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

18 Pages·2015·4.24 MB·English

by  Xavier Delclòs

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