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Ethnographic accounts from around the world have reported the widespread use of insects as food by people1,2,3. In some cases, such as among the Shoshone and other Great Basin tribes of the U.S., swarms of grasshoppers and crickets were driven into pits and blankets4, while among the Paiute the larvae of Pandora moths (Coloradia pandora lindseyi) were smoked out of trees to fall into prepared trenches, where they would be cooked5. Across the world, insects could be mass-harvested, often seasonally, offering high nutritional value especially in fat, protein and vitamins6. The harvesting of insects in the past has ranged from opportunities to feed large communal gatherings during times of plenty, to more individualistic economic pursuits such as in the search for delicacies or the exploitation of low-ranked resources when other foods were scarce or depleted7,8,9. Irrespective of the catch, insects often represented an important component of the diet, and of the reliability and thus dependability of locales as resource zones, with implications for social scheduling and cultural practice. However, a paucity of archaeological studies of insect food remains has resulted in a downplay or omission of the use of insects from archaeological narratives and deep-time community histories10.
In Australia, a wide range of insects is known to have been eaten by Aboriginal groups, in particular the larvae (‘witchetty grubs’) of cossid moths (especially Endoxyla leucomochla) in arid and semi-arid areas11,12,13. Of particular interest to archaeologists and behavioural ecologists has been the seasonal consumption of Bogong moths by mass gatherings of Aboriginal groups in the southern portions of the Eastern Uplands14 (Fig. 1). However, no conclusive archaeological evidence has ever been reported for the processing or use of Bogong moths.
Figure 1

(A) Bogong moth, Agrotis infusa (photo: Ajay Narendra). (B) Thousands of moths per square metre aestivating on a rock surface (photo: Eric Warrant).

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The Cloggs Cave grindstone
Cloggs Cave is located 72 m above sea level in the southern foothills of the Australian Alps, in the lands of the Krauatungalung clan of the GunaiKurnai Aboriginal peoples of southeastern Australia (Fig. 2). The cave is a small, 12 m long × 5 m wide × 6.8 m high limestone karst formation that is today entered through a walk-through opening on the side of a cliff (Fig. 3). Indirect sunlight dimly illuminates the cave for much of the day (Supplementary Fig. S1).
Figure 2

Location of Cloggs Cave and the area of the GunaiKurnai Land and Waters Aboriginal Corporation, at the southern foothills of the Australian Alps. Esri ArcMap 10.5 (https://desktop.arcgis.com/en/arcmap/) and Adobe Illustrator CC 2017 (21.0) (https://helpx.adobe.com/au/illustrator/release-note/illustrator-cc-2017-21-0-release-notes.html) were used by CartoGIS Services, College of Asia and the Pacific at the Australian National University, to create the map.

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Figure 3

Cloggs Cave cliffline above the Buchan River flood plain, showing location of cave entrance (white rectangle) (photo: Bruno David).

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Archaeological excavations were first undertaken in 1971–197214, followed by a new program of excavations in 2019–2020, initiated by the GunaiKurnai Land and Waters Aboriginal Corporation and directed by Bruno David. The new excavations were aimed at better determining the site’s stratigraphy and the antiquity of Aboriginal occupation (Supplementary Fig. S2). An intensive dating programme showed that the oldest excavated evidence for human activity dates to between 19,330–19,730 cal BP (median age of 19,530 cal BP; cal BP = before AD1950) and 20,590–23,530 cal BP (median age of 21,690 cal BP) (all calibrated radiocarbon ages in the text are presented at 95.4% probability range. See “Methods”; Supplementary Fig. S3)15,16,17.
During the 2019 excavations, a small, flat grindstone was found. The finely stratified hearth layers of stratigraphic unit (SU) 2 in which it was found were radiocarbon-dated to 1567–1696 cal BP at their top (uncalibrated: 1724 ± 16 BP; median age of 1632 cal BP) and 2002–2117 cal BP at their base (uncalibrated: 2091 ± 16 BP; median age of 2062 cal BP). The grindstone therefore dates to between 1600 and 2100 years ago (see “Methods”; Supplementary Figs. S3 and S4)17. No other grindstone has been found at Cloggs Cave.
The grindstone is a tabular fragment of sandstone with two flat and parallel ground surfaces (Surfaces A and B), in the form of a flat dish (Fig. 4). It measures 10.5 cm long × 8.3 cm wide × 2.2 cm thick and weighs 304 g. The outer, intact margin is elliptical in plan view; the other three margins indicate old breaks that have been subsequently worn from use. Therefore, prior to its deposition at Cloggs Cave, the grindstone had been used in its current form.
Figure 4

The Cloggs Cave grindstone. (A) Surface A, with the accretion that formed across parts of the surface after its use. (B) Surface B. (C) Margin A. (D) Margin B. (E) Narrow end. The numbers in circles are the residue sample numbers; the ‘control’ samples are in areas where grinding did not take place (photos: Richard Fullagar).

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To understand how the grindstone was used, we undertook use-wear and residue analyses (see “Methods”). The central area of both its surfaces contain fine unidirectional striations (Supplementary Figs. S5A and S5B), a lowered but not levelled topography, and areas of missing or ripped quartz grains (Supplementary Figs. S5C and S5D). Its use to shape ground stone axes is an unlikely function because the Cloggs Cave grindstone surfaces are relatively flat with only very slight concavities, and the lowered surface topography (Fig. 4) lacks broad grooves typical of axe grinding.
When viewed at lower (up to 5 ×) magnification under a stereozoom microscope with a point source of light, each surface appears relatively rough compared with grindstones used for processing seeds, which, in Australia, tend to be highly smoothed and polished18,19. There are numerous ‘pits’ where sand grains have been plucked from the surface during use (Supplementary Fig. S5D). The presence of a lowered surface topography (Supplementary Fig. S5C) with a lack of smooth, developed polish suggests that the stone was not used to process siliceous plants.
The repeated mechanical action of grinding has been shown to force residues into the voids and interstitial spaces of ground surfaces, where they become trapped20,21,22. Residue analyses conducted on grindstones worldwide have relied on microscopic observations of individual residue morphologies. However, visually diagnostic features can be altered by the mechanical forces of grinding, heat, and contact with water and various environmental factors, which can cause residues to swell or become amorphous21,22,23,24. The distinctiveness of residue identifications can be enhanced significantly with the introduction of biochemical staining that can be observed under high-power microscopy and is best used in conjunction with microscopic use-wear analysis and identification of residue morphologies22.
We extracted nine samples, or ‘lifts’, for residue analysis from across Surface A and Surface B of the Cloggs Cave grindstone, including a control sample from an unworked part of each surface (Fig. 4; see “Methods”). These samples were analysed using a recently developed biochemical staining technique that enables residues to be identified from colorimetric changes occurring at a cellular level, rather than relying solely on structural features (see “Methods”)22. We used the collagen stain Picrosirius Red (PSR) to differentiate between plant and animal residues (see “Methods”). When PSR comes into contact with collagen (a protein unique to animals), it reacts to produce clear and distinctive staining and enhanced birefringence in cross-polarised light22,25.
Residues extracted from the grindstone
A range of residues were identified in the lifts, including amorphous collagen, collagen fibres, collagen structures, partially woven collagen, possible bone-like fragments, moth wing segments, a possible moth hind leg, amorphous cellulose, wood-like structures with pits, carbonised material, bordered pits and minerals (see below).
We found collagenous residues in mid-range densities across Samples 1 and 4 from Surface B and across Sample 5 from Surface A (Supplementary Fig. S6). These extractions were taken from central areas across each modified surface. In all cases, the frequency of the collagenous residues was approximately three times greater than the collagenous residues associated with the control samples. Residues include damaged collagen fibres of varying thicknesses, including some reticular fibres.
Woven collagen structures clearly show birefringence in cross-polarised light across Sample 1. Woven collagen, which forms quickly, is mechanically weak and usually associated with immature bone. Although woven collagen may persist as tendon and ligament attachments to bone, it is generally replaced by organised parallel collagen fibre bundles at skeleton maturity26. Collagen fibrils are found in the connective tissues of vertebrates as well as in invertebrates such as insects27, and may be present as individual strands, woven structures or parallel bundles, including among the Lepidoptera (moths and butterflies)28.
The density and combination of collagenous residues on the Cloggs Cave grindstone indicates that it was used to process fauna. A variety of collagenous materials (including woven collagen) were found in association with carbonised residues across Sample 2, which was extracted from a crystalline layer. The residues present on Samples 1 and 2 suggest that an insect or immature vertebrate was prepared and cooked using the grindstone.
We identified a moderate density of carbonised plant residues across Sample 2, in particular, wood-like structures with pits. These ranged from being partially to completely carbonised. Partially carbonised residues were also seen across Sample 4. In addition, bordered pits in small clusters were identified, along with pits within the carbonised structures. Bordered pits are cavities that are essential components in the water-transport system of higher-order plants and are found in the lignified cell walls of xylem conduits (vessels and tracheids). The pit membrane allows water to pass between xylem conduits, but limits the spread of embolism and vascular pathogens in the xylem29. Small quantities of lignin were also present (see “Methods”). Lignin is found in the cell walls of vascular plants (especially in wood and bark) and is responsible for the rigidity of plant structures.
The residues identified via biochemical staining are consistent with the use of twigs and bark as fuel for fires such as those of the microstratified ashy layers in which the grindstone was found (see Supplementary Fig. S3)17. Partially carbonised wood-like material was also noted across Sample 5. The density and distribution of carbonised residues varies across extractions. Our observations suggest either that: (a) the stone has been placed in or near fires; or (b) ash, embers or fires of varying heat were placed or lit across the stone, for varied durations of time.
We identified especially high densities (frequency of residue particles per unit volume of sample) of amorphous cellulose across Samples 1, 2, 4 and 5 (Supplementary Fig. S7). The presence of partially carbonised amorphous cellulose indicates that the plant residues were associated with fire. While the high density is indicative of a plant-processing event, there is no evidence of combinations of plant residues normally expected from plant processing. In particular, no starch grain or phytolith was seen in any of the extractions. While low heat can damage starch and cause its structure to be disrupted and its characteristic extinction-cross to be lost, low heat does not completely destroy starch visibility30. Similarly, phytoliths can be reshaped but not destroyed by fire31. The presence of animal and mineral residues but absence of starches and phytoliths is thus interpreted as a true absence of plant processing activities rather than a taphonomic effect of environmental factors negatively impacting their preservation.
We found a high density of variably carbonised insect wings in Sample 6 (Surface A), and lower densities in Samples 2 and 4. These wing fragments contain regular patterning or structure and exhibit distinct birefringence in cross-polarised light. A portion of proteinaceous material was associated with a ‘tangle’ of these structures (Fig. 5). To assess whether the insect remains were those of the Bogong moth, we compared the residues on Samples 2, 4 and 6 with a comparative reference sample (see “Methods”). All 26 cases of wing segments from the grindstone matched the metrical and morphological characteristics of those from Bogong moths in the reference material. The recorded damage on the archaeological wing segments, such as ripped wing structures, small rectangular wing fragments and tearing in various states of carbonisation, is what would be expected from ethnohistoric accounts of Bogong moth processing. Aboriginal people from across the region are known to have cooked Bogong moths on heated earth during the early and mid-nineteenth century. The moths were stirred during cooking, causing the wings and legs to be broken off by friction and heat. Some of the moths were pounded and ground into a paste which could then be smoked to preserve the food for weeks1,2.
Figure 5

Examples of Bogong moth segments from lifted samples (all at × 400 magnification). (A) Partially carbonised wing structures from Sample 2 (pp). (B) Partially carbonised wing structure and carbonised material from Sample 2 (pp). (C) Partially carbonised moth wing segment from Sample 4 (pp). (D–E) Damaged moth wing segment from Sample 6 (D pp; E xp). (F–G) Damaged moth wing segment from Sample 6 (F pp; G xp). (H) Damaged moth wing segment with proteinaceous material, from Sample 6 (pp). (I) Unburnt moth wing segment from Sample 4 (pp). (J) Damaged moth wing segment with attachment, from Sample 6 (pp). (K) Damaged moth wing segments from Sample 6 (pp). (L–M) Probable moth hind leg from Sample 6 (L pp; M xp). (N) Damaged moth wing segment from Sample 6 (pp). (O) Damaged moth wing segment with attachment, from Sample 6 (pp). Light source = plane (pp), part polarised (part pol) and cross-polarised (xp) (photos: Birgitta Stephenson).

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