The oldest giant lacewings (Neuroptera: Kalligrammatidae) from the Lower Jurassic of Germany
Introduction
The Mesozoic Kalligrammatidae are the most attractive insects in the order Neuroptera. The family contains the largest species within the order, with forewings up to 160 mm long (Bechly and Makarkin, 2016). Most are characterized by conspicuous eye-spots on the fore- and hind wings, and mouthparts with a long proboscis, by which they superficially resemble tropical butterflies (Labandeira et al., 2016).
The first species of Kalligrammatidae was described by Walther (1904) from the famous Upper Jurassic (Tithonian) Solnhofen Lithographic Limestone, i.e., Kalligramma haeckeli. Handlirsch published the first reconstruction of its venation (Handlirsch, 1906, pl. 48, fig. 5) and of the complete insect (Handlirsch, 1911, pl. 10, fig. 17), often reproduced in paleontology and entomology textbooks (e.g., Abel, 1920, fig. 158; Martynova, 1962, fig. 865). Since then, knowledge of fossil kalligrammatids has considerably expanded, especially in the last two decades. The family currently comprises 49 species belonging to 17 genera from the Middle/lowermost Upper Jurassic of China (Daohugou and Yujiagou), the Upper Jurassic of Germany (Solnhofen) and Kazakhstan (Karatau), the Lower Cretaceous of England (Tonbridge and Teffont Evias), China (Yixian Formation), Mongolia (Anda-Khuduk), Russia (Transbaikalia: Baissa), and Brazil (Crato Formation) (Handlirsch, 1906–1908, 1919; Martynova, 1947; Panfilov, 1968; Martins-Neto, 1992, 1997; Ponomarenko, 1992; Lambkin, 1994; Ren and Guo, 1996; Jarzembowski, 2001; Ren and Oswald, 2002; Ren, 2003; Zhang, 2003; Zhang and Zhang, 2003; Engel, 2005; Makarkin et al., 2009; Yang et al., 2009, 2011, 2014a, 2014b; Makarkin, 2010; Jepson et al., 2012; Liu et al., 2014, 2015; Bechly and Makarkin, 2016).
The Cretanallachiinae Makarkin, 2017 (including Burmogramma Liu et al., 2018) from the lowermost Cenomanian Burmese amber was considered a subfamily of Kalligrammatidae by Liu et al. (2018), primarily as each wing of Burmogramma bears an eye-spot like type 1 of Kalligrammatidae (see Labandeira et al., 2016) and the forewing MP (for abbreviations see “2. Material and methods”) is branched as in many Kalligrammatidae, i.e., the posterior trace of MP bears several anteriorly-directed branches. However, similar constructions of eye-spots and MP branching are also present in several undescribed ithonid-like species from the Jurassic of Daohugou (VM, pers. obs.). Other characters of Burmogramma (venation of both the fore- and hind wings, mouthparts, male and female genitalia) do not principally differ from those of other genera of Cretanallachiinae, suggesting that this subfamily is monophyletic. Their wings are characterized in particular by such symplesiomorphies at the order level as the presence of the distal nygma in both fore- and hind wings, and the long, sinuous basal crossvein between R and M in the hind wing, not detected in any Kalligrammatidae. Therefore, we cannot consider Cretanallachiinae to be members of the Kalligrammatidae based on these character states, which might be homoplasious.
The hypothesis that kalligrammatids were specialized pollinators feeding on pollen of Bennettitales and cycads was first proposed by Zherikhin (1978, 1980) and Rasnitsyn (1980) based on the mouthparts of Meioneurites spectabilis Engel, 2005 from the Upper Jurassic of Karatau (Kazakhstan). Since then, many long-proboscid species of Kalligrammatidae have been described. They are now generally thought to have fed on the pollen and secretions of the reproductive organs of extinct gymnosperms (mostly Bennettitales) (Labandeira, 2010; Labandeira et al., 2016).
Together with other Mesozoic long-proboscid insects (such as Mecoptera and Diptera) kalligrammatids not only reveal the high diversity of Mesozoic pollinating insects, but also highlight the diversity of Mesozoic pollinator-dependent plants prior to the dominance of angiosperms (Liu et al., 2018).
Kalligrammatidae in the Lower Jurassic was first reported by Makarkin et al. (2009) based on a photograph of specimen BSPG 2009/XIX, 11 in Tischlinger (1992) from the lower Toarcian Posidonia Shale of Kerkhofen (Oberpfalz, Bavaria). This specimen was acquired in 2009 by Bayerische Staatssammlung für Geologie and Paläontologie (Munich) as part of the private collection of Winfried Hartwig (Nuremberg) (Nose, 2010).
Another specimen of the same species was subsequently found by Michael Klopschar in the former marl pit of Schandelah near Braunschweig, who reported it as Kalligramma sp. (Klopschar, 2006, fig. 19.19).
These two specimens are described here as belonging to Liassopsychops curvatus Bode, 1953, a neuropteran species which was based on a distal wing fragment from the lower Toarcian of Grassel near Braunschweig. A second kalligrammatid species is described here, based on a fragmentary wing from the Schandelah marl pit, also made available to us by Michael Klopschar.
Section snippets
Material and methods
The paper is based on three specimens of Kalligrammatidae from two Lower Jurassic (lower Toarcian) localities of Germany, Schandelah and Kerkhofen (see details below in sections 3.1 and 3.2).
We follow the venational terminology of Breitkreuz et al. (2017). Terminology of wing spaces and details of venation (e.g., traces, veinlets, ORB) follows Oswald (1993).
Venational abbreviations: A1–A3, first to third anal veins; Cu, cubitus; CuA, cubitus anterior; CuP, cubitus posterior; hv, humeral veinlet
Lower Toarcian localities
Fossil insects of the Lower Jurassic of middle and western Europe are abundant in a brief interval at the beginning of the lower Toarcian (ca. 182 Ma). These assemblages are unique for this time. They were deposited in black shales during the Toarcian Oceanic Anoxic Event of the European epicontinental sea (Posidonia Shale) (Jenkyns, 1988). The fossils are extraordinarily well preserved in fine-grained micritic calcareous nodules or layers formed in anoxic, calm water without bioturbation of
Systematic palaeontology
Class Insecta Linnaeus, 1758
Order Neuroptera Linnaeus, 1758
Family Kalligrammatidae Handlirsch, 1906–1908 [1906]
Subfamily Liassopsychopinae n. subfam.
Etymology: From Liassopsychops, a genus-group name. The genitive case of the Greek noun ops is op-os, and its stem is op, and therefore the correct spelling is Liassopsychopinae (not Liassopsychopsinae).
Diagnosis: Forewing with recurrent, branched humeral veinlet; Sc, RA distally not fused; MP, CuA, CuP, A1 are dichotomously branched; eye-spot well
Paleoclimatic conditions in the early Toarcian
It is now well established that the climate in the Early Jurassic dramatically changed from temperate to hot greenhouse (even super-greenhouse) during the Pliensbachian–Toarcian interval (e.g., Hermoso et al., 2013; Hesselbo et al., 2013; Slater et al., 2019; van de Schootbrugge et al., 2019). The early Toarcian is the warmest interval of the Jurassic Period, with an abrupt rise in global temperatures of up to ∼7°C in mid-latitudes at the onset of the Toarcian Oceanic Anoxic Event (Xu et al.,
Acknowledgements
We thank the following people for various help: Michael Klopschar (Braunschweig) for allowing us to study the Schandelah specimens and providing additional information on the Schandelah and Hondelage sections; Mike Reich (BSPG) for allowing us to study the Kerkhofen specimen; Jürgen Vespermann (Roemer-Paeliceus Museum Hildesheim), Carmen Heunisch (Bundesanstalt für Geowissenschaften und Rohstoffe, Hannover) and Günther Schweigert (Staatliches Museum für Naturkunde, Stuttgart) for photographs of
References (114)
- et al.
A new gigantic lacewing species (Insecta: Neuroptera) from the Early Cretaceous of Brazil confirms the occurrence of Kalligrammatidae in the Americas
Cretaceous Research
(2016) - et al.
Lacewings (Insecta: Neuroptera) from the Lower Cretaceous Purbeck Limestone Group of southern England
Cretaceous Research
(2012) - et al.
Palynological evidence of an Early Cretaceous age for the Yixian Formation at Sihetun, western Liaoning
China. Cretaceous Research
(2007) - et al.
High niche diversity in Mesozoic pollinating lacewings
Nature Communication
(2018) New taxa of unusual Dilaridae (Neuroptera) with siphonate mouthparts from the mid-Cretaceous Burmese amber
Cretaceous Research
(2017)- et al.
A comparative overview of the neuropteran assemblage of the Early Cretaceous Yixian Formation (China), with description of a new genus of Psychopsidae (Insecta: Neuroptera)
Cretaceous Research
(2012) - et al.
Polar record of Early Jurassic massive carbon injection
Earth and Planetary Science Letters
(2011) Lehrbuch der Paläozoologie
(1920)Silky lacewings (Neuroptera: Psychopsidae) from the Eocene–Paleocene transition of Denmark with a review of the fossil record and comments on phylogeny and zoogeography
Insect Systematics & Evolution
(2001)Insekten aus dem oberen Lias von Grimmen (Vorpommern, Norddeutschland)
Neue Paläontologische Abhandlungen
(1996)
Revision of the “Trichoptera” described by Geinitz and Handlirsch from the Lower Toarcian of Dobbertin (Germany) based on new material
Insects from the Lower Toarcian of Middle Europe and England
Komplette Libelle Sphenophlebia pommerana Ansorge, 1996
Fossilien (Journal für Erdgeschichte)
Über Insektenfunde beim Kanalbau
Fossilien
Pterosaur dietary hypotheses: a review of ideas and approaches
Biological Reviews
Orthoptera und Neuroptera aus dem oberen Lias von Braunschweig
Jahrbuch der Preussischen Geologischen Landesanstalt
Die Insektenfauna des Ostniedersächsischen Oberen Lias
Palaeontographica: Beiträge zur Naturgeschichte der Vorwelt (A)
Makrofossilführung der “Siemensi-Geoden” (Mittlerer Lias Epsilon, Unteres Toarcium) von Kerkhofen/Oberpfalz (Bayern): Neue Insekten- und Pflanzenfunde
Geologische Blätter für Nordost-Bayern und Angrenzende Gebiete
Wing tracheation in Chrysopidae and other Neuropterida (Insecta): A resolution of the confusion about vein fusion
American Museum Novitates
A plesiosaur from the Lower Jurassic and the taxonomic position of the Sphenodontida
Palaeontographica (A)
Superclass Hexapoda
The Pliensbachian–Toarcian (Early Jurassic) extinction: A North American perspective
Ueber geognostischen Verhältnisse der Umgegend von Dörnten nördlich Goslar, mit besonderer Berücksichtigung der Fauna des oberen Lias
Atlas zu den Abhandlungen zur Geologischen Specialkarte von Preussen und den Thüringischen Staaten. Band 8, Heft 2
A remarkable kalligrammatid lacewing from the Upper Jurassic of Kazakhstan (Neuroptera: Kalligrammatidae)
Transactions of the Kansas Academy of Science
Diverse Neuropterida in Cretaceous amber, with particular reference to the paleofauna of Myanmar (Insecta)
Nova Supplementa Entomologica
Assemblages of small tetrapods from the Early Jurassic of Britain
Middle Jurassic microvertebrate assemblages from the British Isles
Phylogeny and classification of the Stenophlebioptera (Odonata: Epiproctophora)
Annales de la Société Entomologique de France
Die fossilen Insekten und die Phylogenie der rezenten Formen
Ein Handbuch für Palaeontologen und Zoologen
Ueber fossile Insekten
Eine neue Kalligrammide (Neuroptera) aus dem Solnhofener Plattenkalke
Senckenbergiana
Neue Untersuchungen über die fossilen Insekten mit Ergänzungen und Nachträgen sowie Ausblicken auf phylogenetische, palaeogeographische und allgemein biologische Probleme. II. Teil
Annalen des Naturhistorischen Museums in Wien
Black shale deposition during Toarcian super-greenhouse driven by sea level
Climate of the Past
Mochras borehole revisited: a new global standard for Early Jurassic earth history
Scientific Drilling
Das schwarze Jurameer und seine unsichtbaren Bewohner
Insekten und andere Gliederfüßer
Early Cretaceous insects from southern England
Modern Geology
A new Wealden fossil lacewing
The early Toarcian (Jurassic) anoxic event; stratigraphic, sedimentary and geochemical evidence
American Journal Science
Ein Koniferenrest aus dem Posidonienschiefer des Unteren Jura (Schwarzer Jura ε, Unter-Toarcium) von Süddeutschland
Stuttgarter Beiträge zur Naturkunde (B)
Two new lacewings (Neuroptera) from the Upper Jurassic locality of Shar-Teg (Mongolia)
Paleontologicheskii Zhurnal
Jurassic Neuroptera (Insecta) of Central Asia
Jurassic lacewings (Insecta: Neuroptera) of Western Siberia
Paleontologicheskii Zhurnal
Microanatomy and life history in Palaeopleurosaurus (Rhynchocephalia: Pleurosauridae) from the Early Jurassic of Germany
Science of Nature (Naturwissenschaften)
EP 19: Hondelage: Grube im Posidonienschiefer (Lias epsilon)
Mecoptera and Diptera from the early Toarcian (Early Jurassic) deposits of Wolfsburg–Große Kley (Lower Saxony, Germany)
Earth and Environmental Science Transactions of the Royal Society of Edinburgh
Über die Höckerfazies im Lias ε Nordbayerns
Fortschritte der Geologie und Paläontologie
The pollination of Mid Mesozoic seed plants and the early history of long-proboscid insects
Annals of the Missouri Botanical Garden
The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies
Proceedings of the Royal Society (B)
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