB 018805. C, line drawing of holotype. D, Wonnacottella
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pulcherrimma Whalley and Jarzembowski, 1985, hindwing redrawn from Whalley
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and Jarzembowski (1985). Scale bars represent 5 mm.
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S0195-6671(18)30341-0
DOI:
https://doi.org/10.1016/j.cretres.2018.11.019
Reference:
YCRES 4022
To appear in:
Cretaceous Research
Received Date: 14 August 2018 Revised Date:
15 October 2018
Accepted Date: 21 November 2018
Please cite this article as: Li, Y., Jarzembowski, E., Chen, J., Wang, B., New Palaeontinidae (Insecta: Hemiptera) from the Lower Cretaceous of southern England, Cretaceous Research, https:// doi.org/10.1016/j.cretres.2018.11.019. 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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New Palaeontinidae (Insecta: Hemiptera) from the Lower Cretaceous of
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southern England
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Yuling Lia,b, Edmund Jarzembowskic, Jun Chena,d, Bo Wanga,e
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a
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Geology and Palaeontology and Center for Excellence in Life and Paleoenvironment,
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Chinese Academy of Sciences, 39 East Beijing Road, Nanjing 210008, China.
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b
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c
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d
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276000, China.
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e
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Minerals, Shandong University of Science and Technology, Qingdao, Shandong
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266590, China.
University of the Chinese Academy of Sciences, Beijing 100049, China.
Department of Earth Sciences, Natural History Museum, London SW7 5BD, UK.
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Institute of Geology and Paleontology, Linyi University, Shuangling Road, Linyi
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Shandong Provincial Key Laboratory of Depositional Mineralization & Sedimentary
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State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of
Corresponding author: [email protected]
Abstract
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Two new species of butterfly bug, Valdicossus mikewebsteri sp. nov. and
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Ilerdocossus prowsei sp. nov. are described from the Lower Cretaceous of southern
ACCEPTED MANUSCRIPT England. Valdicossus mikewebsteri is the first complete hindwing of a palaeontinid
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from the UK. It is different from the type species in having a hindwing with a
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different colour pattern, vein Sc+R straight, and branch RA2 slightly curved
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posteriorly then recurved; the length of vein RP+MP1 is shorter than that of the type
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species and vein A1 is short. Ilerdocossus prowsei sp. nov. is described based on a
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well-preserved forewing from the Lower Weald Clay (upper Hauterivian) of England;
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it is different from other species of the genus in possessing a narrower forewing and
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vein CuA bifurcating beyond the nodal line; the membrane is pitted distal of the nodal
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line.
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Key words: Valdicossus, Ilerdocossus, Palaeontinidae, Wealden, England
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1. Introduction
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Palaeontinidae (Insect: Hemiptera) is an extinct family of hemipterous insects from the Triassic (Carnian) to Cretaceous (Aptian) (Szwedo, 2018),, and their fossils
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have been discovered in Europe, Central Asia, Russia, China, South Korea, Africa and
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South America (Shcherbakov and Popov, 2002; Menon et al., 2005; Chen et al., 2016;
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Nam et al., 2017). The first palaeontinid was described from the Middle Jurassic of
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England and mistaken for a lepidopteran (Butler, 1873; Tillyard, 1921). So far,
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Cretaceous palaeontinids, belonging to more than 30 species in 12 genera, have been
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discovered in Spain, Russia, China, England and Brazil (Handlirsch, 1906; Martynov,
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1926; Gómez-Pallerola, 1984; Whalley and Jarzembowski, 1985; Shcherbakov, 1988;
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Ren, 1995, 1998; Ueda, 1996, 2008; Martins-Neto, 1998; Menon and Heads, 2005;
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Menon et al., 2005; Wang et al., 2008a).
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The early Cretaceous palaeontinids from Europe are mainly reported from Spain and England. The Lithographic Limestone of Montsech, Lérida province, Spain is a
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facies development of the Caliza con Caraćeas Formation (Garrido-Megias and Rios
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Aragües, 1972) and was deposited in a lagoonal-lacustrine environment which
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became stagnant at times (Schairer and Janicke, 1970). The Spanish palaeontinids
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from the Lithographic Limestone are as follows: Pachepsyche vidali Meunier, 1902;
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Montsecocossus sererai Gómez-Pallerola, 1984; Ilerdocossus villaltai
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Gómez-Pallerola, 1984; Wonnacottella pulcherrima Whalley and Jarzembowski, 1985
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(Meunier, 1902; Gómez-Pallerola, 1984; Whalley and Jarzembowski, 1985). The
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Wealden Supergroup of southern England has yielded a unique and abundant Early
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Cretaceous insect fauna (Jarzembowski, 1995), but palaeontinids are uncommon. The
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first Wealden palaeontinid, a forewing, was reported from the Lower Weald Clay
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Formation which was probably attributable to the genus Ilerdocossus
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Gómez-Pallerola, 1984 (Jarzembowski, 1984). A second Wealden palaeontinid,
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Valdicossus chesteri, was described on the basis of a well-preserved hindwing which
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was the first hindwing of Palaeontinidae from the UK (Wang et al., 2008a). Here we
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describe two new species from two different localities: Valdicossus mikewebsteri sp.
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nov. is from a new palaeontinid locality, Smokejacks brickworks in southern England,
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and is the first complete hindwing of Palaeontinidae from the UK. Ilerdocossus
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prowsei sp. nov. is based on a well-preserved forewing from the former Clock House
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Brickworks, also in southern England.
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2. Material and methods
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The material was found by Mike Webster and one of us (EAJ) in the county of Surrey, southern England (Fig.1), and is deposited in the Natural History Museum
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(London) and Booth Museum of Natural History (Brighton) respectively. The
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specimens were prepared by EAJ with a Burgess vibrotool under a Soviet MBS-1
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binocular microscope.
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The wing venation nomenclature of Palaeontinidae used in this paper is based on the interpretations by Wang B. et al. (2009) and Nel et al. (2012). The line drawings
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were readjusted on photographs using image-editing software (Coreldraw 2017 and
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Adobe Photoshop CS). In the drawings, dashed lines denote the nodal line in the
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forewing, and grey solid lines indicate faint and extrapolated veins.
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3. Systematic palaeontology
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Order: Hemiptera Linnaeus, 1758
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Infraorder: Cicadomorpha Evans, 1946
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Superfamily: Palaeontinoidea Handlirsch, 1906
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Family: Palaeontinidae Handlirsch, 1906
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Genus Valdicossus Wang, Zhang, and Jarzembowski, 2008
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Type species. Valdicossus chesteri Wang, Zhang, and Jarzembowski, 2008; by original designation and monotypy.
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Type horizon and locality. Lower Weald Clay (lower Hauterivian), Cooden, East Sussex, southern England.
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Species included. Two species, Valdicossus chesteri Wang, Zhang and
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Jarzembowski, 2008 and V. mikewebsteri sp. nov, both from the Lower Cretaceous of
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England.
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Genus Valdicossus Wang, Zhang, and Jarzembowski, 2008
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Valdicossus mikewebsteri sp. nov.
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Fig. 2A-C
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Etymology. After Mike Webster, fossil finder.
Holotype. NHMUK II S4046a, b, part and counterpart; hind wing. Deposited in
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The Natural History Museum, London; Upper Weald Clay Formation, below BGS bed
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5c, Lower Cretaceous (lower Barremian), Smokejacks brickworks, Surrey, England.
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Diagnosis. Costal area wide and more than half length of wing. RA1 and RA2 arising from RA1+2 at same level of indentation; branch RA2 slightly curved
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posteriorly then recurved. Vein MP3+4 unbranched and slightly curved; vein CuP
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slightly curved anteriorly; vein Pcu curved posteriorly; A1 short; membrane with
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spotty colour pattern.
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Description. Hindwing oval, with length 16.8 mm long; width 12 mm wide.
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Costal margin with distinct indentation. Anterior margin curved in distal part. Costal
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area wide and more than half length of the wing, length/width ratio 2.9, maximal
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width in basal 1/4th. Vein R fused with vein Sc basally and branching into veins
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Sc+RA and RP. Branch Sc separating from vein Sc+RA and ending in costal margin
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just basad of wing indentation. RA1 and RA2 arising from RA1+2 opposite indentation,
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RA1 ending in costal margin just distad of indentation. Branch RA2 slightly curved
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posteriorly then recurved, ending on margin. Vein RP branching from stem R at 0.2 of
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wing length, then fusing with vein MP1 for a long distance basad of indentation and
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finally bifurcating a little basad of wing indentation. Vein RP+MP1 length 2.4 mm,
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veins MP1, MP2 and MP3+4 arising from stem MP at wing base; branch MP1 almost
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straight; branch MP2 slightly curved anteriorly; vein MP3+4 unbranched and slightly
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curved; stem CuA forking into veins CuA1 and CuA2 basad of indentation; branches
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CuA1 and CuA2 long, branch CuA2 slightly curved. Vein CuP slightly curved
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anteriorly. Vein Pcu curved posteriorly. Vein A1 short. Wing membrane infuscate with
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three transverse light fasciae.
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Genus Ilerdocossus Gómez-Pallerola, 1984
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Type species. Ilerdocossus villaltai Gómez-Pallerola, 1984; by original
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designation.
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Species included. Nine species, I. villaltai and I. pulcherrima (Whalley and
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Jarzembowski, 1985), I. hui (Ren et al., 1998), I. beipiaoensis (Ren et al., 1998) and I.
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exiguus (Ren et al., 1998), I. fengningensis (Ren et al., 1998), I. pingquanensis (Ren
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et al., 1998), and I. ningchengensis (Wang et al., 2008b), and new species I. prowsei
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sp. nov..
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Genus Ilerdocossus Gómez-Pallerola, 1984
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Ilerdocossus prowsei sp. nov.
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Figs. 3A-C
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Etymology. After Alan Prowse, fossil collector. Holotype. BMB 014927, 018805; forewing; Lower Weald Clay Formation
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beneath BGS bed 3/3a, Lower Cretaceous (upper Hauterivian); Clockhouse
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brickworks, near Capel, Surrey, England, UK.
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Diagnosis. Forewing narrow, large size; vein Cu divided into CuA and CuP at
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slightly curved; discal cell about one-third of wing length; stem MP dividing into
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MP1+2 and MP3+4 before nodal line; vein MP1+2 bifurcating distad of indentation; vein
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MP3+4 bifurcating basad of indentation; branch CuA1 strongly curved anteriorly,
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branch CuA2 curved posteriorly then recurved; clavus absent. Membrane pitted distad
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of nodal line.
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Description. Forewing triangular, with length 59 mm, width 25 mm. Costal
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margin slightly curved anteriorly. Nodal indentation distinct, at about basal 0.42 of
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wing length. Vein Sc arising basally, fused with stem R+MP at basal 0.17 of wing
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length. Stem R+MP curved, connected to branch CuA by crossvein r+mp-cua at same
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point where stem Cu branches into veins CuA and CuP. Crossvein r+mp-cua long
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and oblique. Branches RA, RP and M departing from stem R+MP at same point.
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Branch RA slightly curved anteriorly. Branch RP connected to branch MP1 by
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crossvein r-mp1 distad of indentation. Stem M dividing into MP1+2 and MP3+4 before
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nodal line. Branch MP1+2 branching into veins MP1 and MP2 distad of indentation.
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Branch MP3+4 branching into MP3 and MP4 at basal 0.36 of wing length. Branch MP1
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partly directed anteriorly, geniculate at junction with crossvein r-mp1. Branch MP3
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almost straight then curved anteriorly near margin. Crossvein mp4-cua long, oblique
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and slightly curved. Stem Cu branching into veins CuA and CuP at basal 0.09 of wing
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length. Branch CuA slightly curved posteriorly between crossveins r+mp-cua and
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mp4-cua, branching into veins CuA1 and CuA2 after crossvein mp4-cua, basad of nodal
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branch CuA2 curved posteriorly then recurved. Discal cell length 19 mm, width 8.5
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mm at nodal line, about one-third of wing length; antenodal region trapezoidal,
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postnodal region less than half of antenodal region in length. Nodal line traceable as a
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crease across RA and RP to MP1+2, M3+4 and then separating discal cell into two parts;
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it continues crossing CuA just before its initial division, clavus absent. Membrane
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pitted distal of nodal line.
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4. Discussion
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Valdicossus Wang et al., 2008 was erected based on a well-preserved but
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incomplete hindwing from the Lower Cretaceous Wealden Supergroup of southern
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England (Wang et al., 2008a). Valdicossus mikewebsteri sp. nov. was placed in the
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genus Valdicossus based on the following features: hindwing oval, anterior margin
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curved in distal part; costal area wide; vein R fused with vein Sc basally, and
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branching into veins Sc+RA and RP; vein Sc terminating in costal margin just basad
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of wing indentation; branch RP fused with vein MP1 for a long distance basad of level
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of indentation; veins MP1, MP2 and MP3+4 arising from stem MP at wing base; vein
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MP3+4 unbranched. The new species and the type species are from different locations
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and horizons; it differs from the latter in the hindwing having a different colour
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pattern; vein Sc+R straight; branch RA2 slightly curved posteriorly then recurved;
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vein RP bifurcating from vein RP+MP1 basad of vein Sc separating from vein Sc+RA,
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short. They both have the following characters: anterior margin curved in the distal
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part; vein R fused with vein Sc basally and branching into veins Sc+RA and RP; veins
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MP1, MP2 and MP3+4 arising from stem MP at the wing base; vein MP3+4 is
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unbranched and vein RP fused with vein MP1 for a long distance basad of the
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indentation.
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Ilerdocossus Gómez-Pallerola, 1984 was erected based on a forewing from the
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Lower Cretaceous of Lérida, Spain. Menon et al. (2005) considered both
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Wonnacottella and Liaocossus to be the junior synonyms of Ilerdocossus. Wang B. et
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al. (2008b) also considered three genera (Ilerdocossus, Wonnacottella, and Liaocossus)
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as belonging to the same genusand our material also provides further support this
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view.
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We placed Ilerdocossus prowsei sp. nov. in the genus Ilerdocossus based on the following features: forewing triangular, with narrow costal area; reduced clavus;
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nodal indentation distinct; RA, RP and MP separate at the same point; discal cell long;
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antenodal region trapezoid. It mostly resembles Wonnacottella pulcherrimma
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(Fig.3D)(Whalley and Jarzembowski, 1985) from Spain: they both have a narrow
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forewing with a similar venation, and vein MP1+2 bifurcating distad of the indentation;
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vein MP3+4 bifurcates basad of the indentation, branch CuA2 is curved posteriorly
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then recurved; and the membrane is pitted distal of the nodal line. But the former
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differs from the latter in having a much larger and more narrow forewing, vein Sc
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into CuA and CuP at the same point where vein CuA meets crossvein r+mp-cua; stem
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M divides into MP1+2 and MP3+4 before the nodal line, vein CuA bifurcates beyond
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the line, cross vein r-mp1 is long. It is also distinctly different from other species of
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Ilerdocossus in having a forewing with membrane pitted distal of the nodal line, and
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vein CuA bifurcating beyond the nodal line.
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5. Conclusions
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Based on two well-preserved fossils from England, the new species Valdicossus mikewebsteri sp. nov. and Ilerdocossus prowsei sp. nov. are proposed. Our new
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material further confirms that Ilerdocossus was probably widely distributed in Eurasia
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and was diverse in the Early Cretaceous whereas Valdicossus appears to be a Wealden
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endemic.
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Acknowledgements
This research was supported by the National Natural Science Foundation of
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China (41572010, 41622201, 41688103, 41502007), the Strategic Priority Research
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Program (B) of the Chinese Academy of Sciences (XDB26000000), and the State Key
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Laboratory of Palaeobiology and Stratigraphy (Nanjing Institute of Geology and
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Palaeontology, CAS) (183101 and 183105). This is a Leverhulme Emeritus
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Fellowship contribution for EAJ. We thank Terry Keenan (London) for help with
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photography.
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Wang, B., Zhang, H.C., Fang, Y., Wang, D.J., and Ji, S.Z., 2008b. New data on
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Cretaceous Palaeontinidae (Insecta: Hemiptera) from China. Cretaceous
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Research, 29, 551–560.
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Wang, B., Zhang, H. C. and Szwedo, J., 2009. Jurassic Palaeontinidae from the China and
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Cicadomorpha). Palaeontology, 52, 53–64.
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Hemiptera:
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Whalley, P.E.S., and Jarzembowski, E.A., 1985. Fossil insects from the Lithographic
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Limestone of Montsech (late Jurassic-early Cretaceous), Lérida Province, Spain.
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Bulletin of the British Museum (Natural History), Geology, 38, 381–412.
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Figure legends：
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Fig.1. Map showing the location of insect fossil localities. S: Smokejacks brickworks;
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C: Clockhouse brickworks.
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Fig.2. A-C, Valdicossus mikewebsteri sp. nov. A, Holotype NHMUK II S4046a, part;
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B, counterpart of holotype NHMUK II S4046b. C, line drawing of holotype. D,
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Valdicossus chesteri Wang et al., 2008, hindwing redrawn from Wang et al. (2008a).
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Scale bars represent 5 mm.
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Fig.3. A-C, Ilerdocossus prowsei sp. nov. A, Holotype BMB 014927 (CHa5), part; B,
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counterpart of holotype BMB 018805. C, line drawing of holotype. D, Wonnacottella
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pulcherrimma Whalley and Jarzembowski, 1985, hindwing redrawn from Whalley
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and Jarzembowski (1985). Scale bars represent 5 mm.
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