An insect to fill the gap
A complete insect fossil from the Devonian period has long been sought. The finding of a candidate may improve our patchy understanding of when winged insects evolved. See Letter p .82


point speaking to its importance. We can perceive only what the fossil 300 nsects are, in terms of species number, record permits us to perceive. From our the most successful group of animals current viewpoint, the diversification of Carboniferous ever to have lived. But their evolutioninsects and of our own terrestrial verteary origins are a source of controversy, brate ancestors seems to have occurred 325 and will continue to be so until the fossil in two evolutionary bursts9. Between record finally yields up unequivocal evi425 million and 385 million years Palaeodictyoptera dence of insect beginnings. On page 82 of ago, both groups probably originated this issue, Garrouste et al.1 claim to have and underwent an initial evolution350 found precisely this. Although it can ary radiation as they began occupying Hexapoda gap hardly be described as well preserved, the newly available subaerial realm. the fossil shows a six-legged thorax, long There then follows a long period, called single-branched antennae, triangular Romer’s gap (360 million to 345 miljaws and a 10-segmented abdomen (see lion years ago) for the vertebrates and 375 Strudiella devonica Fig. 2 of the paper1). Insects are the only the longer Hexapoda gap for insects known arthropods (joint-legged inver(385 million to 325 million years ago), tebrate animals) with this anatomical during which few, if any, fossils of these Devonian combination, allowing the authors to groups can be found (Fig. 1). Then, 400 make a strong case for the fossil’s insectan with apparent suddenness, an explosive identity. appearance of many new forms takes Rhyniella praecursor The 8-millimetre-long fossil, which place in the second round of diversithe authors named Strudiella devonica, fication. For the insects, large winged 425 was found in a small rock slab excavated species of the major groups (mayflies, at a quarry in Belgium. Strudiella is proto-dragonflies and others, includdated to approximately 370 million years ing extinct types) show up, seemingly old, which places it late in the Devo- Figure 1 | Winged beginnings. The fossil record provides ample without precursors. The insects were examples of winged insects from around 325 million years ago, nian period (Fig. 1). This was the time such as the order Palaeodictyoptera from the Carboniferous off and running on their way to world when terrestrial ecosystems were first period. However, there is little evidence of insect evolution before domination. assembling from their aquatic progeni- this time; only a handful of fossils, including 402-million-year-old These gaps, and the two bouts of evotors2 — the first forests were established Rhyniella praecursor, which appears similar to extant collembolan lution that they create, may or may not and the earliest four-legged vertebrates arthropods, have been found. But none of these few examples be real. There is evidence that a period were crawling out from freshwater pools comes from the period between 385 million and 325 million years of low atmospheric oxygen concentraonto land. So far, only suggestive traces ago, which is referred to as the Hexapoda gap (striped region) tion coincided with the gap period, and of insects have been found in rocks of because of the lack of insect evidence. Now, however, Garrouste this could have suppressed the rate of 1 this age. The famous Rhynie chert, a et al. report the finding of Strudiella devonica, a fossil dated to appearance of novel anatomy10. But a 370 million years ago that shows multiple anatomical features that sedimentary deposit in Scotland that are characteristic of insects. more parsimonious explanation is simis about 402 million years old, contains ply that we have not yet found the right fossils of collembolans3, a class of animal that For example, the fossilized head of a wingless rock formations to reveal fossils that would fill contains today’s ubiquitous springtails, and insect found6,7 in Canadian strata somewhat in the gaps. For example, most of the exposed which is regarded as closely related to insects. older than the New York deposits is almost cer- strata for this period in Europe and North The Rhynie chert has also yielded a pair of tainly a contaminant — a much more recent America are of marine, not land, origin. jaw fossils called Rhyniognatha, which may or contemporary insect lodged in a crack in This brings us to the second reason for be from an advanced, winged insect4. In New the rocks. And Leverhulmia mariae, also from the importance of Strudiella — it is dated to York state, some fossilized scraps of character- Scottish chert near Rhynie, could have been an a time smack in the middle of the Hexapoda istic cuticle and the framework of a single com- insect, a close relative, or neither — it seems gap (Fig. 1). According to Garrouste et al., pound eye, perhaps from a primitive, wingless to have too many legs to be easily classified8. this significantly narrows the gap. And if, as insect, have been found in 385-million-year So although its age makes it too late to be the authors suggest, the fossil came from the old rocks5. But these fragments more or less an insect ancestor, or even the earliest insect, young stage of an animal that would have had complete the picture of all that is known of Strudiella is nonetheless of great potential wings as an adult, their finding would mean insects at this crucial time in Earth’s history. significance as the oldest complete insect that winged insects originated much earlier There have also been some false alarms. fossil yet found. This is the first and primary than fossils have heretofore told us, and that
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Time (millions of years ago)

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the sudden appearance of many winged kinds around 325 million years ago is deceptive. It would also suggest that the Rhyniognatha fossils could indeed be the mandibles of a winged insect, and that the diversification of winged species could have taken place at a much more leisurely pace, over some 45 million years. Considering the crucial role of insects in present-day ecology, the number of people engaged in studying their fossil history is dismally small. Furthermore, current specialists focus mostly on events from the Mesozoic period — the ‘Age of Dinosaurs’ — which began some 70 million years after the time of the first known winged insect fossils, or on even more recent amber-preserved insects,

which are largely indistinguishable from living forms11. The beginnings of the insects are to be found in rocks much older even than those that enclosed Strudiella, but almost no one is looking for them. The paltry few insect fossils contemporary with Strudiella — and indeed Strudiella itself — were serendipitous, not deliberate, finds, and the Hexapoda gap still looms large. ■ William A. Shear is in the Department of Biology at Hampden-Sydney College, Hampden-Sydney, Virginia 23943, USA. e-mail:
1. Garrouste, R. et al. Nature 488, 82–85 (2012). 2. Shear, W. A. & Selden, P. A. In Plants Invade the Land. Evolutionary & Environmental Perspectives

(eds Gensel, P. G. & Edwards, D.) 29–51 (Columbia Univ. Press, 2001). 3. Greenslade, P. J. & Whalley, P. E. S. In Proc. 2nd Int. Semin. Apterygota (ed. Dallai, R.) 319–323 (Univ. Siena, 1986). 4. Engel, M. S. & Grimaldi, D. A. Nature 427, 627–630 (2004). 5. Shear, W. A. et al. Science 224, 492–494 (1984). 6. Labandeira, C. C., Beall, B. S. & Hueber, F. M. Science 242, 913–916 (1988). 7. Jeram, A. J., Selden, P. A. & Edwards, D. Science 250, 658–661 (1990). 8. Fayers, S. R. & Trewin, N. H. Palaeontology 48, 1117–1130 (2005). 9. Labandeira, C. C. & Sepkoski, J. J. Science 261, 310–315 (1993). 10. Ward, P., Labandeira, C. C., Laurin, M. & Berner, R. A. Proc. Natl Acad. Sci. USA 103, 16818–16822 (2006). 11. Grimaldi, D. A. & Engel, M. S. Evolution of the Insects (Cambridge Univ. Press, 2005).


of soil organic matter caused by rising global temperatures8. To assess variations in carbon sinks over the past 50 years, Ballantyne et al. used a strikingly simple approach: they calculated the annual changes in the global atmospheric CO2 inventory from a worldwide network of long-term observations. By subtracting the annual total amount of anthropogenic CO2 emissions from Careful analysis reveals that the global uptake of anthropogenic carbon dioxide these changes, they quantified the net CO2 emissions by carbon sinks has doubled during the past 50 years — but the uptake by the land and oceans each year. This fractions of this absorbed by land and by sea remain unclear. See Letter p .70 fundamental strategy reminds us that from time to time we should step back and carefully consider the basic inputs and outputs of the INGEBORG LEVIN afforestation or increased biomass produc- carbon cycle at a global level. Although similar tion by plants. These carbon sinks have been work has been done before2,9, Ballantyne et al. recisely quantifying the fate of man-made efficiently damping increases in anthropo- go one step further by quantifying an increase carbon dioxide is vital for reliably esti- genic CO2 (Fig. 1), but will not necessarily in global CO2 uptake as worldwide emissions mating future atmospheric CO2 levels continue to do so. Possible mechanisms that are increasing. and the contribution of this greenhouse gas could result in reduced carbon uptake include To determine the uncertainties of their estito global climatic change. There has been changes in carbon chemistry and mixing in mated carbon budget, the authors combined much debate about whether currently active the oceans7, and potential feedback from land the most comprehensive, high-accuracy carbon sinks are likely to change drastically in ecosystems such as increased decomposition measurements of atmospheric CO2 available the near future1, or might already with three inventories of estimated have weakened during the past global fossil-fuel CO2 emissions, 500 few decades2,3. On page 70 of this and with three inventories of issue, Ballantyne and colleagues4 emissions estimated to have been clarify matters. By calculating the caused by land-use change. The 450 worldwide increase in the atmosinventories had been made indeCO2 pheric CO2 burden during the past pendently of each other, and were accumulation 50 years from precise global obserobtained using different methods. Total anthropogenic CO2 in sinks 400 vations, and by carefully accountBringing together all these data ing for anthropogenic carbon allowed the authors to reliably calsources and their uncertainties, culate the errors of their different 350 they find that total global carbon budget terms. Their finding that Atmospheric CO2 sinks have not declined during this the net carbon sink is increasing is period. Rather, they have increased therefore robust, and proves that 300 more than twofold — although fosocean and land sinks have not both 2000 1960 1970 1980 1990 2010 sil-fuel CO2 emissions have risen decreased in the last half century. Year almost fourfold5,6 over this time. How the total sink was divided Roughly half of the anthropobetween oceans and land in the genic CO2 emissions caused by Figure 1 | Partial absorption of anthropogenic carbon dioxide. The past, and whether the oceans’ CO2graph compares atmospheric levels of carbon dioxide since the 1960s the burning of fossil fuels and by (measured at Mauna Loa, Hawaii11) with the levels that would have occurred uptake rate has decreased, as sugland-use change (such as deforest- as a result of the accumulation of anthropogenic CO emissions in the gested by some modelling studies2, 2 ation) are currently taken up by the absence of carbon sinks (as calculated by Ballantyne et al.4). The amount of remain open questions. oceans or re-enter the terrestrial CO2 that has accumulated in sinks is represented by the difference between Why is it so important to know biosphere — for example, through the two curves (green arrow). where half of the carbon that we are

The balance of the carbon budget

CO2 (parts per million)

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