A 100 million-year-old armoured caterpillar supports the early diversification of moths and butterflies
Graphical abstract
Introduction
The caterpillar is probably the most famous type of larva of the group Insecta. Some groups of scorpion flies and saw flies have caterpillar-type larvae (e.g. Williamson, 2009, p. 19901), but more commonly known are the larvae of moths and butterflies (Lepidoptera), not least due to the book of Eric Carle “The very hungry caterpillar”. The group Lepidoptera is one of the “big four”, lineages with more than 100,000 formally described species (the other lineages are Diptera, Coleoptera and Hymenoptera, the groups of flies, beetles and wasps). As the group Lepidoptera is so species-rich and due to the fact that butterflies and moths spend quite a large part of their life span in their larval form, caterpillars must represent quite a significant part of the biomass of modern-day terrestrial ecosystems. Caterpillars naturally are an important food source for many other organisms (e.g. Lin et al., 2019); as defoliaters, caterpillars have economic impact (e.g. Berti Filho, 1981; Dhir et al., 1992; Zanuncio et al., 1994), and some species occurring in high densities are even a factor in human health issues (e.g. Rosen, 1990; Gottschling et al., 2007; Gómez, 2014).
Given the importance of caterpillars in the modern fauna, it is surprising that the fossil record of caterpillars is quite scarce (Sohn et al., 2012, Sohn et al., 2015). In addition, most of the record is restricted to findings in amber and mostly to only 40 million years ago (Eocene) or even younger occurrences (Sohn et al., 2012). Due to the scarceness even from these rather young ambers, also single, relatively young specimens may represent quite significant findings (e.g. Fischer et al., 2019). Cretaceous ambers (c. 100 Ma), otherwise quite productive, have so far only provided very few caterpillar specimens, four of them were depicted. Older deposits lack caterpillars entirely.
The first of the Cretaceous caterpillar fossils is a broken head capsule from Canadian amber (MacKay, 1970, p.380 Fig. 1; Fig. 1A). A much better preserved specimen from Lebanese amber was depicted and discussed in several occasions (Grimaldi, 1996; Grimaldi, 1999, p. 405, fig. 12e; Grimaldi and Engel, 2005, p. 566, fig. 13.21; Fig. 1B), representing the oldest record of the group Glossata (embracing the major diversity of the group Lepidoptera). Xia et al. (2015, pp. 132, 133) figured two well-preserved caterpillar specimens from Myanmar amber (Fig. 1C, D), yet without further description or discussion. There was mentioning of additional specimens from Taimyr amber, yet we could not find any depictions of these specimens (Zherikhin and Sukacheva, 1973, p. 38; Skalski, 1979, p. 63)
All these finds appear to represent rather simple types of caterpillars, more or less appearing “naked”, while many modern caterpillars exhibit a wide range from similarly naked forms over very hairy ones to larvae with strong, defensive spines. Here we report a new lepidopteran caterpillar from 100 million-year-old Myanmar amber with an unusual combination of characters, most prominently it is armoured with numerous dorsal spines. We discuss the implications of this new finding.
Section snippets
Materials and methods
The new fossil specimen reported here comes from Myanmar amber (“Burmese amber”), about 100 million-year-old deposits, Hukawng Valley, Kachin State, Myanmar (Cruickshank and Ko, 2003). The specimen was legally purchased via the internet platform ebay.com, from the trader burmite-miner. The specimen is deposited in the Palaeo-Evo-Devo Research Group Collection of Arthropods, Ludwig-Maximilians-Universität München, Germany under repository number PED 0886. It was documented on a VHX-6000 digital
Description of the new specimen
Body elongate, worm-shaped, with distinct capsulate head and trunk (head partly ripped off the trunk; Figs. 2A, B, 3A). Trunk appears soft without distinct sclerotisations, but numerous distinct folds. Not all folds representing segment boundaries, hence folds partly obscuring trunk segmentation, yet 13 trunk units more or less well apparent. Anterior twelve units representing segments (thorax segments 1–3 and abdomen segments 1–9); last one, trunk end, most likely composite of two former
Identity of the new fossil
The fossil specimen overall has the appearance of a caterpillar. Caterpillar-type larvae occur not only in representatives of the group Lepidoptera, but also of Hymenoptera (e.g. Ross, 1959; Costa, 1997) and Mecoptera (e.g. Somma and Dunford, 2009; Saltin et al., 2016), to a certain degree also of Coleoptera (Chrysomelidae; e.g. Zhang et al., 2020) and to a lesser extent of Trichoptera (e.g. Ross, 1944; see comparison in Pryor, 1951). Yet, the arrangement of the appendages of the abdomen
Credit author statement
Both authors conceptualized the study, investigated the specimens, analysed the data, and wrote the manuscript. JTH documented the specimens, created the figures, and acquired funding.
Declaration of Competing Interest
The authors declare that they have no conflict of interest.
Acknowledgements
We thank Fenja I. Haug for drawing the specimens and Gideon T. Haug for assistance with the imaging. The Volkswagen Foundation kindly funds JTH with a Lichtenberg Professorship. We thank J. Matthias Starck, Munich, for continuous support. We highly appreciate the effort of all people providing open access, open source and low cost software.
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