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• W2801583634 abstract "•A new lacewing species is described from the Cretaceous based on larvae•Distinctive foliate lobes are present on the thorax and abdomen of these larvae•Such morphological modifications grossly match coeval liverworts•The new larvae are the first example of direct mimicry in lacewing larvae Camouflage and mimicry are staples among predator-prey interactions, and evolutionary novelties in behavior, anatomy, and physiology that permit such mimesis are rife throughout the biological world [1Wickler W. Mimicry in plants and animals. McGraw-Hill, New York1968Google Scholar, 2Pasteur G. A classificatory review of mimicry systems.Annu. Rev. Ecol. Syst. 1982; 13: 169-199Crossref Scopus (164) Google Scholar]. These specializations allow for prey to better evade capture or permit predators to more easily approach their prey, or in some cases, the mimesis can serve both purposes. Despite the importance of mimesis and camouflage in predator-avoidance or hunting strategies, the long-term history of these traits is often obscured by an insufficient fossil record. Here, we report the discovery of Upper Cretaceous (approximately 100 million years old) green lacewing larvae (Chrysopoidea), preserved in amber from northern Myanmar, anatomically modified to mimic coeval liverworts. Chrysopidae are a diverse lineage of lacewings whose larvae usually camouflage themselves with a uniquely constructed packet of exogenous debris, conveying greater stealth upon them as they hunt prey such as aphids as well as evade their own predators [3Grimaldi D.A. Engel M.S. Evolution of the Insects. Cambridge University Press, Cambridge2005Google Scholar, 4Canard M. Semeria Y. New T.R. Biology of Chrysopidae. Junk Publishers, The Hague1984Google Scholar]. However, no lacewing larvae today mimic their surroundings. While the anatomy of Phyllochrysa huangi gen. et sp. nov. allowed it to avoid detection, the lack of setae or other anatomical elements for entangling debris as camouflage means its sole defense was its mimicry, and it could have been a stealthy hunter like living and other fossil Chrysopoidea or been an ambush predator aided by its disguise. The present fossils demonstrate a hitherto unknown life-history strategy among these “wolf in sheep’s clothing” predators, one that apparently evolved from a camouflaging ancestor but did not persist within the lineage. Camouflage and mimicry are staples among predator-prey interactions, and evolutionary novelties in behavior, anatomy, and physiology that permit such mimesis are rife throughout the biological world [1Wickler W. Mimicry in plants and animals. McGraw-Hill, New York1968Google Scholar, 2Pasteur G. A classificatory review of mimicry systems.Annu. Rev. Ecol. Syst. 1982; 13: 169-199Crossref Scopus (164) Google Scholar]. These specializations allow for prey to better evade capture or permit predators to more easily approach their prey, or in some cases, the mimesis can serve both purposes. Despite the importance of mimesis and camouflage in predator-avoidance or hunting strategies, the long-term history of these traits is often obscured by an insufficient fossil record. Here, we report the discovery of Upper Cretaceous (approximately 100 million years old) green lacewing larvae (Chrysopoidea), preserved in amber from northern Myanmar, anatomically modified to mimic coeval liverworts. Chrysopidae are a diverse lineage of lacewings whose larvae usually camouflage themselves with a uniquely constructed packet of exogenous debris, conveying greater stealth upon them as they hunt prey such as aphids as well as evade their own predators [3Grimaldi D.A. Engel M.S. Evolution of the Insects. Cambridge University Press, Cambridge2005Google Scholar, 4Canard M. Semeria Y. New T.R. Biology of Chrysopidae. Junk Publishers, The Hague1984Google Scholar]. However, no lacewing larvae today mimic their surroundings. While the anatomy of Phyllochrysa huangi gen. et sp. nov. allowed it to avoid detection, the lack of setae or other anatomical elements for entangling debris as camouflage means its sole defense was its mimicry, and it could have been a stealthy hunter like living and other fossil Chrysopoidea or been an ambush predator aided by its disguise. The present fossils demonstrate a hitherto unknown life-history strategy among these “wolf in sheep’s clothing” predators, one that apparently evolved from a camouflaging ancestor but did not persist within the lineage. Camouflage and mimicry are pervasive throughout the biological world as part of the usual interactions between predators and their prey, allowing both to avoid detection [1Wickler W. Mimicry in plants and animals. McGraw-Hill, New York1968Google Scholar, 2Pasteur G. A classificatory review of mimicry systems.Annu. Rev. Ecol. Syst. 1982; 13: 169-199Crossref Scopus (164) Google Scholar]. Among insects, the icons of mimicry include familiar stick and leaf insects, leaf-like moths or katydids, or the varied convergent patterns of coloration in the Müllerian and Batesian complexes of species made famous by heliconiine and other butterflies [3Grimaldi D.A. Engel M.S. Evolution of the Insects. Cambridge University Press, Cambridge2005Google Scholar]. More simplified means of camouflage are precursors to more elaborate forms of mimicry and often evolve in the form of cryptic patterns of coloration but at their most intricate include the active construction of concealment. One of the more remarkable examples of such strategies are the larvae of green lacewings (Neuroptera: Chrysopidae), who collect exogenous materials from their environment and use them to construct a packet of debris that is characteristically placed on the dorsal surface of the body to form a protective covering [4Canard M. Semeria Y. New T.R. Biology of Chrysopidae. Junk Publishers, The Hague1984Google Scholar, 5Tauber C.A. Tauber M.J. Albuquerque G.S. Debris-carrying in larval Chrysopidae: unraveling its evolutionary history.Ann. Entomol. Soc. Am. 2014; 107: 295-314Crossref Scopus (52) Google Scholar]. The debris packet is held in place by a series of tubercles beset with frequently elongate and sometimes modified setae, and the entanglement of the material amid these hair-like structures serves to keep affix the covering to the body [5Tauber C.A. Tauber M.J. Albuquerque G.S. Debris-carrying in larval Chrysopidae: unraveling its evolutionary history.Ann. Entomol. Soc. Am. 2014; 107: 295-314Crossref Scopus (52) Google Scholar]. The larva actively collects material from its environment, which it adds to the back of the body by arching its flexible head backward to gradually push more and more material into the growing packet of debris [6Jones D.T. Further notes on the snail-collecting aphis-lion larva (Neuroptera: Chrysopidae).Entomol. News. 1941; 52: 39-44Google Scholar, 7Eisner T. Silberglied R.E. A chrysopid larva that cloaks itself in mealybug wax.Psyche (Stuttg.). 1988; 95: 15-19Crossref Scopus (26) Google Scholar]. The material collected is not random, and different species will seek different materials, ranging from plant fragments to the carcasses of their prey [8Eisner T. Hicks K. Eisner M. Robson D.S. “Wolf-in-sheep’s-clothing” strategy of a predacious insect larva.Science. 1978; 199: 790-794Crossref PubMed Scopus (116) Google Scholar]. Lacewing larvae are efficient predators of small, soft-bodied arthropods such as aphids, hence the fact that they are sometimes referred to as aphis-lions or aphis-wolves and are often employed in biological control programs to control populations of agricultural and horticultural pests [4Canard M. Semeria Y. New T.R. Biology of Chrysopidae. Junk Publishers, The Hague1984Google Scholar, 9McEwan P.K. New T.R. Whittington A.E. Lacewings in the Crop Environment. Cambridge University Press, Cambridge2001Crossref Google Scholar]. In fact, the debris packet used by lacewing larvae not only protects the larva from its own predators but allows for them to minimize detection when approaching their own prey in what has been dubbed a “wolf in sheep’s clothing” strategy of active hunting [8Eisner T. Hicks K. Eisner M. Robson D.S. “Wolf-in-sheep’s-clothing” strategy of a predacious insect larva.Science. 1978; 199: 790-794Crossref PubMed Scopus (116) Google Scholar]. The use of debris as a form of camouflage is ancient among green lacewings, having arisen once and apparently been lost relatively few times [10Haruyama N. Mochizuki A. Duelli P. Naka H. Nomura M. Green Lacewing phylogeny, based on three nuclear genes (Chrysopidae, Neuroptera).Syst. Entomol. 2008; 33: 275-288Crossref Scopus (31) Google Scholar, 11Dai Y.T. Winterton S.L. Garzón-Orduña I.J. Liang F.Y. Liu X.Y. Mitochondrial phylogenomic analysis resolves the subfamily placement of enigmatic green lacewing genus Nothancyla (Neuroptera: Chrysopidae).Aust. Entomol. 2017; 56: 322-331Crossref Scopus (11) Google Scholar, 12Jiang Y. Garzón-Orduña I.J. Winterton S.L. Yang F. Liu X. Phylogenetic relationships among tribes of the green lacewing subfamily Chrysopinae recovered based on mitochondrial phylogenomics.Sci. Rep. 2017; 7: 7218Crossref PubMed Scopus (11) Google Scholar]. In fact, such behavior is well known among the stem group of Chrysopidae, demonstrating that camouflage extends deep into the lineage’s history [13Pérez-de la Fuente R. Delclòs X. Peñalver E. Speranza M. Wierzchos J. Ascaso C. Engel M.S. Early evolution and ecology of camouflage in insects.Proc. Natl. Acad. Sci. USA. 2012; 109: 21414-21419Crossref PubMed Scopus (78) Google Scholar, 14Wang B. Xia F. Engel M.S. Perrichot V. Shi G. Zhang H. Chen J. Jarzembowski E.A. Wappler T. Rust J. Debris-carrying camouflage among diverse lineages of Cretaceous insects.Sci. Adv. 2016; 2: e1501918Crossref PubMed Scopus (80) Google Scholar]. Fossil larvae of stem-group chrysopoids preserved with their covering of debris are known from as far back as the Early Cretaceous [13Pérez-de la Fuente R. Delclòs X. Peñalver E. Speranza M. Wierzchos J. Ascaso C. Engel M.S. Early evolution and ecology of camouflage in insects.Proc. Natl. Acad. Sci. USA. 2012; 109: 21414-21419Crossref PubMed Scopus (78) Google Scholar, 14Wang B. Xia F. Engel M.S. Perrichot V. Shi G. Zhang H. Chen J. Jarzembowski E.A. Wappler T. Rust J. Debris-carrying camouflage among diverse lineages of Cretaceous insects.Sci. Adv. 2016; 2: e1501918Crossref PubMed Scopus (80) Google Scholar], and Chrysopoidea as a whole is known from fossils as early as the mid-Jurassic [3Grimaldi D.A. Engel M.S. Evolution of the Insects. Cambridge University Press, Cambridge2005Google Scholar, 15Nel A. Delclòs X. Hutin A. Mesozoic chrysopid-like Planipennia: a phylogenetic approach (Insecta: Neuroptera).Ann. Soc. Entomol. Fr. (N.S.). 2005; 41: 29-68Crossref Scopus (55) Google Scholar, 16Engel M.S. Winterton S.L. Breitkreuz L.C.V. Phylogeny and evolution of Neuropterida: where have wings of lace taken us?.Annu. Rev. Entomol. 2018; 63: 531-551Crossref PubMed Scopus (77) Google Scholar]. It is abundantly clear that camouflage has been an important element in the life history of Chrysopoidea and one of the many factors that has likely helped fuel their successful diversification over the last 165 million years [5Tauber C.A. Tauber M.J. Albuquerque G.S. Debris-carrying in larval Chrysopidae: unraveling its evolutionary history.Ann. Entomol. Soc. Am. 2014; 107: 295-314Crossref Scopus (52) Google Scholar]. In fact, Chrysopidae are today one of the most diverse lineages of Neuroptera, with nearly 1,500 species [16Engel M.S. Winterton S.L. Breitkreuz L.C.V. Phylogeny and evolution of Neuropterida: where have wings of lace taken us?.Annu. Rev. Entomol. 2018; 63: 531-551Crossref PubMed Scopus (77) Google Scholar, 17Brooks S.J. Barnard P.C. The green lacewings of the world: a generic review (Neuroptera: Chrysopidae).Bull. Br. Mus. Nat. Hist. Entomol. 1990; 59: 117-286Google Scholar], second only to the antlions whose own larvae have evolved dramatic means to avoid detection during prey capture (albeit as ambush predators rather than active hunting) by way of their characteristic funnel pits [3Grimaldi D.A. Engel M.S. Evolution of the Insects. Cambridge University Press, Cambridge2005Google Scholar, 16Engel M.S. Winterton S.L. Breitkreuz L.C.V. Phylogeny and evolution of Neuropterida: where have wings of lace taken us?.Annu. Rev. Entomol. 2018; 63: 531-551Crossref PubMed Scopus (77) Google Scholar]. Interestingly, the earliest of known chrysopoid larvae with debris packets and morphological specializations for carrying such materials demonstrate a once-greater range of anatomical variation for the lineage [13Pérez-de la Fuente R. Delclòs X. Peñalver E. Speranza M. Wierzchos J. Ascaso C. Engel M.S. Early evolution and ecology of camouflage in insects.Proc. Natl. Acad. Sci. USA. 2012; 109: 21414-21419Crossref PubMed Scopus (78) Google Scholar, 14Wang B. Xia F. Engel M.S. Perrichot V. Shi G. Zhang H. Chen J. Jarzembowski E.A. Wappler T. Rust J. Debris-carrying camouflage among diverse lineages of Cretaceous insects.Sci. Adv. 2016; 2: e1501918Crossref PubMed Scopus (80) Google Scholar]. The tubercles of the larvae are often greatly elongate, and the setae are capitate or exhibit other modifications for entanglement [13Pérez-de la Fuente R. Delclòs X. Peñalver E. Speranza M. Wierzchos J. Ascaso C. Engel M.S. Early evolution and ecology of camouflage in insects.Proc. Natl. Acad. Sci. USA. 2012; 109: 21414-21419Crossref PubMed Scopus (78) Google Scholar], extending the total range of morphological variety between living and fossil green lacewings and demonstrating that larvae of crown-group Chrysopidae are less disparate. Nonetheless, they all fall well within the “camouflage strategy” for evading detection rather than outright mimicry, whereby the body itself is modified to assume the character of the environment (either in terms of shape, coloration, or chemistry). A recent discovery of lacewing larvae from the Cenomanian of southern Asia demonstrates for the first time that Chrysopoidea once included also mimetic species. A series of larvae preserved in amber from northern Myanmar are morphologically modified such that the dorsal protuberances of the body, rather than specialized for the entanglement of debris, are instead modified into distinctive broad, foliate lobes, and themselves apically trilobed or bilobed, that greatly match coeval liverworts frequently as inclusions in the same deposits. These larvae are the first example of direct mimicry in lacewing larvae, an evolutionary innovation subsequently lost within the lineage. Order Neuroptera Linnaeus, 1758. Superfamily Chrysopoidea Schneider, 1851. Phyllochrysa huangi gen. et sp. nov. ZooBank LSID (Phyllochrysa): urn:lsid:zoobank.org:act:9855A5A3-95AD-4F9C-8FFA-228E14A5E9F2. ZooBank LSID (Phyllochrysa huangi): urn:lsid:zoobank.org:act:3B061C47-9883-419E-9D52-44B51DA51002. Holotype. Larva, NIGP167955; Upper Cretaceous (earliest Cenomanian) amber (ca. 100 million years ago), Tanai Township, Myitkyina, District of Kachin State, northern Myanmar; the amber piece preserves a complete larva of P. huangi; it is polished in the form of nearly semicircular, clear and transparent, with length × width about 16.0 × 14.0 mm, height about 6.6 mm. Paratype. Larva, LPAM BA17003; same locality and age as holotype; the amber piece preserves a complete larva of P. huangi; it is polished in the form of elliptical, clear and transparent, with length × width about 17.0 × 12.0 mm, height about 5.0 mm. The genus-group name is a combination of phyllon (Greek, meaning, “leaf”) and chrysos (Greek, meaning, “gold” and a common suffix for chrysopoid genera). The specific epithet honors Yiren Huang, who kindly donated the holotype for our study. Larva (Figure 1): Body flat, with extensively projected, foliate, lateral plates from prothorax to abdominal segment V. Body setation absent. Head capsule reduced, retracted under prothorax; mandible-maxilla complex (“jaws”) long, thin, feebly curved distally, without dentition; antenna long, terminal flagellomere distinctly expanded, with a narrow longitudinal groove (probably a sensory slit). Legs moderate in length and thickness, length among fore-, mid-, and hind legs nearly equal; a long, trumpet-shaped, pretarsal empodium present, pretarsal claw simple, slightly recurved, much shorter than empodium. Thorax dorsally with several rows of elevated ridges at middle; thoracic lateral plates much larger than abdominal lateral plates, each distally bearing a few small lobes. Abdomen slightly narrower than thorax, and segments I–V much wider and longer than segments VI–X, which lack lateral plates; abdominal lateral plates distally tapering. Refer to online Supplemental Information for the complete species description (Data S1) and additional figures (Figures S1 and S2). The general bauplan, together with the posteriorly gradually narrowed head (Figure S1A), the toothless, slightly curved jaws (Figures 1D and S1B), the numerous flagellomeres (Figure 1D), and the presence of trumpet-shaped empodia (Figures 1E and S1D), demonstrates that the larva belongs to the superfamily Chrysopoidea [4Canard M. Semeria Y. New T.R. Biology of Chrysopidae. Junk Publishers, The Hague1984Google Scholar], a group that includes crown-group green lacewings as well as numerous extinct stem groups that are either unassigned to family or that have at times in the past been recognized as their own families or subfamilies outright [15Nel A. Delclòs X. Hutin A. Mesozoic chrysopid-like Planipennia: a phylogenetic approach (Insecta: Neuroptera).Ann. Soc. Entomol. Fr. (N.S.). 2005; 41: 29-68Crossref Scopus (55) Google Scholar]. Since most of these higher groups are characterized on the basis of adults, particularly traits of wing venation [15Nel A. Delclòs X. Hutin A. Mesozoic chrysopid-like Planipennia: a phylogenetic approach (Insecta: Neuroptera).Ann. Soc. Entomol. Fr. (N.S.). 2005; 41: 29-68Crossref Scopus (55) Google Scholar], it is not presently possible to assign the fossil larvae to any one of these extinct subfamilies or families, and they are therefore left as incertae sedis among stem-group Chrysopoidea. The concept of this superfamily has shifted dramatically over the years, ranging from the inclusion of numerous families [15Nel A. Delclòs X. Hutin A. Mesozoic chrysopid-like Planipennia: a phylogenetic approach (Insecta: Neuroptera).Ann. Soc. Entomol. Fr. (N.S.). 2005; 41: 29-68Crossref Scopus (55) Google Scholar, 18Ren D. Makarkin V.N. Ascalochrysidae – a new lacewing family from the Mesozoic of China (Insecta: Neuroptera: Chrysopoidea).Cretac. Res. 2009; 30: 1217-1222Crossref Scopus (15) Google Scholar], many of which subsequently proved to render the group paraphyletic or even polyphyletic [19Yang Q. Makarkin V.N. Winterton S.L. Khramov A.V. Ren D. A remarkable new family of Jurassic insects (Neuroptera) with primitive wing venation and its phylogenetic position in Neuropterida.PLoS ONE. 2012; 7: e44762Crossref PubMed Scopus (90) Google Scholar]. Accordingly, many of these families have been removed to other groups, while others (e.g., Mesochrysopidae, Corydasialidae) have been demoted as subfamilies forming a grade of stem groups within the base of Chrysopidae [16Engel M.S. Winterton S.L. Breitkreuz L.C.V. Phylogeny and evolution of Neuropterida: where have wings of lace taken us?.Annu. Rev. Entomol. 2018; 63: 531-551Crossref PubMed Scopus (77) Google Scholar]. These are either united with hemerobiids when they have been considered sister groups, as superfamily Hemerobioidea [16Engel M.S. Winterton S.L. Breitkreuz L.C.V. Phylogeny and evolution of Neuropterida: where have wings of lace taken us?.Annu. Rev. Entomol. 2018; 63: 531-551Crossref PubMed Scopus (77) Google Scholar], or, if chrysopids are not sister to Hemerobiidae [20Winterton S.L. Lemmon A.R. Gillung J.P. Garzón-Orduña I.J. Badano D. Bakkes D.K. Breitkreuz L.C.V. Engel M.S. Moriarity Lemmon E. Liu X. et al.Evolution of lacewings and allied orders using anchored phylogenomics (Neuroptera, Megaloptera, Raphidioptera).Syst. Entomol. 2017; 43: 330-354Crossref Scopus (121) Google Scholar], then as their own superfamily, Chrysopoidea, with only a single family (as is adopted herein). Regardless, monophyly of Chrysopidae has been well supported [11Dai Y.T. Winterton S.L. Garzón-Orduña I.J. Liang F.Y. Liu X.Y. Mitochondrial phylogenomic analysis resolves the subfamily placement of enigmatic green lacewing genus Nothancyla (Neuroptera: Chrysopidae).Aust. Entomol. 2017; 56: 322-331Crossref Scopus (11) Google Scholar, 20Winterton S.L. Lemmon A.R. Gillung J.P. Garzón-Orduña I.J. Badano D. Bakkes D.K. Breitkreuz L.C.V. Engel M.S. Moriarity Lemmon E. Liu X. et al.Evolution of lacewings and allied orders using anchored phylogenomics (Neuroptera, Megaloptera, Raphidioptera).Syst. Entomol. 2017; 43: 330-354Crossref Scopus (121) Google Scholar], and certain apomorphic traits such as the trumpet-shaped empodia of larvae or the fusion of CuA (cubitus anterior) and MP (media posterior) (to form a pseudocubitus) in the forewing serve to unite various fossils with modern chrysopids. The new lacewing larvae possess a number of distinctive characters that may be informative to infer its familial affinity. The presence of the numerous flagellomeres and the trumpet-shaped empodia are shared with the larvae of crown-group Chrysopidae as well as Hallucinochrysa diogenesi, a camouflaging chrysopoid larva from the Lower Cretaceous (Albian) amber of Spain [13Pérez-de la Fuente R. Delclòs X. Peñalver E. Speranza M. Wierzchos J. Ascaso C. Engel M.S. Early evolution and ecology of camouflage in insects.Proc. Natl. Acad. Sci. USA. 2012; 109: 21414-21419Crossref PubMed Scopus (78) Google Scholar]. In Hemerobiidae (brown lacewings), a family long considered as sister to Chrysopoidea (but see recent phylogenomic results, which cast some doubt on this conclusion [20Winterton S.L. Lemmon A.R. Gillung J.P. Garzón-Orduña I.J. Badano D. Bakkes D.K. Breitkreuz L.C.V. Engel M.S. Moriarity Lemmon E. Liu X. et al.Evolution of lacewings and allied orders using anchored phylogenomics (Neuroptera, Megaloptera, Raphidioptera).Syst. Entomol. 2017; 43: 330-354Crossref Scopus (121) Google Scholar]), the larvae lack such flagella and only possess trumpet-shaped empodia during the first instar [21New T.R. The immature stages of Drepanacra binocula (Neuroptera: Hemerobiidae), with notes on the relationships of the genus.J. Aust. Entomol. Soc. 1975; 14: 247-250Crossref Scopus (8) Google Scholar, 22Miller G.L. Lambdin P.L. Redescription of the larval stage of Hemerobius stigma Stephens (Neuroptera: Hemerobiidae).Fla. Entomol. 1984; 67: 377-382Crossref Google Scholar]. It is notable that many brown lacewing larvae have the head only partially retracted into the prothorax [21New T.R. The immature stages of Drepanacra binocula (Neuroptera: Hemerobiidae), with notes on the relationships of the genus.J. Aust. Entomol. Soc. 1975; 14: 247-250Crossref Scopus (8) Google Scholar, 23Makarkin V.N. Wedmann S. Weiterschan T. First record of a fossil larva of Hemerobiidae (Neuroptera) from Baltic amber.Zootaxa. 2012; 3417: 53-63Crossref Scopus (12) Google Scholar], while the larva of Phyllochrysa gen. nov. has the head completely retracted into the prothorax. This character state is likely autapomorphic in relation and the result of the extreme expansion of the thorax via the foliate lobes, with the superficial resemblance to the condition of brown lacewings being convergent and resulting from different morphological modifications. The immature stages of Mesozoic chrysopoids are poorly known. Hitherto, only two named species have their larval forms described, including Pedanoptera arachnophila from Upper Cretaceous (Cenomanian) amber of Myanmar and H. diogenesi from the Lower Cretaceous (Albian) amber of Spain [13Pérez-de la Fuente R. Delclòs X. Peñalver E. Speranza M. Wierzchos J. Ascaso C. Engel M.S. Early evolution and ecology of camouflage in insects.Proc. Natl. Acad. Sci. USA. 2012; 109: 21414-21419Crossref PubMed Scopus (78) Google Scholar, 24Liu X. Zhang W. Winterton S.L. Breitkreuz L.C.V. Engel M.S. Early morphological specialization for insect-spider associations in Mesozoic lacewings.Curr. Biol. 2016; 26: 1590-1594Abstract Full Text Full Text PDF PubMed Scopus (42) Google Scholar]. Similar to the present larvae, the larvae of these two species are also spectacular in morphology and paleobiology. The larva of P. arachnophila had extremely elongate legs and specialized pretarsi that are composed of serrate pretarsal claws and a reduced empodium, with such morphological specializations considered traits used for preying on spiders [24Liu X. Zhang W. Winterton S.L. Breitkreuz L.C.V. Engel M.S. Early morphological specialization for insect-spider associations in Mesozoic lacewings.Curr. Biol. 2016; 26: 1590-1594Abstract Full Text Full Text PDF PubMed Scopus (42) Google Scholar]. The larva of H. diogenesi bears specialized cuticular processes forming a dorsal basket that carries a dense debris packet for the purpose of camouflage [13Pérez-de la Fuente R. Delclòs X. Peñalver E. Speranza M. Wierzchos J. Ascaso C. Engel M.S. Early evolution and ecology of camouflage in insects.Proc. Natl. Acad. Sci. USA. 2012; 109: 21414-21419Crossref PubMed Scopus (78) Google Scholar, 25Pérez-de la Fuente R. Delclòs X. Peñalver E. Engel M.S. A defensive behavior and plant-insect interaction in Early Cretaceous amber--The case of the immature lacewing Hallucinochrysa diogenesi.Arthropod Struct. Dev. 2016; 45: 133-139Crossref PubMed Scopus (31) Google Scholar]. Unnamed larvae with similarly modified cuticular processes to H. diogenesi are also found in the Lower Cretaceous (Barremian) amber of Lebanon and the Upper Cretaceous (Cenomanian) amber of Myanmar [14Wang B. Xia F. Engel M.S. Perrichot V. Shi G. Zhang H. Chen J. Jarzembowski E.A. Wappler T. Rust J. Debris-carrying camouflage among diverse lineages of Cretaceous insects.Sci. Adv. 2016; 2: e1501918Crossref PubMed Scopus (80) Google Scholar]. The new larvae differ from the aforementioned species owing to the lack of those particular specializations, and instead exhibits its own peculiar and unique morphologies associated with a unique mode of life among chrysopoids. Considering the antennae, it is notable that the flagellum in all of these larval species is more or less swollen distally, although each differs in the particular details of such augmented morphology. By contrast, the flagellum in all known larvae of crown-group Chrysopidae tapers distally. Thus, the distally swollen larval flagellum could be a noteworthy character state to assign H. diogenesi and P. huangi together with P. arachnophila. The adult of P. arachnophila is known and indicates relationship to the former family Mesochrysopidae (now considered only a stem group to Chrysopidae: [16Engel M.S. Winterton S.L. Breitkreuz L.C.V. Phylogeny and evolution of Neuropterida: where have wings of lace taken us?.Annu. Rev. Entomol. 2018; 63: 531-551Crossref PubMed Scopus (77) Google Scholar]), suggesting that the former two species also belong in this grade. It remains uncertain whether such a trait is apomorphic (supporting mesochrysopines or a subset of them as monophyletic) or plesiomorphic (and that this group could be a paraphyletic grade to progressively more derived chrysopoids). Considerable phylogenetic work remains to be undertaken and with a greater swath of fossil diversity. The morphological specialization in the new chrysopoid larva is unique and is unknown among any living or fossil lacewings. The most striking feature refers to the development of the broadly foliate thoracic and abdominal lateral plates, which resemble the lobes of some liverworts (Marchantiophyta). There is a high diversity of liverworts in Burmese amber. Their stems range from 2 to 10 mm in length, and their leaves are orbicular-ovate, broadly obovate, or subtriangular, with the length ranging from 0.3 to 1.5 mm (Figure S2). Several specimens are approximately 6 mm long and 2 mm wide, and their leaves are on either side of the stem, subtriangular or ovate with pointed apex, 1 mm long and 0.5–1.0 mm wide (Figure 2). The larvae of P. huangi closely approximate the morphology of these liverworts in size and gross morphology, including the size and shape of leaves, strongly implicating mimicry of the liverwort by these chrysopoid larvae. As noted, plant mimicry is well known in insects, and foliate expansions such as those observed in P. huangi can be found in nymphs and adults of lineages as diverse as leaf insects (Phasmatodea: Phylliidae) or leaf-mimicking mantises (Mantodea: Empusidae, Mantidae) [3Grimaldi D.A. Engel M.S. Evolution of the Insects. Cambridge University Press, Cambridge2005Google Scholar]. However, fossil evidence of leaf mimesis among insects is scarce, particularly from the Mesozoic. Prior to the discovery of P. huangi, only two groups putatively exhibit such leaf mimicry, one of which is a group of lacewings among the stem group to lance lacewings (Osmylidae). Species of Bellinympha (Osmyloidea: “Saucrosmylidae”) were found to mimic gymnosperms of the Cycadales, Bennettitales, and Ginkgoales [26Wang Y. Liu Z. Wang X. Shih C. Zhao Y. Engel M.S. Ren D. Ancient pinnate leaf mimesis among lacewings.Proc. Natl. Acad. Sci. USA. 2010; 107: 16212-16215Crossref PubMed Scopus (77) Google Scholar, 27Wang Y. Labandeira C.C. Shih C. Ding Q. Wang C. Zhao Y. Ren D. Jurassic mimicry between a hangingfly and a ginkgo from China.Proc. Natl. Acad. Sci. USA. 2012; 109: 20514-20519Crossref PubMed Scopus (80) Google Scholar]. Nonetheless, the organ associated with mimetic function in these species is confined to the wings of the adults, largely the result of mimetic pigmentation on the wing" @default.
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