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Home , Aleuropteryginae, Ascalaphidae, Burmese amber, New Jersey amber, Ululodes

This is anopenaccess article undertheCCBY-NC-SAThis México. of de Autónoma responsibility Nacional the Universidad under Reviewing Peer Manuscript accepted: Corrected manuscript received: Manuscript received: BSGM2021v73n3a271220 73 (3), A271220. http://dx.doi.org/10.18268/ Mexicana, Geológica Sociedad la de Boletín larvae: lacewing of record the and in Mexican preserved owlfly An Solórzano-Kraemer, M.M.,Haug, J. T., 2021, Haug, C.,Haug, G. T., Baranov, V. A., How tocitethisarticle: [email protected] * Corresponding author:(C.Haug) many. Ger Main, am Frankfurt 60325 25, ganlage 3 ner-Str. 10,80333München, Germany. 2 tinsried, Germany. Planegg-Mar 82152 2, Straße Großhaderner 1 Joachim T. Carolin del Mioceno Una larva de mosca búho preservada en ámbar mexicano y el registro de de crisopa larvae An owlfly larvapreserved inMexicanamber and the Miocenerecord of lacewing license(https://creativecommons.org/licenses/by-nc-sa/4.0/) LMU Munich, GeoBio-Center, Richard-Wag Senckenberg ResearchSenckenber SenckenbergInstitute, M Mnc, eatet f Booy II, Biology of Department Munich, LMU Haug Haug 1,2* , Gideon T. 1,2 Haug Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - - - 1 , Viktor A. Mexico; Neogene. ; ; Keywords: ; amber are discussed here. Miocene in larvae lacewing of rarity the explaining possibly factors taphonomic and Ecological . and Eocene in numbers the to compared low astonishingly is number overall collec the tions, private in specimens additional be to seem there While far. so literature the in specimens eight only with scarce, rather remains and amber Dominican in preserved It only otherwise larvae. includes lacewing of record the Miocene review we Additionally, larva. first lacewing the a and amber, Mexican Miocene of from report second the only represents to specimen Remarkably, the debris it. camouflaging attach to used been have could which setae, numerous carrying protrusions segments finger-like Trunk possess short. rather is trunk The each. least (stemmata) eyes at simple with three hills eye distinct has that view) dorsal (in head square-shaped less or more a from arise stylets These teeth. three with each with stylets, prominent (Ascalaphidae) large owlfly an of larva a likely most amber, Mexican Miocene from larva lacewing a of a specimen new report we Here amber. in preserved predominantly fossils, as also sometimes but fauna, modern the in only not found lower jaws. These impressive larvae can be ven and byupper formed stylets om-injecting prominent with larvae mostly predatory have Insecta of These representatives Holometabola. of lineage small rather a today is (lacewings) Neuroptera ABSTRACT Baranov 1 , MónicaM. Solórzano-Kraemer - - /73(3)A271220/2021 minentes estiletes formados por las mandíbulas minentes estiletes formados son principalmente depredadoras, tienenpro que Estos representantes de Insecta, con larvas bastante pequeño entre los insectos holometábolos. Neuróptera (crisopas) día es hoyen un linaje RESUMEN México; Neógeno. Neuróptera; Myrmeleontiformia; Palabras clave: Ascalaphidae; aquí discutidos. de crisopa en elámbar del Mioceno, son también que posiblemente la rareza expliquen de las larvas Cretácico. Los factores ecológicos y tafonómicos, cantidad de ejemplaresámbar en del Eocenoy sorprendentementecomparaciónbajo en con la colecciones nales en privadas, elnúmero total es ahora. Sibienparecehaber especímenes adicio especímenes mencionados en la literatura hasta y que sigue siendo bastante escaso, con solo ocho incluye en ámbar dominicano fósiles preservados de crisopa, que únicamente Mioceno de larvas Además,primera larva. del revisamos el registro delMiocenoyla crisopa ámbar en mexicano de una reporte representael segundo únicamente para su camuflaje. Notablemente, este espécimen haber sido utilizadas para acumular escombros de dedos con numerosas setas que podrían forma del tronco poseen protuberanciassegmentos en mata) cada una. El tronco es más bien corto. Los que presentan al menos tres ojos simples (stem cuadrada vista (en dorsal) con colinas visibles más o menos provienende unacabeza deforma estiletes, cada uno contres dientes. Estos estiletes y prominentes “moscas búho”) con grandes también denominadas (a veces de Ascalaphidae del Mioceno, queesmuyprobablemente una larva en ámbar mexicano de crisopa preservada larva espécimende una Aquí presentamos un nuevo fósiles, en ámbar.mayoritariamente preservados también como sino a veces en la fauna moderna, no solo se puedenencontrar impresionantes larvas superior e inferior que inyectan veneno. Estas 3 , 1 - - -

Miocene lacewing larvae including specimen in Mexican amber ABSTRACT Miocene lacewing larvae including specimen in Mexican amber INTRODUCTION 1. Introduction 2 2 or , lacewing larvae are quite special in dragonflies, special larvae areor quite lacewing predators. ambush butterflies certain resemble lacewings adult While fierce generally are larvae Lacewing larva. of type unique rather a by ised character is lacewings, of group the Neuroptera, eod a rcnl smaie b Przd la Pérez-de by summarised recently was record amber The resins. fossil to restricted largely is and scarcer much is larvae lacewing of record ( preservation of potential havehigh to a seem wings whoseadults, as found commonly compression been have As lacewings fossils, 2020). Haug in discussion see their highly specialised eyes (for more information, to due ‘larvae’ as addressed forms immature with Insecta of ingroup the 335), p. 2005, Engel and (Grimaldi Holometabola of lineage ly-appearing benefits (Tauber ecological and effectiveness their to due also ago, years some control pest as introduced successfully latteraresevere havepests,lions plant aphid been ( aphids feeding on wings). They are, as the name suggests, specialised lace (brownHemerobiidae and lacewings) (green groups the of larvae the lions, aphid of that is larvae lacewing of fraction well-known Another 2012). (Badano ground sandy in bur rowed funnels trap with cases many in prey their catch which , of those possibly are larvae 1999, 2007). Aspöck and Aspöck 1984; Gepp 1964; (MacLeod palps the parts, distal its by recognisable mainly is and capsule head the into integrated lip, deeply is lower or labium, The palps. lack or parts, maxillae distal The ingroups. lacewing some larva in the of length entire the of third one to up reachcan largerepresentsizestylets a and of pair This tool. sucking and venom-injecting combined maxillae,the a stylet, a lower jaws, form to of one with interlocked jaw,is upper or mandible, Each prominent. very and (prognath) directed forward strongly are mouthparts Their morphology. their / / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín erpea s eeal cniee a ear an considered generally is Neuroptera lacewing of examples prominent most The et al. e.g. e.g. aakn 08. h fossil The 2018). Makarkin 2000). Tauber et al. http://dx.doi.org/10.18268/BSGM2021v73n3a271220 2003). As the As 2003). /73(3)A271220/2021 - - - - /73(3)A2712202021 09, or pcmn fo Ceaeu Spanish Cretaceous from specimens four 2019), Fuente amber la Pérez-de Lebanese 1980; (Whalley Cretaceous from originated specimens 11 least at ambers: Cretaceous cially have specimens, providedespe quite a number of 1995). the Eocene Green River (Dayvaultformation from specimen single a and specimen) presumed et al. (Martins-Neto mation for Crato Cretaceous the from specimen a single include fossils compression from occurrences et al. Fuente of the group Magnoliopsida (“angiosperms”; (“angiosperms”; Magnoliopsida Seyfullah group the of representativesare resin-producingtrees Miocene group the of whilethe during Coniferopsida(“gymnosperms”), representatives are Eocene and Cretaceous the during trees resin-producing The (Solórzano-Kraemer account carefully,into taken be to has bias ecological since treated be must However,comparisons declined. have could larvae lacewing understand of diversity the better when to ambers younger the with interesting to compare the larval diversity forms of is it regard, that On fauna. modern the to pared com ago years million 100 than more lacewings of forms larval of diversity enormous an indicate al. et al. Badano 2017; Zhang Wichard2017; 2018; 2016, (Xia larvae lacewing 2020c) yielded an astonishing amount of amber,this Haug see of background political as “”; for a recent the discussion of p.fig. known284, (also from Amber 3). specimen from amber (Grimaldi 2000, dian amber (Engel and Grimaldi 2008) and a single Cana Cretaceous from specimen single a 2016), Perrichot 2003; (Perrichot amber French Cretaceous 2020), from specimens 2016, three 2012, Fuente la (Pérez-de amber 2018,2019a,2019b,2019c, 2020a,2020b). Ambers Ambers that have so far yielded lacewing larvae Especially these findings from the Cretaceous the from findings these Especially 2018;Lu 2019a, 2019b, 2020a, 2020b). The only two only The 2020b). 2020a, 2019b, 2019a, 2020, including a discussion about another about discussion a including 2020, et al. et al. (2020) and remains growing ( et al. et al. 08, hs anl assemblages faunal thus 2018), 2015; Wang2015; 2018;Makarkin 2018;Haug et al. et al. 2007; Herrera-Flórez Herrera-Flórez 2007; et al. et al. 2010; Wang 2010; 2016; Liu et al et Liu 2016; 05 2018). 2015, et al. e.g. 2018, Haug et al. et al. et al. et - - - -

~6 y) ae o a orlto o foramin of correlation a on amber based Dominican Mya) to (~16 age similar a for argued (2007) Solórzano-Kraemer discussion; in still is fromChiapas amber Mexican of age whereasthe cene (Iturralde-Vinent and MacPhee 1996, 2019), Mio Middle early as dated been has amber can and Domini 2007). Grimaldi Grimaldi and Engel 2005; Engel 1999; Poinar and Poinar 1996; (Wu amber Dominican Miocene from been reported have specimens eight only far So ambers. hitherto notyieldedlacewinglarvae. have ambers Eocene Moreover,Myanmar.other amber from from than findings fewer significantly Haug 2020; et al. Makarkin 2004; Wichard Scheven 2009; Weitschat 2002; Wichard and Weitschat (Weidner1958; MacLeod 1970; Larsson 1978; specimens Janzen 2002; lacewing larval of number al. ( different be could trees the inhabiting aeig ave rm oiia abr and amber Dominican from larvae lacewing Calvillo-Canadell naea (genus) group species extant the of species amber-producing tree in both instances belongs to environments,fromsimilar come ambers, Dominican the and the and Mexican the ambers, Both (Alvarado-Ortega Formation Quinta La the of also sands quartz correspond coarse Miocene those here, to described amber the of origin the Palenque, Municipality,near Agua de Ambers from Estrella de Belén deposit in the Salto Granja La Campo Formation.Quinta La the mines,to whichbelong namely deposit, strontium one and from microfossils mollusks, , of biostratigraphythe yearson million based 22.8 of Serrano-Sánchez However, Rico. Puerto and Hispaniola, Mexico, Chiapas, in formations different from pollen and mollusks, crustaceans, decapodan corals, iferans, 2017a). vn cre i te eod rm Miocene from record the is scarcer Even a quite provided has amber Baltic Eocene ee e umrs te icn rcr o of record Miocene the summarise we Here . Pia 19; onr n Bon 2002; Brown and Poinar 1991; (Poinar L. 2013; Gröhn 2015; Pérez-de la Fuente la Pérez-de 2015; Gröhn 2013; et al. et al. 2012; Wunderlich 2012; Wedmann et al. et al. 00, ) Yt tee r still are there Yet, b). 2020a, 2010). 21) rud o a age an for argued (2015) Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín et al. http://dx.doi.org/10.18268/BSGM2021v73n3a271220 09 Ol 2011; Ohl 2009; et al. e.g. Peris 2018). Hyme et al. et - - - -

2.1. MATERIAL 2. MaterialandMethods 2.3. IMAGE PROCESSING 2.2. DOCUMENTATION METHODS h acsin ubr M B 62. Preserved 642a. Be SMF number accession the under (SMF) Main am Frankfurt Naturmuseum, ForschungsinstitutSenckenberg the at lection und col amber the in stored is specimen Mexico.The Chiapas, in Palenque to Municipality,near Agua de Salto the in deposit Belén de Estrella the from In the centrethis studyof is a single amber piece of amber from theEstrella deBeléndeposit. Mexican Miocene from specimen new a report tion was used to optimise the images (cf. Haug used to be presented here. The built-in HDR func was contrast best the providing combination The background. black a and white was a with performed documentation illuminations, both Under et al. (Haug light cross-polarised coaxial by once ways, different two in illuminated was It inaccessible. documented from one side only, the other side was Keyence a digital microscope.6000 VHX was specimen The on performed was Documentation remains andquitesomedebris. plant as well as (), louse bark one dae), of one Formici(: and one ), , of four , of one (Diptera: six (Collembola), springtail one lacewing larva, there are numerous the syn-inclusions: beside 642, Be SMF piece amber the inside tl irsoe ee rcse atmtcly by automatically processed were microscope ital dig Keyence6000 the VHX on recorded Images view (Haug of field in limitationsovercome to recorded were limitations in field; depth severalof adjacent stacks overcome to recorded were stacks Image 2013b). 03) n oc b uplrsd ig light. ring unpolarised by once and 2013a) et al. /73(3)A271220/2021 / 73(3)A271220/2021 2008,2011). et al. 3 3 - - - -

INTRODUCTION / MATERIAL Miocene lacewing larvae including specimen in Mexican amber AND METHODS MATERIAL AND METHODS / Miocene lacewing larvae including specimen in Mexican amber RESULTS 3. Results 2.4. PRESENTATION 4 4 iil tutrso the fossil were colour-marked Visible structures of grams, saturation, sharpness). (histo images all optimising for used were CS3 and CS2 Photoshop Adobe software. built-in the 110, 111; p. 129, fig. 128) was interpreted as the as interpreted was 128) fig. 129, p. 111; 110, pp. between plates (figure specimen second The provided. was size of indication view.No ventral This in visiblewell is 1B) Figure 2, (specimen specimen (Myrmeleontidae). an of larva the as interpreted was 111) 110, pp. between plates 94; figure fig. 86, (p. specimen first The amber. Dominican preservedlarvaein lacewing of imens lacewing(Chrysopidae). green a of larva a as interpretation an with compatible based. The overall appearance is in our view more is interpretation this characters which on unclear remains Yet().it lacewing brown a well of larva a as it is interpreted author The accessible. stylets prominent the with head the but curled, slightly is body the view; dorso-lateral in (p.specimen fig. 136, The 1A). Figure seen is 224) larva preserved in Dominican amber (specimen 1; occurrences. independent two as specimen same the preting cross-referencing, avoiding inter complete wayof most the represent should redundancy,it certain a includes this While occurrence. original the to reference with chronologically included also are re-figured been has specimen same the which in chronologically.listed Cases are ambers Miocene lacewing larvae depictions All of in occurrences of corresponding structures. to assigned were colours same simplified; slightly were drawings The literature. from larvae wing lace redraw to structures. used was the CS2 Illustrator Adobe of interpretation an provide to / / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín 2) 1) Wu (1996) depicted a a specimen lacewing of Poinar and Poinar (1999) twodepicted spec http://dx.doi.org/10.18268/BSGM2021v73n3a271220 /73(3)A271220/2021 - - - - /73(3)A2712202021 accessible view. indorsal stated to be 2 mm in total length. specimen The is was 1D) Figure 4; (specimen specimen The 142). (p. amber Dominican in preserved larva lacewing limited. these overallspecimens is thereforeinformation of The number. accession any provide not did and also position, systematic the of description rate accu more a for details providenecessary not the did (1999) Poinar and Poinar provided. of was size indication No apparent. well are mandibles visible due to with debris, yet the large well less is 1C) of Figure 3; (specimen specimen this body The (Ascalaphidae). owlfly an of larva ican amber. The first one is a re-figure (p. 32, 32, fig. (p. re-figure a is one first amber. The ican lacewing larvae specimens preservedof in Domin (2007). Grimaldi and Engel havere-figured in mens been speci three All fig. 9.31). 352, (p. setae long with protrusions short view,exhibiting latero-dorsal in wellshowsthatpreservedis specimen it the of ing 2B). Figure 7; Length was stated to be 1.3 mm.Adetailed draw (specimen lion aphid an hence number), further without (AMNH Chrysopidae 38, fig.group the 9.24).The third one is a larva of (p. view ventral in accessible well is it that shows specimen the of photomicrograph A mm. 8 be to the of representativegroup a as interpreted was 2A) Figure 6; (specimen specimen The M3550). tion Collec Morone number (accession Ascalaphidae group the of representative a is specimen second The fig. 9.20). 346, p. 2005, Engel view and (Grimaldi latero-dorsal in accessible amber, preserved clear well is in it that shows photomicrograph specimen the A mm. of 4.5 be to stated was group the of tative represen possible a as interpreted was 1E) Figure Morone 5; (specimen specimen The number M1282). Collection, (accession Myrmeleontidae group the of larva a is one first amber. The ican lacewing larvae specimens preservedof in Domin 3) 5) 4) Scheven (2004) depicted one specimen of a of specimen one depicted (2004) Scheven ne ad rmli 20) eitd five depicted (2007) Grimaldi and Engel three depicted (2005) Engel and Grimaldi ure 10. egh a stated was Length 1900. Currie, Porrerus Navas, 1913. Length 1913. Navas, ------fig. 42and p.40,fig.43),totalsizeabout4.5mm. dominicanus 4, owlfly larva (Ascalaphidae),based onScheven(2004,p.142),sizeunknown. E.Specimen5,antlionlarva,possiblyof 2 (p. 86,fig. 94and figure between pp. 110,111).C.Specimen3(figurebetween plates pp. 110,111;p.129,fig.128).D.Specimen plates (Chrysopidae), based onWu(1996,p.136,fig. 224),sizeunknown. B,C.Based onPoinarand Poinar(1999),sizesunknown. B.Specimen values indicatecorresponding structures.Not toscale,specimenswithmoreavailabledetailsdepictedlarger. A.Specimen1,aphid lion Figure 1 drawings of the habitus and the head (p. 46, fig. 46, (p. head the and habitus the of drawings interpretive and 47) fig. 45, (p. photomicrograph group the of representative a as interpreted was 2C) Figure 8; (specimen specimen The (accession MACT-1200). number Ascalaphidae group the of larva a is specimen third The provided. was 43) fig. 40, (p. Poinar, 1996. Additionally an interpretive drawing species the of tive specimen was interpreted as a possible representa numberaccession givenwas MACT-1282.as The larva depicted in Grimaldi and Engel (2005). Here the specimen photomicrograph of 5, of the antlion The second one is a grey-scale version (p. 39, fig. 42) an accession number (DR-10-1454) was provided. Here (2005). Engel and Grimaldi in depicted lion 7, specimen of drawing the of 34) Lacewinglarvaepreserved inMioceneDominicanamber fromthe literature.Allsimplified interpretive drawings; similargrey Stangeand Poinar.MACT-1282,basedonGrimaldiand Engel (2005,p.346,fig. 9.20;alsoEngel and Grimaldi2007,p.39, Ululodes Porrerus dominicanus Porrerus , of a distinct species sp.A species 1. distinct a of , Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín http://dx.doi.org/10.18268/BSGM2021v73n3a271220 i.e. Stange and Stange the aphid the - rwn fthehead(p. 50,fig. 51)wasprovided.drawing of Additionally,interpretive 2. sp.an species distinct as a the representativegroup of interpreted was Figure2A) 6; (specimen specimen numberaccession givenwas MACT-3550.as The larva owlfly depicted in Grimaldi and Engel the (2005). Here 6, specimen of photomicrograph the of 50) fig. 49, (p. version grey-scale a is one fifth The mm. stated1.5 was be capsule to head the of Length 49). fig. 47, (p. provided was head the of drawing interpretive An 1. sp. species distinct the group the of representative a as interpreted was 2D) Figure 9; (specimen men speci The DR-10-2010). AMNH number sion the group a Ascalaphidae representative(acces of also is (p.fourthspecimen mm 5.9 The about 44). be to stated was length Body provided. were 48) /73(3)A271220/2021 / 73(3)A271220/2021 Ululodes Ululodes adas fof also and ado the and of Porrerus 5 5 - -

Miocene lacewing larvae including specimen in Mexican amber RESULTS Miocene lacewing larvae including specimen in Mexican amber RESULTS (Ascalaphidae), 46, fig.48),owlfly larva(Ascalaphidae), lion (Chrysopidae), DR-10-1454,totalsizeabout 1.3mm.C,D.Based onEngel and Grimaldi (2007).C.Specimen8(p.45,fig.47and p. Ululodes on Grimaldiand Engel (2005).A.Specimen 6 (p.38,fig.9.24;alsoEngel and Grimaldi 2007,p.49,fig. 50),owlfly larva (Ascalaphidae), similar greyvaluesindicatecorresponding to scale,specimenswithmore structures.Not availabledetailsdepicted larger.A,B.Based Figure 2 6 6 holders whether we could depict their images in images their depict could we copyright whether holders the asked we available, images these Haug 2019; Haug websites as scientific source in Haug and use of of online certain resources as additional data sources (see used discussion we amber, Miocene in vae 6) / / Lacewing larvae preserved in Miocene Dominican amber from the literature, continued. All simplified interpretive drawings;interpretivesimplified continued. All literature, the from preservedin MioceneDominicanamber larvae Lacewing sp.2,MACT-3550,totalsizeabout8mm.B.Specimen 7 (p.352,fig. 9.31;alsoEngel and Grimaldi2007,p.32,fig.34),aphid Given the scarceness of data on lacewing lar lacewing on data of scarceness the Given Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Ululodes sp.1,AMNH DR-10-2010,headcapsule length about1.5mm. et al. 2020a). In order to make to order In 2020a). Ululodes http://dx.doi.org/10.18268/BSGM2021v73n3a271220 sp. 1, MACT-1200, body length about 5.9 mm. D. Specimen 9 (p. 47, fig. 49), owlfly larva owlfly 49), fig. 47, (p. Specimen 9 D. mm. 5.9 about length body MACT-1200, 1, sp. /73(3)A271220/2021 - /73(3)A2712202021 specimen is unclear. The second the these of specimens of Size (terratreasures.com). 101 Adventures TerraTreasures and by (Figure provided kindly image was 3A) The (Ascalphidae). owlfly an of the larva is specimens these of first The amber. six such specimens, all are preserved of in Dominican images present we total, In contribution. this Figure 3 Images fromGlobalNaturalResources. C.Antlion.D,E.Owlfly.to scale. F.Aphid lion.Not TerraTreasures and Adventures101 (terratreasures.com). B.Antlion;imagefrom Huang Yi-Jen (amber boy, huangyijen0708). C–F. r E sas av fanowlfly (Ascalaphidae). ure 3E)isalsoalarva of slightly longer than 4 mm. The fifth specimen is (Fig It (Ascalaphidae). owlfly an of larva the is 3D) long.mm 3 than less (Figurefourthspecimen The slightly is It (Myrmeleontidae). antlion an larvaof the is 3C) (Figure specimen third Resources. The specimen were kindly provided by GlobalNatural sixth to third the of Images unclear. is specimen the of Size boy, huangyijen0708). (amber Yi-Jen Huang byprovided kindly was 3B) (Figure image The (Myrmeleontidae). antlion an of larva the is Images of additional specimens of lacewing larvae preserved in Miocene Dominican amber. A. Owlfly; image from image Owlfly; A. Dominican amber. in Miocene preserved larvae lacewing of specimens additional of Images Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín http://dx.doi.org/10.18268/BSGM2021v73n3a271220 - - to bediscovered. still remains that valuablediversity the about certain information to contribute sources internet cene lacewing larvae, specimens found in different Mio of scarcity distinct the to available.Due are available for further detailed study, only the images not are specimens six long.These mm 4 about is It (Chrysopidae). lion aphid an is 3F) (Figure men speci sixth The (istockphoto.com). Images Getty by iStock repository online the in available is also image This long. mm 3 than less slightly is It /73(3)A271220/2021 / 73(3)A271220/2021 7 7 - -

Miocene lacewing larvae including specimen in Mexican amber RESULTS Miocene lacewing larvae including specimen in Mexican amber RESULTS (compound structure formed bymandible and maxilla);ta=trunk appendage= trunk (“leg”);te end. abdomen (posterior trunk); or protrusion;pt=prothorax;sy stylet hc=headcapsule;msmesothorax; mt=metathorax;pr process ring light.B.Colour-marked interpretation of structures, arrowheads indicate stemmata.Abbreviations:1–3=teethon stylet;ad= Figure 4 8 8 h Snkneg oshnsnttt n Natur und Forschungsinstitut of Senckenberg collection the amber the of part is and Mexico, Chiapas, Municipality,in PalenqueAgua near de Salto the in deposit Belén de Estrella from amber 4B, and 5). The specimen originates from Mexican 4A, (Figures here reported is (Ascalaphidae) larva 7) and thick. large rather is piece amber The 642a). Be SMF number (accession Main am Frankfurt museum, / / New owlfly larva from Miocene Chiapas amber, specimen SMF Be 642a. A. Composite image, blackbackground, NewowlflylarvafromMiocene Chiapasamber, specimenSMF Be642a.A.Composite unpolarised Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín nw osl pcmn f a rsmd owlfly presumed a of specimen fossil new A http://dx.doi.org/10.18268/BSGM2021v73n3a271220 /73(3)A271220/2021 - /73(3)A2712202021 nent setae. Ocular segment recognisablebypair setae.segment a nent Ocular in view.dorsal Lateral rims with numerous promi square-shaped roughly capsule Head trunk. than longer structures protruding and capsule Head capsule. head distinct with Head 1–5. segments Head: differentiated intoheadandtrunk. and segments) post-ocular 19 + segment (ocular segments 20 (presumably) into organised is Body Description: formed byformed ocular segment and post-ocular The larva is visible in dorsal view. dorsal in visible is larva The - Miocene Chiapas amber,based onspecimenSMF Be642a. Figure 5 r cre i te itl ein te i i facing inwards. is prominentareThree presentteeth along tip the region, distal the They in curved (prognath). are forward protruding capsule, head as long as times 1.5 large,almost Mandibles mandibles. accessible, 3 segment post-ocular of compound structures (stylets), yet only appendages post-ocular segments 3 and 4 presumably forming of Appendages apparent. 2 and post-oc 1 segments ular of structures external No protrusions hills”). distinct (“eye on capsule, head on erally antero-lat Groups (stemmata). eyes simple three eyes.simple atEachleast with group of groups of Simplified restoration of thenew owlfly larvafrom Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín http://dx.doi.org/10.18268/BSGM2021v73n3a271220 - - 4.1. IDENTITYOFTHESPECIMEN 4. Discussion rn sget 41 (boe sget 1–7; segments (abdomen 4–10 segments Trunk shorter.slightly Protrusionssegment. preceding to sub-similar 8) segment post-ocular (metathorax; 3 segment Trunksetae. long numerous with sion protru Each side. each on protrusions finger-like two With head. the as wide as about 7) segment post-ocular (mesothorax; 2 segment Trunk setae. a With head. than smaller protrusion on each side, narrower bearing numerous 6) segment ular post-oc (prothorax; 1 Trunksegment (abdomen). ones posterior further than larger (thorax) ments Trunk: the mandible. of edge outer and inner the on present are setae the the mandible.inside of Additionally numerous ontiformia (Haug ontiformia been present Myrmele in other representatives of also have teeth three (Ascalaphidae). Yet, owlflies and only (Myrmeleontidae) antlions of larvae fauna in found modern the in are teeth prominent Three lacewings). antlion-like all with group (the Myrmeleontiformia of and Neuroptera of larva a as specimen the identify to allow mandible morphology the especially and shape overall The subdivision onlypartlyapparent. (legs) are apparent. Details remain largely unclear, appendages trunk regionsMoreor distal end. rior poste the along setae long numerous With wide. than slender,longer 17–19) segments post-ocular 9–11; segments undifferentiated abdomen senting with protrusion numerous long setae. Each Trunk end (most likely side. repre each on protrusion narrowing towards the posterior. With a finger-like strongly segments, 16) preceding than longer segment slightly post-ocular 8; 11 segment segment (abdomen Trunk setae. long numerous with protrusion Each side. each on protrusion ger-like fin a with segment Each posterior. the towards narrower Progressively segments. preceding than Shorter sub-similar. 9–15) segments post-ocular ogr hn ie Atro tre seg three Anterior wide. than longer /73(3)A271220/2021 / 73(3)A271220/2021 et al. 2019c). 9 9 ------

Miocene lacewing larvae including specimen in Mexican amber RESULTS / DISCUSSION Miocene lacewing larvae including specimen in Mexican amber DISCUSSION CRETACEOUS AMBER 4.2. THEAMBERRECORDOFLACEWING LARVAE: 10 10 bbl. lo ohr etrs hrceiig the characterising features other Also, bubble. a under concealed partly but visible, well not is it fossil there is a the tibio-tarsus, this the region appendage of In whether unclear partly remains leg). it here described (metathorax trunk on 3 (“fusion”) appendage tibio-tarsus a the of share presence Ascalaphidae and Myrmeleontidae upto lacewing larvae (Wang output of the concerning unparalleled is amber Myanmar represent others forms. aberrant-appearing different, astonishingly but forms, modern resemble larvae these larvae.of lacewing Some of diversity France and Myanmar have yielded an astonishing Jersey,New Spain, from Canada, ambers taceous (Pérez-de la Fuente from the Lebanon represent the oldest such larvae larvae Lacewing larvae. lacewing for true also is ambers have been given quite some attention. This al. (Schmidt mites eriophyoidean of species two and (Diptera), midge a Insecta, of representative groupa specific isquitesignificant. of fossil oldest the finding regard,thatrecord. On fossil the in preserved rarely otherwise are which behaviour their and organisms of history tionary years old and on the evolu ecosystems millions of important especially forestbecause they provide data on the of ecology as considered been have time geological deep from Ambers entomologists. especially scientists, various fascinated has Amber (Ascalaphidae) owlfly an of report first the possibly amber,and larva from lacewing this a of report first the 4–8), figs. 10–13, pp. 2007; Grimaldi and Engel pidae; (the other one is an adult lacewing, Mantis a lacewing from Mexican amber second report of the represents this case, any In Ascalaphidae. of species a of larva the is specimen the that likely visible. Still the overall makes it most morphology Badano by forward put (as Ascalaphidae + Myrmeleontidae group / / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín 02. hrfr, h isc-ih Cretaceous -rich the Therefore, 2012). Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín rasc mes ae o a ol yedd one yielded only far so have ambers . et al. et al. 2018, 2019). Younger Cre 07 p 10 ae not are 100) p. 2017, http://dx.doi.org/10.18268/BSGM2021v73n3a271220 et al. 2016: about et /73(3)A271220/2021 - - - /73(3)A2712202021 BALTIC AMBER 4.3. THEAMBERRECORDOFLACEWING LARVAE: atc me, ae ieie ile lacewing provides yielded amber Baltic lineages, Forsome larvae. likewise have especially amber, Eocene, Baltic the from ambers Younger theiceberg. the tipof net by various amber traders, this seems to be only inter via offered specimens additional numerous Given collections. amber Cretaceous different in available currently are specimens 200 than more Fuente la Pérez-de also see 2020b; 2020a, 2019c, 2019b, Haug 2018; Makarkin 2018; 2016, Liu in specimens single mostly specimens; 30 hs s qie ihr ubr oprd o the Miocene record. to compared number higher quite a is this still lacewings), (silky and 07.19a) seven Wichardspecimens; larval with piece one (especially by influenced strongly is number This amber. Baltic in preserved larvae lacewing of specimens et al. remains affinity unclear ( exact the but Neuroptera, of larvae clearly are that specimens some also and Haug specimens; twelve least (at Psychopsidae 1970), MacLeod specimen; (one Weitschat1970; Wichardand 2002), MacLeod specimens; two least (at Ascalaphidae Makarkin specimen; (one erobiidae 2004, pp. 68, 69; Weitschat 2009, p. 254, fig. 45). Hem Scheven specimens: (three Chrysopidae Wedmann specimens; five least (at 2011), Ohl specimen; (one tispidae Man of larvae include finds Further traders). of far; Haug so specimens 20 least (at Nevrorthidae example for amber,as Cretaceous the for than specimens of numbers concerning extensive more is record to be present). For some other lineages the Eocene seem more 2002; 1998, Wichard and Weitschat far; so specimen one least (at Wichard specimens; two (at least example for as record, oldest the 2020a). In total, there are (at least) about 50 about least) (atarethere total, In 2020a). et al. e.g. et al. Larsson 1978, see discussion in Haug 2020). It appears that overall clearly overall that appears It 2020). 2020b; seen more in the possession et al. 2009, p. 113, fig.p.113, 2009, et al. et al. et al. et al. 09 and 2009) et al. 2020a) 2019a, 2013), 2012), et al. - - -

LOWER FORTHEMIOCENERECORD? 4.4. WHYISTHENUMBEROFLACEWING LARVAE of unknown age from Madagascar and the Afri the and Madagascar from age unknown of resinhaveterygidae or fromcopal described been 2013, p. 55, group thefig. Coniop 206). Adults of Colombian copal or resin (age unknown) (Kobbert from known is larva lacewing a of case one only resin, or copal younger In small. still is inclusions relatively recentamber with so that the amount of (Schmidt Zealand New (Antoine Peru from 2014), Ethiopia (Shi China fromPerrichot 2020), and (Bouju from known are amber, Mexican the besides bioinclusions, with deposits amber cene Mio However, amber. Dominican in preserved specimens to restricted far so recordwas Miocene extensive. Besidesthe specimen reported here, the than the Cretaceous one. The Miocene is even less extensive less The is larvae lacewing of deposits.record Eocene fossil younger to Cretaceous the forestsfromamber in larvae lacewing of presence the in decrease apparent an shows internet the in and literature the in available record fossil The imens dueto several (Haug factors overall spec amber has provided ahighoutput of Myanmar Especially Myanmar). (Lebanon, Asia (Canadian and America amber) Spanish (Frenchand Europe amber), North including ambers) Eocene and Miocene for case the also is this (yet coverage geographic wide a with ambers several sampling or ecological bias? The Cretaceous offers a not and pattern true a represent this could far 6). Inhow (Figure the Miocene in and lowest Eocene, finally the in lower is number the Creta ceous, the for available is number highest The overtime: larvae available lacewing of decrease a larvae. prising that there are so lacewing few specimens of sur somehow view,is this it of light the in 2016); erous” amber ( amber has been considered to be a “highly fossilif Eocene in (Grimaldi amber Cambay and amber Dominican Miocene in reported also group a 2004), (Engel mainland can The simple numbers give a pattern that shows that pattern a give numbers simple The e.g. Arillo and Ortuño 2005; Penney et al. Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín et al. et al. 2018), all discoveredall 2018), http://dx.doi.org/10.18268/BSGM2021v73n3a271220 2013). Dominican 2013). 2006), and from and 2006), et al. 2020c). et al. ------

ursaee Cerlpdaee r or Cheirolepidiaceae , ingroups the of also but the ingroup of mostly (“gymnosperms”), Coniferopsida of representatives were ambers, Cretaceous the generated later which trees, resin-producing The Fuente Haug 2020; 2019, 2018, 2016, 2012, la Pérez-de 2008; Grimaldi and (Engel specimens significant of number a providedquite 2020). Yet, the other Cretaceous have ambers also 2019b, Herrera-Flórez 2020b; 2020a, 2019c, 2017; Badano 2015; Wang Xia 2008; Grimaldi and (Engel amber mar Myan from reported been have larvae lacewing It may therefore not be surprising that so many new gnr) ti icue lcwns f te ingroup (Grimaldi theAleuropteryginae of lacewings includes this (genera); groups species 18 total in amber, Dominican in preserved forms fossil havewhich but Australasia, to restricted representatives have nowadays that al. (Peng Australia and Africa southern Asia, East South- in today occur Psychopsidae of sentatives Repre 2010). Aspöck and (Aspöck Australia and days occur in Mediterranean nowaEurope, China, Japan Nevrorthidae of Representatives ambers. American Miocene in them find to unlikely more it makes thatoccurrence of rangesrestricted quite larvae. lacewing other than easily more resin in trapped PsychopsidaeNevrorthidaebecame and of larvae Nymphidae that speculateWe might 1970). (MacLeod known of larva single is, a there only as example, rare, for quite are groups other of vae Lar above). (see Nevrorthidae and Psychopsidae lacewings, within lineages specific to due mostly is larvae of number high relatively the closer, look (Wolfe Sciadopityaceae or Pinaceae ingroup the of Coniferopsida group the the of representative a also was amber Baltic of Eocene tree resin-producing The amber. Baltic from known larvae lacewing some quite are there that surprising be not should it Hence tradition. (Seyfullah 01. hr ae eea eape o groups, of examples several are There 2011). The Eocene Baltic amber has a long research long a has amber Baltic Eocene The oa Nvotia ad scosde show Psychopsidae and Nevrorthidae Today et al. et al. /73(3)A271220/2021 / 73(3)A271220/2021 et al. 2018). 2016; Liu 2018;Haug et al. et al. et al. 2016, 2018; Zhang 2016). Yet, if we Yet,if 2016). et al. 03. tl, we Still, 2013). 2018,2019a, et al. 2020a). et al. et al. 11 11 et - - - -

Miocene lacewing larvae including specimen in Mexican amber DISCUSSION Miocene lacewing larvae including specimen in Mexican amber DISCUSSION in theEocene. Coniferopsida (“gymnosperms”) andMagnoliopsida (“angiosperms”).the dominanceofgroups Note Psychopsidae and Nevrorthidae Ascalaphidae (owlflies; lowerright image).Thedifferent sources of amber areindicated of byschematicrepresentatives thegroups Chrysopidae; greenlacewings, aphidlions;upper rightimage)and alarger unnamed group including Myrmeleontidae(antlions)and for someingroups:Neuroptera,as of aswell Nevrorthidae,Psychopsidae(silkylacewings), Chrysopoideaingroup (including its Figure 6 12 12 these two lineages might, in any case, be the part of of absence The 2006). (Grund larvaeaquatic with Insecta, of groups merolimnic of tatives represen concerning poor notoriously is deposit in Dominican amber is furthermore unlikely as the The representativesoccurrence Nevrorthidae of of it. exclude fully not also can Nevrorthidae,we but and Psychopsidae of larvae find to expect cannot / / Depiction of Depiction number of larval lacewing specimens preserved inamber per timeslice.Numbers givenfortheentire group Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín http://dx.doi.org/10.18268/BSGM2021v73n3a271220 i.e. those /73(3)A271220/2021 - /73(3)A2712202021 bnac fsternorrhynchan hemipterans. abundance of ing their abundance and diversity in ambers to the general byrelatdiscussed (2007) was Grimaldi and Engel in lacewings of bias ecological An bias. sampling by explained easily be not may ambers from this amber israther low. larvae lacewing known of number the that effect h hg nme o ave rm Cretaceous from larvae of number high The - the different samples. in 3.7% to 0.3 from common, more were wings Zherikhin by collected “gymnosperms”) Coniferopsida, of of species from Pinus originating resins modern in (Solórzano-Kraemer nized fromEuarthropoda 3,923 representatives of recog been has (0.02%) Neuroptera group the of adult one only 2013, in collected resin Malagasy were collected (Solórzano-Kraemer adult) one and larvae (four lacewings five imens), spec (89 abundant lightly were sternorrhynchans the in mer (Solórzano-Krae Madagascar in and Mexico in “angiosperm”) Magnoliopsida, forestsof (ingroup in traps sticky and resin ern-day than larvae) and sternorrhynchan adults hemipterans in mod more are there abundance low the within however adults, and (larvae lacewings of lowabundancethe support also studies phonomic Actuo-ta ambers. Miocene in rare are latter The ih re o h group Fabaceae butand Magnoliopsida, “angiosperms”) the of trees with associated and abundant today are Miocene the specific faunas. owntheir with (“angiosperms”) Magnoliopsida of ingroups groups both Dipterocarpaceae, the and of Fabaceae trees by produced were ambers in preserved (Solórzano-Kraemer therefore amber and resin in entombed resin-probecome to probability the higher a to have tree ducing relation closer in or in living Eocene ambers. frequent in Miocene amber than inCretaceous or more be to seem larvae antlion The organism. of the the resin drip in of the soil to preserve this kind styles. implies This resin production at soil level or life soil-dwelling their to amber in meleontidae) (Myr larvae antlion of scarcity the related (2020) Fuente la larvae. Pérez-de of specific not are However, it has to be noticed that these data these that noticed be to has However, it Some elements of the fauna abundant during abundant fauna the of elements Some It is well known that the more active organisms et al. . and L. Hymenaea 2015, 2018). Notably, in the sticky traps Notably,sticky 2018). the 2015, in et al. Picea (2009) in Northern Eurasia lace Eurasia Northern in (2009) tp frs i Mxc, where Mexico, in forest -type . it, 84 bt ingroups (both 1824 Dietr, A. et al. Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín 2018). On the contrary,the On 2018). Hymenaea et al. http://dx.doi.org/10.18268/BSGM2021v73n3a271220 2018). Miocene 2018). Hymenaea et al. igop f of (ingroup 2015). In -type et al. ------

5. Outlook Hitherto, there is no study of differing ecological differing of study no is there Hitherto, ambers. Cretaceous in also antlions of case the in and ambers, Eocene in rare quite are larvae lion three aphid-lion specimens). Owlfly(-like) and ant additionally,specimens; (four antlions and imens) spec (nine owlflies of larvae are ambers Miocene in forms common most the that interesting is it (Peris Magnoliopsida undergo an evolutionary the shift group to trees of not did lacewings, including pollinators, as such Cretaceous, the during Coniferopsida group the 2015, 2017a). (“gymnosperms”; Coniferopsida group the of trees by dominated forests in absent these ambers intolargerthese ambers quantitative analyses. ambers. In this way, we hope to be able to integrate Miocene from larvae lacewing more on report to to serve encourage more also people, professional and amateur, can contribution present the that Wesurface.not hope do present, fact in being yet larvae, that effect the to leads simply that adults this aspect. in challenging partly is larvae lacewing available overallof yetnumberthe lowcomparison, a such to slice time another add potentially could cene Mio The general. in diversity of losses also but particular,in diversity lacewing of changes the of understanding our improving for patterns esting time through larvae (Haug lacewing of morphology comparing quantitatively for used been already have amber Eocene and Cretaceous from Fossils from unbiasedamber, copalandresin collections. data quantitative more need will we aspect, this relatedbias.ecological an to For exploringfurther is larvae lacewing of abundance the whether test could study actuo-taphonomic an Such opsida. vs. Magnoliopsida groups the of trees resin-producing on larvae on concentrated biases oeohrfua lmnsrltdt re fof trees to related elements faunal other Some It may still be a matter of focussed interest in interest focussed of matter a be still may It et al. 2020a). Such studies can reveal inter reveal can studies Such 2020a). /73(3)A271220/2021 / 73(3)A271220/2021 et al. 2017b). In this regard, this In 2017b). e.g. Peris et al. 13 13 - - - - -

Miocene lacewing larvae including specimen in Mexican amber DISCUSSION / OUTLOOK ACKNOWLEDGEMENTS / Miocene lacewing larvae including specimen in Mexican amber REFERENCES References Acknowledgements 14 14 com), Huang Yi-Jen (amber boy, huangyijen0708), (terratreasures. 101 Adventures and raTreasures Ter thank We comments. helpful for reviewers Serrano Lourdes México,de Sánchez,twoanonymousCiudad and de María to grateful are We Arillo, A., Ortuño, V.M., fossil 2005, Catalogue of J.J., Flynn, D., Franceschi, De P.O., Antoine, M., Cuevas-García, J., Alvarado-Ortega, software. isLEONpublication This #18. in providing open access, open source and low cost involved people all of effort the appreciate highly Wesupport. continuous for Munich, Starck,J.M. the LMU excellent via Junior Researcher Fund. funded We thank was CH CGL2017-84419). UE AEI/FEDER, Spanish “CRE”, (project Spain Competitiveness of and Economy of Ministry the thanks also MMSK 90946). N. (Project MMSK and Professorship Lichtenberg a with JTH funds kindly Foundation Volkswagen The images. ing provid kindly for GlobalNaturalResources and / / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín u Ntrud – ei B Gooi und (Geologie Beiträge Palaontologie), 352,1–68. B Serie – Stuttgarter Naturkunde zur (Miocene): Dominican amber from described species insect pnas.0605801103 https://doi.org/10.1073/ 103(37), America, 13595–13600. of States United of Sciences the of Academy National the of Proceedings Miocene: middle diversity the during Neotropical Salas- reveals A., Amazonia western Goswami, from Amber 2006, R., Gismondi, N., Espurt, Y., Baby,Calderón, A., P.,M., Nel, Benammi, https://doi.org/10.1016/j. jasrep.2017.11.029 17, Reports, 462–476. 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