Termite activity in the mid-Cretaceous of Australia
Article number: 27.3.a48
https://doi.org/10.26879/1420
Copyright Palaeontological Association, September 2024
Author biographies
Plain-language and multi-lingual abstracts
PDF version
Appendices
Submission: 13 June 2024. Acceptance: 13 September 2024.
ABSTRACT
In this paper we provide the oldest evidence of termites in Australia, based on an aggregation of several hundred morphologically distinctive faecal pellets preserved as opalized casts from the Griman Creek Formation (Albian-Cenomanian: c. 100 Ma) at Lightning Ridge, New South Wales, Australia. This trace fossil extends the record of isopterans in Australia around 40 to 50 million years earlier than previously identified termite wing impressions, and indicates that this group was an active component of the detritivorous community in eastern Gondwanan terrestrial ecosystems by the mid-Cretaceous. The distinctive prismatic faecal pellets with hexagonal cross-sections (referable to Microcarpolithes hexagonalis Vangerow) were probably produced by kalotermitid or mastotermitid termites. The associated fossil plant assemblage indicates that the producers of the faecal pellets likely fed on conifer wood. Based on the distribution of extant termites, the climate of the Lightning Ridge area (Surat Basin) was probably warm and moist during the mid-Cretaceous. Recognition that termites were well established in Australian terrestrial ecosystems by the Albian-Cenomanian implies that vicariance may have been just as influential as trans-oceanic dispersal in determining the early distribution of major termite clades. Opalization of these delicate faecal pellets highlights the potential for further discoveries of three-dimensionally preserved soft or friable body and trace fossils in the Lightning Ridge opal deposits.
Stephen McLoughlin. Department of Palaeobiology, Swedish Museum of Natural History, P.O. Box 50007, S-104 05 Stockholm, Sweden. (Corresponding author) steve.mcloughlin@nrm.se
Artai A. Santos. Department of Palaeobiology, Swedish Museum of Natural History, P.O. Box 50007, S-104 05 Stockholm, Sweden. ArtaiAnton.SantosLopez@nrm.se, artaisl29@gmail.com
Sherri Donaldson. Palaeoscience Research Centre, University of New England, Armidale, NSW, 2351, Australia. sdonal24@myune.edu.au
Christian Pott. LWL-Museum für Naturkunde, Westfälisches Landesmuseum mit Planetarium, Sentruper Straße 285, D-48161 Münster, Germany. Christian.Pott@lwl.org
Matthew McCurry. Earth and Sustainability Science Research Centre, School of Biological, Earth and Environmental Sciences (BEES), University of New South Wales, Kensington, New South Wales 2052, Australia; Australian Museum Research Institute, 1 William Street, Sydney, New South Wales 2010, Australia. Matthew.McCurry@Australian.Museum
Keywords: Isoptera; coprolites; opalization; plant-insect interactions; Kalotermitidae, Mastotermitidae
Final citation: McLoughlin, Stephen, Santos, Artai A., Donaldson, Sherri, Pott, Christian, and McCurry, Matthew. 2024. Termite activity in the mid-Cretaceous of Australia. Palaeontologia Electronica, 27(3):a48.
https://doi.org/10.26879/1420
palaeo-electronica.org/content/2024/5339-opalized-termite-coprolites
Copyright: October 2024 Palaeontological Association.
This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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INTRODUCTION
Termites (infraorder Isoptera) are detritivorous insects that play a major role in nutrient cycling in various terrestrial ecosystems on all continents except Antarctica (Eggleton, 2011). They are eusocial insects, known for constructing some of the most complex colonial habitats in the animal kingdom (Noirot, 1970). In turn, numerous ancient and modern invertebrates and vertebrates have adapted to feed specifically or predominantly on termites (Sheppe, 1970; Redford, 1987; Luo and Wible, 2005). Termites and termitophagous animals are prominent components of northern Australian ecosystems; yet, despite their key ecological roles in the landscape, very little is known about the origins of Australasian representatives of this group.
On a global basis, the sedimentary record has yielded numerous occurrences of termite body fossils, coprolites, nests, and borings structures from a broad array of Cenozoic and Cretaceous deposits (e.g., Colin et al., 2011; Engel et al., 2011; Jouault et al., 2021), indicating that the importance of Isoptera in terrestrial ecosystems extends far back in time. Nevertheless, the fossil record highlights some important palaeogeographical and temporal gaps that inhibit reconstruction of the evolutionary history of this group. The origin of termites remains controversial. Classically, a Late Jurassic origin has been proposed for the divergence of Isoptera from their cryptocercid cockroach sister group (Engel et al., 2009; Bourguignon et al. 2014; Legendre et al., 2015). Some controversial features interpreted as termite nests from Upper Triassic and Lower Jurassic deposits have been used to suggest a more ancient origin (Hasiotis and Dubiel, 1995; Bordy et al., 2004, 2009, 2010) but the attribution of these structures to Isoptera is disputed (Genise et al. 2005; Genise, 2017) and some may represent megarhizoliths (Alonso-Zarza et al., 2008; Genise, 2017). An Early Jurassic age for the stem-termite origin and a Late Jurassic age for the crown-termite origin have been proposed based on combined molecular and fossil data (Legendre et al., 2015). However, other molecular and fossil data have suggested the possibility of a crown isopteran origin around the Triassic-Jurassic boundary (c. 200 Ma) or even older (Davis et al., 2009; Ware et al., 2010; Jouault et al., 2021). The disparity in results regarding the origin and diversification of termites makes the fossil record crucial for understanding the evolutionary history of this important group of insects. Thus far, the oldest convincing body fossils of termites were recovered from Lower Cretaceous deposits of Europe (see Appendix 1), specifically from the Berriasian Doronino Formation (Vršanský and Aristov, 2014) and Zaza Formation (Engel et al., 2007), Russia. The earliest evidence of isopteran coprolites derives from the Berriasian Missão Velha Formation of Brazil (Pires and Sommer, 2009), and over 40 other examples of fossil termite faecal pellets have been documented (see Appendix 2).
Cretaceous body fossils or coprolites of termites are relatively common in Europe (Jarzembowski, 1981; Lacasa-Ruiz and Martínez-Delclós, 1986; Schlüter, 1989; Engel et al., 2007, 2011; Engel and Delclòs, 2010; Colin et al., 2011; Engel, 2014; Sánchez-García et al., 2020), Asia (Ren, 1995; Krishna and Grimaldi, 2003; Engel et al., 2007, 2011, 2016; Colin et al, 2011; Zhao et al., 2020; Jouault et al., 2021, 2022a, 2022b; Jouault and Nam, 2023; Engel and Jouault, 2024: Jiang et al., 2024), and the Americas (Krishna, 1990; Fontes and Vulcano, 1998; Krishna and Grimaldi, 2000; Martins-Neto et al., 2006; Bechly, 2007; Grimaldi et al., 2008). There is also evidence of hexagonal isopteran coprolites in various Cretaceous deposits across Africa (Colin et al., 2011 and references therein). By contrast, the Australasian fossil record of Isoptera is depauperate. Previously, the oldest fossil isopteran from this region was a single fragmentary termite forewing described by Riek (1952) from the Paleocene-Eocene (58.5-55 Ma: Langford et al., 1995) Redbank Plains Formation near Ipswich, Queensland, and assigned to Blattotermes neoxenus Riek (Mastotermitidae). Kaulfuss et al. (2010) described another solitary forewing from Lower Miocene deposits of Foulden Maar, Otago, New Zealand, assigning it to Stolotermes kupe Kaulfuss, Harris et Lee (Stolotermitidae). McCurry et al. (2022) reported two additional termite (Mastotermitidae) wings from a thinly laminated goethitic siltstone of latest Early to earliest Late Miocene age (c. 16-11 Ma) near Talbragar in central New South Wales. Thus far, these are the only body fossil records of Isoptera from Australasia. In the absence of a rich body fossil record, trace fossils (coprolites, tunnels in wood, and nest structures) offer scope for elucidating the ancient history of termites in this region.
Australia has an extensive record of Cretaceous woods, albeit that few of these materials have been studied and most investigations have focused on the taxonomy and palaeoclimatic implications of these remains. Silicified conifer wood is abundant in eastern Australian Cretaceous basins (White, 1961; Frakes and Francis, 1990; Dettmann et al., 1992; Philippe et al., 2004; Fletcher et al., 2014, 2015; Tosolini et al., 2018). This abundance is probably linked to the release of permineralizing silica from the weathering of volcanic ash derived from the Whitsunday Igneous Province that was emplaced during the breakup of Australia and New Zealand.
In some marine deposits, the wood is entombed in calcareous concretions (Frakes and Francis, 1990) and, in others, (e.g., Western Australian marginal basins) the wood is preserved by silicification and phosphatization via the decomposition of glauconite and apatite in greensands (Simpson, 1912; McLoughlin et al., 1995b; McLoughlin, 1996; McLoughlin and McNamara, 2001; Mory and Hocking, 2017). Mummified, coalified, and charcoalified woods are also represented in various Australian basins but this material has been understudied (Backhouse et al., 1995; McLoughlin et al., 2002; Carpenter et al., 2016; Tosolini et al., 2018). Thus, there is considerable potential for investigation of termite traces in Australian fossil wood preserved in multiple styles.
Woody axes can have long taphonomic trajectories, acquiring biogenic (bacterial, fungal, arthropod, and molluscan) damage while the tree is alive, as deadwood in the form of stumps and logs on land, during transport in freshwater systems, and as driftwood or submerged log grounds after being washed into marine or lacustrine environments (Philippe et al., 2022). Although some of the published Australian Cretaceous woods contain evidence of fungal attack, especially in the form of pocket rot (McLoughlin, 1996), and other examples host borings from teredinid bivalves (McLoughlin et al., 1995b) and oribatid mites (Fletcher and Salisbury, 2014), none of the Cretaceous axes has yet revealed termite damage.
Australasian Cenozoic woods have yielded at least two examples of termite faecal pellets within tunnels excavated in wood that is now permineralized. Well-preserved examples include Neotermes -like fossil frass documented from mid-Cenozoic permineralized plant litter in central Queensland, Australia (Rozefelds and De Baar, 1991), and Kalotermes faecal pellets recorded in Miocene Avicennia wood from North Island New Zealand (Sutherland, 2003). Sub-fossil (Pleistocene) termite mounds have also been documented from stabilized sand dune systems in the Simpson Desert, central Australia (Miller, 1989).
There are over 360 extant species of termites in Australia within about five families (Watson and Abbey, 1993). Many Australian species are endemic. They are important ecosystem engineers as various groups have selective feeding patterns and abilities to consume and degrade leaf litter, grass, wood, fungi and diffuse organic matter in soil (Clement et al., 2021), liberating nitrogen, phosphorus and potassium that plants can exploit (Griffiths et al., 2021) and, in the process, contributing to c. 1-3% of the atmospheric CH4 budget (Sanderson, 1996; Nauer et al., 2018). Extant wood and humic soil-feeding termites are roughly constrained to latitudes between 45° N and 45° S (Hellemans et al., 2022). They are particularly abundant and diverse in moist, warm (tropical) climates, where they constitute c. 10% of animal biomass and are responsible for >55% of organic matter decomposition (Bignell, 2006; Jones and Eggleton, 2011; Griffiths et al., 2019, 2021). They do not live in cool or cold habitats (Sanderson, 1996; Atlas of Living Australia, 2024).
Here, we document opalized casts of clusters of prismatic coprolites from the Albian-Cenomanian Griman Creek Formation at Lightning Ridge that are interpreted to represent the oldest evidence for termites in Australia. We evaluate the probable producers of the coprolites, their importance for interpreting the early radiation of termites, and their role in nutrient recycling in mid-Cretaceous ecosystems.
GEOLOGICAL SETTING
Opal is Australia’s official national gemstone, with a significant percentage of commercial extraction occurring at Lightning Ridge in northern New South Wales (Department of Industry, Science and Resources, Commonwealth of Australia, 2022). Both precious and non-precious opal has been exploited from this area for over 100 years, with many examles representing replacements of the hard parts of Cretaceous organisms.
The area around Lightning Ridge (centred on 29°25′39″S, 147°58′44″E) is characterized by a low-lying NW-SE trending mesa system in the southern part of the Surat Basin (Herrmann and Maas, 2022; Figure 1A, B). The Griman Creek Formation (Rolling Downs Group) is a sedimentary succession up to 400 m thick, comprising generally thin-bedded, interlaminated, fine- to medium-grained sandstone, siltstone, mudstone, and minor coals, intercalated with thicker (c. 1-1.5 m thick) cross-bedded sandstones and conglomerates (Green et al., 1997; Payenberg and Reilly, 2004; Bell et al., 2019). The Griman Creek Formation at Lightning Ridge has been divided into two members: the Wallangulla Sandstone Member, comprising chiefly fine-grained, laminated, clay-rich sandstone with discontinuous claystone lenses of the informal ‘Finch clay facies’; and the overlying Coocoran Claystone Member, a claystone unit ≤10 m thick (Moore, 2002; Bell et al., 2019). The formation was deposited in low-energy freshwater fluvial and lacustrine systems (coastal plain environments) draining into the epeiric Eromanga Sea to the west (Bell et al., 2019). Laser Ablation Multicollector Inductively Coupled Plasma Mass Spectrometry U-Pb-Th radioisotopic dating of detrital zircons recovered from a volcanogenic bentonitic clay bed overlying the ‘Finch clay facies’ at the Hard Hill locality, yielded a range of Cretaceous depositional ages, with opalization of the formation occurring much later, during the Neogene (Bell et al., 2019; Mustoe and Smith, 2023; Figure 1C). More specifically, the suite of clear euhedral to elongate prismatic zircon grains with oscillatory magmatic zoning yielded 238U/206 Pb ages of 117±1.75 to 96.9±1.49 Ma (mid-Aptian to mid-Cenomanian), with the youngest coherent set of grains dated as 98.08+1.1/−2 Ma (early Cenomanian: at 95% confidence) and the youngest graphical peak in the population being 101 Ma (latest Albian: Bell et al., 2019). Thus, we interpret the age of the underlying ‘Finch clay facies’ of the Wallangulla Sandstone Member to approximate that of the Albian-Cenomanian boundary (hereafter, informally designated mid-Cretaceous; Figure 1CFigure 1C).
Although there is very little surface expression of the Griman Creek Formation in this region, fossils have been discovered in lenses of the ‘Finch clay facies’, extracted from small-scale subsurface mining activities since the early 1900s (Meakin, 2011). Many of the opal occurrences fill tectonic or dissolution voids in the ‘Finch clay facies’ but some represent casts of invertebrate shells vertebrate bones or robust plant organs. These do not preserve the calcium carbonate, calcium phosphate, or cellulose/lignin of the original animal and plant remains but represent infilling of voids left by dissolution of the organisms’ hard parts. Dissolution and subsequent infilling of the voids by opaline silica is inferred to be primarily a Neogene phenomenon associated with the development of deep weathering profiles across the region (Herrmann and Maas, 2022; Mustoe and Smith, 2023).
The opalized biota of the Griman Creek Formation is considered to represent one of the most important Cretaceous terrestrial fossil assemblages in Australia, containing a diverse array of vertebrate, invertebrate, and plant fossils preserved as opalized casts (Bell et al., 2019). The rich array of vertebrate remains includes both freshwater and amphibious animals: amiid fishes, dipnoans, lamniform chondrichthyans, chelids, crocodylomorphs, and elasmosaurid plesiosaurs (Kemp and Molnar, 1981; Kear, 2006; Smith, 2010; Smith and Kear, 2013; Hart et al., 2019; Kemp and Berrell, 2020; Berrell et al., 2023). Terrestrial vertebrate fossils include theropods (Bell et al., 2016; Birch et al., 2019, 2020), sauropods (Frauenfelder et al., 2021), ornithopods (Molnar and Galton, 1986; Bell et al., 2018a, 2018b; Kitchener et al., 2019), enantiornithine birds (Molnar, 1999), and monotreme mammals (Archer et al., 1985; Clemens et al., 2003; Flannery et al., 2024). Freshwater invertebrate remains include bivalve and gastropod molluscs, and a single clade of decapod crustaceans (Newton, 1915; Hocknull, 2000; Hamilton-Bruce et al., 2002, 2004; Hamilton-Bruce and Kear, 2006, 2010; Kear and Godthelp, 2008; Bell et al., 2020). The fossil plant suite from the Griman Creek Formation is currently under study by the authors and includes various equisetaleans, ferns, and cupressacean and araucarian conifers, but lacks obvious angiosperm remains (Smith and Smith, 1999).
MATERIAL AND METHODS
The single available specimen was photographed using an Apple iPhone 12 dual camera system with primary 12 megapixel sensor with 1.4 µm pixels and 26 mm equivalent f/1.6 lens. SEM micrographs were acquired of the uncoated specimen using a JEOL NeoScope JCM-7000 Benchtop SEM at the Australian Museum, Sydney, employing an acceleration voltage of 15.0 kv under low-vacuum mode, with EDS acquisition under high-vacuum mode. Tomography was undertaken using a Nikon XT H 225 ST X-ray CT system at 130 kV and 70 µA with a resulting voxel size of 6.069 µm that was reconstructed in VGstudio Max V2023.3. The studied specimen is registered as AM F 128064 and stored at the Australian Museum, Sydney, where it was received as a donation by E. Smith in 2004.
SYSTEMATIC PALEONTOLOGY
IchnogenusMICROCARPOLITHES Vangerow, 1954
Type ichnospecies. Microcarpolithes hexagonalis Vangerow, 1954; by subsequent designation of Hall (1963).
Microcarpolithes hexagonalis Vangerow, 1954
Figure 2A-J
Description. The specimen consists of an aggregation of several hundred hexagonally prismatic opaline pellets; the block is 27 mm long, 16 mm wide, and 10 mm thick (Figure 2A-D). Individual pellets are oblong, 0.97-(1.33)-1.62 mm long (n=40), 0.59-(0.76)-1.00 mm in transverse diameter (n=40), and conspicuously hexagonal in cross-section (Figure 2E-G), with smoothly faceted surfaces (Figure 2H-J), and truncate to broadly rounded ends. Tomographic analysis reveals that individual coprolites have either a uniform finely granular internal composition (Figure 2E) or have an equivalent uniform outer rind and an internal cavity or porous region constituting one- to two-thirds of the pellet diameter (Figure 2F-G). The cavities locally contain irregularly arranged minute flecks of detritus. The coprolites lack any obvious regular organization or common orientation (Figure 2C). They are bound together by porous interstitial clays or opaline silica (Figure 2D, I, Figure 3).
Remarks. The structures described in this study differ from replacements of euhedral quartz crystals by their bluntly rounded termini (at both ends of the pellets) and irregular arrangement. These mid-Cretaceous structures are interpreted to be termite coprolites based on their equivalency in shape and cross-section to termite coprolites assigned to Microcarpolithes hexagonalis Vangerow, from the Late Cretaceous of the Netherlands (Vangerow, 1954; Colin et al., 2011). Microcarpolithes hexagonalis was originally considered to represent angiosperm seeds but later reinterpreted by Kovach and Dilcher (1988) as termite coprolites (Colin et al., 2011; Moreau et al., 2019). Here we follow the criteria of Colin et al. (2011) who proposed that Microcarpolithes hexagonalis “must be used to uniquely designate termite coprolites that are cylindrical in shape and with a conspicuous hexagonal section, making it clearly distinct from some coprolites of curculionid weevil larvae that occasionally produce subhexagonal pellets.” Although the Australian Cretaceous coprolites are slightly longer than the type specimen of M. hexagonalis from Europe, we do not regard this small difference (less than 1 mm) to be taxonomically significant for a trace fossil, which is in line with the proposal that size should not be considered primary criteria for differentiating ichnotaxa (Bertling et al., 2022).
DISCUSSION
Prismatic faecal pellets with sharply hexagonal cross-sections are atypical among insects, being produced by some groups of termites and some anobiid beetles, although those produced by coleopterans are typically much larger and less regular than pellets produced by isopterans (Weidner, 1956; Sutherland, 2003). Termopsidae, Kalotermitidae Archotermopsidae, and Mastotermitidae termites generate oblong pellets (Rohr et al., 1986; Colin et al., 2011; Moreau et al., 2019; Dong et al., 2022), whereas some other termite clades (e.g., Rhinotermitidae) produce amorphous, liquid, or gelatinous faeces (Arquette and Rodriguez, 2011; Colin et al., 2011). Termopsid pellets are generally larger and less regular than those of kalotermitids and mastotermitids (Light, 1934), and they lack surficial striate. Kalotermitids and mastotermitids are the most common producers of prismatic/faceted (hexagonal in cross-section) faecal pellets (Colin et al., 2011; Lewis et al., 2014) and, consequently, are the most likely producers of the Australian Cretaceous coprolites. The fecal pellets of some extant Kalotermitidae ( Kalotermes ) are smaller than those produced by the single extant species of Mastotermitidae ( Mastotermes darwinensis Froggatt). Although the mid-Cretaceous coprolites described here are more consistent with the size of the pellets produced by M. darwinensis (1-1.2 mm long according to Colin et al. 2011), the dimensional range and shapes of the Lightning Ridge coprolites are not sufficiently distinctive to exclude production by either mastotermitid or kalotermitid termites.
The hexagonal cross-section of termite dung is generated via pressure from six muscle bands that squeeze moisture out of the faecal mass in the rectum in order to collect water in the rectal grooves for retention by the insect (Child, 1934). The precise shape of the faecal pellet may also be influenced by the diet items and their moisture content (Lance, 1946). The cavities within some of the Lightning Ridge coprolites (Figure 2E-G) probably reflect incomplete opal casting of the pellet rather than constituting a primary textural or compositional character.
We interpret the dense arrangement of the Lightning Ridge coprolites to be the result of packing within a confined space, e.g., in a gallery formerly excavated within wood or within a termite mound or subterranean tunnel. The kalotermitid faecal pellets described from mid-Cenozoic (?Oligocene) deposits of central Queensland occur as masses in galleries within conifer wood or as free masses preserved in a silicified rainforest leaf litter assemblage (Rozefelds and De Baar, 1991). Those pellets are similar to the Lightning Ridge coprolites in being oblong and hexagonal in cross-section, but they are slightly larger (2.0-2.6 mm long and 0.8-1.0 mm wide). However, pellet size may be influenced by the availability and type of food or other ecological constraints (Rozefelds and De Baar, 1991). Apart from the slightly smaller size and more sharply defined hexagonally prismatic form of the Lightning Ridge coprolites, there is little to differentiate these from the central Queensland material, which supports a kalotermitid affinity for the new material and denotes a long (>100-million-year) history for this isopteran family in Australia.
The fossil flora of the Lightning Ridge opal fields is currently under study by the authors. Preliminary results indicate that the Albian-Cenomanian woody vegetation of this area was dominated by cupressacean and araucariacean conifers. The Lightning Ridge plant assemblage shares several taxa with the slightly younger Winton Formation flora of the Eromanga Basin to the north, in western Queensland (McLoughlin et al., 1995a, 2010). The former differs mainly in the apparent absence of angiosperms; hence, we infer that, like the Oligocene examples from Queensland (Rozefelds and De Baar, 1991), the Lightning Ridge mid-Cretaceous termites were probably feeding primarily on coniferous wood.
The Lightning Ridge specimen is not the only example of an opalized mass of termite coprolites. Rogers (1928, 1938) documented termite coprolites in Pliocene opalized wood from Santa Barbara, California. They are of similar size and shape to the Lightning Ridge material but more loosely aggregated and bound by interstitial opal and colloform lussatite.
Given that extant termites do not live in cool or cold habitats, it is likely that the climate experienced at Lightning Ridge during the late Early Cretaceous was fairly warm, despite the region’s relatively high (c. 60° S) palaeolatitude (Burgener et al., 2023). In addition, permineralized woods preserved at Lightning Ridge have marked growth ring boundaries suggesting strong seasonality in productivity. A relatively warm climate is supported by the presence of crocodilians and the diverse range of fossil organisms found in the opalized assemblages at Lightning Ridge (Smith and Smith, 1999; Bell et al., 2019). This is consistent with inferred global stepwise warming through the Albian-Turonian interval (Dettmann et al., 1992; Forster et al., 2007; Bottini and Erba, 2018; Huber et al., 2018; Burgener et al., 2023).
In North America, at least one archaic mammal ( Fruitafossor windscheffeli Luo and Wible, 2005) had, by the Late Jurassic, developed forelimb adaptations for digging and specialized dentition consistent with feeding on termites, other insects or soft plant matter. However, similar adaptations can develop among vermivorous mammals (Charles et al., 2013). Among extant Australian animals, the numbat ( Myrmecobius fasciatus : Marsupialia), short-beaked echidna ( Tachglossus aculeatus : Monotremata), and at least 16 species of lizards specialize in feeding on termites (Abensperg-Traun and Steven, 1997). As yet, no lizard or marsupial fossils have been recovered from the Griman Creek Formation at Lightning Ridge. Recent work has uncovered six species of monotremes from this deposit (Flannery et al., 2024). Among these, Opalios splendens Flannery et al. 2024 has some features reminiscent of adaptations to termitivory (e.g., long, slender jaws, reduced teeth with simplified crown shape, and inter-molar or -premolar diastemata) although these might, alternatively, have been adaptations to feeding on vermiform invertebrates. Although isopterans were clearly widespread by this time, whether any vertebrates targeted termites as a food source as early as the mid-Cretaceous in Australia remains speculative.
Mastotermitids and kalotermitids are relatively basal in at least some phylogenies of Isoptera, suggesting divergence early in the history of this order (Legendre et al., 2008; Lo and Eggleton, 2011). Owing to the relatively poor body fossil record of termites and recent discoveries of isopteran body and trace fossils extending back to, at latest, the Early Cretaceous on multiple continents (Jarzembowski, 1981; Rohr et al., 1986; Martínez-Delclòs and Martinell, 1995; Francis and Harland, 2006; Pires and Sommer, 2009; Colin et al., 2011; Dong et al., 2022; Greppi et al., 2023), we suggest that the ancient distribution of many termite clades remains inadequately constrained and poorly dated. Thus, biogeographic hypotheses invoking long-distance dispersal for the distribution of many termite groups (Emerson, 1955, 1969; Bourguignon et al., 2016; Buček et al., 2022) are probably premature. It is clear that isopterans have a fossil record extending back to the mid-Mesozoic, before the major phase of Pangean breakup (McLoughlin, 2001) and, on that basis, vicariance related to continental separation may have played an important role in the distribution of termite groups through time.
CONCLUSIONS
The distinctive opal-replaced, clustered, hexagonally prismatic coprolites with blunt termini from the Griman Creek Formation at Lightning Ridge, Surat Basin, New South Wales, are tentatively assigned to kalotermitid or mastotermitid termites. Being dated to c. 100 Ma, they represent the oldest evidence of termites in Australasia, pre-dating previous body and trace fossil records by c. 40-50 million years. These coprolites demonstrate the presence of termite activity in mid-Cretaceous continental ecosystems of eastern Australia, suggesting that these organisms were already important detritivores of dead, probably coniferous wood before the diversification of angiosperms. Mineral replacement of this delicate frass cluster, presumably representing faeces backfilling a bored cavity within wood, highlights the possibility of discovering additional opalized delicate body and trace fossils at Lightning Ridge that will provide a more complete understanding of the structure, biotic interactions and energy flow within the mid-Cretaceous coastal plain ecosystems of eastern Australia. Based on the modern climatic preferences of kalotermitid and mastotermitid termites, along with co-preserved crocodilians, a relatively warm climate is invoked for the high-latitude southern Surat Basin around the Albian-Cenomanian transition. The growing number of Mesozoic body and trace fossil records from multiple continents suggests that isopterans were widely established around the world before the major phase of Pangean breakup.
ACKNOWLEDGEMENTS
This research was funded by a Swedish Research Council VR grant (number 2022-03920) to SM. Specimen AM F 128064 was donated to the Australian Museum by E. Smith. We thank the editor and reviewers for their constructive comments on the manuscript.
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