Boletín de la Sociedad Geológica Mexicana / 73 (3) / A271220/ 2021 / 1 An owlfly larva preserved in Mexican amber and the Miocene record of lacewing larvae Una larva de mosca búho preservada en ámbar mexicano y el registro de larvas de crisopa del Mioceno Carolin Haug1,2*, Gideon T. Haug1, Viktor A. Baranov1, Mónica M. Solórzano-Kraemer3, Joachim T. Haug1,2 ABSTRACT 1 LMU Munich, Department of Biology II, ABSTRACT RESUMEN Großhaderner Straße 2, 82152 Planegg-Mar- tinsried, Germany. Neuroptera (lacewings) is today a rather Neuróptera (crisopas) es hoy en día un linaje 2 LMU Munich, GeoBio-Center, Richard-Wag- small lineage of Holometabola. These bastante pequeño entre los insectos holometábolos. ner-Str. 10, 80333 München, Germany. representatives of Insecta have mostly Estos representantes de Insecta, con larvas que predatory larvae with prominent ven- son principalmente depredadoras, tienen pro- 3 Senckenberg Research Institute, Senckenber- om-injecting stylets formed by upper and minentes estiletes formados por las mandíbulas ganlage 25, 60325 Frankfurt am Main, Ger- lower jaws. These impressive larvae can be superior e inferior que inyectan veneno. Estas many. found not only in the modern fauna, but impresionantes larvas no solo se pueden encontrar * Corresponding author: (C. Haug) sometimes also as fossils, predominantly en la fauna moderna, sino a veces también como [email protected] preserved in amber. Here we report a fósiles, mayoritariamente preservados en ámbar. new specimen of a lacewing larva from Aquí presentamos un nuevo espécimen de una Miocene Mexican amber, most likely a larva de crisopa preservada en ámbar mexicano larva of an owlfly (Ascalaphidae) with del Mioceno, que es muy probablemente una larva large prominent stylets, each with three de Ascalaphidae (a veces también denominadas teeth. These stylets arise from a more or “moscas búho”) con grandes y prominentes less square-shaped head (in dorsal view) estiletes, cada uno con tres dientes. Estos estiletes that has distinct eye hills with at least provienen de una cabeza de forma más o menos three simple eyes (stemmata) each. The cuadrada (en vista dorsal) con colinas visibles trunk is rather short. Trunk segments que presentan al menos tres ojos simples (stem- possess finger-like protrusions carrying mata) cada una. El tronco es más bien corto. Los numerous setae, which could have been segmentos del tronco poseen protuberancias en used to attach camouflaging debris to forma de dedos con numerosas setas que podrían it. Remarkably, the specimen represents haber sido utilizadas para acumular escombros only the second report of a lacewing para su camuflaje. Notablemente, este espécimen How to cite this article: from Miocene Mexican amber, and the representa únicamente el segundo reporte de una Haug, C., Haug, G. T., Baranov, V. A., first larva. Additionally, we review the crisopa en ámbar mexicano del Mioceno y la Solórzano-Kraemer, M. M., Haug, J. T., 2021, Miocene record of lacewing larvae. It primera larva. Además, revisamos el registro del An owlfly larva preserved in Mexican amber includes otherwise only fossils preserved Mioceno de larvas de crisopa, que únicamente and the Miocene record of lacewing larvae: in Dominican amber and remains rather incluye fósiles preservados en ámbar dominicano Boletín de la Sociedad Geológica Mexicana, scarce, with only eight specimens in the y que sigue siendo bastante escaso, con solo ocho 73 (3), A271220. http://dx.doi.org/10.18268/ literature so far. While there seem to be especímenes mencionados en la literatura hasta BSGM2021v73n3a271220 additional specimens in private collec- ahora. Si bien parece haber especímenes adicio- tions, the overall number is astonishingly nales en colecciones privadas, el número total es low compared to the numbers in Eocene sorprendentemente bajo en comparación con la and Cretaceous ambers. Ecological and cantidad de ejemplares en ámbar del Eoceno y del Manuscript received: taphonomic factors possibly explaining Cretácico. Los factores ecológicos y tafonómicos, Corrected manuscript received: the rarity of lacewing larvae in Miocene que posiblemente expliquen la rareza de las larvas Manuscript accepted: amber are discussed here. de crisopa en el ámbar del Mioceno, son también aquí discutidos. Keywords: Ascalaphidae; Peer Reviewing under the responsibility of Myrmeleontiformia; Neuroptera; Palabras clave: Ascalaphidae; Universidad Nacional Autónoma de México. Mexico; Neogene. Myrmeleontiformia; Neuróptera; México; Neógeno. This is an open access article under the CC BY-NC-SA license(https://creativecommons.org/licenses/by-nc-sa/4.0/) Miocene lacewing larvae including specimen in Mexican amber http://dx.doi.org/10.18268/BSGM2021v73n3a271220 2 / Boletín de la Sociedad Geológica Mexicana // 7373 (3)(3) // A271220/A271220 / 2021 2021 1. Introduction Fuente et al. (2020) and remains growing (e.g. Haug et al. 2019a, 2019b, 2020a, 2020b). The only two Neuroptera, the group of lacewings, is character- occurrences from compression fossils include a ised by a rather unique type of larva. Lacewing single specimen from the Cretaceous Crato for- INTRODUCTION larvae are generally fierce ambush predators. mation (Martins-Neto et al. 2007; Herrera-Flórez While adult lacewings resemble certain butterflies et al. 2020, including a discussion about another or dragonflies, lacewing larvae are quite special in presumed specimen) and a single specimen from their morphology. Their mouthparts are strongly the Eocene Green River formation (Dayvault et al. forward directed (prognath) and very prominent. 1995). Each mandible, or upper jaw, is interlocked with Ambers that have so far yielded lacewing larvae one of the maxillae, lower jaws, to form a stylet, a have provided quite a number of specimens, espe- combined venom-injecting and sucking tool. This cially Cretaceous ambers: at least 11 specimens pair of stylets can reach a large size and represent originated from Cretaceous Lebanese amber up to one third of the entire length of the larva (Whalley 1980; Pérez-de la Fuente et al. 2018, in some lacewing ingroups. The maxillae lack 2019), four specimens from Cretaceous Spanish distal parts, or palps. The labium, or lower lip, amber (Pérez-de la Fuente 2012, 2016, 2020), is deeply integrated into the head capsule and is three specimens from Cretaceous French amber mainly recognisable by its distal parts, the palps (Perrichot 2003; Perrichot et al. 2010; Wang et al. (MacLeod 1964; Gepp 1984; Aspöck and Aspöck 2016), a single specimen from Cretaceous Cana- 1999, 2007). dian amber (Engel and Grimaldi 2008) and a single The most prominent examples of lacewing specimen from New Jersey amber (Grimaldi 2000, larvae are possibly those of antlions, which catch p. 284, fig. 3). Amber from Myanmar (also known their prey in many cases with trap funnels bur- as “Burmese amber”; for a recent discussion of the rowed in sandy ground (Badano 2012). Another political background of this amber, see Haug et al. well-known fraction of lacewing larvae is that of 2020c) yielded an astonishing amount of lacewing aphid lions, the larvae of the groups Chrysopidae larvae (Xia et al. 2015; Wang et al. 2016; Liu et al (green lacewings) and Hemerobiidae (brown lace- 2016, 2018; Wichard 2017; Zhang 2017; Badano wings). They are, as the name suggests, specialised et al. 2018; Lu et al. 2018; Makarkin 2018; Haug et on feeding aphids (e.g. Tauber et al. 2003). As the al. 2018, 2019a,2019b,2019c, 2020a, 2020b). latter are severe plant pests, aphid lions have been Especially these findings from the Cretaceous successfully introduced as pest control some years indicate an enormous diversity of larval forms of ago, also due to their effectiveness and ecological lacewings more than 100 million years ago com- benefits (Tauberet al. 2000). pared to the modern fauna. On that regard, it is Neuroptera is generally considered an ear- interesting to compare the diversity of larval forms ly-appearing lineage of Holometabola (Grimaldi with the younger ambers to better understand and Engel 2005, p. 335), the ingroup of Insecta when the diversity of lacewing larvae could have with immature forms addressed as ‘larvae’ due to declined. However, comparisons must be treated their highly specialised eyes (for more information, carefully, since ecological bias has to be taken into see discussion in Haug 2020). As compression account (Solórzano-Kraemer et al. 2015, 2018). fossils, lacewings have been commonly found as The resin-producing trees during the Cretaceous adults, whose wings seem to have a high potential and Eocene are representatives of the group of preservation (e.g. Makarkin 2018). The fossil Coniferopsida (“gymnosperms”), while during the record of lacewing larvae is much scarcer and is Miocene resin-producing trees are representatives largely restricted to fossil resins. The amber fossil of the group Magnoliopsida (“angiosperms”; record was recently summarised by Pérez-de la Seyfullah et al. 2018), thus faunal assemblages Miocene lacewing larvae including specimen in Mexican amber http://dx.doi.org/10.18268/BSGM2021v73n3a271220 Boletín de la Sociedad Geológica Mexicana / 73 (3) / A271220/ 2021 / 3 inhabiting the trees could be different (e.g. Peris et report a new specimen from Miocene Mexican al. 2017a). amber from the Estrella de Belén deposit. Eocene Baltic amber has provided quite a number of larval lacewing specimens (Weidner 1958; MacLeod 1970; Larsson 1978; Janzen 2002; 2. Material and Methods INTRODUCTION / MATERIAL AND METHODS