Thursday, January 24, 2013

Butterflies Before Butterflies, Flowers Before Flowers

Minute Tree Fungus Beetle - Strigocis opacicollis - male
Strigocis opacicollis (Ciidae), photographed by Tom Murray
Giving insects common names is a luxury a systematic entomologist can rarely afford: with ~1,200,000 species described, I think you can see why. Thanks to Linnaean binomial nomenclature, we can avoid the confusion inherent in non-scientific verbiage. It could be argued that it's difficult for a layman to memorize, for instance, the names of the beetle families Endomychidae, Erotylidae, Cryptophagidae, Leiodidae, Derodontidae, Mycetophagidae, Archeocrypticidae, Tetratomidae, and Ciidae; but these beetles' respective vernacular sobriquets are even more confusing (handsome fungus beetles, pleasing fungus beetles, silken fungus beetles, round fungus beetles, tooth-necked fungus beetles, hairy fungus beetles, cryptic fungus beetles, polypore fungus beetles, and minute tree-fungus beetles). And those are just the common names in English. With all that in mind, it's rather surprising that entomologists have recently gotten into the habit of giving entirely prehistoric insect taxa their own colloquial names.

Meganeura monyi, a famous crow-sized Carboniferous griffinfly
This all began with the members of the order Meganisoptera, which perished (along with so many other creatures) in the Permo-Triassic mass extinction 251 million years past (there is one debatable record from the Triassic; Anderson & Anderson, 1983). They were regarded for many years as a primitive lineage of the extant order Odonata (dragonflies and damselflies), which they matched in ecology if not in detail (Laurentiaux-Vieira et al., 1952). You are probably familiar with the bigger ones as the notorious "giant dragonflies" of the Carboniferous Period's torrid coal-forming swamps: the largest known meganisopteran (and, indeed, the most gigantic insect of all time)the 279-million-year-old Kansan Meganeuropsis permiana (Carpenter, 1939)had a 28-inch wingspan. However, later taxonomists agreed this lineage deserved a separate ordinal status (whilst still closely related; Bechly et al., 2001), and discarded the hitherto-used term "Protodonata" in favor of Meganisoptera (Riek & Kukalova-Peck, 1984); hence, calling the likes of Meganeuropsis "dragonflies" is inaccurate. Knowing that the Meganisoptera remain very much a fixture of the public imagination and would continue to be called "dragonflies" unless something were done, the new name "griffinfly" was proposed (Grimaldi & Engel, 2005), and it seems to have caught on among paleoentomologists (Prokop & Nel, 2010).

Reconstruction of Kalligramma haeckeli; antennae inaccurately long (Makarkin & Menon, 2007)
Whether giving entirely extinct organisms their own common names is advisable or not, I would like to propose one for a distinctive family of extinct lacewings: the Kalligrammatidae, restricted to Eurasia 164-132 m.y.a. (Makarkin et al., 2009; Makarkin, 2010; Zherikhin et al., 1999; Liu et al., 2006). ("Lacewing", as you may know from a previous post"Mantidflies: Chimeras of the Insect World"is a general term for any member of the order Neuroptera, unless otherwise specified.) Kalligrammatids were some of the largest neuropterans of all time, with broad, often colorful wings that suggest ponderously fluttering flight; some species bear eye-spots a lá modern silkmoths and butterflies (Grimaldi & Engel, 2005). One kalligrammatid even imitated contemporary cycads' leaves: the earliest evidence of plant mimicry (so common today) in insects (Wang et al., 2010).

Psychopsis mimica, photographed by Graeme V.
Now for some phylogenetic background: the Kalligrammatidae's closest modern kin are the Psychopsidae (silky lacewings), which are today restricted to tropical Asia, southern Africa and Meganesia*; they share their Mesozoic cousins' ridiculously dense wing venation (making students draw psychopsid wings by hand is a disciplinary action used by some stricter professors of entomology at graduate schools). Silky lacewings, kalligrammatids and some other extinct families (such as the diverse but little-known Osmylopsychopidae and Brongniartiellidae) are generally referred to as "psychopsoid" (Greek for "how many veins does one wing need?"), but—despite being sometimes classified as Psychopsoidea, or even as an independent suborder (Krivokhatsky, 1998)—are doubtlessly polyphyletic (Makarkin & Archibald, 2005). The fossil families' study is hampered by the fact that most are known only from wings. Yet, through all that, the silky lacewings and the Kalligrammatidae remain convincingly linked (Schlüter, 1986).

Photograph of Chasmoptera huttii (Nemopteridae) by Fred Hort
Before the recent massive revision of neuropteran taxonomy, silky lacewings (and their sibling taxa by extension) were regarded as akin to the Osmylidae, a family mainly noteworthy in that its members are cosmopolitan with the inexplicable exception of North America; but the "psychopsoids" (at least the ones whose anatomy we understand) have been reclassified in the suborder Myrmeleontiformia, the most famous members of which are the sand-dwelling, pit-making antlions (Myrmeleontidae). Among the myrmeleontiforms, the next of kin to Kalligrammatidae (after Psychopsidae) is apparently the beautiful Nemopteridae (spoon-winged lacewings), which have lengthy and ornately spatulate hind wings (as can be seen at right) (Aspöck et al., 2001).

But I digress from the purpose of this postthat of giving kalligrammatids a common name. As it so happens, these fluttering lacewings of the mid-Mesozoic Era deserved the epithet "butterflies of the Jurassic" for more reason than their appearance (butterflies would not appear until the Cenozoic Era): they (and the related Aetheogrammatidae; Yang et al., 2012) were, like their modern simulacra, solely nectar-feeders, as can be readily seen by the elongated form of their mouth-parts (Engel, 2005). Since kalligrammatids were superficially like butterflies—but not the real thing—I would propose calling them "margarineflies".

The fact that the so-called margarineflies were clear nectarivores is interesting, given that the flowering plants (Angiospermae) do not appear in the fossil record until 10 million years after these lacewings' evident extinction (Qiang et al., 2004). But, just as butterflies converged with their margarinefly antecedents, so some groups of plants that were not strictly "flowering" evolved reproductive organs of the same phenotype as flowers: namely, the Bennettitales (Speer, 2000), a taxon whose members disingenuously resemble cycads to such a degree that they are called cycadeoids. ("Bennettitaleans" is the taxonomically correct name for these plants, but "cycadeoid" is much easier to pronounce.) Cycadeoids were an integral part of our planet's flora for much of the Mesozoic Era, with the mass extinction that killed off the dinosaurs dealing the cycadeoids a hard blow: thereafter they lingered on only in Tasmania until the cooling of climate that came at the Paleogene Period's end (McLoughlin et al., 2011). 

Illustration of an arborescent cycadeoid; blossoms were located on the trunk
The nature of the bennettitalean/angiosperm relationship has been debated almost since paleobotany's beginning; but that controversy has no bearing on margarinefly ethology. Structurally speaking, cycadeoid "inflorescences" are drastically different from true flowers: but they were obviously entomophilous (pollinated by insects) just the same, and it's reasonable to deduce that margarineflies could have been some of the pollinators that assisted in cycadeoid reproduction.

Notice that I say "some": margarineflies were part of an entire ecological guild of pollinators that predated the great angiosperm radiation, a guild that was interdependent with the Bennettitales, Gnetophyta, and some Cycadophyta. Certain mid-Cretaceous gingkos were also entomophilous, being pollinated by thrips; this is in contrast to modern ginkgos, which are exclusively anemophilous (Peñalver et al., 2012). The margarineflies' compatriots in non-ginkgo gymosperm pollination were the extinct fly families Rhagionemestriidae and Archisargidae, as well as an entire suborder of scorpionflies (Aneuretopsychina). Tellingly, all of these taxa disappeared in the Early Cretaceous concurrently with the turnover from domination by gymnosperms to a floral regime of angiosperms (Labandeira, 2010): they were utter specialists. 

For ecologists, the co-radiation of pollinating insects and angiosperms is the archetype of co-evolution—but it was not anything new under the sun: other plants (and other insects) had already managed something very like to it.

*The continent comprised of Australia, Tasmania, New Guinea, and adjacent islands.
Not descended from a common ancestor that was itself a member of the group.

Anderson, J. M. and Anderson, H. M. (1983). Palaeoflora of Southern Africa Molteno Formation (Triassic). [H. Sims/H. Lindon/H. Lindon].

Aspöck, U.; Plant, J. D.; and Nemeschkal, H. L. (2001). Cladistic analysis of Neuroptera and their systematic position within Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera). Systematic Entomology, 26, 73-86.

Bechly, G.; Brauckmann, C.; Zessin, W.; and Gröning, E. (2001). New results concerning the morphology of the most ancient dragonflies (Insecta: Odonatoptera) from the Namurian of Hagen-Vorhalle (Germany). J. Zool. Syst. Evol. Res., 39(2001), 209-226.

Carpenter, F. M. (1939). The Lower Permian Insects of Kansas. Part 8: Additional Megasecoptera, Protodonata, Odonata, Homoptera, Psocoptera, Protelytroptera, Plectoptera and Protoperlaria. Proceedings of the American Academy of Arts and Sciences, 73(3), 29-70.

Engel, M. S. (2005). A remarkable kalligrammatid lacewing from the Upper Jurassic of Kazakhstan (Neuroptera: Kalligrammatidae). Transactions of the Kansas Academy of Science, 108, 59-62.

Grimaldi, D. and Engel, M. S. (2005). Evolution of the Insects. Cambridge: Cambridge University Press.

Krivokhatsky, V. A. (1998). Antlions (Neuroptera: Myrmeleontidae) of the Palaearctic Region (morphology, classification, zoogeography). Ph.D. dissertation. St. Petersburg: Zoological Institute, Russian Academy of Sciences. 

Labandeira, C. C. (2010). The Pollination of Mid-Mesozoic Seed Plants and the Early History of Long-proboscid Insects. Annals of the Missouri Botanical Garden, 97(4), 469-513.

Laurentiaux-Vieira, F.; Ricour, J.; and Laurentiaux, D. (1952). Un Protodonate du Trias de la Dent de Villard (Savoie). Bulletin de la Societe geologique de France, 6(2), 319-325.

Liu, Y.; Liu, Y.; Ji, S.; and Yang, Z. (2006). U-Pb zircon age for the Daohugou Biota at Ningcheng of Inner Mongolia and comments on related issues [electronic version]. Chinese Science Bulletin, 51(21), 2,634-2,664. Retrieved 1/22/13 from

Makarkin, V. N. (2010). New psychopsoid Neuroptera from the Early Cretaceous of Baissa, Transbaikalia. Annals of the Entomological Society of France, 46 (1-2), 254-261. 

Makarkin, V. N. and Archibald, S. B. (2005). Substitute names for three genera of fossil Neuroptera, with taxonomic notes. Zootaxa, 1054, 15–23.

Makarkin, V. N. and Menon, F. (2007). First record of the fossil 'rapismatid-like' Ithonidae (Insecta, Neuroptera) from the Lower Cretaceous Crato Formation of Brazil. Cretaceous Res., 28, 743-753.

Makarkin, V. N.; Ren, D; and Yang, Q. (2009). Two new species of Kalligrammatidae (Neuroptera) from the Jurassic of China, with comments on venational homologies. Annals of the Entomological Society of America, 102, 964-969.

McLoughlin, S.; Carpenter, R. J.; and Pott, C. (2011). Ptilophyllum muelleri (Ettingsh.) comb. nov. from the Oligocene of Australia: Last of the Bennettitales? International Journal of Plant Sciences, 172(4), 574-585. Retrieved 1/22/13 from

Peñalver, E.; Labandeira, C. C.; Barrón, E.; Delclòs, X.; Nel, P.; Nel, A.; Tafforeau, P.; and Soriano, C. (2012). Thrips pollination of Mesozoic gymnosperms. Proceedings of the National Academy of Sciences of the United States of America. Retrieved 1/23/13 from

Prokop, J. and Nel, A. (2010). New griffenfly, Bohemiatupus elegans from the Late Carboniferous of western Bohemia in the Czech Republic (Odonatoptera: Meganisoptera: Meganeuridae). Annales de la Société Entomologique de France, 46(1-2), 183-188.

Qiang, J. I.; Hongqi, L. I.; Bowe, L. M.; Liu, Y.; and Taylor, D. W. (2004). Early Cretaceous Archaefructus eoflora sp. nov. with Bisexual Flowers from Beipiao, Western Liaoning, China [electronic version]. Acta Geologica Sinica, 78(4), 883-896. Retrieved 1/22/13 from 

Riek, E. F. and Kukalova-Peck, J. (1984). A new interpretation of dragonfly wing venation based upon Early Upper Carboniferous fossils from Argentina (Insecta: Odonatoidea) and basic character states in pterygote wings. Canadian Journal of Zoology, 62, 1,150-1,166. 

Schlüter, T. (1986). The fossil Planipennia – a review. In: Gepp, J.; Aspöck, H.; and Hölzel, H. (eds.). Recent Research in Neuropterology. Proceedings of the 2nd International Symposium on Neuropterology (pp. 103-111). Graz: privately printed. 

Speer, B. R. (2000). Introduction to the Bennettitales: the Cycadophytes. Retrieved 1/23/13 from

Wang, Y.; Liu, Z.; Wang, X.; Shih, C.; Zhao, Y.; Engel, M. S.; and Ren, D. (2010). Ancient pinnate leaf mimesis among lacewings. Proceedings of the National Academy of Sciences [electronic version]. Retrieved 1/19/13 from  

Yang, Q.; Makarkin, V. N.; Winterton, S. L.; Khramov, A. V.; and Ren, D. (2012). A Remarkable New Family of Jurassic Insects (Neuroptera) with Primitive Wing Venation and Its Phylogenetic Position in Neuropterida. PLOS One. Retrieved 1/22/13 from

Zherikhin, V. V.; Mostovski, M. B.; Vršansky, P.; Blagoderov, V. A.; and Lukashevich, E. D. (1999). The unique Lower Cretaceous locality Baissa and other contemporaneous fossil insect sites in North and West Transbaikalia. Proceedings of the First Palaeontological Conference, Moscow, 1998, 185-191. 

1 comment: