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New mantises (Insecta: Mantodea) in Cretaceous ambers from Lebanon, Spain, and Myanmar PDF

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CretaceousResearch60(2016)91e108 ContentslistsavailableatScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes New mantises (Insecta: Mantodea) in Cretaceous ambers from Lebanon, Spain, and Myanmar Xavier Delclo(cid:1)s a,*, Enrique Pen~alver b, Antonio Arillo c, Michael S. Engeld, Andre(cid:3) Nel e, Dany Azar f,g, Andrew Ross h aDepartamentd'Estratigrafia,PaleontologiaiGeoci(cid:1)enciesMarines,andInstitutdeRecercadelaBiodiversitat(IRBio),FacultatdeGeologia,Universitatde Barcelona,MartíiFranqu(cid:1)ess/n,08028Barcelona,Spain bMuseoGeominero,InstitutoGeolo(cid:3)gicoyMinerodeEspan~a,RíosRosas23,28003Madrid,Spain cDepartamentodeZoologíayAntropologíaFísica,FacultaddeBiología,UniversidadComplutense,28040Madrid,Spain dDivisionofEntomology(Paleoentomology),NaturalHistoryMuseum,andDepartmentofEcology&EvolutionaryBiology,1501CrestlineDriveeSuite140, UniversityofKansas,Lawrence,Kansas66045,USA eInstitutdeSyst(cid:3)ematique,E(cid:3)volution,Biodiversit(cid:3)e,ISYEBeUMR7205eCNRS,MNHN,UPMC,EPHE,Mus(cid:3)eumnationald'Histoirenaturelle, SorbonneUniversit(cid:3)es,57rueCuvier,CP50,Entomologie,75005Paris,France fLebaneseUniversity,FacultyofSciencesII,DepartmentofNaturalSciences,Fanar,MatnP.O.Box26110217,Lebanon gNanjingInstituteofGeologyandPalaeontology,ChineseAcademyofSciences,Nanjing210008,PRChina hDepartmentofNaturalSciences,NationalMuseumsScotland,ChambersSt.,EdinburghEH11JF,UK a r t i c l e i n f o a b s t r a c t Articlehistory: DiversenewmaterialofmantisesfoundintheCretaceousamber-bearingdepositsfromLebanon(Bar- Received28July2015 remian),Spain(Albian),andMyanmar(AlbianeCenomanian)aredescribedandfigured.TheLebanese Receivedinrevisedform andSpanishformsarenymphs;whiletheonefromMyanmarisanadultspecimen.TheLebanesenymph 28October2015 corresponds to a new specimen of Burmantis lebanensis Grimaldi, 2003 while the adult Burmese Acceptedinrevisedform1November2015 (Myanmar)specimenbelongstothenewspeciesBurmantiszherikhini.TheSpanishspecimenrepresents Availableonlinexxx anewgenusandspeciesandisestablishedasAragonimantisaenigma,butisconsideredfamilyincertae sedis. The Spanish specimen is the first record of Mesozoic mantises from western-European amber Keywords: deposits.ArevisedphylogenetichypothesisforCretaceousmantisesisproposed. Amber Mantodea ©2015ElsevierLtd.Allrightsreserved. Newspecies Phylogeny Cretaceous 1. Introduction The monophyly of Mantodea is well supported by several characterssuchasthepresenceofraptorialforelegs,presenceofan Mantodea (mantises) are a lineage of polyneopteran insects, ultrasound “ear” on the metathorax (not present in Cretaceous comprising approximately 2400 described species distributed in mantises), and a femoral brush, among others traits (Roy, 1999; nearly 434 genera (Ehrmann, 2002; Svenson and Whiting, 2004; Svenson and Whiting, 2004; Grimaldi and Engel, 2005). Man- Wieland,2013),andareamongthemorefamiliarofinsectgroups todea is phylogenetically related to the clade of crown-group owing to their characteristic raptorial forelegs, large eyes, and Blattaria and Isoptera in the more inclusive Dictyoptera (Kevan, distinctive stance and habitus (Zherikhin, 2002; Grimaldi and 1977; Lo et al., 2000; Grimaldi and Engel, 2005), which evolved Engel,2005).Mantisesarepredatoryandoccupyawidedistribu- fromroach-likeinsectswithreducedovipositors.Theoldestknown tion across generally warmer biomes, mainly in intertropical re- definitive representatives of the Mantodea date from the Late gions,andhavingdiversifiedintoaconsiderablevarietyofhabitats JurassicandEarlyCretaceous(Grimaldi,1997;Zherikhin,2002;Lo fromAfricandesertstoAsianrainforests. etal.,2003),althoughsomeLateCarboniferousandPermiantaxa havebeenarguedtorepresentstem-groupMantodea(e.g.,Be(cid:3)thoux and Wieland, 2009; Be(cid:3)thoux et al., 2010). Among the roaches, * Correspondingauthor.Tel.:þ34934021381. Vr(cid:4)sanský (2005) proposed that Mantodea evolved from the E-mail addresses: [email protected] (X. Delclo(cid:1)s), [email protected] Jurassic,free-livingfamilyLiberiblattinidaeVr(cid:4)sanský,2002a,andas (E. Pen~alver), [email protected] (A. Arillo), [email protected] (M.S. Engel), a resultof a shift toa predaceous mode of life, a hypothesis that [email protected](A.Nel),[email protected](D.Azar),[email protected](A.Ross). http://dx.doi.org/10.1016/j.cretres.2015.11.001 0195-6671/©2015ElsevierLtd.Allrightsreserved. 92 X.Delclo(cid:1)setal./CretaceousResearch60(2016)91e108 impliesthatthefamilyisparaphyleticandofnoclassificatoryvalue Mantodea originating on Gondwanaland during the Early Creta- (see further comments regarding this hypothesis in Be(cid:3)thoux and ceous (Svenson and Whiting, 2009). The first major divergence Wieland, 2009). Kukalova(cid:3)-Peck and Beutel (2012) and Gorochov amongthelineageputativelyoccurringasaresultof theAtlantic (2013) denied the hypothesis proposed by Be(cid:3)thoux and Wieland breakup, separating Africa from South America. According to (2009) and Be(cid:3)thoux et al. (2010) regarding the relationship be- SvensonandWhiting(2007),thebreakupofGondwanalandpro- tween Mantodea and the Paleozoic Strephocladidae Martynov, duced numerous divergences within the order, although under- 1938 (a junior synonym of the family Anthracoptilidae standablythedegreetowhichthisaccordswiththefossilrecordis Handlirsch,1922; after Guan et al. 2015), a Paleozoic group that unknowngiventhescantdirectevidenceofmantisdiversityduring similarly possesses raptorial forelegs (see Be(cid:3)thoux and Wieland, the Mesozoic. A recent molecular phylogenetic analysis of the 2009).Kukalova(cid:3)-PeckandBeutel(2012)consideredthisfamilyas entire Dictyoptera proposed a putative age of Late Carbon- stem-Holometabola while Gorochov (2013) proposed a relation- iferouseEarlyPermianforthestem-mantodeans,arangeinterest- shipwiththeEoblattida(¼CnemidolestodeasensuBe(cid:3)thoux,2005). inglyinlinewiththoseagesarguedbysomeauthorsalthoughtheir ThehypothesisputforwardbyBe(cid:3)thouxandco-workerswasbased specific taxa were likely not stem-Mantodea (e.g., Be(cid:3)thoux and on wing-venational organization of some “protorthopteran” spe- Wieland,2009;Be(cid:3)thouxetal.,2010),andaJurassicdiversification cies, and regards that some of these Carboniferous and Permian forcrown-groupmantodeans(Legendreetal.,2015). species belonged to stem-group Mantodea, distant relatives of Over the last two decades, diverse phylogenetic hypotheses modernmantises,andatanageofabout175Myearlierthanpre- havebeenproposedforDictyoptera,basedonmorphologicaland viousevidencesuggestedfortheclade.Recently,Guanetal.(2015) molecular sources of data as well as a growing appreciation and proposed that the Anthracoptilidae belong to the clade Paoliida incorporationoffossilevidence.Notsurprisingly,withthegrowth (sensu Prokop et al., 2014), themselves a sister group to or stem of data and methods of analysis, diverse and not necessarily groupoftheDictyoptera,andthisseemstobethebestsupported mutually reconcilably results have been obtained, although some conclusion based on available evidence, and considered Strepho- significant advances have been made. Some of these studies cladidae a junior synonymy of Anthracoptilidae. Vr(cid:4)sanský (2012) recoveredMantodeawith Blattaria and these assister totheIso- erected the family Mutoviidae for species from the Permian of ptera (e.g., Thorne and Carpenter,1992; Kambhampati,1995), or Russiaand whichheregardedas Blattaria;however, membersof Mantodea with Isoptera collectively as the sister of Blattaria thisfamilyshowacleardivisionofveinsR1andRs,andwhichis (DeSalle,1994),MantodeaassistertoBlattidaeandIsoptera(e.g., moretypicalofMantodeathanofBlattaria.Accordingly,thisfamily Klass,1997,2000;Loetal.,2000,2003;Deitzetal.,2003;Pellens should be regarded as of uncertain placement within the Dic- etal.,2007;Misofetal.,2014;Legendreetal.,2015),and/orwith tyoptera until more complete material is discovered, particularly Isoptera nested among Blattaria, thereby resurrecting a 19th and thestructureoftheforelegs. early 20th concept for the affinity of termites (e.g., Grimaldi and Vr(cid:4)sanský and Bechly (2015) recently erected the family Engel, 2005; Inward et al., 2007; Engel et al., 2009; Ware et al., Manipulatoridae for one interesting new species, Manipulator 2010;Krishnaetal.,2013). modificaputis, from Burmese amber. They regarded it as ‘an early The first formal quantitative analysis of extant mantodean sidebranchofthestemgroupofMantodea’thoughrejectitfrom phylogeny was provided by Svenson and Whiting (2004), who Mantodeaduetotheabsenceofaforewingpseudoveinandmedian consideredBlattariaassistertoMantodeaandwithIsopterasub- ocellus, undifferentiated mouthparts, cockroach-like hind wing ordinateintheformer.Theirdatasupportedthenotionthatpre- venation, and only partly differentiated forelegs. They puzzlingly vious phylogenetic estimates and classifications included a large dubbed it a ‘cockroach’, which they go on to use for all crown number of paraphyletic families and subfamilies (i.e., Roy,1999; Dictyoptera.Firstly,werejecttheuseoftheterm‘cockroach’inthis Ehrmann, 2002). Presently, relationships among the various con- wayasitcausesconfusionanddiffersutterlyfromalmostuniversal stituentlineages, as well as definitive evidence formonophylyof usage,andpreferthemoretraditionalapplicationofthiscommon those groups, remain unclear. Beier (1968) split Mantodea into name for all crown-group Blattaria excluding Isoptera (termites), eight living families: Chaeteessidae, Mantoididae, Metallyticidae, andprefertouse‘roachoids’forallothercockroach-likeDictyop- Amorphoscelididae, Eremiaphilidae, Empusidae, Hymenopodidae, tera.TheyplaceditwithinBlattariabasedontheformofthehind- and Mantidae, considering the first three as the most basal and wingvenation,whichalthoughcockroach-likeisnotthatdissimilar withmostspeciesclusteredinMantidae.Thecurrentclassification fromthetwoearlymantishindwingsdescribedbyGratshevand recognizes more than 18 families (Ehrmann, 2002), but as Zherikhin(1993),andwhichalsohaveabranchingR.Itshouldbe mentioned above it seems clear that several are paraphyletic as expectedthatthehindwingsofearlymantisesandcontemporary currently circumscribed (Svenson and Whiting, 2004, 2009). For cockroaches would not be very different so this reason alone is recent mantises it is generally well supported that the family insufficient to place M. modificaputis in Blattaria. Secondly, it is Mantoididae is sister to all other crown-group Mantodea (Klass, describedasa‘pursuitpredator’giventheabsenceofspinesonits 1997; Svenson and Whiting, 2004). Nonetheless, Wieland (2010) walkinglegs.Certainlyitcouldhavebeenpredatoryowingtothe proposed that the basal dichotomies within Mantodea were mantis-likeshapeofthehead,withlarge,exophthalmiccompound [Chaeteessa] þ ([Metallyticus] þ [Mantoida þ remaining Man- eyesandapparentlyraptorialforelegs,thoughitdoesnotappearto todea]).Thefirstseriousanalysisthatintegratedfossilspecieswas have specialist spines for prey capture. Its ability as an ‘active that of Grimaldi (2003) who recovered the earliest Cretaceous runnerandpursuitpredator’ishighlydubiousasmantisesalsodo genusBaissomantis(Baissomantidae)assistertothetruemantises not have spines on their mid- and hind-legs yet hunt by stealth (order Mantodea), and that the majority of Cretaceous species ratherthanactivepursuit.WeprefertoregardM.modificaputisasa formed a basal grade to all extant clades. Grimaldi (2003) also mantis-like dictyopteran, and one that was more likely a stealth consideredAmbermantis(Ambermantidae)andalllivingmantises predator. which have a profemur with discoidal spines (Eumantodea) to Generally,aJurassic/Cretaceousagehasbeenarguedforcrown- comprise the clade Neomantodea, with Chaeteessa as the sister group Mantodea (e.g., Zherikhin, 2002; Vr(cid:4)sanský, 2002a, 2002b; group to all other Eumantodea. The analysis of Grimaldi (2003) Grimaldi,2003;Loetal.,2003;GrimaldiandEngel,2005,among further indicated that mantises were basal to living roaches and others). Model-based estimates based strictly on molecular data termitesowingtothepresenceofvariousplesiomorphiesrelative have even hypothesized an Early Jurassic age, and with modern tothelatter. X.Delclo(cid:1)setal./CretaceousResearch60(2016)91e108 93 Mantis fossils are comparatively rare (Ehrmann, 1999, 2002; Table1 KnownFossilMantodea. GrimaldiandEngel,2005;Wieland,2013),andthishashampered considerably our understanding of the historical evolution of the Familyincertaesedis group (Table 1). Given that fossils have the potential to radically Ambermantis recastournotionsofrelationships,biogeographicpatterns,andthe (cid:2) AmbermantiswozniakiGrimaldi,2003;TuronianfromNewJersey.Grimaldi originsofevolutionarynoveltiesandbiologicalphenomena;thisis (2003)erectedthefamilyAmbermantidaetoincludethisspecies,butsee a lamentable state of affairs. Although several mantodean speci- Wieland(2013). mens have been found from amber-bearing deposits throughout theworld(Ehrmann,2002),uptothepresentonly29fossilspecies Amorphoscelites (cid:2) Amorphoscelites sharovi Gratshev and Zherikhin, 1993; Valanginian havebeendescribed(Table1).Remarkably,fromamongthistotal eHauterivianfromSiberia.Fragmentaryforeleg;originallyplacedinAmor- 21 have been found in Cretaceous deposits, both in limestones phoscelidae:uncertainfamilialplacementsensuGrimaldi(2003). (mainly wings) and amber (some complete adults but mainly unwingednymphs)(GratshevandZherikhin,1993;Grimaldi,2003; Aragonimantis Ho€rnigetal.,2013).FurtherspecimensfromtheLowerCretaceous (cid:2) Aragonimantisaenigmagen.andsp.nov.;upperAlbianfromSanJust(Spain). amber of Japan (http://news.nationalgeographic.com/news/2008/ Burmantis(¼GryllomantisGorochov,2006) 04/080425-amber-mantis.html), Upper Cretaceous of Canada (cid:2) Burmantis asiatica Grimaldi, 2003; upper Albianelower Cenomanian from (Pike, 1995), and the Lower Cretaceous limestones of Spain and Myanmar. Mongolia (Vr(cid:4)sanský, 2002a, 2005), etc., remain unstudied and (cid:2) B. lebanensis Grimaldi, 2003; Barremian from Bcharreh Mountain and Al- without formal description. Grimaldi (2003) considered that the Rihan/Jezzine(Lebanon).¼Gryllomantislebanensis(Grimaldi,2003)sensu Gorochov(2006)inGryllomantidaesensuGorochov(2006). Mesozoic genera Amorphoscelites Gratshev and Zherikhin, 1993, (cid:2) B.zherikhinisp.nov.;upperAlbianelowerCenomanianfromMyanmar. Burmantis Grimaldi, 2003, Chaeteessites Gratshev and Zherikhin, 1993, Cretophotina Gratshev and Zherikhin, 1993, Electromantis Chaeteessites Gratshev and Zherikhin, 1993, Jersimantis Grimaldi, 1997, Kaza- (cid:2) Chaeteessites minutissimus Gratshev and Zherikhin, 1993; Santonian from khophotina Gratshev and Zherikhin, 1993, and Vitimiphotina TaymyrPeninsula(Siberia)-UncertainfamilialplacementsensuGrimaldi (2003). GratshevandZherikhin,1993wereofuncertainfamilialposition. Grimaldi (2003) noted that the Cretaceous mantises largely Cretophotina[originallyincludedinChaeteessidaebutuncertainfamilial possess plesiomorphic characters, particularly in regard to their placementsensuGrimaldi(2003)] pattern of wing venation, the profemoral brush, the profemoral (cid:2) Cretophotina mongolica Gratshev and Zherikhin, 1993; BarremianeAptian spines, and/or the protibial spur. It appears as though mantises fromMongolia. (cid:2) C.santanensisLee,2014;AptianfromSantanadoCarirí,Brazil. were in a ‘nascent’ phase of their evolution during the Early (cid:2) C.serotinaGratshevandZherikhin,1993;TuronianfromKazakhastan. Cretaceous,andthattrueMantodea,completewithraptorialfore- (cid:2) C. tristriata Gratshev and Zherikhin, 1993; ValanginianeHauterivian from legs,probablyappearedintheLateJurassic. Siberia. Enigmatically,Gorochov(2006)excludedthegeneraBurmantis (cid:2) C.selenginesisVr(cid:4)sanský,2002;LowerCretaceousfromSharin-Gol(Mongolia). (cid:2) Cretophotinasp.;BarremianfromLasHoyas(Spain).InVr(cid:4)sanský(2002). andJersimantisnotonlyfromMantodeabutevenfromDictyoptera, (cid:2) Cretophotina sp.; BerriasianeValanginian from Sharin-Gol (Mongolia). In andmainlyowingtotheshortlengthoftheirprocoxae(butreferto Vr(cid:4)sanský(2005). theemendeddiagnosisofBurmantisbelow).Suchaconclusionis notsupportedbyabroaderswathofcharacterevidenceandthereis Electromantis noreasontoremovethesetaxafromtheMantodea,andfarlessto (cid:2) Electromantis sukatshevae Gratshev and Zherikhin, 1993; Santonian from TaymyrPeninsula(Siberia)-OriginallyplacedinCretomantidae:uncertain excludethemfromDictyoptera. familialplacementsensuGrimaldi(2003). HerewedescribevariousnewmantisesfromtheLowerCreta- ceousofSpain,Lebanon,andMyanmar.Thediscoveryofthisnew Jersimantis materialpermitsustoreviewalsorelationshipsamongtheseand (cid:2) JersimantisluzziiGrimaldi,1997;TuronianfromNewJersey. otherMesozoictaxaandinrelationtothelivingmantises.Overall (cid:2) J.burmiticusGrimaldi,2003;upperAlbianelowerCenomanianfromMyan- mar¼Burmantisburmitica(Grimaldi,2003)sensuGorochov(2006). the new material further highlights the diversity of Mantodea during the Cretaceous and the importance of fossils, despite the Kazakhophotina paucityofmaterial,foradvancingknowledgeofmantisevolution. (cid:2) Kazakhophotina corrupta Gratshev and Zherikhin, 1993; Turonian from Kazakhstan.Fragmentofthewing.OriginallyincludedinChaeteessidae: 2. Geologicalandpalaeontologicalsettings uncertainfamilialplacementsensuGrimaldi(2003). Vitimophotina 2.1. Spanishamber (cid:2) Vitimophotina corrugata Gratshev and Zherikhin, 1993; Valanginian eHauterivianfromSiberia.Fragmentarywing.OriginallyincludedinChae- The newSpanish species was found in the amber-bearing de- teessidae:uncertainfamilialplacementsensuGrimaldi(2003). positofSanJust(Utrillas,Teruel).Spainisrichinamberoutcropsof Genusunknown EarlyCretaceousage(mainlyAlbian),butonlynineofthemhave (cid:2) Unclassified Mantis. Santonian from Kuji, Iwate Prefecture, NE Japan. See provided fossil arthropods as bioinclusions (Delclo(cid:1)s et al., 2007; http://news.nationalgeographic.com/news/2008/04/080425-amber-mantis. Pen~alver and Delclo(cid:1)s, 2010). The richest fossil associations have html(2015feb.). been found in Pen~acerrada (Alonso et al., 2000) and El Soplao (Najarro et al., 2009, 2010), in the BasqueeCantabrian Basin, and Baissomantidaey SanJust(Pen~alveretal.,2007),intheMaestrazgoBasin(Fig.1).The (cid:2) eBaHisasuotmerainvtiaisnfrmoamcuSliabteariaG. ratshev and Zherikhin, 1993; Valanginian amber piece comes from a grey-black claystone level with abun- (cid:2) B.pictaGratshevandZherikhin,1993;ValanginianeHauterivianfromSiberia. dantplantmacroremains,suchasfernsofthegenusCladophlebis, several conifers such as Arctiopitys, Brachyphyllum, Glenrosa, and Cretomantidaey Frenelopsis, and ginkgoales such as Eretmophyllum (¼Nehvizdya) (cid:2) CretomantislarvalisGratshevandZherikhin,1993;ValanginianeHauterivian fromSiberia. andPseudotorellia(B.Gomez,pers.com.2013).Thedepositissit- uated in the Utrillas Group (sensu Rodríguez-Lo(cid:3)pez et al., 2009) (continuedonnextpage) 94 X.Delclo(cid:1)setal./CretaceousResearch60(2016)91e108 Table1(continued) Fuenteetal.,2012;Saupeetal.,2012;Engeletal.,2013;Perisetal., Santanmantidaey 2014). (cid:2) SantanmantisaxelrodiGrimaldi,2003;AptianfromSantanadoCariri,inBrazil. SeeHo€rnigetal.(2013). 2.2. Lebaneseamber Chaeteessidae(¼Archephemeridae) The Lebanese amber-bearing deposits with bioinclusions are (cid:2) ArvernineurainsignisPiton,1940;PaleocenefromMenat(France).SeeNeland fromtheLowerCretaceous,underthe“FalaisedeBlanche”datedas Roy(1996). (cid:2) LithophotinacostalisCockerell,1914;EocenefromColorado.InMeyer(2003). Ante-Jezzinian(lateBarremanianeearlyAptian,inMaksoudetal., (cid:2) LithophotinafloccosaCockerell,1908;EocenefromColorado.InMeyer(2003). 2014),andthustheamberisofBarremianage.Anumberofout- (cid:2) Megaphotina sichotensis Gratshev and Zherikhin, 1993; Oligocene from crops (more than 400) have yielded amber but only 22 of these Sikhote-Alin. haveprovidedbioinclusions(Azaretal.,2010;Azar,2012).Thenew (cid:2) Chaeteessasp.;AptianfromSantanadoCariri,inBrazil.InLee(2011),butsee specimencomesfromtheoutcropofAl-Rihan,CazaJezzine(Jezzine alsoLee(2014). (cid:2) Chaeteessasp.;MiocenefromtheDominicanRepublic.InGrimaldi(2003). Department), Mohafazat Loubnan El-Janoubi (South Lebanon (cid:2) Chaeteessidaesp.;EocenefromBalticamber.InErhmann(1999). Governorate), in southern Lebanon, and where approximately 40 (cid:2) Chaeteessidaesp.;OligocenefromGermany.InErhmann(1999). bioinclusionshavebeenfoundtodate.Theamberpiececomesfrom (cid:2) Chaeteessidaesp.;CampanianfromCanada.InErhmann(1999). (cid:2) Chaeteessidaesp.;EocenefromIndia.InRustetal.(2010). agreysandstonelevel,whiletheoutcrophasbeendatedasAnte- Jezzinian (Maksoud et al., 2014). Based on geological and palae- PossiblyinChaeteessidae ontologicalcorrelation,theoutcropissituatedintheChoufSand- (cid:2) ArchaeophlebiaenigmaticaPiton,1940;PaleocenefromMenat(France).See stone Formation (¼Gre(cid:1)s de Base or C1 in older usages), and NelandRoy(1996). corresponds to a Barremian fluvial delta deposit. The Al-Rihan amberoutcropwasdiscoveredonlyrecently(in2012byD.A.).Up Mantidae (cid:2) EobruneriatessellataCockerell,1913;EocenefromColorado.InMeyer(2003). to now the following arthropod orders have been found as in- (cid:2) Mantisreligiosa(Linne(cid:3),1758);PliocenefromWillershausen(Germany).In clusionsinthisamber:Acari,Araneae,Archaeognatha,Hemiptera, Beier(1967). Blattaria, Mantodea, Orthoptera, Psocoptera, Thysanoptera, Cole- (cid:2) Mantidaesp.;MiocenefromtheDominicanRepublic.InGrimaldi(2003). optera,Hymenoptera,Neuroptera,andDiptera. (cid:2) Mantidaesp.;OligocenefromGermany.InEhrmann(1999). (cid:2) Mantidaesp.;EocenefromBalticamber.InGrimaldi(2003). (cid:2) ProchaeradodisenigmaticusPiton,1940;PaleoceneofMenat(France).SeeNel 2.3. Burmeseamber(Myanmar) andRoy(1996). AlthoughBurmeseamberhasbeenfoundfromseveraldistricts, Mantoididae such as Shwebo, Thayetmyo, Pakoku, and Pegu, it has only been (cid:2) MantoidamatthiasglinkiZompro,2005;EocenefromBalticamber. (cid:2) Mantoidasp.;MiocenefromtheDominicanRepublic.InGrimaldi(2003). minedandcommercializedintheHukawngValley(Myitkyinaand (cid:2) Mantoididaesp.;EocenefromBalticamber.InWeitschatandWichard(2002). UpperChindwindistricts)ofnorthernMyanmar(Rossetal.,2010). ExcludedfromthisfamilybyWieland(2013). ThenewspecimencomesfromtheNoijeBumHillsintheHukawng Valleyandwherediversebioinclusionshavebeendiscoveredover Liturgusidae theyears(e.g., Ross etal., 2010;Bardenand Grimaldi,2014). The (cid:2) Liturgusidaesp.;EocenefromBalticamber.InErhmann(1999). (cid:2) Liturgusidaesp.;MiocenefromtheDominicanRepublic.InErhmann(1999). mantisoriginatedfromageologicalsectioncharacterizedbyinter- beddedsandstones,siltstones,shales,micriticlimestones,andcoal. Tarachodidae The age of Burmese amber was considered to be late Albian by (cid:2) Tarachodidaesp.;MiocenefromtheDominicanRepublic.InErhmann(1999). Cruikshank and Ko (2003) based on palynomorphs and an ammonite, or lower Cenomanian by Shi et al. (2012) based on Vatidae (cid:2) Vatidaesp.;MiocenefromtheDominicanRepublic.InErhmann(1999). volcanic zircons(but considerNicholson et al., 2015; Ross,2015). ThephylogeneticanalysisofantsstudiedbyBardenandGrimaldi ExcludedfromMantodea (2014)seemstocorroboratetheageproposedbyShietal.(2012). Juramantidaey Ithasbeensuggestedthattheresinproducerwasanaraucariacean (cid:2) JuramantisinitialisVr(cid:4)sanský,2002;UpperJurassicfromShar-TeginMongolia. treeclosetothemodernAgathis. NotconsideredasaMantodeabyGrimaldi(2003)becauseoftheabsenceof charactersofMantodea. Up to now, those arthropod orders found as inclusions are (Rasnitsyn and Ross, 2000; Grimaldi et al., 2002; Ross and York, Jantarimantidaey(¼ArchimantidaeVr(cid:4)sanský,2002) 2004;Rossetal.,2010;BardenandGrimaldi,2014;Bonatoetal., (cid:2) Jantarimantiszherikhini(Vr(cid:4)sanský,2002)(¼ArchimantiszherikhiniVr(cid:4)sanský, 2014; Engel and Grimaldi, 2014; Brolyet al., 2015; Dunlop et al., 2002[Archimantispraeocc.Saussure,1869]).TuronianfromNewJersey.Ac- 2015; Engel et al., 2016a, 2016b; Wunderlich, 2015): Acari, Ara- cordingtoGrimaldi(2003)theholotypeofthisspeciesisaroachbelongingto thefamilyUmenocoleidae(nowPonopterixidae).Asecondspecimenstudied neae, Amblypygi, Solifugae, Thelyphonida, Ricinulei, Opiliones, byVr(cid:4)sanský(butnotconsideredasaparatype)waslaterconsideredapara- Pseudoscorpiones, Scorpiones, Geophilomorpha, Scolopen- typeofAmbermantiswozniakiGrimaldi,2003. dromorpha, Polyxenida, Siphonophorida, Isopoda, Collembola, Zygentoma, Archaeognatha, Ephemeroptera, Odonata, Blattaria, Isoptera, Mantodea, Orthoptera, Phasmatodea, Plecoptera, Der- which corresponds in this area to a fluvial delta swamp deposit maptera, Embiodea, Zoraptera, Hemiptera, Psocoptera, Thysa- (Queroletal.,1992),orwetareasinafore-ergsystem(Rodríguez- noptera, Raphidioptera, Megaloptera, Mecoptera, Coleoptera, Lo(cid:3)pezetal.,2009).SanJustamberwasdiscoveredduringthelast Strepsiptera, Hymenoptera, Neuroptera, Trichoptera, Lepidoptera, decade(seePen~alveretal.,2007)anduptonowthosearthropod andDiptera. orders found as inclusions include Acari, Araneae, Blattaria, Iso- ptera, Orthoptera, Hemiptera, Thysanoptera, Coleoptera, Hyme- 3. Materialstudiedandmethods noptera,NeuropteraandDiptera(Arilloetal.,2008,2009a,2009b, 2010, 2012; Engeland Delclo(cid:1)s, 2010; Pen~alverand Delclo(cid:1)s, 2010; The Spanish specimen (SJ-10-17) was embedded in a high- Pen~alver and Nel, 2010; Pen~alver and Szwedo, 2010; Pen~alver qualitycastingepoxy(Epo-tek301),accordingtotheprotocolsof etal.,2010;Ortega-Blancoetal.,2011a,2011b,2011c;Pe(cid:3)rez-dela Corraletal.(1999)andNascimbeneandSilverstein(2000),which X.Delclo(cid:1)setal./CretaceousResearch60(2016)91e108 95 Fig.1. Palaeogeographicmapshowingtheprovenanceofthestudiedspecimens(mapafterBlakey,2011).Inthetexttherearedetailedinformationfromgeologyoftheoutcrops, andthefossilrecordassociatedtothenewtaxa. allowed physical protection and optimal viewing (dorsal and (2003),KlassandEulitz(2007),andBe(cid:3)thouxandWieland(2009) ventralviews).TheLebanesespecimen(RIH-1E)waspreparedina are observable in fossil specimens, such as: 1) presence of the glasscoffinwithamediumofCanadabalsamandfollowingthose interantennal sulcus bordering the ‘scutellum’ (a profoundly ill- protocolsdescribedinAzar(2000).TheBurmesespecimen(NMS, namedarea)of the frons (absentinBlattodea); 2) raptorial fore- Anderson Collection, National Museums Scotland) was only legs; 3) the profemora with a “femoral brush” (a specialized partially polished and otherwise left untreated. The holotype of groomingdevicelocatedontheantero-distalsurfaceoftheprofe- Chaeteessites minutissimus Gratshev and Zherikhin, 1993, was mora),absentinCretaceousspecies;4)presenceofthesupracoxal restudied and new descriptive information provided herein. The sulcusthatdividestheprothoraxintoprozonaandmetazona,and piece of amber was polished on both sides without being 5)thepartialfusionofveinsRPandMintheforewing.Mantodeans embedded. also exhibit several plesiomorphic characters: 1) three ocelli; 2) Descriptions are provided here in the philosophical under- pentameroustarsi(i.e.,fivetarsomeres);and3)multi-segmented standing that descriptive work forms the fundamental basis of cerci (Wieland, 2013). Grimaldi (2003) characterized the order comparativesciencesandrepresentsthecriticaldatafromwhich Mantodea and based his classification on the following set of broader patterns are derived (Grimaldi and Engel, 2007). charactersobservableinfossilspecimens:1)pronotumquadrate, Morphological terminology generally follows that of Grimaldi saddle-shaped,notcoveringthehead;2)forelegsspinous,raptorial, (2003) and Wieland (2013), with various updates to standardize andwithfullymoveableprocoxae,andwiththeprotibiabearinga termsacrossotherinsectorders(whereapplicable).Thefollowing largeapicalspineorspur;3)mid-andhindlegslong,slender,and abbreviationsareusedforspecimensandtheirofficialrepositories: usedinwalking,and4)forewingwithapseudovein(thepseudo- SJ-10-17,representsthematerialfoundintheSanJustoutcropand veinisnotpresentinallrecentMantodea(A.R.,pers.obs.),butthis housed at the Fundacio(cid:3)n Conjunto Paleontolo(cid:3)gico de Teruel- doesnotrenderitasinvalidasasynapomorphy,merelythatitis Dino(cid:3)polis,Teruel,Spain;RIH-1E,isforthatmaterialhousedinthe subsequentlylostamongsomehighermantises). NaturalHistoryMuseumofLebaneseUniversity,FacultyofSciences GenusAragonimantisgen.nov. II, Fanar, Lebanon; and NMS, is used for National Museums Scot- Typespecies:Aragonimantisaenigmasp.nov. land,Edinburgh,UK. All specimens were drawn using an Olympus U-DA drawing Typelocality.SanJustoutcrop,LowerCretaceous(upperAlbian)of tubeattachedtoanOlympusBX51compoundmicroscope.Photo- Teruel(Spain). micrography relied on a digital camera, ColorView III & Soft Im- Etymology.ThenewgenericnameisacombinationofArago(cid:3)n,the agingSystems,attachedtothesamemicroscope. Autonomous Community where the San Just amber fossil site is located,andthegenericnameMantis,acommonstemfornamesin 4. Systematicpaleontology Mantodea.Thegenderofthenameisfeminine. Diagnosis. Distinguished from other genera known as nymphs in Order:MantodeaBurmeister,1838 Cretaceous ambers (i.e., Chaeteessites, Electromantis, Jersimantis, Family:Incertaesedis Burmantis)mainlyonforelegstructure:profemurwithventromesal Mantodea are considered monophyletic and supported by rowofeightstout,shortspines,alternatingwithninealsostoutbut extensivemorphologicalandmoleculardata.Afewoftheautapo- shorter spines; three relatively short spines (not stiff setae) on morphies proposed by Boudreaux (1979), Klass and Ehrmann ventrolateral edge (the two distal spines are closer together). 96 X.Delclo(cid:1)setal./CretaceousResearch60(2016)91e108 Protibiawithmesal(anteroventral)rowofthickspines,increasing Aragonimantisaenigmasp.nov. insizedistad,withwell-definedarticulation;atapexatleastone Figs.2and3 terminal,thick,posteroventralspinebutmuchshorter(lessthan1/ Typelocalityandhorizon.Thespecimenwasfoundinamberfrom 5)thanapicalanteroventralspine.Probasitarsomereshorterthan grey-black claystones with abundant plant remains in the protibia. Coxae covered by spicules. Femoral and tibial cuticles UtrillasGroup(sensuRodríguez-Lo(cid:3)pezetal.,2009;Barro(cid:3)netal., entirely covered with fine, scale-like microsculpture (at least on 2015), Lower Cretaceous, middleeupper Albian (sensu fore-andmidlegs).Inaddition,ocellipresent. Villanueva-Amadoz et al., 2010), upper Albian (E. Barro(cid:3)n, pers. Fig.2. Aragonimantisaenigmagen.andsp.nov.,holotype:SJ-10-17,inventralhabitusanddorsalviewofheadandanteriormarginofpronotum. X.Delclo(cid:1)setal./CretaceousResearch60(2016)91e108 97 Fig.3. Aragonimantisaenigmagen.andsp.nov.,holotype:SJ-10-17.1)habitus,2)maxillarypalp,3)leftraptorialforeleg,4)distalpartofprotibiaanditsdistalspines,5)detail ofprotibiaandbaseoftarsus,6)aroliumanddistalclawsofforelegpretarsus,7)midlegsurface,8)distalspineofmesotibia.Scalebars:1and3:1.5mm;5:1mm;2,4,6e8: 200mm. 98 X.Delclo(cid:1)setal./CretaceousResearch60(2016)91e108 com. 2015), which corresponds to a deposit of a fluvial deltaic protibial length; probasitarsus and protarsomere 2 with a small swamp. The outcrop of San Just (Pen~alver et al., 2007; Pen~alver dorsallobe,andprotarsomere3withaventrallobe.Propretarsus and Delclo(cid:1)s, 2010) is located in the municipality of Utrillas with prominent, triangle-shaped arolium; paired claws well (Teruel Province, Arago(cid:3)n Autonomous Community, eastern developed,withabasalwidening(Fig.3.6).Femoralandtibialcu- Spain). ticles entirely covered with fine scale-like microsculpture (on at leastfore-andmidlegs)(Fig.3.7).Cursorialmidlegspresent.Esti- Holotype.SJ-10-17(bodyfossil,anteriorhalfofanymph)fromSan mated mesofemoral length: 2.77, width: 0.72; mesofemur Just amber in a prism 23 (cid:3) 16 (cid:3) 3 mm. The amber fragment approximatelyaswideasprofemur,withoutspinesalongitslength. (17(cid:3)8(cid:3)3mm)isdark,withsomebubblesanddesiccatedsurfaces Estimatedmesotibiallength:2.31,width:0.22;mesotibiawithtwo brownincolour,containingdebrisandabundantspecimensiden- largeapicalspurs,each0.32mminlength(Fig.3.8),andaventral tical to those found in Cretaceous French ambers which were rowofshort,thickspine-likesetae.Mesobasitarsusstronglyelon- identified as aerial hyphae of sooty moulds of the genus Meta- gate.Abdomennotpreserved. capnodium (Metacapnodiaceae) (see Girard et al., 2009, 2011). Housed at the Fundacio(cid:3)n Conjunto Paleontolo(cid:3)gico de Teruel- Remarks.Theprofemoralgroomingdevice,orprofemoralbrush,is Dino(cid:3)polis,Spain. presentinallmodernMantodeaandinsomeCretaceoustaxa,such Etymology.Latinaenigma,referringtotheinabilitytoassignittoa as Burmantis, but absent in Jersimantis and Aragonimantis, and is givenfamily. considered an autapomorphy of Neomantodea. The profemoral brushconsistsofslightlythickenedsetaeinBurmantis,whereasin Diagnosis. As for genus, with the following additional characters: morederivedtaxa,thesetaearescale-likeandflattened(Wieland, Antero-ventralrowofprotibialspinespresentondistaltwo-thirds 2013). The profemoral brush is also present in the most basal of tibia and comprising ten thick spines having fine longitudinal genera of recent Mantodea such as Chaeteessa, Metallyticus, and striation. All anteroventral profemoral spines stout, although of Mantoida.Considerablemorphologicalchangesoccurintheonto- differentsizes. geneticdevelopmentofthisstructure.Itispresentinbothnymphal Description (all measures in mm). Head globular as observed in instarsandadults,andhasfeather-shapedsetae;theirshapedoes ventralanddorsalviews,2.20widthincludingcompoundeyes;no notchangeinadults,butsetaearedistinctlylongerandincreasein processes or ridges observed. Compound eyes large,moreorless number. globular(dimensions0.76(cid:3)0.68),protrudinglaterallyfromhead The profemorahavean antero-ventralrowof17 spines (char- capsule(exophthalmic),withalargefrontalfieldandinnermargins acter“45”,state3inWieland,2013),andthisisalsofoundin‘basal’ closetoscape,andconsistingoffine,abundantommatidia.Ocelli moderngenera,suchasChaeteessaandMantoida,butalsoinBlat- present(atleastonelateralposteriorocellusvisible)(Fig.2).Frons tariaandinsomefossilspeciessuchasBurmantisssp.andCreto- and clypeus largely obscured. Mandible with three teeth (as mantislarvalis. figured).Labialpalpsshort,thin,onlytwodistalpalpomerespre- Mantodeanshaveabasal,obliquerowofonetofivespinesbe- served. Maxillary palps well-developed (ca.1.20 length), with at tween the anterior- and posterior-ventral profemoral rows of leastfoursubequalpalpomeres(baseobscure)(Fig.3.2).Antenna spines, termed the “discoidal” spines. The modern genus Metal- filiform, incomplete, with symmetrical antennomeres; flag- lyticus exhibits a single discoidal spine and species of Chaeteessa ellomere1long(longerthanscapeþpedicel);basalflagellomeres only two (Wieland, 2013). Discoidal spines are absent in the shortandcompact(lengthlessthanwidth),graduallylengthened CretaceousgeneraChaeteessites,Ambermantis,Burmantis,Aragoni- apically(Fig.2). mantis,Cretomantis,andJersimantis,aswellasacrossBlattaria. Pronotumlargelyincompleteandpoorlypreserved,apparently Spines on the cursorial meso- and metafemora have been withoutspicule-likesetulae;notcoveringposteriorofhead.Only described in the present species as well as Burmantis lebanensis, fore-andmidlegspreserved.Raptorialforelegscompleteandwell- Jersimantisburmiticus,Cretomantislarvalis,Ambermantis wozniaki, preserved, unlike midlegs. Left foreleg visible, preserved without andSantanmantisaxelrodi(Ho€rnigetal.,2013). deformation, but ventromesal row of spines on profemur not GenusBurmantisGrimaldi,2003 completely visible. Right foreleg cleared, slightly deformed but Typespecies:BurmantisasiaticaGrimaldi,2003,Myanmar. with entire profemoral ventromesal row preserved (thus it has beenpossibletoreconstructtheforelegincompletedetail:Figs.2, Remarks.Grimaldi(2003)didnotindicateintheoriginalaccount 3.3e3.6).Procoxallength:1.10,width:0.43;procoxawithadorsal thelengthoftheprocoxa,althoughheconsideredthemtobeshort andaventralantero-apicaldiverginglobe,withtworowsofspines; inB.asiaticaandB.lebanensis.Basedonthenewmaterialdiscussed ventralsurfacewithoutdiscoidalspines.Profemora;basalthirdof herein,theprocoxaisasisfoundelsewhereamongMantodea,and profemurslightlyinflatedandbulbous,lackingasmallbasalpatch its purportedly short stature may have been overstated or mis- ofsensillae;withanantero-ventralrowofeightstout,shortspines, interpretedintheoriginaldescription(furtherrenderingdoubtful alternating with nine shorter spines; with dense, fine pilosity in Gorochov's (2006) assertions regarding the placement of the ventralfurrow;aprofemoralgroomingdeviceor“femoralbrush” genus). notvisible,thusdistinctionbetweenslightlythickenedorflattened BurmantislebanensisGrimaldi,2003 andscale-likesetaenotpossible.Protibiallength:1.46,width:0.17; Figs.4and6 protibiawithantero-ventralrowoftenthickspinesincreasingin sizedistad(Fig.3.3),ondistaltwo-third,having finelongitudinal Newspecimen.SpecimenRIH-1E(completebodyfossilofanymph) striation; postero-ventral row of spines notvisible, exceptdistal- fromLebanese amber (Figs. 4, 6.1e6.2) mounted in a glass prism most spine (of thin setae); distal spines of both antero- and 12(cid:3)8(cid:3)6mmfilledwithCanadabalsam.Theamberfragmentis postero-ventralrowsthick,withdifferentdegreesofdevelopment transparentandlightlyyellow.Housedintheambercollectionof (the last posteroventral spine is less than 1/5 the size of the last theNaturalHistoryMuseumoftheLebaneseUniversity,Facultyof anteroventralspine:0.04mmvs.0.24mm),andwithwell-defined SciencesII(Fanar),Lebanon. articulationsbutnotspur-like(Fig.3.4);apicalmostspine(¼tibial spur)ofanteroventralrowaslongas3(cid:3)widthofprotibia(Fig.3.5). Locality and horizon. Lower Cretaceous, amber of Al-Rihan, Caza Protarsuslength:2.53(probasitarsus:1.15);probasitarsus0.7e0.8(cid:3) Jezzine (Jezzine Department), Mohafazat Loubnan El-Janoubi X.Delclo(cid:1)setal./CretaceousResearch60(2016)91e108 99 Fig.4. BurmantislebanensisGrimaldi,specimenRIH-1Edorsalhabitus,showingcompletepreservedchaetotaxyandbodycolourpattern,andlateralhabituslessdetailed. (South Lebanon Governorate), southern Lebanon (Azar and Nel, (Fig.5.2);galeaobscured.Antennafiliform,incomplete(onlythetwo 2013). The holotype of B. lebanensis was found at the Bcharreh scapes, one pedicel, and a fragment of one flagellum preserved): outcrop,closetotheHasrounvillageinBarremianclay-sandstones scape(0.18length(cid:3)0.09width)withthreestrongdistalsetae. (Maksoudetal.,2014). Pronotum,mesonotumandmetanotumsurfaceswithscale-like microsculpturehaving distal margins moresclerotizedand finely Original diagnosis. Differs from B. asiatica by its fewer (4 vs.10) denticulate(Fig.5.5);surelytheycorrespondtothesmallirregular small spines on the profemur alternating among thicker spines; tuberclesobservedbyGrimaldi(2003).Pronotumquadrate,com- pronotumandsomescleritescoveredwithsmalltubercles,instead pleteandwell-preserved(0.87length,0.91width,0.40high);not ofminutespiculelikesetulae;cercishorterandwith9e10(vs.12) coveringheadandwithchaetotaxyconstitutedby19pairsofshort cercomeresandwithoutelongatesetaeapically. setae(sixpairsaremarginal)asfigured(Fig.5.5).Mesonotumca. 0.90 length including the anterior covered portion,1.21 greatest Description (measures in mm). Body colouration: pronotum with width and 0.45 high; chaetotaxyconstituted by20 pairs of short twomid-longitudinalbandsandmesonotumplusmetanotumwith setae(11pairsaremarginal)asfigured.Metanotumca.0.80length mid-level maculations and latero-longitudinal bands; abdomen includingtheanteriorcoveredportion,1.24greatestwidthand0.42 with two pairs of longitudinal bands (one centro-lateral and one highwith14pairsofshortsetae(11aremarginal)asfigured. lateral);alllegswithmaculatedpattern,mainlyproximalanddistal Raptorialforelegscompleteandwell-preserved,exceptforsome maculationsasfigured(Figs.4,6.1). portions of tarsi (Fig. 6). Coxae and some indeterminate thoracic Headroundedinfrontal,dorsal,ventral,andlateralviews,width scleritescoveredbyspiculesandwithsomestronglateralsetaeas 1.38includingcompoundeyesand1.20high.Compoundeyeslarge, figured.Procoxallength:1.00,width:0.35.Profemorallength:1.42, withbroadfrontalfield(compoundeyedimensionsinfrontalview: width: 0.34; profemur basal third slightly inflated and bulbous, 0.48 width, 0.69 high; see Fig. 5.1), prominent (exophthalmic); lacking a small, ventro-basal patch of sensillae, but covered by ommatidianotdiscernibleaspreserved.Unpairedanteriorocellus dense, fine pubescence on ventral surface and two longitudinal present(havingstrongocellarsetae,seeFig.5.2),butlateralposterior rows of short spines on dorsal surface (Fig. 5.2), with an ante- ocellinotapparent.Frons,clypeus,andlabrumpreserved(asfigured). roventralrowoffive(maybefour)stout,shortspines,alternating Mandible with fine teeth (as figured). Labial palps short, thin. with slender setae (Figs. 5.2e5.3, 6.4e6.5); three relatively short Maxillary palps well-developed(ca. 0.60 length), with five palpo- spines on postero-ventral edge (the two distal spines are closer meres,subequalinlengthexceptpalpomereIIlongerthanremainder together)andaminutespinedistally;profemoralbrushnotvisible 100 X.Delclo(cid:1)setal./CretaceousResearch60(2016)91e108 Fig.5. BurmantislebanensisGrimaldi,specimenRIH-1E.1)headandraptorialleftforeleginfrontalview,2)sameinventralview,3)rightprofemurinexterno-lateralview(five arrowsindicatefivestout,shortspinesofventromesalrow),4)genitaliainlateralviewshowingcercalcolourpattern(stylihavebeenhighlighted),5)thoracicnotaindorsalview, showingcolourpattern,completechaetotaxy,andtwodetailsofscale-likemicrosculpture. (present in Burmantis ssp.), thus distinction between slightly (Figs. 6.4e6.5). Protarsus: ca. 1.44 long (probasitarsomere 0.72 thickenedorflattenedandscalelikesetaenotpossible. long);probasitarsus0.9(cid:3)protibiallength.[Thenewspecimenhas Protibial length 0.75, width 0.13, with a depression in antero- themesotarsuslongerthanthemesotibia(consideredequalinthe basal position (identical to those observed in Aragonimantis n. originaldescriptionofthespecies,butthishasnotbeenconfirmed gen. and in holotype of C. minutissimus from Santonian Siberian through re-examination of the holotype)]. Pretarsus with promi- amber, E.P., pers. obs.); anteroventral row of 10 thick spines nentarolia,pairedclawswithslightwideningbasally.Mesofemoral increasinginsizedistadindistalthree-quartersoflength(Fig.6.3), length2.77,greatestwidth0.28;mesofemurapproximatelyaswide with fine longitudinal striation (Figs. 5.1e5.2); apicalmost spine asprofemur,withoutspinesalongitslength.Mesotibiallength1.15, (length: 0.36, width: 0.04) nearly 2(cid:3) width of protibia; poster- width 0.15; mesotibia with two large spurs, each 0.15 long (as oventralrowofspineswell-visible,composedbyatleastsixthin longer as mesotibial width), and an anterior row of short, thick spines, distalmost spine thick (apicalmost spine length 0.17, 0.02 spine-like setae. Mesotarsus 1.51 long. Metatibia 1.68 long wide)(Fig.5.2);distalspinesofantero-andpostero-ventralrows (approximately1.5(cid:3)aslongasmesotibia).Femoralandtibialcu- thick,withdifferentdegreesofdevelopment(onelarge,onesmall) ticlesentirelycoveredwithfinescale-likemicrosculpture(Fig.5.2). and with well-defined points of articulation but not spurlike Abdomencompleteandwell-preserved:short,broad,tergiteswith

Description:
Wieland, 2013), and are among the more familiar of insect groups owing to their characteristic raptorial forelegs, large eyes, and distinctive stance and habitus (Zherikhin, 2002; Grimaldi and. Engel, 2005). Mantises are predatory and occupy a wide distribu- tion across generally warmer biomes, mai
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