copy Koninklijke Brill NV Leiden 2011 DOI 101163187631211X578415

Insect Systematics amp Evolution 42 (2011) 161ndash177 brillnlise

Th e fi rst Progonocimicidae (Insecta Hemiptera Coleorrhyncha) from Lower Cretaceous Lebanese amber

Jacek Szwedo a Dany Azar b Kamil Ziadeacute c a Museum and Institute of Zoology Polish Academy of Sciences

64 Wilcza Street PL00-679 Warsaw Poland b Lebanese University Faculty of Sciences II Department of Biology

PO Box 90656 Fanar-Matn Lebanon c Pfi zer-Lebanon Awkar-Mectapharm Building PO Box 90-674 Jdeideh Lebanon

Corresponding autor e-mail szwedomiizwawpl Published 1 July 2011

Abstract Ilahulgabalus endaidus gen spn (Progonocimicidae Cicadocorinae) the fi rst representative of Coleor-rhyncha from the Lower Cretaceous amber of Lebanon is described Th e placement of the new taxon within Coleorrhyncha and the evolutionary history of the suborder are discussed

Keywords Ilahulgabalus genn Ilahulgabalus endaidus spn Lower Cretaceous amber Lebanon morphology taxonomy

Introduction

Th e suborder Coleorrhyncha Myers et China 1929 also known as the lsquomoss bugsrsquo is of special interest within the Hemiptera because of its long evolutionary history interesting morphological features and the limited distribution of its recent representa-tives (Evans 1982 Popov amp Shcherbakov 1996 Burckhardt 2000 Grimaldi amp Engel 2005 Burckhardt 2009 Wang et al 2009) Th e relationship of the Coleorrhyncha to the remainder of the Hemiptera is of particular interest as this group was postulated to be the sister-taxon to the suborder Heteroptera (Schlee 1969 Sorensen et al 1995 Ouvrard et al 2000 Bourgoin amp Campbell 2002 Yang 2002 Schaefer 2003 Brożek 2007 Xie et al 2008 Burckhardt 2009 Wang et al 2009) Palaeontological interpre-tations of the Coleorrhyncha traceable via the Progonocimicidae to the latest Permian (255 million years ago) suggest that the group derived from Cicadomorpha Pros-boloidea Ingruidae Th erefore the Coleorrhyncha evolved in parallel to the true bugs (Heteroptera) acquiring some superfi cial similarities but retaining basic diff erences

162 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

(Popov amp Shcherbakov 1991 1996 Shcherbakov amp Popov 2002) Th e Heteroptera which appeared much later in the Middle Triassic share the costal fracture and fore wingndashthoracic coupling device with and doubtless derived from a superfamily of primitive Cicadomorpha mdash the Scytinopteroidea which like Coleorrhyncha is descended from Ingruidae (Shcherbakov 1996 2000)

Th e Coleorrhyncha comprises three families the extant Peloridiidae Breddin 1897 with 17 genera and 32 described species (Burckhardt 2009) and the extinct families Karabasiidae Popov 1985 placed within the Peloridioidea Breddin 1897 and Progonocimicidae Handlirsch 1906 placed within the Progonocimicoidea Handlirsch 1906 (Evans 1982 Popov amp Shcherbakov 1991 1996)

Th e oldest known fossil of the Progonocimicidae is Actinoscytina Tillyard 1926 from the Upper Permian of Belmont NSW Australia Th is family is not considered a monophyletic unit (Grimaldi amp Engel 2005) but rather a paraphyletic assemblage of the stem-group Coleorrhyncha leading to the lineages of Karabasiidae (Middle JurassicndashUpper Cretaceous) mdash comprising Karabasia and Hoploridium mdash and Peloridiidae (Recent) Th e Progonocimicidae were widely spread in Eurasia Australia South America and South Africa from the Upper Permian through to the Lower Cretaceous (Wootton 1963 Popov amp Wootton 1977 Jarzembowski 1991 Popov amp Shcherbakov 1991 Klimaszewski amp Popov 1993 Martins-Neto et al 2003 Grimaldi amp Engel 2005 Bechly amp Szwedo 2007 Heads 2008 Wang et al 2009 in press)

Th e family comprises over 20 genera split in two subfamilies Progonocimicinae and Cicadocorinae Progonocimicidae would have been small- to medium-sized insects with good jumping ability but already somewhat dorsoventrally fl attened (Shcherbakov amp Popov 2002) Th eir hind tibiae (at least in the Cicadocorinae) have one to four lat-eral and sometimes apical movable spurs Hind tarsi are three-segmented with the basitarsomere the largest segment and the basitarsomere and midtarsomere having apical pectens of teeth-bearing macrosetae

Permian and some Triassic genera of Progonocimicinae ( Actinoscytina Heteroscytina Evans 1956 Triscytina Evans 1956 and probably Platyscytinella Evans 1956) were still similar to the ancestral Ingruidae whereas in the other Triassic genera the head structure and tegminal venation were considerably modifi ed meriting suprageneric separation Late Progonocimicinae and some Cicadocorinae demonstrate noteworthy trends culminating in the Peloridioidea their antennae became incrassate with 4ndash6 segments (eg genus near Progonocimex Handlirsch 1906 in Progonocimicinae and Onokhoia Yu Popov 1988 in Cicadocorinae) the anteclypeus and postclypeus merge (eg Woottonia Popov et Shcherbakov 1991 of Progonocimicinae and Popovus Oumlzdikmen et Demir 2007 of Cicadocorinae) and the anteclypeus and lora became apically concealed by the preepisterna (eg Woottonia ) Th e basal cell of the tegmen is expanded in some representatives of Progonocimicinae and branch M 3+4 is basally fused with branch CuA 1 leaving only a short free apex in Pelorisca Yu Popov et Shcherbakov 1991 Some Progonocimicinae demonstrate the tendency towards a fl at-ter body and more broadened paranota and tegmen costal areas with additional small membranizations of metepisterna that could indicate reduction of jumping ability

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 163

On the contrary Cicadocorinae having powerful hind legs with movable tibial spurs and apical teeth and broad membranization in front of hind coxae were the best hop-pers among Coleorrhyncha It is assumed that the nymphs of Progonocimicidae were probably non-jumping and both nymphs and imagines were phloem-feeders (Popov amp Shcherbakov 1991 1996)

Materials and Methods

Th e specimen described here originates from the Daychouniyyeh (Dayshouniyeh) locality of the Early Cretaceous Lebanese amber (Figs 1ndash3) Th e amber outcrop is Neocomian aged (ValanginianndashBerremian) amber is associated with lignite and plant debris in a layer of dark clay Few inclusions are presently reported from this fossil site and to date no hemipterans have been recovered (Azar et al 2010)

Th e botanical origin of Lebanese amber is still under discussion Trees of the Araucariaceae ( Agathis levantensis Poinar et Milki 2001 and Araucaroxylon sp) or Cheirolepidiaceae families ( Protocarpoxylon sp) have both been proposed in the past although a relationship with resin-exuding Cycadaceae or a new unknown family were also suggested (Poinar amp Milki 2001 Koteja amp Azar 2008 Azar et al 2010)

Th e piece of amber containing the inclusion have been prepared as proposed by Azar (2000) and Azar et al (2003) mdash ie dissected ground polished and embedded in Canada balsam between two cover slips An Olympus SZH10 stereoscopic micro-scope equipped with a camera lucida and a Nikon Microphot-FX microscope were used for microscopic examination with changeable direct and transmitted normal and polarized light Feature measurements are considered approximate due to the eff ects of shriveling and possible optical deformations Photographs were taken using Olympus Camedia C-5060 and Nikon Eclipse E 600 digital cameras under control of Lucia software and adjusted using Combine ZP and Adobe Photoshop Elements 60 Venation terminology follows the general scheme of Kukalovaacute-Peck (1991) with inter-pretation proposed by Popov amp Shcherbakov (1991)

Systematic Palaeontology

Hemiptera Linnaeus 1758 Coleorrhyncha Myers et China 1929 Progonocimicoidea Handlirsch 1906 Progonocimicidae Handlirsch 1906 Cicadocorinae Becker-Migdisova 1958

Ilahulgabalus genn

Type species

Ilahulgabalus endaidus spn by present designation and monotypy

164 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

Diagnosis

Tegmen with venation pattern resembling Yuripopovia Jarzembowski 1991 but clearly diff ering in the following features three terminals of vein R (four terminals in Yuri-popovia ) veinlet r-m distinctly basad of M 1+2 forking ( r-m apicad of M 1+2 forking in Yuripopovia ) and radio-medial cell length 43times width (about 53times as long as wide in Yuripopovia ) Hind wing venation diff ers distinctly from Absoluta Becker-Migdisova 1962 in short dScRA terminal reaching margin distinctly basad of wing apex and absence of branch RA 2 (dScRA 1 reaching near apex and RA 2 parallel to margin but not reaching it in Absoluta ) and fused apical portion of stems CuP and Pcu mdash common stalk CuP+Pcu (CuP and Pcu separate in Absoluta or reaching margin at the same point) Hind tibia with single lateral spine (2ndash4 lateral spines among other Cicadocorinae genera)

Etymology

Th e generic name is derived from the name of god mdash Ilāh hul-Gabal (meaning lsquothe God of the Mountainrsquo) mdash of Phoenician and Syrian mythology Gender masculine

Figs 1ndash3 Daychouniyyeh outcrop Lebanon (1) Geological map scale bar=1 km (2) satellite image (3) detail of outcrop with its discoverer (KZ) Th is fi gure is published in colour in the online edition of this journal which can be accessed via httpwwwbrillnlise

162 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

(Popov amp Shcherbakov 1991 1996 Shcherbakov amp Popov 2002) Th e Heteroptera which appeared much later in the Middle Triassic share the costal fracture and fore wingndashthoracic coupling device with and doubtless derived from a superfamily of primitive Cicadomorpha mdash the Scytinopteroidea which like Coleorrhyncha is descended from Ingruidae (Shcherbakov 1996 2000)

Th e Coleorrhyncha comprises three families the extant Peloridiidae Breddin 1897 with 17 genera and 32 described species (Burckhardt 2009) and the extinct families Karabasiidae Popov 1985 placed within the Peloridioidea Breddin 1897 and Progonocimicidae Handlirsch 1906 placed within the Progonocimicoidea Handlirsch 1906 (Evans 1982 Popov amp Shcherbakov 1991 1996)

Th e oldest known fossil of the Progonocimicidae is Actinoscytina Tillyard 1926 from the Upper Permian of Belmont NSW Australia Th is family is not considered a monophyletic unit (Grimaldi amp Engel 2005) but rather a paraphyletic assemblage of the stem-group Coleorrhyncha leading to the lineages of Karabasiidae (Middle JurassicndashUpper Cretaceous) mdash comprising Karabasia and Hoploridium mdash and Peloridiidae (Recent) Th e Progonocimicidae were widely spread in Eurasia Australia South America and South Africa from the Upper Permian through to the Lower Cretaceous (Wootton 1963 Popov amp Wootton 1977 Jarzembowski 1991 Popov amp Shcherbakov 1991 Klimaszewski amp Popov 1993 Martins-Neto et al 2003 Grimaldi amp Engel 2005 Bechly amp Szwedo 2007 Heads 2008 Wang et al 2009 in press)

Th e family comprises over 20 genera split in two subfamilies Progonocimicinae and Cicadocorinae Progonocimicidae would have been small- to medium-sized insects with good jumping ability but already somewhat dorsoventrally fl attened (Shcherbakov amp Popov 2002) Th eir hind tibiae (at least in the Cicadocorinae) have one to four lat-eral and sometimes apical movable spurs Hind tarsi are three-segmented with the basitarsomere the largest segment and the basitarsomere and midtarsomere having apical pectens of teeth-bearing macrosetae

Permian and some Triassic genera of Progonocimicinae ( Actinoscytina Heteroscytina Evans 1956 Triscytina Evans 1956 and probably Platyscytinella Evans 1956) were still similar to the ancestral Ingruidae whereas in the other Triassic genera the head structure and tegminal venation were considerably modifi ed meriting suprageneric separation Late Progonocimicinae and some Cicadocorinae demonstrate noteworthy trends culminating in the Peloridioidea their antennae became incrassate with 4ndash6 segments (eg genus near Progonocimex Handlirsch 1906 in Progonocimicinae and Onokhoia Yu Popov 1988 in Cicadocorinae) the anteclypeus and postclypeus merge (eg Woottonia Popov et Shcherbakov 1991 of Progonocimicinae and Popovus Oumlzdikmen et Demir 2007 of Cicadocorinae) and the anteclypeus and lora became apically concealed by the preepisterna (eg Woottonia ) Th e basal cell of the tegmen is expanded in some representatives of Progonocimicinae and branch M 3+4 is basally fused with branch CuA 1 leaving only a short free apex in Pelorisca Yu Popov et Shcherbakov 1991 Some Progonocimicinae demonstrate the tendency towards a fl at-ter body and more broadened paranota and tegmen costal areas with additional small membranizations of metepisterna that could indicate reduction of jumping ability

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 163

On the contrary Cicadocorinae having powerful hind legs with movable tibial spurs and apical teeth and broad membranization in front of hind coxae were the best hop-pers among Coleorrhyncha It is assumed that the nymphs of Progonocimicidae were probably non-jumping and both nymphs and imagines were phloem-feeders (Popov amp Shcherbakov 1991 1996)

Materials and Methods

Th e specimen described here originates from the Daychouniyyeh (Dayshouniyeh) locality of the Early Cretaceous Lebanese amber (Figs 1ndash3) Th e amber outcrop is Neocomian aged (ValanginianndashBerremian) amber is associated with lignite and plant debris in a layer of dark clay Few inclusions are presently reported from this fossil site and to date no hemipterans have been recovered (Azar et al 2010)

Th e botanical origin of Lebanese amber is still under discussion Trees of the Araucariaceae ( Agathis levantensis Poinar et Milki 2001 and Araucaroxylon sp) or Cheirolepidiaceae families ( Protocarpoxylon sp) have both been proposed in the past although a relationship with resin-exuding Cycadaceae or a new unknown family were also suggested (Poinar amp Milki 2001 Koteja amp Azar 2008 Azar et al 2010)

Th e piece of amber containing the inclusion have been prepared as proposed by Azar (2000) and Azar et al (2003) mdash ie dissected ground polished and embedded in Canada balsam between two cover slips An Olympus SZH10 stereoscopic micro-scope equipped with a camera lucida and a Nikon Microphot-FX microscope were used for microscopic examination with changeable direct and transmitted normal and polarized light Feature measurements are considered approximate due to the eff ects of shriveling and possible optical deformations Photographs were taken using Olympus Camedia C-5060 and Nikon Eclipse E 600 digital cameras under control of Lucia software and adjusted using Combine ZP and Adobe Photoshop Elements 60 Venation terminology follows the general scheme of Kukalovaacute-Peck (1991) with inter-pretation proposed by Popov amp Shcherbakov (1991)

Systematic Palaeontology

Hemiptera Linnaeus 1758 Coleorrhyncha Myers et China 1929 Progonocimicoidea Handlirsch 1906 Progonocimicidae Handlirsch 1906 Cicadocorinae Becker-Migdisova 1958

Ilahulgabalus genn

Type species

Ilahulgabalus endaidus spn by present designation and monotypy

164 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

Diagnosis

Tegmen with venation pattern resembling Yuripopovia Jarzembowski 1991 but clearly diff ering in the following features three terminals of vein R (four terminals in Yuri-popovia ) veinlet r-m distinctly basad of M 1+2 forking ( r-m apicad of M 1+2 forking in Yuripopovia ) and radio-medial cell length 43times width (about 53times as long as wide in Yuripopovia ) Hind wing venation diff ers distinctly from Absoluta Becker-Migdisova 1962 in short dScRA terminal reaching margin distinctly basad of wing apex and absence of branch RA 2 (dScRA 1 reaching near apex and RA 2 parallel to margin but not reaching it in Absoluta ) and fused apical portion of stems CuP and Pcu mdash common stalk CuP+Pcu (CuP and Pcu separate in Absoluta or reaching margin at the same point) Hind tibia with single lateral spine (2ndash4 lateral spines among other Cicadocorinae genera)

Etymology

Th e generic name is derived from the name of god mdash Ilāh hul-Gabal (meaning lsquothe God of the Mountainrsquo) mdash of Phoenician and Syrian mythology Gender masculine

Figs 1ndash3 Daychouniyyeh outcrop Lebanon (1) Geological map scale bar=1 km (2) satellite image (3) detail of outcrop with its discoverer (KZ) Th is fi gure is published in colour in the online edition of this journal which can be accessed via httpwwwbrillnlise

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 163

On the contrary Cicadocorinae having powerful hind legs with movable tibial spurs and apical teeth and broad membranization in front of hind coxae were the best hop-pers among Coleorrhyncha It is assumed that the nymphs of Progonocimicidae were probably non-jumping and both nymphs and imagines were phloem-feeders (Popov amp Shcherbakov 1991 1996)

Materials and Methods

Th e specimen described here originates from the Daychouniyyeh (Dayshouniyeh) locality of the Early Cretaceous Lebanese amber (Figs 1ndash3) Th e amber outcrop is Neocomian aged (ValanginianndashBerremian) amber is associated with lignite and plant debris in a layer of dark clay Few inclusions are presently reported from this fossil site and to date no hemipterans have been recovered (Azar et al 2010)

Th e botanical origin of Lebanese amber is still under discussion Trees of the Araucariaceae ( Agathis levantensis Poinar et Milki 2001 and Araucaroxylon sp) or Cheirolepidiaceae families ( Protocarpoxylon sp) have both been proposed in the past although a relationship with resin-exuding Cycadaceae or a new unknown family were also suggested (Poinar amp Milki 2001 Koteja amp Azar 2008 Azar et al 2010)

Th e piece of amber containing the inclusion have been prepared as proposed by Azar (2000) and Azar et al (2003) mdash ie dissected ground polished and embedded in Canada balsam between two cover slips An Olympus SZH10 stereoscopic micro-scope equipped with a camera lucida and a Nikon Microphot-FX microscope were used for microscopic examination with changeable direct and transmitted normal and polarized light Feature measurements are considered approximate due to the eff ects of shriveling and possible optical deformations Photographs were taken using Olympus Camedia C-5060 and Nikon Eclipse E 600 digital cameras under control of Lucia software and adjusted using Combine ZP and Adobe Photoshop Elements 60 Venation terminology follows the general scheme of Kukalovaacute-Peck (1991) with inter-pretation proposed by Popov amp Shcherbakov (1991)

Systematic Palaeontology

Hemiptera Linnaeus 1758 Coleorrhyncha Myers et China 1929 Progonocimicoidea Handlirsch 1906 Progonocimicidae Handlirsch 1906 Cicadocorinae Becker-Migdisova 1958

Ilahulgabalus genn

Type species

Ilahulgabalus endaidus spn by present designation and monotypy

164 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

Diagnosis

Tegmen with venation pattern resembling Yuripopovia Jarzembowski 1991 but clearly diff ering in the following features three terminals of vein R (four terminals in Yuri-popovia ) veinlet r-m distinctly basad of M 1+2 forking ( r-m apicad of M 1+2 forking in Yuripopovia ) and radio-medial cell length 43times width (about 53times as long as wide in Yuripopovia ) Hind wing venation diff ers distinctly from Absoluta Becker-Migdisova 1962 in short dScRA terminal reaching margin distinctly basad of wing apex and absence of branch RA 2 (dScRA 1 reaching near apex and RA 2 parallel to margin but not reaching it in Absoluta ) and fused apical portion of stems CuP and Pcu mdash common stalk CuP+Pcu (CuP and Pcu separate in Absoluta or reaching margin at the same point) Hind tibia with single lateral spine (2ndash4 lateral spines among other Cicadocorinae genera)

Etymology

Th e generic name is derived from the name of god mdash Ilāh hul-Gabal (meaning lsquothe God of the Mountainrsquo) mdash of Phoenician and Syrian mythology Gender masculine

Figs 1ndash3 Daychouniyyeh outcrop Lebanon (1) Geological map scale bar=1 km (2) satellite image (3) detail of outcrop with its discoverer (KZ) Th is fi gure is published in colour in the online edition of this journal which can be accessed via httpwwwbrillnlise

164 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

Diagnosis

Tegmen with venation pattern resembling Yuripopovia Jarzembowski 1991 but clearly diff ering in the following features three terminals of vein R (four terminals in Yuri-popovia ) veinlet r-m distinctly basad of M 1+2 forking ( r-m apicad of M 1+2 forking in Yuripopovia ) and radio-medial cell length 43times width (about 53times as long as wide in Yuripopovia ) Hind wing venation diff ers distinctly from Absoluta Becker-Migdisova 1962 in short dScRA terminal reaching margin distinctly basad of wing apex and absence of branch RA 2 (dScRA 1 reaching near apex and RA 2 parallel to margin but not reaching it in Absoluta ) and fused apical portion of stems CuP and Pcu mdash common stalk CuP+Pcu (CuP and Pcu separate in Absoluta or reaching margin at the same point) Hind tibia with single lateral spine (2ndash4 lateral spines among other Cicadocorinae genera)

Etymology

Th e generic name is derived from the name of god mdash Ilāh hul-Gabal (meaning lsquothe God of the Mountainrsquo) mdash of Phoenician and Syrian mythology Gender masculine

Figs 1ndash3 Daychouniyyeh outcrop Lebanon (1) Geological map scale bar=1 km (2) satellite image (3) detail of outcrop with its discoverer (KZ) Th is fi gure is published in colour in the online edition of this journal which can be accessed via httpwwwbrillnlise

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 165

Description

Pronotum slightly wider than mesonotum estimated as about twice as wide as long in midline Mesonotum about as long as wide at base with distinct lsquocellularrsquo sculpture

Tegmen translucent with distinct venation about 225times as long as wide at widest point 238times as long as wide at claval apex Costal margin distinctly thickened slightly widened at base with distinct ventral epipleura (veins Pc+CP) Costal margin not strongly curved at base arcuate Delicate ridge present on dorsal margin reaching level of terminal RA 2 apex costal margin thickened to the level of terminal M 3+4 apex Vein bSc relatively short reaching half of basal cell length adhering to basal cell Basal cell elongate about 45times as long as wide arculus absent Stem Sc+R leaving basal cell basad of M- and CuA-stem origin points Branch dScRA 1 forked apicad of claval vein junc-tion basad of half tegmen length distinctly oblique reaching margin at level of claval apex apical portion of terminal dScRA 1 dilated Branch RA+RP about as long as ter-minal dScRA 1 terminal RA 2 subparallel to terminal dScRA 1 dilated near apex Forking of RP basad of veinlet r-m terminal RP slightly dilated at apex Stem M leaving basal cell from same point as stem CuA forked apicad of claval apex branch M 1+2 forked distinctly apicad of r-m veinlet terminal M 3+4 single parallel to terminals of CuA Stem CuA strongly curved at base where it leaves the basal cell then arcuate terminal CuA 1 straight parallel to terminal M 3+4 terminal CuA 2 more arcuate subparallel to postclaval margin Claval furrow (vein CuP) distinct transecting fused veins Pcu+A 1 and weakly visible in apical portion Apex of clavus exceeding half tegmen length claval veins Pcu and A 1 fused at frac34 of clavus length Veinlet r-m placed distinctly distad of veinlet m-cua Appendix distinctly narrower than adjoining apical cubital cell Radio-medial cell elongate about 43times as long as wide mediocubital cell distinctly shorter about 08times as long as radio-medial cell

Wing membranous about 22times as long as wide Costal margin distinctly angularly curved at base thickened Vein bSc present short adhering to basal cell Basal cell nar-row Stem dSc+R+M leaving basal cell with a short common stem Branch dScRA short reaching margin distinctly basad of wing apex at about half wing length branch RP single Stem M separated at about 02 of wing length forked distinctly basad of veinlet r-m at about ⅔ of wing length Vein CuA forked basad of stem M forking branch CuA 1 completely fused with branch M 3+4 branch CuA 2 reaching apical margin at same distance as branch M 3+4 CuA 1 and fused CuP+Pcu stalk Vein CuP placed close to the vein CuA parallel in apical ⅓ curved to join Pcu and fuse as short common stalk CuP+Pcu Vein Pcu slightly curved fused in apical portion with CuP Vein A 1 single slightly thickened at base Veinlet r-m placed distinctly apicad of stem M fork-ing Anal lobe distinct separated by a fold

Hind tibia with lateral spine at about half its length fi ve elongate apical teeth Basitarsomere longer than combined length of mid and apical tarsomeres six small subapical teeth plus median lobe Mid tarsomere shortest segment with distinct median lobe Apical tarsomere elongate tarsal claws distinct arolium wide bilobate with distinct median incision

Pregenital portion of abdomen wider than long tapering to the apex posterior mar-gins of sternites arcuate posterior margin of sternite VII distinctly cuneately incised

166 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

posterior margin of sternite VIII slightly convex Pygofer relatively short in ventral portion dorsal portion elongate with a pair of short apical processes Genital styles in apical portion slightly curved mediad Aedeagus with phallobase sclerifi ed distinctly elongate canaliculate distinctly exceeding tips of styles

Ilahulgabalus endaidus spn (Figs 4ndash25 28ndash32)

Holotype

Male specimen inclusion no DAYndash1C Azar collection preserved in Canada balsam Deposited provisionally in the Museacuteum national drsquoHistoire naturelle Paris France

Figs 4ndash7 Ilahulgabalus endaidus gen spn (4) General dorsal view in amber (5) general ventral view in amber (6) pronotum and mesonotum in dorsal aspect (7) sculpture of mesonotum Th is fi gure is pub-lished in colour in the online edition of this journal which can be accessed via httpwwwbrillnlise

166 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

posterior margin of sternite VIII slightly convex Pygofer relatively short in ventral portion dorsal portion elongate with a pair of short apical processes Genital styles in apical portion slightly curved mediad Aedeagus with phallobase sclerifi ed distinctly elongate canaliculate distinctly exceeding tips of styles

Ilahulgabalus endaidus spn (Figs 4ndash25 28ndash32)

Holotype

Male specimen inclusion no DAYndash1C Azar collection preserved in Canada balsam Deposited provisionally in the Museacuteum national drsquoHistoire naturelle Paris France

Figs 4ndash7 Ilahulgabalus endaidus gen spn (4) General dorsal view in amber (5) general ventral view in amber (6) pronotum and mesonotum in dorsal aspect (7) sculpture of mesonotum Th is fi gure is pub-lished in colour in the online edition of this journal which can be accessed via httpwwwbrillnlise

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 167

Inclusion in Lower Cretaceous Lebanese amber Specimen without head ventral por-tion of thorax and fore and mid legs pronotum partly destroyed base of left and tip of right tegmen destroyed base and tip of left hind wing not preserved

Age and outcrop

Early Cretaceous Neocomian (ValanginianndashHauterivian) Daychouniyyeh Caza El-Mten (El-Matn District) Mouhafazit Jabal Loubnan (Mont Lebanon Governorate) Lebanon Th is outcrop was discovered by one of the present authors (KZ) in April 2005

Etymology

Specifi c epithet is derived from Ancient Greek lsquoendaidosrsquo meaning resinous full of resin

Diagnosis

Tegmen membranous slightly smokey-coloured with dark markings at base arculus and claval apex Apical cells elongate but shorter than prenodal cells Radio-medial

Figs 8ndash11 Ilahulgabalus endaidus gen spn (8) Left tegmen and hind wing dorsal view (9) left tegmen and hind wing ventral view (10) right tegmen and hind wing dorsal view (11) right tegmen and hind wing ventral view Th is fi gure is published in colour in the online edition of this journal which can be accessed via httpwwwbrillnlise

168 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

cell with apex straight not sinuate Costal area about as wide as adjoining radial cell First interradial cell (between dScRA 1 and RA 2 ) diamond-shaped second interra-dial cell (cell C1) narrow about 39times as long as wide Postnodal portion of branch M 1+2 relatively long with two terminals M 1 and M 2 forked distinctly apicad of veinlet r-m Veinlet m-cua and nodal portion of branch CuA 2 at the same line Appendix narrower than frac14 of adjoining apical cubital cell Single lateral spine of hind tibia placed at half tibia length hind tibia with fi ve elongate apical teeth Tibio-tarsal formula 56

Figs 12ndash17 Ilahulgabalus endaidus gen spn (12) Left hind tibia and tarsus (13) apical portion of tibia and tarsal segments (14) apex of tibia (15) basitarsomere and mid tarsomere (16) mid tarsomere and apical tarsomere with tarsal claws and arolium dorsal view (17) mid tarsomere and apical tarsomere with tarsal claws and arolium ventral view Th is fi gure is published in colour in the online edition of this journal which can be accessed via httpwwwbrillnlise

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 169

Description

Large coleorrhynchan estimated length approximately 6 mm Pronotum wider than base of mesonotum Mesonotum about as wide as long in midline with distinct cel-lular sculpture Tegmina membranous slightly smokey-coloured with three small dark markings at base of tegmen arculus and claval apex Venation distinct costal margin curved thickened terminals dScRA 1 RA 2 and RP dilated at apex Hind wing mem-branous with clearly visible venation common stalk CuP+Pcu relatively long Hind tibia with single lateral spine at half of its length fi ve apical teeth Tibio-tarsal formula 56 Metatarsal claws large arolium large bilobate Posterior margin of abdominal sternite VII cuneately incised posterior margin of abdominal sternite VIII slightly convex medially Ventral posterior margin of pygofer straight apical portion of pygofer prolonged posteriad bearing two apical short processes Anal tube tubular with two apical processes Styles with apical portions curved mediad with bluntly angular api-ces Phallobase sclerotized canaliculate in apical portion somewhat tubuliform slightly tapering apicad 325times as long as wide at base

Preserved portion of body 46 mm long preserved fragment with tegmina about 547 mm long Pronotum about 1 mm long 18 mm wide Mesonotum 143 mm long mesonotum 15 mm wide at base Tegmen length 44 mm tegmen width at claval apex

Figs 18ndash21 Ilahulgabalus endaidus gen spn (18) Apical portion of abdomen dorsal view (19) apical portion of abdomen ventral view (20) male genital organs ventrolateral view (21) male genital organs dorsal view Th is fi gure is published in colour in the online edition of this journal which can be accessed via httpwwwbrillnlise

170 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

185 mm maximum tegmen width 195 mm Hind tibia 128 mm long tarsus length 08 mm with claws basitarsomere 039 mm long midtarsomere 016 mm long apical tarsomere 024 mm long claws and arolium about 01 mm long

Discussion

Th e new genus and species Ilahulgabalus endaidus described above is the fi rst known representative of Coleorrhyncha from a fossil resin It diff ers from other Cicadocorinae known from the Cretaceous Period ( Ildavia Yu Popov 1993 Valdiscytina Yu Popov 1993 and Yuripopovia ) from England and Popovus and Onokhoia from Mongolia and Transbaikalia respectively) in several features of tegmen venation In Ilahulgabalus bSc is very short not reaching the apex of basal cell and the thickening of costal margin reaches up to M 1 terminal whereas in the other genera this thickening does not usually exceed terminal RA 2 (eg Jurassic Eocercopis Handlirsch 1939) Ilahulgabalus also diff ers from other Cicadocorinae in its narrow appendix distinctly narrower than

Figs 22ndash27 Ilahulgabalus endaidus gen spn (22) Right tegmen (23) left tegmen (24) right hind wing (25) left hind wing (26) Ilahulgabalus reconstruction of tegmen venation and vein nomenclature (27) Absoluta minima Becker-Migdisova 1962 right hind wing (after Popov amp Shcherbakov 1991) with vein nomenclature Scale bar=1 mm

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 171

Figs 28ndash32 Ilahulgabalus endaidus gen spn (28) Left hind tibia and tarsus (29) hind tarsus (30) male genital organs dorsal view (31) male genital organs lateroventral view (32) male genital organs ventral view Scale bars (28) 05 mm (29) 01 mm (30ndash32) 025 mm

adjoining cubital cell and in that the terminals of branches dScRA 1 RA 2 and RP of tegmen are dilated at the apex mdash features not known among other Cicadocorinae It also diff ers from the Jurassic genera ( Archicercopis Handlirsch 1929 Eocercopis from Germany and Mesocimex Hong 1983 and Cicadocoris Becker-Migdisova 1958 from Asia) in details of the tegmen structure and its venation

172 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

Fig 33 Phylogram of Coleorrhyncha based on Popov amp Shcherbakov (1991 1996) Heads (2008) and Wang et al (2009 in press) Reliable synapomorphies marked with circles and numbers (1ndash10) Progonocimicidae 1 head lacking areolae 2 suprantennal ledges (nearly) meeting above the postclypeus 3 tegmina and hind wings apically overlapping in repose with enlarged posterior apical cells 4 sinistral overlap of the tegmina (left tegmen always over sometimes markedly diff ering in apical venation from the right tegmen) 5 tegmen with short transverse CuA 2 and appendix (parallel to situation in Cicadellidae) 6 interalar coupling of heteropterous type 7 fore and middle tarsi with two segments 8 pygofer barrel-shaped styles protruding elbowed 9 nymphs cryptic on thick stems 10 imagines jumping on

(Continued )

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 173

With respect to hind wing venation Ilahulgabalus diff ers from the only other known Cicadocorinae hind wing Absoluta in its short branch dScRA and fused terminal CuP+Pcu reaching the margin Th e hind wing of Ilahulgabalus diff ers from the mod-ern Peloridium Breddin 1897 by having forked stems ScR M and CuA which are single in the latter in addition the apex of stem ScR exceeds the hind wing apex in Peloridium reaching nearly to the apex of terminal M In the genus Peloridium hind wing venation is simplifi ed (China 1961 drsquoUrso 1993) lacking stem Sc+R with only portion of branch RA 2 retained (branch ScRA 1 is lost as interpreted by Popov amp Shcherbakov 1991) plus transverse veinlets or fused terminals of stems M and CuA however like Absoluta CuP is close to Pcu in Peloridium whereas in Ilahulgabalus the short common stalk of CuP and Pcu is present

Th e new genus also diff ers from other known Cicadocorinae in having a single lat-eral spine on the hind tibia (2ndash4 in known taxa) Th e male genital segment of Ilahulgabalus somewhat resembles in general structure male genital segments in recent Peloridiidae (Myers amp China 1929 Evans 1982 Yang amp Chang 2000 Burckardt 2009) A barrel-shaped pygofer is a general feature of the Coleorrhyncha but in Ilahulgabalus it is lacking the ventral processes and lateral projections (a feature of the Progonocimicoidea) present in the modern Peloridiidae (Peloridioidea)

Although Cicadocorinae is a blind off shoot within the coleorrhynchan evolution-ary lineages some general comparisons can be made to the modern moss-bugs

Fig 33 (Cont) larger plants (11ndash15) Progonocimicinae 11 suprantennal ledge continuous below median ocellus 12 tegmina and wings held fl at in repose 13 tegmen with precostal carina defl ected dorsad or obsolete 14 antefurcal dScRA 1 at least partly concave 15 stem R not continued by RP (some Triassic members of Progonocimicinae still similar to Ingruiidae whereas the others present considerably modifi ed head structure and tegminal venation pattern mdash these individuals possibly merit suprageneric separation) (16ndash18) Cicadocorinae 16 tegmen with clavus shortened 17 hind coxae enlarged 18 hind tibia with lateral spines (19ndash22) Pelorisca 19 basal cell large 20 dScRA 1 convex 21 stem CuA S-shaped beyond basal cell 22 M 3+4 confl uent with CuA 1 except apex ( Pelorisca characters fi t with the hypothetical transitional form between Progonocimicinae and Karabasiidae and possibly merits suprageneric separa-tion but this opinion needs confi rmation by the still unknown body structures) (23ndash27) Karabasiidae 23 head with clear-cut temples close to paranota 24 tegmen with precostal carina narrow to reduced 25 short free Sc distinct along stems R+M obscure distally not reaching costal margin 26 basal cell enlarged elongate 27 claval fracture always retained (28ndash33) Karabasiinae 28 head with a pair of are-olae 29 antennae three-segmented incrassate 30 basal cell with arculus longitudinal 31 terminals M 3+4 and CuA 1 fused 32 hind wing lacking closed cells 33 hind tarsomere with lateroapical teeth (34ndash42) Hoploridiinae 34 body very fl at and densely punctate 35 paranota wide 36 tegmina with precostal carina reduced membranized with venation reticulate 37 tegmina not covering sides of abdomen in repose 38 metepisterna entirely sclerotized 39 hind legs unarmed 40 two hind tarsomeres 41 adults not jumping 42 adults and possibly nymphs cryptic on bark (43ndash54) Peloridiidae 43 body with dor-sum fl atter than venter 44 paranota wide areolate 45 tegmen with wide areolate costal fi eld developed 46 tegmen with Sc distinct distally 47 in the macropterous form hind wing with reduced stem Sc+R and terminal ScRA 1 48 hind wing with only apical portion of RA 2 retained RA 2 apex exceeding hind wing apex 49 stems M and CuA single 50 CuP and Pcu close but not fused in apical portions 51 hind legs slender unarmed 52 hind tarsi two-segmented (parallel to Hoploridiinae) 53 imagines jumping 54 both nymphs and imagines cryptic on moss

174 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

In Pelori diidae the anal tube is two-segmented and the lsquoretractile fl aprsquo corresponds to true abdominal segment XI (Popov amp Shcherbakov 1996 Yang amp Chang 2000) Th e situation in Ilahulgabalus is not clear but the anal tube seems to be tubular and some-what fl attened provided with two apical processes additionally it seems that segments X and XI are fused or that segment XI is invaginated into segment X as in Peloridiidae A sclerotized phallobase distinctly separated and not invaginated into the pygofer clearly diff erentiates Ilahulgabalus from the modern Peloridiidae in which the phal-lobase (as interpreted by Yang amp Chang 2000) seems to be at least partly invaginated into the pygofer Genital styles in the modern Peloridiidae are moderately long some-times shortened and variously shaped the shortened styles of Ilahulgabalus resemble somewhat these but very little else is known about genital structures among the Cicadocorinae Prolongation of apical portion of pygofer relatively short styles and a sclerotized phallobase seem to be characters particular to this fossil representative of cicadocorine Progonocimicidae

As mentioned above the Cicadocorinae is a blind evolutionary lineage (Fig 33) having only remote relationships with the modern Peloridioidea Cicadocorinae likely split off from their Progonocimicinae ancestors in the Late Triassic as the oldest known fossils comes from the Lower Jurassic of Europe and Asia (Popov amp Shcherbakov 1991 Wang et al 2009) and the group was in decline by the Lower Cretaceous Th e general trend in morphological evolutionary changes in this compact subfamily includes the shortening of the clavus and development of jumping abilities as sug-gested by broad membranization in front of the hind coxae transverse mid and hind coxae the latter enlarged and powerful hind legs with lateral spines and apical teeth (Popov amp Shcherbakov 1991 1996) Ilahulgabalus comes from the late period of this subfamilyrsquos evolution being remnants of the group dominating in the Jurassic together with Onokhoia Yu Popov 1988 and Popovus Oumlzdikmen et Demir 2007 from Transbaikalai and Mongolia respectively Shcherbakov amp Popov (1996) suggested that the decline of the Cicadocorinae is no mere chance being synchronous with the rise and radiation of modern leafhoppers (Cicadomorpha Clypeata Membracoidea Cicadellidae) Th is statement is supported by the fact that the rapid evolution of leafhoppers (and other hemipteran families) and decline of the other bugs seems to relate to the Mid-Cretaceous biotic crisis and reorganization of the biosphere eg the extinction of numerous gymnosperm and pro-angiosperm lineages and the rapid radiation and dominance of modern angiosperm plants (Rasnitsyn 1988 Zherikhin 2002 Krassilov 2003 Taylor et al 2009) promoted diversifi cation of lsquomodernrsquo hemip-teran groups

Acknowledgements

We wish to acknowledge Dr Yuri A Popov (Paleontological Institute RAN Moscow) for his valuable discussions and support during studies of fossil Coleorrhyncha We wish also acknowledge two anonymous reviewers for their valuable comments and

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 175

suggestions Th is paper is a contribution to a project ldquoInsects in the Lower Cretaceous Lebanese amber mdash their taxonomy palaeobiodiversity and evolutionrdquo under agree-ment between the Lebanese University in Beirut and Museum and the Institute of Zoology Polish Academy of Sciences in Warsaw ndash ldquoOrigin and evolution of biodiver-sity of Europe and Middle Eastrdquo

References

Azar D ( 2000 ) Les ambres meacutesozoiumlques du Liban Th egravese de Doctorat de lrsquoUniversiteacute Paris XI France 164+144 p annexes

Azar D Gezeacute R amp Acra F ( 2010 ) Lebanese amber In Penney D (Ed) Biodiversity of fossils in amber from the major world deposits Siri Scientifi c Press Manchester pp 271 ndash 298

Azar D Perrichot V Neacuteraudeau D amp Nel A ( 2003 ) New psychodid fl ies from the Cretaceous ambers of Lebanon and France with a discussion about Eophlebotomus connectens Cockerell 1920 (Diptera Psychodidae) Annals of the Entomological Society of America 96 117minus127

Bechly G amp Szwedo J ( 2007 ) 1114 Coleorrhyncha moss bugs In Martill DM Bechly G amp Loveridge RF (Eds) Th e Crato fossil beds of Brazil Window into an ancient world Cambridge University Press Cambridge pp 313 ndash 317

Bourgoin Th amp Campbell BC ( 2002 ) Inferring a phylogeny for Hemiptera Falling into the lsquoAutapomorphic Traprsquo Denisia 4 zugleich Kataloge des OOuml Landesmuseums Neue Folge Nr 176 67 ndash 82

Brożek J ( 2007 ) Labial sensillae and the internal structure of the mouthparts of Xenophyes cascus (Bergroth 1924) (Peloridiidae Coleorrhyncha Hemiptera) and their signifi cance in evolutionary studies on the Hemiptera Monograph Aphids and other hemipterous Insects 13 35 ndash 42

Burckhardt D ( 2000 ) Peloridiids (Hemiptera Peloridiidae) and Gondwana biogeography In III Southern Connections Congress 17ndash22 January 2000 Lincoln University Canterbury New Zealand Pro-gramme and abstracts p 24

Burckhardt D ( 2009 ) Taxonomy and phylogeny of the Gondwanan moss bugs or Peloridiidae (Hemiptera Coleorrhyncha) Deutsche Entomologische Zeitschrift 56 173 ndash 235

China WE ( 1961 ) South American Peloridiidae (Hemiptera-Homoptera Coleorrhyncha) Transactions of the Royal Entomological Society of London 114 131 ndash 161

DrsquoUrso V ( 1993 ) Th e wing coupling apparatus in Peloridium hammoniorum Breddin 1897 (Insecta Rhynchota) Spixiana 16 133 ndash 139

Evans JW ( 1982 (1981) ) A review of present knowledge of the family Peloridiidae and new genera and new species from New Zealand and New Caledonia (Hemiptera Insecta) Records of the Australian Museum 34 381 ndash 406

Grimaldi D amp Engel MS ( 2005 ) Evolution of the insects Cambridge University Press Cambridge 755 p Heads SW ( 2008 ) A new species of Yuripopovia from the Early Cretaceous of the Isle of Wight

(Coleorrhyncha Progonocimicidae) British Journal of Entomology and Natural History 21 247minus253 Jarzembowski EA ( 1991 ) New insects from the Weald Clay of the Weald Proceedings of the Geologistsrsquo

Association 102 93 ndash 108 Klimaszewski SM amp Popov YuA ( 1993 ) New fossil hemipteran insects from southern England

(Hemiptera Psyliina + Coleorrhyncha) Annals of the Upper Silesian Museum Entomology 1 (Suppl) 13 ndash 36

Koteja J amp Azar D ( 2008 ) Scale insects from Lower Cretaceous amber of Lebanon (Hemiptera Sternorrhyncha Coccinea) Alavesia 2 133 ndash 167

Krassilov VA ( 2003 ) Terrestrial Paleoecology and Global Change Russian Academic Monographs Volume 1 Pensoft Sofi a xvi+464 pp

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 171

Figs 28ndash32 Ilahulgabalus endaidus gen spn (28) Left hind tibia and tarsus (29) hind tarsus (30) male genital organs dorsal view (31) male genital organs lateroventral view (32) male genital organs ventral view Scale bars (28) 05 mm (29) 01 mm (30ndash32) 025 mm

adjoining cubital cell and in that the terminals of branches dScRA 1 RA 2 and RP of tegmen are dilated at the apex mdash features not known among other Cicadocorinae It also diff ers from the Jurassic genera ( Archicercopis Handlirsch 1929 Eocercopis from Germany and Mesocimex Hong 1983 and Cicadocoris Becker-Migdisova 1958 from Asia) in details of the tegmen structure and its venation

172 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

Fig 33 Phylogram of Coleorrhyncha based on Popov amp Shcherbakov (1991 1996) Heads (2008) and Wang et al (2009 in press) Reliable synapomorphies marked with circles and numbers (1ndash10) Progonocimicidae 1 head lacking areolae 2 suprantennal ledges (nearly) meeting above the postclypeus 3 tegmina and hind wings apically overlapping in repose with enlarged posterior apical cells 4 sinistral overlap of the tegmina (left tegmen always over sometimes markedly diff ering in apical venation from the right tegmen) 5 tegmen with short transverse CuA 2 and appendix (parallel to situation in Cicadellidae) 6 interalar coupling of heteropterous type 7 fore and middle tarsi with two segments 8 pygofer barrel-shaped styles protruding elbowed 9 nymphs cryptic on thick stems 10 imagines jumping on

(Continued )

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 173

With respect to hind wing venation Ilahulgabalus diff ers from the only other known Cicadocorinae hind wing Absoluta in its short branch dScRA and fused terminal CuP+Pcu reaching the margin Th e hind wing of Ilahulgabalus diff ers from the mod-ern Peloridium Breddin 1897 by having forked stems ScR M and CuA which are single in the latter in addition the apex of stem ScR exceeds the hind wing apex in Peloridium reaching nearly to the apex of terminal M In the genus Peloridium hind wing venation is simplifi ed (China 1961 drsquoUrso 1993) lacking stem Sc+R with only portion of branch RA 2 retained (branch ScRA 1 is lost as interpreted by Popov amp Shcherbakov 1991) plus transverse veinlets or fused terminals of stems M and CuA however like Absoluta CuP is close to Pcu in Peloridium whereas in Ilahulgabalus the short common stalk of CuP and Pcu is present

Th e new genus also diff ers from other known Cicadocorinae in having a single lat-eral spine on the hind tibia (2ndash4 in known taxa) Th e male genital segment of Ilahulgabalus somewhat resembles in general structure male genital segments in recent Peloridiidae (Myers amp China 1929 Evans 1982 Yang amp Chang 2000 Burckardt 2009) A barrel-shaped pygofer is a general feature of the Coleorrhyncha but in Ilahulgabalus it is lacking the ventral processes and lateral projections (a feature of the Progonocimicoidea) present in the modern Peloridiidae (Peloridioidea)

Although Cicadocorinae is a blind off shoot within the coleorrhynchan evolution-ary lineages some general comparisons can be made to the modern moss-bugs

Fig 33 (Cont) larger plants (11ndash15) Progonocimicinae 11 suprantennal ledge continuous below median ocellus 12 tegmina and wings held fl at in repose 13 tegmen with precostal carina defl ected dorsad or obsolete 14 antefurcal dScRA 1 at least partly concave 15 stem R not continued by RP (some Triassic members of Progonocimicinae still similar to Ingruiidae whereas the others present considerably modifi ed head structure and tegminal venation pattern mdash these individuals possibly merit suprageneric separation) (16ndash18) Cicadocorinae 16 tegmen with clavus shortened 17 hind coxae enlarged 18 hind tibia with lateral spines (19ndash22) Pelorisca 19 basal cell large 20 dScRA 1 convex 21 stem CuA S-shaped beyond basal cell 22 M 3+4 confl uent with CuA 1 except apex ( Pelorisca characters fi t with the hypothetical transitional form between Progonocimicinae and Karabasiidae and possibly merits suprageneric separa-tion but this opinion needs confi rmation by the still unknown body structures) (23ndash27) Karabasiidae 23 head with clear-cut temples close to paranota 24 tegmen with precostal carina narrow to reduced 25 short free Sc distinct along stems R+M obscure distally not reaching costal margin 26 basal cell enlarged elongate 27 claval fracture always retained (28ndash33) Karabasiinae 28 head with a pair of are-olae 29 antennae three-segmented incrassate 30 basal cell with arculus longitudinal 31 terminals M 3+4 and CuA 1 fused 32 hind wing lacking closed cells 33 hind tarsomere with lateroapical teeth (34ndash42) Hoploridiinae 34 body very fl at and densely punctate 35 paranota wide 36 tegmina with precostal carina reduced membranized with venation reticulate 37 tegmina not covering sides of abdomen in repose 38 metepisterna entirely sclerotized 39 hind legs unarmed 40 two hind tarsomeres 41 adults not jumping 42 adults and possibly nymphs cryptic on bark (43ndash54) Peloridiidae 43 body with dor-sum fl atter than venter 44 paranota wide areolate 45 tegmen with wide areolate costal fi eld developed 46 tegmen with Sc distinct distally 47 in the macropterous form hind wing with reduced stem Sc+R and terminal ScRA 1 48 hind wing with only apical portion of RA 2 retained RA 2 apex exceeding hind wing apex 49 stems M and CuA single 50 CuP and Pcu close but not fused in apical portions 51 hind legs slender unarmed 52 hind tarsi two-segmented (parallel to Hoploridiinae) 53 imagines jumping 54 both nymphs and imagines cryptic on moss

174 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

In Pelori diidae the anal tube is two-segmented and the lsquoretractile fl aprsquo corresponds to true abdominal segment XI (Popov amp Shcherbakov 1996 Yang amp Chang 2000) Th e situation in Ilahulgabalus is not clear but the anal tube seems to be tubular and some-what fl attened provided with two apical processes additionally it seems that segments X and XI are fused or that segment XI is invaginated into segment X as in Peloridiidae A sclerotized phallobase distinctly separated and not invaginated into the pygofer clearly diff erentiates Ilahulgabalus from the modern Peloridiidae in which the phal-lobase (as interpreted by Yang amp Chang 2000) seems to be at least partly invaginated into the pygofer Genital styles in the modern Peloridiidae are moderately long some-times shortened and variously shaped the shortened styles of Ilahulgabalus resemble somewhat these but very little else is known about genital structures among the Cicadocorinae Prolongation of apical portion of pygofer relatively short styles and a sclerotized phallobase seem to be characters particular to this fossil representative of cicadocorine Progonocimicidae

As mentioned above the Cicadocorinae is a blind evolutionary lineage (Fig 33) having only remote relationships with the modern Peloridioidea Cicadocorinae likely split off from their Progonocimicinae ancestors in the Late Triassic as the oldest known fossils comes from the Lower Jurassic of Europe and Asia (Popov amp Shcherbakov 1991 Wang et al 2009) and the group was in decline by the Lower Cretaceous Th e general trend in morphological evolutionary changes in this compact subfamily includes the shortening of the clavus and development of jumping abilities as sug-gested by broad membranization in front of the hind coxae transverse mid and hind coxae the latter enlarged and powerful hind legs with lateral spines and apical teeth (Popov amp Shcherbakov 1991 1996) Ilahulgabalus comes from the late period of this subfamilyrsquos evolution being remnants of the group dominating in the Jurassic together with Onokhoia Yu Popov 1988 and Popovus Oumlzdikmen et Demir 2007 from Transbaikalai and Mongolia respectively Shcherbakov amp Popov (1996) suggested that the decline of the Cicadocorinae is no mere chance being synchronous with the rise and radiation of modern leafhoppers (Cicadomorpha Clypeata Membracoidea Cicadellidae) Th is statement is supported by the fact that the rapid evolution of leafhoppers (and other hemipteran families) and decline of the other bugs seems to relate to the Mid-Cretaceous biotic crisis and reorganization of the biosphere eg the extinction of numerous gymnosperm and pro-angiosperm lineages and the rapid radiation and dominance of modern angiosperm plants (Rasnitsyn 1988 Zherikhin 2002 Krassilov 2003 Taylor et al 2009) promoted diversifi cation of lsquomodernrsquo hemip-teran groups

Acknowledgements

We wish to acknowledge Dr Yuri A Popov (Paleontological Institute RAN Moscow) for his valuable discussions and support during studies of fossil Coleorrhyncha We wish also acknowledge two anonymous reviewers for their valuable comments and

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 175

suggestions Th is paper is a contribution to a project ldquoInsects in the Lower Cretaceous Lebanese amber mdash their taxonomy palaeobiodiversity and evolutionrdquo under agree-ment between the Lebanese University in Beirut and Museum and the Institute of Zoology Polish Academy of Sciences in Warsaw ndash ldquoOrigin and evolution of biodiver-sity of Europe and Middle Eastrdquo

References

Azar D ( 2000 ) Les ambres meacutesozoiumlques du Liban Th egravese de Doctorat de lrsquoUniversiteacute Paris XI France 164+144 p annexes

Azar D Gezeacute R amp Acra F ( 2010 ) Lebanese amber In Penney D (Ed) Biodiversity of fossils in amber from the major world deposits Siri Scientifi c Press Manchester pp 271 ndash 298

Azar D Perrichot V Neacuteraudeau D amp Nel A ( 2003 ) New psychodid fl ies from the Cretaceous ambers of Lebanon and France with a discussion about Eophlebotomus connectens Cockerell 1920 (Diptera Psychodidae) Annals of the Entomological Society of America 96 117minus127

Bechly G amp Szwedo J ( 2007 ) 1114 Coleorrhyncha moss bugs In Martill DM Bechly G amp Loveridge RF (Eds) Th e Crato fossil beds of Brazil Window into an ancient world Cambridge University Press Cambridge pp 313 ndash 317

Bourgoin Th amp Campbell BC ( 2002 ) Inferring a phylogeny for Hemiptera Falling into the lsquoAutapomorphic Traprsquo Denisia 4 zugleich Kataloge des OOuml Landesmuseums Neue Folge Nr 176 67 ndash 82

Brożek J ( 2007 ) Labial sensillae and the internal structure of the mouthparts of Xenophyes cascus (Bergroth 1924) (Peloridiidae Coleorrhyncha Hemiptera) and their signifi cance in evolutionary studies on the Hemiptera Monograph Aphids and other hemipterous Insects 13 35 ndash 42

Burckhardt D ( 2000 ) Peloridiids (Hemiptera Peloridiidae) and Gondwana biogeography In III Southern Connections Congress 17ndash22 January 2000 Lincoln University Canterbury New Zealand Pro-gramme and abstracts p 24

Burckhardt D ( 2009 ) Taxonomy and phylogeny of the Gondwanan moss bugs or Peloridiidae (Hemiptera Coleorrhyncha) Deutsche Entomologische Zeitschrift 56 173 ndash 235

China WE ( 1961 ) South American Peloridiidae (Hemiptera-Homoptera Coleorrhyncha) Transactions of the Royal Entomological Society of London 114 131 ndash 161

DrsquoUrso V ( 1993 ) Th e wing coupling apparatus in Peloridium hammoniorum Breddin 1897 (Insecta Rhynchota) Spixiana 16 133 ndash 139

Evans JW ( 1982 (1981) ) A review of present knowledge of the family Peloridiidae and new genera and new species from New Zealand and New Caledonia (Hemiptera Insecta) Records of the Australian Museum 34 381 ndash 406

Grimaldi D amp Engel MS ( 2005 ) Evolution of the insects Cambridge University Press Cambridge 755 p Heads SW ( 2008 ) A new species of Yuripopovia from the Early Cretaceous of the Isle of Wight

(Coleorrhyncha Progonocimicidae) British Journal of Entomology and Natural History 21 247minus253 Jarzembowski EA ( 1991 ) New insects from the Weald Clay of the Weald Proceedings of the Geologistsrsquo

Association 102 93 ndash 108 Klimaszewski SM amp Popov YuA ( 1993 ) New fossil hemipteran insects from southern England

(Hemiptera Psyliina + Coleorrhyncha) Annals of the Upper Silesian Museum Entomology 1 (Suppl) 13 ndash 36

Koteja J amp Azar D ( 2008 ) Scale insects from Lower Cretaceous amber of Lebanon (Hemiptera Sternorrhyncha Coccinea) Alavesia 2 133 ndash 167

Krassilov VA ( 2003 ) Terrestrial Paleoecology and Global Change Russian Academic Monographs Volume 1 Pensoft Sofi a xvi+464 pp

172 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

Fig 33 Phylogram of Coleorrhyncha based on Popov amp Shcherbakov (1991 1996) Heads (2008) and Wang et al (2009 in press) Reliable synapomorphies marked with circles and numbers (1ndash10) Progonocimicidae 1 head lacking areolae 2 suprantennal ledges (nearly) meeting above the postclypeus 3 tegmina and hind wings apically overlapping in repose with enlarged posterior apical cells 4 sinistral overlap of the tegmina (left tegmen always over sometimes markedly diff ering in apical venation from the right tegmen) 5 tegmen with short transverse CuA 2 and appendix (parallel to situation in Cicadellidae) 6 interalar coupling of heteropterous type 7 fore and middle tarsi with two segments 8 pygofer barrel-shaped styles protruding elbowed 9 nymphs cryptic on thick stems 10 imagines jumping on

(Continued )

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 173

With respect to hind wing venation Ilahulgabalus diff ers from the only other known Cicadocorinae hind wing Absoluta in its short branch dScRA and fused terminal CuP+Pcu reaching the margin Th e hind wing of Ilahulgabalus diff ers from the mod-ern Peloridium Breddin 1897 by having forked stems ScR M and CuA which are single in the latter in addition the apex of stem ScR exceeds the hind wing apex in Peloridium reaching nearly to the apex of terminal M In the genus Peloridium hind wing venation is simplifi ed (China 1961 drsquoUrso 1993) lacking stem Sc+R with only portion of branch RA 2 retained (branch ScRA 1 is lost as interpreted by Popov amp Shcherbakov 1991) plus transverse veinlets or fused terminals of stems M and CuA however like Absoluta CuP is close to Pcu in Peloridium whereas in Ilahulgabalus the short common stalk of CuP and Pcu is present

Th e new genus also diff ers from other known Cicadocorinae in having a single lat-eral spine on the hind tibia (2ndash4 in known taxa) Th e male genital segment of Ilahulgabalus somewhat resembles in general structure male genital segments in recent Peloridiidae (Myers amp China 1929 Evans 1982 Yang amp Chang 2000 Burckardt 2009) A barrel-shaped pygofer is a general feature of the Coleorrhyncha but in Ilahulgabalus it is lacking the ventral processes and lateral projections (a feature of the Progonocimicoidea) present in the modern Peloridiidae (Peloridioidea)

Although Cicadocorinae is a blind off shoot within the coleorrhynchan evolution-ary lineages some general comparisons can be made to the modern moss-bugs

Fig 33 (Cont) larger plants (11ndash15) Progonocimicinae 11 suprantennal ledge continuous below median ocellus 12 tegmina and wings held fl at in repose 13 tegmen with precostal carina defl ected dorsad or obsolete 14 antefurcal dScRA 1 at least partly concave 15 stem R not continued by RP (some Triassic members of Progonocimicinae still similar to Ingruiidae whereas the others present considerably modifi ed head structure and tegminal venation pattern mdash these individuals possibly merit suprageneric separation) (16ndash18) Cicadocorinae 16 tegmen with clavus shortened 17 hind coxae enlarged 18 hind tibia with lateral spines (19ndash22) Pelorisca 19 basal cell large 20 dScRA 1 convex 21 stem CuA S-shaped beyond basal cell 22 M 3+4 confl uent with CuA 1 except apex ( Pelorisca characters fi t with the hypothetical transitional form between Progonocimicinae and Karabasiidae and possibly merits suprageneric separa-tion but this opinion needs confi rmation by the still unknown body structures) (23ndash27) Karabasiidae 23 head with clear-cut temples close to paranota 24 tegmen with precostal carina narrow to reduced 25 short free Sc distinct along stems R+M obscure distally not reaching costal margin 26 basal cell enlarged elongate 27 claval fracture always retained (28ndash33) Karabasiinae 28 head with a pair of are-olae 29 antennae three-segmented incrassate 30 basal cell with arculus longitudinal 31 terminals M 3+4 and CuA 1 fused 32 hind wing lacking closed cells 33 hind tarsomere with lateroapical teeth (34ndash42) Hoploridiinae 34 body very fl at and densely punctate 35 paranota wide 36 tegmina with precostal carina reduced membranized with venation reticulate 37 tegmina not covering sides of abdomen in repose 38 metepisterna entirely sclerotized 39 hind legs unarmed 40 two hind tarsomeres 41 adults not jumping 42 adults and possibly nymphs cryptic on bark (43ndash54) Peloridiidae 43 body with dor-sum fl atter than venter 44 paranota wide areolate 45 tegmen with wide areolate costal fi eld developed 46 tegmen with Sc distinct distally 47 in the macropterous form hind wing with reduced stem Sc+R and terminal ScRA 1 48 hind wing with only apical portion of RA 2 retained RA 2 apex exceeding hind wing apex 49 stems M and CuA single 50 CuP and Pcu close but not fused in apical portions 51 hind legs slender unarmed 52 hind tarsi two-segmented (parallel to Hoploridiinae) 53 imagines jumping 54 both nymphs and imagines cryptic on moss

174 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

In Pelori diidae the anal tube is two-segmented and the lsquoretractile fl aprsquo corresponds to true abdominal segment XI (Popov amp Shcherbakov 1996 Yang amp Chang 2000) Th e situation in Ilahulgabalus is not clear but the anal tube seems to be tubular and some-what fl attened provided with two apical processes additionally it seems that segments X and XI are fused or that segment XI is invaginated into segment X as in Peloridiidae A sclerotized phallobase distinctly separated and not invaginated into the pygofer clearly diff erentiates Ilahulgabalus from the modern Peloridiidae in which the phal-lobase (as interpreted by Yang amp Chang 2000) seems to be at least partly invaginated into the pygofer Genital styles in the modern Peloridiidae are moderately long some-times shortened and variously shaped the shortened styles of Ilahulgabalus resemble somewhat these but very little else is known about genital structures among the Cicadocorinae Prolongation of apical portion of pygofer relatively short styles and a sclerotized phallobase seem to be characters particular to this fossil representative of cicadocorine Progonocimicidae

As mentioned above the Cicadocorinae is a blind evolutionary lineage (Fig 33) having only remote relationships with the modern Peloridioidea Cicadocorinae likely split off from their Progonocimicinae ancestors in the Late Triassic as the oldest known fossils comes from the Lower Jurassic of Europe and Asia (Popov amp Shcherbakov 1991 Wang et al 2009) and the group was in decline by the Lower Cretaceous Th e general trend in morphological evolutionary changes in this compact subfamily includes the shortening of the clavus and development of jumping abilities as sug-gested by broad membranization in front of the hind coxae transverse mid and hind coxae the latter enlarged and powerful hind legs with lateral spines and apical teeth (Popov amp Shcherbakov 1991 1996) Ilahulgabalus comes from the late period of this subfamilyrsquos evolution being remnants of the group dominating in the Jurassic together with Onokhoia Yu Popov 1988 and Popovus Oumlzdikmen et Demir 2007 from Transbaikalai and Mongolia respectively Shcherbakov amp Popov (1996) suggested that the decline of the Cicadocorinae is no mere chance being synchronous with the rise and radiation of modern leafhoppers (Cicadomorpha Clypeata Membracoidea Cicadellidae) Th is statement is supported by the fact that the rapid evolution of leafhoppers (and other hemipteran families) and decline of the other bugs seems to relate to the Mid-Cretaceous biotic crisis and reorganization of the biosphere eg the extinction of numerous gymnosperm and pro-angiosperm lineages and the rapid radiation and dominance of modern angiosperm plants (Rasnitsyn 1988 Zherikhin 2002 Krassilov 2003 Taylor et al 2009) promoted diversifi cation of lsquomodernrsquo hemip-teran groups

Acknowledgements

We wish to acknowledge Dr Yuri A Popov (Paleontological Institute RAN Moscow) for his valuable discussions and support during studies of fossil Coleorrhyncha We wish also acknowledge two anonymous reviewers for their valuable comments and

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 175

suggestions Th is paper is a contribution to a project ldquoInsects in the Lower Cretaceous Lebanese amber mdash their taxonomy palaeobiodiversity and evolutionrdquo under agree-ment between the Lebanese University in Beirut and Museum and the Institute of Zoology Polish Academy of Sciences in Warsaw ndash ldquoOrigin and evolution of biodiver-sity of Europe and Middle Eastrdquo

References

Azar D ( 2000 ) Les ambres meacutesozoiumlques du Liban Th egravese de Doctorat de lrsquoUniversiteacute Paris XI France 164+144 p annexes

Azar D Gezeacute R amp Acra F ( 2010 ) Lebanese amber In Penney D (Ed) Biodiversity of fossils in amber from the major world deposits Siri Scientifi c Press Manchester pp 271 ndash 298

Azar D Perrichot V Neacuteraudeau D amp Nel A ( 2003 ) New psychodid fl ies from the Cretaceous ambers of Lebanon and France with a discussion about Eophlebotomus connectens Cockerell 1920 (Diptera Psychodidae) Annals of the Entomological Society of America 96 117minus127

Bechly G amp Szwedo J ( 2007 ) 1114 Coleorrhyncha moss bugs In Martill DM Bechly G amp Loveridge RF (Eds) Th e Crato fossil beds of Brazil Window into an ancient world Cambridge University Press Cambridge pp 313 ndash 317

Bourgoin Th amp Campbell BC ( 2002 ) Inferring a phylogeny for Hemiptera Falling into the lsquoAutapomorphic Traprsquo Denisia 4 zugleich Kataloge des OOuml Landesmuseums Neue Folge Nr 176 67 ndash 82

Brożek J ( 2007 ) Labial sensillae and the internal structure of the mouthparts of Xenophyes cascus (Bergroth 1924) (Peloridiidae Coleorrhyncha Hemiptera) and their signifi cance in evolutionary studies on the Hemiptera Monograph Aphids and other hemipterous Insects 13 35 ndash 42

Burckhardt D ( 2000 ) Peloridiids (Hemiptera Peloridiidae) and Gondwana biogeography In III Southern Connections Congress 17ndash22 January 2000 Lincoln University Canterbury New Zealand Pro-gramme and abstracts p 24

Burckhardt D ( 2009 ) Taxonomy and phylogeny of the Gondwanan moss bugs or Peloridiidae (Hemiptera Coleorrhyncha) Deutsche Entomologische Zeitschrift 56 173 ndash 235

China WE ( 1961 ) South American Peloridiidae (Hemiptera-Homoptera Coleorrhyncha) Transactions of the Royal Entomological Society of London 114 131 ndash 161

DrsquoUrso V ( 1993 ) Th e wing coupling apparatus in Peloridium hammoniorum Breddin 1897 (Insecta Rhynchota) Spixiana 16 133 ndash 139

Evans JW ( 1982 (1981) ) A review of present knowledge of the family Peloridiidae and new genera and new species from New Zealand and New Caledonia (Hemiptera Insecta) Records of the Australian Museum 34 381 ndash 406

Grimaldi D amp Engel MS ( 2005 ) Evolution of the insects Cambridge University Press Cambridge 755 p Heads SW ( 2008 ) A new species of Yuripopovia from the Early Cretaceous of the Isle of Wight

(Coleorrhyncha Progonocimicidae) British Journal of Entomology and Natural History 21 247minus253 Jarzembowski EA ( 1991 ) New insects from the Weald Clay of the Weald Proceedings of the Geologistsrsquo

Association 102 93 ndash 108 Klimaszewski SM amp Popov YuA ( 1993 ) New fossil hemipteran insects from southern England

(Hemiptera Psyliina + Coleorrhyncha) Annals of the Upper Silesian Museum Entomology 1 (Suppl) 13 ndash 36

Koteja J amp Azar D ( 2008 ) Scale insects from Lower Cretaceous amber of Lebanon (Hemiptera Sternorrhyncha Coccinea) Alavesia 2 133 ndash 167

Krassilov VA ( 2003 ) Terrestrial Paleoecology and Global Change Russian Academic Monographs Volume 1 Pensoft Sofi a xvi+464 pp

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 173

With respect to hind wing venation Ilahulgabalus diff ers from the only other known Cicadocorinae hind wing Absoluta in its short branch dScRA and fused terminal CuP+Pcu reaching the margin Th e hind wing of Ilahulgabalus diff ers from the mod-ern Peloridium Breddin 1897 by having forked stems ScR M and CuA which are single in the latter in addition the apex of stem ScR exceeds the hind wing apex in Peloridium reaching nearly to the apex of terminal M In the genus Peloridium hind wing venation is simplifi ed (China 1961 drsquoUrso 1993) lacking stem Sc+R with only portion of branch RA 2 retained (branch ScRA 1 is lost as interpreted by Popov amp Shcherbakov 1991) plus transverse veinlets or fused terminals of stems M and CuA however like Absoluta CuP is close to Pcu in Peloridium whereas in Ilahulgabalus the short common stalk of CuP and Pcu is present

Th e new genus also diff ers from other known Cicadocorinae in having a single lat-eral spine on the hind tibia (2ndash4 in known taxa) Th e male genital segment of Ilahulgabalus somewhat resembles in general structure male genital segments in recent Peloridiidae (Myers amp China 1929 Evans 1982 Yang amp Chang 2000 Burckardt 2009) A barrel-shaped pygofer is a general feature of the Coleorrhyncha but in Ilahulgabalus it is lacking the ventral processes and lateral projections (a feature of the Progonocimicoidea) present in the modern Peloridiidae (Peloridioidea)

Although Cicadocorinae is a blind off shoot within the coleorrhynchan evolution-ary lineages some general comparisons can be made to the modern moss-bugs

Fig 33 (Cont) larger plants (11ndash15) Progonocimicinae 11 suprantennal ledge continuous below median ocellus 12 tegmina and wings held fl at in repose 13 tegmen with precostal carina defl ected dorsad or obsolete 14 antefurcal dScRA 1 at least partly concave 15 stem R not continued by RP (some Triassic members of Progonocimicinae still similar to Ingruiidae whereas the others present considerably modifi ed head structure and tegminal venation pattern mdash these individuals possibly merit suprageneric separation) (16ndash18) Cicadocorinae 16 tegmen with clavus shortened 17 hind coxae enlarged 18 hind tibia with lateral spines (19ndash22) Pelorisca 19 basal cell large 20 dScRA 1 convex 21 stem CuA S-shaped beyond basal cell 22 M 3+4 confl uent with CuA 1 except apex ( Pelorisca characters fi t with the hypothetical transitional form between Progonocimicinae and Karabasiidae and possibly merits suprageneric separa-tion but this opinion needs confi rmation by the still unknown body structures) (23ndash27) Karabasiidae 23 head with clear-cut temples close to paranota 24 tegmen with precostal carina narrow to reduced 25 short free Sc distinct along stems R+M obscure distally not reaching costal margin 26 basal cell enlarged elongate 27 claval fracture always retained (28ndash33) Karabasiinae 28 head with a pair of are-olae 29 antennae three-segmented incrassate 30 basal cell with arculus longitudinal 31 terminals M 3+4 and CuA 1 fused 32 hind wing lacking closed cells 33 hind tarsomere with lateroapical teeth (34ndash42) Hoploridiinae 34 body very fl at and densely punctate 35 paranota wide 36 tegmina with precostal carina reduced membranized with venation reticulate 37 tegmina not covering sides of abdomen in repose 38 metepisterna entirely sclerotized 39 hind legs unarmed 40 two hind tarsomeres 41 adults not jumping 42 adults and possibly nymphs cryptic on bark (43ndash54) Peloridiidae 43 body with dor-sum fl atter than venter 44 paranota wide areolate 45 tegmen with wide areolate costal fi eld developed 46 tegmen with Sc distinct distally 47 in the macropterous form hind wing with reduced stem Sc+R and terminal ScRA 1 48 hind wing with only apical portion of RA 2 retained RA 2 apex exceeding hind wing apex 49 stems M and CuA single 50 CuP and Pcu close but not fused in apical portions 51 hind legs slender unarmed 52 hind tarsi two-segmented (parallel to Hoploridiinae) 53 imagines jumping 54 both nymphs and imagines cryptic on moss

174 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

In Pelori diidae the anal tube is two-segmented and the lsquoretractile fl aprsquo corresponds to true abdominal segment XI (Popov amp Shcherbakov 1996 Yang amp Chang 2000) Th e situation in Ilahulgabalus is not clear but the anal tube seems to be tubular and some-what fl attened provided with two apical processes additionally it seems that segments X and XI are fused or that segment XI is invaginated into segment X as in Peloridiidae A sclerotized phallobase distinctly separated and not invaginated into the pygofer clearly diff erentiates Ilahulgabalus from the modern Peloridiidae in which the phal-lobase (as interpreted by Yang amp Chang 2000) seems to be at least partly invaginated into the pygofer Genital styles in the modern Peloridiidae are moderately long some-times shortened and variously shaped the shortened styles of Ilahulgabalus resemble somewhat these but very little else is known about genital structures among the Cicadocorinae Prolongation of apical portion of pygofer relatively short styles and a sclerotized phallobase seem to be characters particular to this fossil representative of cicadocorine Progonocimicidae

As mentioned above the Cicadocorinae is a blind evolutionary lineage (Fig 33) having only remote relationships with the modern Peloridioidea Cicadocorinae likely split off from their Progonocimicinae ancestors in the Late Triassic as the oldest known fossils comes from the Lower Jurassic of Europe and Asia (Popov amp Shcherbakov 1991 Wang et al 2009) and the group was in decline by the Lower Cretaceous Th e general trend in morphological evolutionary changes in this compact subfamily includes the shortening of the clavus and development of jumping abilities as sug-gested by broad membranization in front of the hind coxae transverse mid and hind coxae the latter enlarged and powerful hind legs with lateral spines and apical teeth (Popov amp Shcherbakov 1991 1996) Ilahulgabalus comes from the late period of this subfamilyrsquos evolution being remnants of the group dominating in the Jurassic together with Onokhoia Yu Popov 1988 and Popovus Oumlzdikmen et Demir 2007 from Transbaikalai and Mongolia respectively Shcherbakov amp Popov (1996) suggested that the decline of the Cicadocorinae is no mere chance being synchronous with the rise and radiation of modern leafhoppers (Cicadomorpha Clypeata Membracoidea Cicadellidae) Th is statement is supported by the fact that the rapid evolution of leafhoppers (and other hemipteran families) and decline of the other bugs seems to relate to the Mid-Cretaceous biotic crisis and reorganization of the biosphere eg the extinction of numerous gymnosperm and pro-angiosperm lineages and the rapid radiation and dominance of modern angiosperm plants (Rasnitsyn 1988 Zherikhin 2002 Krassilov 2003 Taylor et al 2009) promoted diversifi cation of lsquomodernrsquo hemip-teran groups

Acknowledgements

We wish to acknowledge Dr Yuri A Popov (Paleontological Institute RAN Moscow) for his valuable discussions and support during studies of fossil Coleorrhyncha We wish also acknowledge two anonymous reviewers for their valuable comments and

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 175

suggestions Th is paper is a contribution to a project ldquoInsects in the Lower Cretaceous Lebanese amber mdash their taxonomy palaeobiodiversity and evolutionrdquo under agree-ment between the Lebanese University in Beirut and Museum and the Institute of Zoology Polish Academy of Sciences in Warsaw ndash ldquoOrigin and evolution of biodiver-sity of Europe and Middle Eastrdquo

References

Azar D ( 2000 ) Les ambres meacutesozoiumlques du Liban Th egravese de Doctorat de lrsquoUniversiteacute Paris XI France 164+144 p annexes

Azar D Gezeacute R amp Acra F ( 2010 ) Lebanese amber In Penney D (Ed) Biodiversity of fossils in amber from the major world deposits Siri Scientifi c Press Manchester pp 271 ndash 298

Azar D Perrichot V Neacuteraudeau D amp Nel A ( 2003 ) New psychodid fl ies from the Cretaceous ambers of Lebanon and France with a discussion about Eophlebotomus connectens Cockerell 1920 (Diptera Psychodidae) Annals of the Entomological Society of America 96 117minus127

Bechly G amp Szwedo J ( 2007 ) 1114 Coleorrhyncha moss bugs In Martill DM Bechly G amp Loveridge RF (Eds) Th e Crato fossil beds of Brazil Window into an ancient world Cambridge University Press Cambridge pp 313 ndash 317

Bourgoin Th amp Campbell BC ( 2002 ) Inferring a phylogeny for Hemiptera Falling into the lsquoAutapomorphic Traprsquo Denisia 4 zugleich Kataloge des OOuml Landesmuseums Neue Folge Nr 176 67 ndash 82

Brożek J ( 2007 ) Labial sensillae and the internal structure of the mouthparts of Xenophyes cascus (Bergroth 1924) (Peloridiidae Coleorrhyncha Hemiptera) and their signifi cance in evolutionary studies on the Hemiptera Monograph Aphids and other hemipterous Insects 13 35 ndash 42

Burckhardt D ( 2000 ) Peloridiids (Hemiptera Peloridiidae) and Gondwana biogeography In III Southern Connections Congress 17ndash22 January 2000 Lincoln University Canterbury New Zealand Pro-gramme and abstracts p 24

Burckhardt D ( 2009 ) Taxonomy and phylogeny of the Gondwanan moss bugs or Peloridiidae (Hemiptera Coleorrhyncha) Deutsche Entomologische Zeitschrift 56 173 ndash 235

China WE ( 1961 ) South American Peloridiidae (Hemiptera-Homoptera Coleorrhyncha) Transactions of the Royal Entomological Society of London 114 131 ndash 161

DrsquoUrso V ( 1993 ) Th e wing coupling apparatus in Peloridium hammoniorum Breddin 1897 (Insecta Rhynchota) Spixiana 16 133 ndash 139

Evans JW ( 1982 (1981) ) A review of present knowledge of the family Peloridiidae and new genera and new species from New Zealand and New Caledonia (Hemiptera Insecta) Records of the Australian Museum 34 381 ndash 406

Grimaldi D amp Engel MS ( 2005 ) Evolution of the insects Cambridge University Press Cambridge 755 p Heads SW ( 2008 ) A new species of Yuripopovia from the Early Cretaceous of the Isle of Wight

(Coleorrhyncha Progonocimicidae) British Journal of Entomology and Natural History 21 247minus253 Jarzembowski EA ( 1991 ) New insects from the Weald Clay of the Weald Proceedings of the Geologistsrsquo

Association 102 93 ndash 108 Klimaszewski SM amp Popov YuA ( 1993 ) New fossil hemipteran insects from southern England

(Hemiptera Psyliina + Coleorrhyncha) Annals of the Upper Silesian Museum Entomology 1 (Suppl) 13 ndash 36

Koteja J amp Azar D ( 2008 ) Scale insects from Lower Cretaceous amber of Lebanon (Hemiptera Sternorrhyncha Coccinea) Alavesia 2 133 ndash 167

Krassilov VA ( 2003 ) Terrestrial Paleoecology and Global Change Russian Academic Monographs Volume 1 Pensoft Sofi a xvi+464 pp

174 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

In Pelori diidae the anal tube is two-segmented and the lsquoretractile fl aprsquo corresponds to true abdominal segment XI (Popov amp Shcherbakov 1996 Yang amp Chang 2000) Th e situation in Ilahulgabalus is not clear but the anal tube seems to be tubular and some-what fl attened provided with two apical processes additionally it seems that segments X and XI are fused or that segment XI is invaginated into segment X as in Peloridiidae A sclerotized phallobase distinctly separated and not invaginated into the pygofer clearly diff erentiates Ilahulgabalus from the modern Peloridiidae in which the phal-lobase (as interpreted by Yang amp Chang 2000) seems to be at least partly invaginated into the pygofer Genital styles in the modern Peloridiidae are moderately long some-times shortened and variously shaped the shortened styles of Ilahulgabalus resemble somewhat these but very little else is known about genital structures among the Cicadocorinae Prolongation of apical portion of pygofer relatively short styles and a sclerotized phallobase seem to be characters particular to this fossil representative of cicadocorine Progonocimicidae

As mentioned above the Cicadocorinae is a blind evolutionary lineage (Fig 33) having only remote relationships with the modern Peloridioidea Cicadocorinae likely split off from their Progonocimicinae ancestors in the Late Triassic as the oldest known fossils comes from the Lower Jurassic of Europe and Asia (Popov amp Shcherbakov 1991 Wang et al 2009) and the group was in decline by the Lower Cretaceous Th e general trend in morphological evolutionary changes in this compact subfamily includes the shortening of the clavus and development of jumping abilities as sug-gested by broad membranization in front of the hind coxae transverse mid and hind coxae the latter enlarged and powerful hind legs with lateral spines and apical teeth (Popov amp Shcherbakov 1991 1996) Ilahulgabalus comes from the late period of this subfamilyrsquos evolution being remnants of the group dominating in the Jurassic together with Onokhoia Yu Popov 1988 and Popovus Oumlzdikmen et Demir 2007 from Transbaikalai and Mongolia respectively Shcherbakov amp Popov (1996) suggested that the decline of the Cicadocorinae is no mere chance being synchronous with the rise and radiation of modern leafhoppers (Cicadomorpha Clypeata Membracoidea Cicadellidae) Th is statement is supported by the fact that the rapid evolution of leafhoppers (and other hemipteran families) and decline of the other bugs seems to relate to the Mid-Cretaceous biotic crisis and reorganization of the biosphere eg the extinction of numerous gymnosperm and pro-angiosperm lineages and the rapid radiation and dominance of modern angiosperm plants (Rasnitsyn 1988 Zherikhin 2002 Krassilov 2003 Taylor et al 2009) promoted diversifi cation of lsquomodernrsquo hemip-teran groups

Acknowledgements

We wish to acknowledge Dr Yuri A Popov (Paleontological Institute RAN Moscow) for his valuable discussions and support during studies of fossil Coleorrhyncha We wish also acknowledge two anonymous reviewers for their valuable comments and

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 175

suggestions Th is paper is a contribution to a project ldquoInsects in the Lower Cretaceous Lebanese amber mdash their taxonomy palaeobiodiversity and evolutionrdquo under agree-ment between the Lebanese University in Beirut and Museum and the Institute of Zoology Polish Academy of Sciences in Warsaw ndash ldquoOrigin and evolution of biodiver-sity of Europe and Middle Eastrdquo

References

Azar D ( 2000 ) Les ambres meacutesozoiumlques du Liban Th egravese de Doctorat de lrsquoUniversiteacute Paris XI France 164+144 p annexes

Azar D Gezeacute R amp Acra F ( 2010 ) Lebanese amber In Penney D (Ed) Biodiversity of fossils in amber from the major world deposits Siri Scientifi c Press Manchester pp 271 ndash 298

Azar D Perrichot V Neacuteraudeau D amp Nel A ( 2003 ) New psychodid fl ies from the Cretaceous ambers of Lebanon and France with a discussion about Eophlebotomus connectens Cockerell 1920 (Diptera Psychodidae) Annals of the Entomological Society of America 96 117minus127

Bechly G amp Szwedo J ( 2007 ) 1114 Coleorrhyncha moss bugs In Martill DM Bechly G amp Loveridge RF (Eds) Th e Crato fossil beds of Brazil Window into an ancient world Cambridge University Press Cambridge pp 313 ndash 317

Bourgoin Th amp Campbell BC ( 2002 ) Inferring a phylogeny for Hemiptera Falling into the lsquoAutapomorphic Traprsquo Denisia 4 zugleich Kataloge des OOuml Landesmuseums Neue Folge Nr 176 67 ndash 82

Brożek J ( 2007 ) Labial sensillae and the internal structure of the mouthparts of Xenophyes cascus (Bergroth 1924) (Peloridiidae Coleorrhyncha Hemiptera) and their signifi cance in evolutionary studies on the Hemiptera Monograph Aphids and other hemipterous Insects 13 35 ndash 42

Burckhardt D ( 2000 ) Peloridiids (Hemiptera Peloridiidae) and Gondwana biogeography In III Southern Connections Congress 17ndash22 January 2000 Lincoln University Canterbury New Zealand Pro-gramme and abstracts p 24

Burckhardt D ( 2009 ) Taxonomy and phylogeny of the Gondwanan moss bugs or Peloridiidae (Hemiptera Coleorrhyncha) Deutsche Entomologische Zeitschrift 56 173 ndash 235

China WE ( 1961 ) South American Peloridiidae (Hemiptera-Homoptera Coleorrhyncha) Transactions of the Royal Entomological Society of London 114 131 ndash 161

DrsquoUrso V ( 1993 ) Th e wing coupling apparatus in Peloridium hammoniorum Breddin 1897 (Insecta Rhynchota) Spixiana 16 133 ndash 139

Evans JW ( 1982 (1981) ) A review of present knowledge of the family Peloridiidae and new genera and new species from New Zealand and New Caledonia (Hemiptera Insecta) Records of the Australian Museum 34 381 ndash 406

Grimaldi D amp Engel MS ( 2005 ) Evolution of the insects Cambridge University Press Cambridge 755 p Heads SW ( 2008 ) A new species of Yuripopovia from the Early Cretaceous of the Isle of Wight

(Coleorrhyncha Progonocimicidae) British Journal of Entomology and Natural History 21 247minus253 Jarzembowski EA ( 1991 ) New insects from the Weald Clay of the Weald Proceedings of the Geologistsrsquo

Association 102 93 ndash 108 Klimaszewski SM amp Popov YuA ( 1993 ) New fossil hemipteran insects from southern England

(Hemiptera Psyliina + Coleorrhyncha) Annals of the Upper Silesian Museum Entomology 1 (Suppl) 13 ndash 36

Koteja J amp Azar D ( 2008 ) Scale insects from Lower Cretaceous amber of Lebanon (Hemiptera Sternorrhyncha Coccinea) Alavesia 2 133 ndash 167

Krassilov VA ( 2003 ) Terrestrial Paleoecology and Global Change Russian Academic Monographs Volume 1 Pensoft Sofi a xvi+464 pp

J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177 175

suggestions Th is paper is a contribution to a project ldquoInsects in the Lower Cretaceous Lebanese amber mdash their taxonomy palaeobiodiversity and evolutionrdquo under agree-ment between the Lebanese University in Beirut and Museum and the Institute of Zoology Polish Academy of Sciences in Warsaw ndash ldquoOrigin and evolution of biodiver-sity of Europe and Middle Eastrdquo

References

Azar D ( 2000 ) Les ambres meacutesozoiumlques du Liban Th egravese de Doctorat de lrsquoUniversiteacute Paris XI France 164+144 p annexes

Azar D Gezeacute R amp Acra F ( 2010 ) Lebanese amber In Penney D (Ed) Biodiversity of fossils in amber from the major world deposits Siri Scientifi c Press Manchester pp 271 ndash 298

Azar D Perrichot V Neacuteraudeau D amp Nel A ( 2003 ) New psychodid fl ies from the Cretaceous ambers of Lebanon and France with a discussion about Eophlebotomus connectens Cockerell 1920 (Diptera Psychodidae) Annals of the Entomological Society of America 96 117minus127

Bechly G amp Szwedo J ( 2007 ) 1114 Coleorrhyncha moss bugs In Martill DM Bechly G amp Loveridge RF (Eds) Th e Crato fossil beds of Brazil Window into an ancient world Cambridge University Press Cambridge pp 313 ndash 317

Bourgoin Th amp Campbell BC ( 2002 ) Inferring a phylogeny for Hemiptera Falling into the lsquoAutapomorphic Traprsquo Denisia 4 zugleich Kataloge des OOuml Landesmuseums Neue Folge Nr 176 67 ndash 82

Brożek J ( 2007 ) Labial sensillae and the internal structure of the mouthparts of Xenophyes cascus (Bergroth 1924) (Peloridiidae Coleorrhyncha Hemiptera) and their signifi cance in evolutionary studies on the Hemiptera Monograph Aphids and other hemipterous Insects 13 35 ndash 42

Burckhardt D ( 2000 ) Peloridiids (Hemiptera Peloridiidae) and Gondwana biogeography In III Southern Connections Congress 17ndash22 January 2000 Lincoln University Canterbury New Zealand Pro-gramme and abstracts p 24

Burckhardt D ( 2009 ) Taxonomy and phylogeny of the Gondwanan moss bugs or Peloridiidae (Hemiptera Coleorrhyncha) Deutsche Entomologische Zeitschrift 56 173 ndash 235

China WE ( 1961 ) South American Peloridiidae (Hemiptera-Homoptera Coleorrhyncha) Transactions of the Royal Entomological Society of London 114 131 ndash 161

DrsquoUrso V ( 1993 ) Th e wing coupling apparatus in Peloridium hammoniorum Breddin 1897 (Insecta Rhynchota) Spixiana 16 133 ndash 139

Evans JW ( 1982 (1981) ) A review of present knowledge of the family Peloridiidae and new genera and new species from New Zealand and New Caledonia (Hemiptera Insecta) Records of the Australian Museum 34 381 ndash 406

Grimaldi D amp Engel MS ( 2005 ) Evolution of the insects Cambridge University Press Cambridge 755 p Heads SW ( 2008 ) A new species of Yuripopovia from the Early Cretaceous of the Isle of Wight

(Coleorrhyncha Progonocimicidae) British Journal of Entomology and Natural History 21 247minus253 Jarzembowski EA ( 1991 ) New insects from the Weald Clay of the Weald Proceedings of the Geologistsrsquo

Association 102 93 ndash 108 Klimaszewski SM amp Popov YuA ( 1993 ) New fossil hemipteran insects from southern England

(Hemiptera Psyliina + Coleorrhyncha) Annals of the Upper Silesian Museum Entomology 1 (Suppl) 13 ndash 36

Koteja J amp Azar D ( 2008 ) Scale insects from Lower Cretaceous amber of Lebanon (Hemiptera Sternorrhyncha Coccinea) Alavesia 2 133 ndash 167

Krassilov VA ( 2003 ) Terrestrial Paleoecology and Global Change Russian Academic Monographs Volume 1 Pensoft Sofi a xvi+464 pp

176 J Szwedo et al Insect Systematics amp Evolution 42 (2011) 161ndash177

Kukalovaacute-Peck J ( 1991 ) Fossil history and evolution of hexapod structures In Naumann ID (Ed) Th e Insects of Australia A textbook for students and research workers 2nd edn Vol 1 Melbourne University Press Melbourne VIC pp 141 ndash 179

Martins-Neto RG Gallego OF amp Melchor RN ( 2003 ) Th e Triassic insect fauna from South America (Argentina Brazil and Chile) a checklist (except Blattoptera and Coleoptera) and description of new taxa Acta Zoologica Cracoviensia 46 ( suppl-Fossil Insects ) 229 ndash 256

Myers JG amp China WE ( 1929 ) Th e systematic position of the Peloridiidae as elucidated by a further study of the external anatomy of Hemiodoecus leai China Annals and Magazine of Natural History 3 282 ndash 294

Ouvrard D Campbell BC Bourgoin T amp Chan KL ( 2000 ) 18S rRNA secondary structure and phylogenetic position of Peloridiidae (Insecta Hemiptera) Molecular Phylogenetics and Evolution 16 403 ndash 417

Popov YuA amp Shcherbakov DE ( 1991 ) Mesozoic Peloridioidea and their ancestors (Insecta Hemiptera Coleorrhyncha) Geologica et Palaeontologica 25 215 ndash 235

Popov YuA amp Shcherbakov DE ( 1996 ) Origin and evolution of the Coleorrhyncha as shown by the fossil record In Schaefer CW (Ed) Studies on Hemipteran Phylogeny Entomological Society of America Lanham MD pp 9 ndash 30

Popov YuA amp Wootton RJ ( 1977 ) Th e Upper Liassic Heteroptera of Mecklenburg and Saxony Systematic Entomology 2 333 ndash 351

Rasnitsyn AP ( 1988 ) Problema globalrsquonogo krizisa nazemnykh biotsenozov v seredinie melovogo peri-oda [Problem of global crisis in the non-marine biocenoses of MidndashCretaceous] In Ponomarenko AG (Ed) Melovoiuml biotsenoticheskiiuml krizis i evolyutsiya nasekomykh [Cretaceous biocoenotic crisis and insect evolution] Nauka Moscow pp 191 ndash 207 (in Russian)

Schaefer CW ( 2003 ) Prosorrhyncha (Coleorrhyncha+ Heteroptera) In Resh VH amp Cardeacute RT (Eds) Encyclopedia of Insects Academic Press San Diego CA pp 947 ndash 965

Schlee D ( 1969 ) Morphologie und symbiose ihre bewertskraft fuumlr die verwandtschaftsbeziehungen der Coleorrhyncha Phylogenetische Studien an Hemiptera IV Heteropteroidea (Heteroptera+Coleorrhyncha) als monophyletische Gruppe Stuttgarter Beitraumlge zur Naturkunde 210 1 ndash 27 (in German)

Shcherbakov DE ( 1996 ) Origin and evolution of the Auchenorrhyncha as shown by the fossil record In Schaefer CW (Ed) Studies on Hemipteran Phylogeny Entomological Society of America Lanham MD pp 31 ndash 45

Shcherbakov DE ( 2000 ) Permian faunas of Homoptera (Hemiptera) in relation to phytogeography and the Permo-Triassic crisis Paleontological Journal 34 (Suppl 3) S251 ndash S267

Shcherbakov DE amp Popov YuA ( 2002 ) 22125 Superorder Cimicidea Laicharting 1781 Order Hemiptera Linneacute 1758 Th e bugs cicadas plantlice scale insects etc (=Cimicida Laicharting 1781 =Homoptera Leach 1815+Heteroptera Latreille 1810) In Rasnitsyn AP amp Quicke DLJ (Eds) History of Insects Kluwer Dordrecht pp 143 ndash 157

Sorensen JT Campbell BC Gill RJ amp Steff en-Campbell JD ( 1995 ) Non-monophyly of Auchenorrhyncha (ldquoHomopterardquo) based upon 18S rDNA phylogeny Ecoevolutionary and cladistic implications within pre-Heteropterodea Hemiptera (sl) and a proposal for new monophyletic sub-orders Pan-Pacifi c Entomologist 71 31 ndash 60

Taylor TN Taylor EL amp Krings M ( 2009 ) Paleobotany Th e biology and evolution of fossil plants 2 nd edn Academic Press Amsterdam xxi+1230 pp

Wang B Szwedo J amp Zhang H ( 2009 ) Jurassic Progonocimicidae (Hemiptera) from China and phy-logenetic evolution of Coleorrhyncha Science in China (Earth Sciences) 52 1953 ndash 1961

Wang B Szwedo J amp Zhang H ( in pr e ss) Th e fi rst Peloridioidea (Hemiptera Coleorrhyncha) from the Lower Jurassic of China Acta Palaeontologica Sinica

Wootton RJ ( 1963 ) Actinoscytinidae (Hemiptera Heteroptera) from the Upper Triassic of Queensland Annals and Magazine of Natural History (Series 13) 6 249 ndash 255

Xie Q Tian Y Zheng L amp Bu W ( 2008 ) 18S rRNA hyper-elongation and the phylogeny of Euhemiptera (Insecta Hemiptera) Molecular Phylogenetics and Evolution 47 463 ndash 471

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Yang C-T ( 2002 ) Preliminary thoughts on the phylogeny of ColeorrhynchandashHeteroptera (Hemiptera) Formosan Entomologist 22 297 ndash 305

Yang C-T amp Chang T-Y ( 2000 ) Th e external male genitalia of Hemiptera (Homoptera ndash Heteroptera) Shih Way Publishers Taipei Taiwan 746 pp

Zherikhin VV ( 2002 ) Ecological history of the terrestrial insects In Rasnitsyn AP amp Quicke DLJ (Eds) History of Insects Kluwer Dordrecht pp 331 ndash 388