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Comprehensive survey of Early
to Middle Triassic Gondwanan
floras reveals under-
representation of plant–
arthropod interactions
Holly-Anne Turner
1
*, Stephen McLoughlin
2
and Chris Mays
1
1
School of Biological, Earth and Environmental Sciences, Environmental Research Institute, University
College Cork, Distillery Fields, Cork, Ireland,
2
Department of Palaeobiology, Swedish Museum of
Natural History, Svante Arrhenius, Stockholm, Sweden
Plants and arthropods are primary drivers of terrestrial ecosystem function. Trace
fossils of plant–arthropod interactions (PAIs) provide a unique window into
assessing terrestrial ecosystem states through geological time and evaluating
changes in herbivorous arthropod feeding guilds in the wake of global biotic
crises. The end-Permian event (EPE; c. 252 Ma) resulted in the loss of keystone
plant species from humid tropical and high-latitude ecosystems and the extinction
of several major insect groups. The subsequent Early to Middle Triassic evinced
diminished terrestrial productivity, punctuated by a series of second-order biotic
crises that hindered recovery. Here, we survey records of Gondwanan Early to
Middle Triassic floral assemblages for evidence of PAIs as an indication of
ecosystem recovery following the EPE. We compiled a comprehensive dataset of
fossil plant taxa and PAIs for lower Mesozoic strata of Gondwana, revealing an
increase in specific and generic floral diversity from the Early to Middle Triassic. We
noted a lack of PAIs reported from many localities with abundant fossil leaves, which
might be interpreted to be a consequence of a post-EPE delay in the recovery of
arthropod feeding guilds compared to the flora. However, by comparing floral
assemblages between regions of Gondwana, our results also partly attribute the
absence of PAIs to the relative paucity of palaeoichnological and palaeobotanical
studies of this interval. To test for potential under-reporting of PAIs in the Triassic,
we present a case study of the well-described Australian Middle Triassic Benolong
Flora. In contrast to existing Australian Early to Middle Triassic PAI reports on only
three plant specimens, this systematic investigation revealed 44 PAI traces
comparable to published examples, hosted by 40 fossil plant fragments (7.77% of
fragments assessed; N= 591). Margin-feeding traces constituted the dominant
Functional Feeding Group (FFG) identified (23 examples: 3.72% of fragments
assessed). Our review highlights several Early and Middle Triassic Gondwanan
plant fossil-rich successions and existing collections that require further
Frontiers in Ecology and Evolution frontiersin.org01
OPEN ACCESS
EDITED BY
Sandra Schachat,
Stanford University, United States
REVIEWED BY
Barbara Cariglino,
National Scientific and Technical Research
Council (CONICET), Argentina
Zhuo Feng,
Yunnan University, China
*CORRESPONDENCE
Holly-Anne Turner
hollyaturner@ucc.ie
RECEIVED 17 April 2024
ACCEPTED 05 August 2024
PUBLISHED 11 September 2024
CITATION
Turner H-A, McLoughlin S and Mays C (2024)
Comprehensive survey of Early to
Middle Triassic Gondwanan floras
reveals under-representation of
plant–arthropod interactions.
Front. Ecol. Evol. 12:1419254.
doi: 10.3389/fevo.2024.1419254
COPYRIGHT
© 2024 Turner, McLoughlin and Mays. This is
an open-access article distributed under the
terms of the Creative Commons Attribution
License (CC BY). The use, distribution or
reproduction in other forums is permitted,
provided the original author(s) and the
copyright owner(s) are credited and that the
original publication in this journal is cited, in
accordance with accepted academic
practice. No use, distribution or reproduction
is permitted which does not comply with
these terms.
TYPE Original Research
PUBLISHED 11 September 2024
DOI 10.3389/fevo.2024.1419254
examination. We predict that investigations of these assemblages will greatly
elucidate the relationships between rapidly changing environments during the
Early and Middle Triassic and their effects on the plant and arthropod
communities in the Southern Hemisphere.
KEYWORDS
End-Permian extinction, fossil, Gondwana, palaeobotany, palaeoecology, plant-
herbivore interactions, post-extinction recovery, Triassic
1 Introduction
1.1 Plant–arthropod interactions in the
fossil record
Plants (c. 380,000 extant species; Christenhusz and Byng, 2016)
and terrestrial arthropods (c. 7 million extant species; Stork, 2018)
constitute the most significant proportion of the lower trophic levels
on which all higher levels of life on land depend, with their
interactions playing a crucial role in terrestrial ecosystem nutrient
cycling. Understanding and quantifying the relationships between
arthropods and the plants with which they are associated can,
therefore, be used to indicate the condition and evolution of
terrestrial ecosystems (Labandeira, 2013). Today, arthropods can
be observed directly interacting with plant organs, and specialized
associations between particular plant and arthropod taxa can be
identified. In fossil plant assemblages, traces of herbivory
(consumption of live plant tissue), detritivory (feeding on dead
tissues) and oviposition (egg-laying) left on or within plant organs
provide evidence of fundamental behavioral interactions that
formed the basis of ancient terrestrial food webs and ecosystem
function (Labandeira and Currano, 2013).
Plant–arthropod interactions (PAIs), captured as trace fossils
on plant tissues, record behaviors of arthropods that may not be
deducible from their body fossils. Moreover, plants are relatively
common in the fossil record and an arthropod may interact with
many plants during its lifetime. So, the likelihood of a plant
fragment exhibiting a PAI being preserved is greater than the
likelihood of preserving an arthropod body fossil. Consequently,
PAIs are important indicators of the richness and abundance of
arthropods and herbivore guilds present in an ancient ecosystem
(Labandeira and Wappler, 2023). Furthermore, fossil leaves are
utilized in many palaeoecological studies as they are consistently
more abundant and widespread than most other plant organs, such
as wood and seeds and, therefore, provide more accessible records
(Wilf, 2008). Morphological deterrents against arthropod
herbivory, such as spines and hairs, have a much higher potential
for preservation in the fossil record than chemical defenses. Their
presence on plant fossils is evidence that arthropod interactions
provided sufficient evolutionary pressure on plant taxa to favor
selection of those plants investing resources in physical defenses
(War et al., 2018).
Although detritivory is interpreted to have occurred since the
initial establishment of complex terrestrial ecosystems in the late
Silurian to Early Devonian (Kenrick et al., 2012;Garwood et al.,
2020), the progression to herbivory is not extensively documented
in the fossil record (Labandeira and Currano, 2013;Labandeira and
Wappler, 2023). Probable arthropod coprolites (fossilized fecal
pellets) containing spores and other fragments associated with
late Silurian (412 Ma) and Early Devonian (390 Ma) land plants
have been identified as the earliest documented evidence for the
consumption of live tissues alongside dead plant matter (Edwards
et al., 1995;Hagström and Mehlqvist, 2012;Labandeira and
Wappler, 2023). The extraction of nutrients from plants via
piercing and sucking of cortical stem tissues has been reported in
Early Devonian vascular plants displaying lesions resulting from
repeated insertion of stylet mouthparts (Banks and Colthart, 1993).
By the Middle Devonian, several types of herbivory were
represented on simple land plants: external tissue feeding, galling,
and piercing and sucking (Labandeira et al., 2014).
Labandeira et al. (2007) published a guide focused primarily on
trace fossil evidence of damage to foliage, consisting initially of 150
distinct Damage Types (DTs) almost all of which are probably
linked to arthropods. The latest available updated compendium (as
yet unpublished; C. Labandeira, pers. comm.) defines over 420
distinct damage types. These DTs are classified into a series of
Functional Feeding Groups (FFGs): 1, hole feeding; 2, margin
feeding; 3, skeletonization; 4, surface feeding; 5, oviposition; 6,
piercing and sucking; 7, mining; 8, galling; 9, seed predation; 10,
pathogenic damage; 11, boring; and 12, domatia construction. To
these feeding and other interactive behaviors can be added
palynophagy and nectarivory, which can be difficult to detect in
the fossil record.
With a standardized classification scheme, the quantification of
fossil leaves with invertebrate traces through time can be utilized to
infer the state of health of the lower trophic levels, e.g., terrestrial
ecosystem development and recovery before and after major
environmental changes (Labandeira et al., 2018). The majority of
mass extinction events were associated with worldwide changes in
the climate, the most common and lethal of which were rapid
Turner et al. 10.3389/fevo.2024.1419254
Frontiers in Ecology and Evolution frontiersin.org02
warming episodes, or ‘hyperthermal events’(Bond and Grasby,
2017). An understanding of how plant–arthropod interactions have
responded to cases of past environmental destabilization will
elucidate how terrestrial ecosystems evolve (Wilf et al., 2001;
Labandeira, 2006;Wilf, 2008;Labandeira and Currano, 2013;
Currano et al., 2016;Labandeira and Wappler, 2023;Zambon
et al., 2023). Moreover, assessing the traits common to mass
extinction survivors, and observing the progression of ecosystem
inhabitantsovertime,enablestheidentification of potential
patterns of recovery between plants and their dependent
invertebrates throughout evolutionary history (Labandeira, 2006;
Cariglino et al., 2021). This may then inform our predictions of how
extant terrestrial ecosystems may be affected by modern rapid
climate changes (Currano et al., 2016;Zambon et al., 2023).
1.2 Latest Permian to Middle Triassic
climatic events
The transition from the Paleozoic to Mesozoic eras was punctuated
by a series of climate-driven ecosystem shifts (Figure 1). The first of
these was the end-Permian event (EPE, c. 252.2 Ma) that has been
clearly linked to a major magmatic episode of the Siberian Traps Large
Igneous Province (Burgess et al., 2017;Joachimski et al., 2019;Shen
et al., 2023). The EPE hyperthermal event involved increases in
atmospheric CO
2
(Cui et al., 2021;Wu et al., 2023), temperature
(Sun et al., 2012;Cui and Kump, 2015), wildfires (Vajda et al., 2020;Lu
et al., 2022;Mays and McLoughlin, 2022), freshwater microbial blooms
(Mays et al., 2021) and enhanced seasonality in precipitation (Fielding
et al., 2019;Frank et al., 2021). A combination of these (and related) kill
mechanisms during the EPE led to the highest biodiversity loss of any
mass extinction within both the marine and continental realms
(Stanley, 2016;Viglietti et al., 2021) including a globally significant
floral and insect turnover (Anderson et al., 1999;Labandeira, 2005;
Cascales-Miñana et al., 2016;Gastaldo et al., 2020;Schachat and
Labandeira, 2021). Therefore, the Triassic fossil flora, entomofauna
and their associated interactions can be used to inform patterns of
recovery and continental ecosystem evolution in the aftermath of
Earth’s greatest extinction event.
The post-EPE recovery of ecosystems is complicated by a series
of Early Triassic climatic events (Figure 1). Three of these events
have demonstrated major biotic impacts (in chronological order): 1,
the late Smithian thermal maximum (c. 250 Ma; Sun et al., 2012;Du
et al., 2022); 2, the Smithian–Spathian event (c. 249.2 Ma; Hermann
et al., 2012a;Lindström et al., 2020); and 3, the Early–Middle
Triassic event (c. 247.2 Ma; Hermann et al., 2012b). At least one of
these (the late Smithian thermal maximum) has been linked to
enhanced global warming (Du et al., 2022), possibly as a result of
renewed Siberian Traps Large Igneous Province magmatism (Shen
et al., 2019) although this is contested (Widmann et al., 2020). The
latter two events (the Smithian–Spathian event and the Early–
Middle Triassic event), however, seem to be associated with
cooling trends (Romano et al., 2013;Widmann et al., 2020).
Although magmatism has been indicated as the ultimate cause of
both (Du et al., 2022;Saito et al., 2023), the precise mechanisms
remain obscure. Regardless of the causes, it is highly likely that the
climatic fluctuations hindered ecological recovery for millions of
years following the EPE (Grauvogel-Stamm and Ash, 2005;
Friesenbichler et al., 2021). A demonstration of this hindrance is
the global absence of thick, economic coal deposits until the Middle
Triassic (Veevers et al., 1994;Retallack et al., 1996); the likely oldest
post-EPE coals in Gondwana are in the c. upper Anisian (Dolby and
Balme, 1976) lower Mungaroo Formation on the Australian
northwest shelf (Retallack et al., 1996;Zeng et al., 2019). The
absence of dense vegetation represented by the coal gap was a
probable contributor to the Early Triassic ‘charcoal depression’,an
interval of relatively low wildfire prevalence (Figure 1;Mays and
McLoughlin, 2022). These signs indicate that, although the main
plant constituents of Gondwanan Middle and Upper Triassic coals
(Umkomasiales, e.g., Dicroidium) had spread across the continent
soon after the EPE, their abundances remained low until long after
the major climatic fluctuations of the Early Triassic.
1.3 Ecosystem evolution after the end-
Permian event (EPE)
Permian floral assemblages of Gondwana are overwhelmingly
dominated by the remains of one group of broadleafed, arborescent
seed plants, the glossopterids, alongside subsidiary lycopsids,
sphenopsids, ferns, cycadaleans, ginkgoaleans, cordaitaleans,
conifers and several seed plants of uncertain affinity (Anderson
and Anderson, 1985;McLoughlin, 1992,1994a,b;Prevec et al.,
2009,2010). With the extinction of glossopterids near the end of the
Permian, new terrestrial ecosystems were slowly established and
dominated by peltaspermalean, voltzialean conifer, pleuromeian/
isoetalean lycophyte, and, eventually, umkomasialean (primarily
Dicroidium)floras through the Early and Middle Triassic (Figure 1;
Lele, 1974;Retallack, 1977;McLoughlin et al., 1997;Anderson et al.,
1998). By the Middle Triassic, Dicroidium had filled the canopy
plant niche left by the glossopterids until their own eventual
extinction across Gondwana at the close of the Triassic, apart
from a few potential relictual communities that persisted into the
Early Jurassic in East Antarctica (Bomfleur et al., 2018).
Evidence for PAIs through the Permian of Gondwana has been
broadly surveyed based on published literature (McLoughlin et al.,
2021b) but there have been few quantitative studies of individual
assemblages (Prevec et al., 2009,2010). Labandeira (2006) identified
four distinct phases of land plant–arthropod associations through
deep time, with the EPE as the boundary between Herbivore
Expansion Phases 2 and 3 (Figure 1). Most of the major
functional feeding groups (FFGs) had evolved by the EPE,
specifically during Herbivore Expansion 2, with the exception of
mining and nectarivory (Labandeira, 2006;Schachat et al., 2014).
Although putative evidence for leaf mining, has been reported on
Glossopteris leaves from the Permian of South America (Adami-
Rodrigues et al., 2004;Cariglino, 2018), these records are equivocal
and, if present, mining was evidently extremely rare prior to the
Triassic (McLoughlin and Santos, 2024). The oldest unequivocal
leaf mining has been identified in leaves of the peltaspermalean seed
fern Vjaznikopteris rigida from Lower Triassic strata of the Volga
Basin in central Russia (Figure 1;Krassilov and Karasev, 2008).
Turner et al. 10.3389/fevo.2024.1419254
Frontiers in Ecology and Evolution frontiersin.org03
Parallel to the floral timeline, the late Permian to Early Triassic
was a pivotal evolutionary interval for insects when the Paleozoic
Evolutionary Fauna was largely replaced by the Modern
Evolutionary Fauna (terms sensu Labandeira, 2005). The floral
and faunal turnovers of the EPE resulted in novel plant–
arthropod associations being established by the Middle Triassic
that characterize Herbivore Expansion Phase 3, but recorded
instances of PAIs for the Early Triassic are sparse (Labandeira,
2006;Labandeira et al., 2018). Comparatively little is known about
the transitional recovery flora and associated herbivores of the first
five million years of the Triassic because there are relatively few
well-preserved or well-documented fossil floras in continental
deposits from the Early Triassic globally (Grauvogel-Stamm and
Ash, 2005;Wappler et al., 2015). The Early Triassic spans a c. five-
FIGURE 1
A summary of latest Permian to early Late Triassic global climatic, entomofaunal and Gondwanan floristic events. The widths of each bubble are
indicative of semi-quantitative, relative abundances within each group, but not between groups. Plant–arthropod associations from Labandeira
(2006). Evidence of Early Triassic mining from Krassilov and Karasev (2008), but precise age is unknown. Previously, Palaeodictyopteroidea were
considered to have been totally extinguished at the EPE (Labandeira and Sepkoski, 1993), but Late Triassic occurrences of probable
palaeodictyopterans have since been recovered (see Labandeira, 2005). Evidence of pre-EPE Dicroidium from Kerp et al. (2006) and Abu Hamad
et al. (2008); for evidence of relictual glossopterids, see main text. Other floristic trends of Gondwana from Retallack (1980a),Anderson et al. (1999,
2007),Hermann et al. (2011),Schneebeli-Hermann et al. (2015) and Mays et al. (2020). Floristic stages after Spalletti et al. (2003);Mays et al. (2020)
and Vajda et al. (2020). Fire regime interpretations from Mays and McLoughlin (2022). Freshwater events from Vajda et al. (2020) and Mays et al.
(2021) but have only been reported from eastern Australia. Global climatic events: EPE, end-Permian event; LSTM, late Smithian thermal maximum;
SSE, Smithian-Spathian event; EMTE, Early-Middle Triassic event. Additional abbreviations: Ca, Carnian Age; Chang., Changhsingian Age; In., Induan
Age; LT, Late Triassic Epoch.
Turner et al. 10.3389/fevo.2024.1419254
Frontiers in Ecology and Evolution frontiersin.org04
million-year interval for which published records of PAIs are,
consequently, scarce (Labandeira et al., 2016;Cariglino et al.,
2021;Labandeira and Wappler, 2023). Furthermore, the majority
of existing PAI data from the Early Triassic come from deposits of
the Northern Hemisphere (Krassilov and Karasev, 2008;
Kustatscher et al., 2014;Labandeira and Wappler, 2023).
With the numerous and extreme environmental changes
through the late Permian to the Middle Triassic, the plant and
invertebrate components of terrestrial ecosystems were altered
accordingly, and this is predicted to be reflected in the
interactions recorded in fossil plant assemblages. Extinction and
turnover of plant and invertebrate lineages, and their corresponding
trophic interactions, are likely to have resulted in a reduced
frequency of associated trace fossils prior to the recovery of these
groups (Labandeira and Currano, 2013).
Building upon recent detailed surveys of Gondwanan Permian
(Prevec et al., 2009,2010;McLoughlin et al., 2021b), Late Triassic
(Scott et al., 2004;Labandeira et al., 2018;Cariglino et al., 2021) and
Jurassic (McLoughlin et al., 2015)PAIs,thisreviewaimsto
document and quantify our current knowledge of PAIs on fossil
leaves during the post-EPE recovery interval of the Early and
Middle Triassic in the Southern Hemisphere. To this end, we
conducted a comprehensive survey of published literature on
austral fossil plant assemblages, with special emphasis on leaf
floras, given their relative abundance and suitability as substrates
of PAIs. A comparison of this data compilation to the results of a
case study on a well-described Middle Triassic floral assemblage
provides an independent gauge of the rate of PAI reporting. The
review of the literature, and additional new case study, will: 1, assess
the reliability of the present state of the fossil record as a gauge of
terrestrial ecosystem health; 2, summarize the broad ecological
stages represented by the plant–arthropod trace fossil records of
the Early and Middle Triassic of Gondwana (with regional
comparisons); 3, assess PAI presence in the eastern Australian
Middle Triassic Benolong Flora, for which no herbivore damage
has yet been reported; 4, establish a series of hypotheses regarding
terrestrial ecosystem collapse and replacement that are testable with
the late Paleozoic to early Mesozoic fossil record of plant–arthropod
interactions; and 5, identify fossil assemblages for further research
to address the gaps in our collective knowledge of Early to Middle
Triassic Gondwanan floras and arthropod interactions.
2 Materials and methods
2.1 Data selection and database taxonomy
The literature evaluation and database creation (Supplementary
Table S1) involved the following inclusion criteria: 1, fossil
assemblages yielding plant material; 2, Early and Middle Triassic
interpreted ages (including those with ages that may extend into the
late Permian or Late Triassic); and 3, localities from the Gondwanan
supercontinent. Some temporally relevant stratigraphic units
bearing fossil flora from across the Southern Hemisphere were
initially identified from the reports of Retallack (1980a),Anderson
and Anderson (1993a,b),Escapa et al. (2011),Cariglino et al. (2021)
and Moisan (2024). For every fossil plant assemblage identified,
additional searches were conducted to identify any existing PAI
records. The primary means of finding relevant papers was by using
varied combinations of search terms in Google and Google Scholar.
The search terms consisted of: “XX Formation”+“XX Basin”+
“Triassic”+“fossil”+“plant”+“leaf”/“leaves”+“flora”+
“palaeobotany”+“Dicroidium”+“insect”/“arthropod”+
“damage”/“herbivory”; searches consisted of a combination of
keywords with and without quotation marks.
Some of the identified fossil assemblages have poor age controls,
and the chronostratigraphic boundaries of some lithostratigraphic
units may extend beyond the target interval (Middle to Upper
Triassic) but contain some beds of relevant age. These instances are
indicated in the database (Supplementary Tables S1,S2). Some
sources report Lower Triassic stratigraphic units containing
specimens typical of pre-EPE late Permian plant assemblages,
such as Glossopteris (in Australia, Jordan, Madagascar and South
Asia). Although some of these recorded fossil assemblages are likely
to be of Early Triassic age, the formations that contain these beds
may: 1, have poor age constraints; 2, span the Permian-Triassic
boundary; and/or 3, the identified fossils may be stratigraphically
misplaced. Some specimens documented by previous authors may
also have been misidentified. Resolving these possibilities is beyond
the scope of the present study. Some records of plant assemblages
have not been assigned to a particular lithostratigraphic unit;
instead, they were listed according to their positions relative to
identifiable units, or the localities at which they were found. For
some units indicated by previous authors to contain plant
macrofossils, it was not possible to locate all the primary
literature. In such cases, non-peer reviewed conference abstracts,
and unpublished doctoral and bachelor’s theses were utilized, and
these were indicated in the database. Despite these efforts, some
units included in the database host very few confirmed species- or
genus-level records, and the authors that have indicated additional
macrofossils have been cited.
Lithostratigraphic units were grouped into time bins: latest
Permian (Lopingian)–Early Triassic (LP–ET), Early Triassic only
(ET), Early to Middle Triassic (ET–MT), Middle Triassic only (MT)
andMiddletoLateTriassic(MT–LT). Assemblages of more
uncertain age spanning the Early to Late Triassic (ET–LT) were
not included in analyses (Supplementary Table S2).
In addition to leaves, all macrofossil plant materials reported in
the compiled literature were entered into the database, including
wood, reproductive organs, seeds, and unidentifiable fragments
(Supplementary Table S1); plant microfossil records (micro-/
megaspores and pollen) were not included. Owing to their
prevalence in the fossil record, and their diverse and diagnostic
forms, this study focused primarily on fossil leaf damage types.
Records of Early to Middle Triassic woods and seeds are sparse, and
these data were not compiled systematically herein. Similarly, this
review deals with damage caused primarily by terrestrial
invertebrate species and excludes any diagnostic herbivory,
detritivory or incidental damage caused to plant remains by
aquatic invertebrate taxa (Philippe et al., 2022).
For every stratigraphic unit, each distinct report of a plant taxon
was entered as one record in the database. Duplicate reports
Turner et al. 10.3389/fevo.2024.1419254
Frontiers in Ecology and Evolution frontiersin.org05
attributed to the same formation (e.g., reports of the same species
collected in separate sampling expeditions) were combined into one
record (Supplementary Table S1). In an attempt to standardize the
database with other eastern Australian Early to Middle Triassic
floras and minimize the potential for taxonomic inflation,
classifications of taxa higher than genus rank follow the scheme
used for description of the Nymboida flora (eastern Australia),
which has been documented extensively (Holmes, 2000,2001a,
2003;Holmes and Anderson, 2005a,b,2007;2008,2013a,2022;
Holmes et al., 2010). Other plant species, genera and higher rank
taxa were included in the database with the original designations
utilized by the cited authors; however, designations were updated in
cases where typographical errors were identified. Consequently,
there is some potential for taxonomic inflation as some genera and
species may have been reassigned in subsequent studies. For
example, Holmes and Anderson (2005a) designated the following
genera as junior synonyms of Dicroidium:Dicroidiopsis,
Diplasiophyllum,Johnstonia,Tetraptilon,Xylopteris and Zuberia.
Misidentifications of specimens and leaf features (incorrect taxa or
pseudotraces) by previous authors may also be included.
For temporal and spatial comparison of floral composition,
records were organized into fourteen groups in accordance with the
aforementioned taxonomic designations: bryophytes, lycophytes,
ferns and sphenophytes (spore producers); bennettitaleans,
conifers, cordaitaleans (sensu the original authors), cycads,
ginkgophytes, gnetopsids, ‘seed ferns’(i.e., Umkomasiales,
Peltaspermales and affiliated groups), gymnosperm incertae sedis
(seed producers); and other taxa of uncertain affinity incertae sedis.
In contrast to the “incertae sedis”group that was defined by
uncertainty in higher taxonomic affinity of the genera by the
original authors, higher taxonomic affinities for 61 plant records,
reported only as a genus or genus + species by the cited sources,
were not able to be identified using the search protocol outlined
above and were placed into an additional group denoted by a
question mark “?”.
In many cases, the plant fossils were not identifiable to species
level and, instead, are only reported to genus or even higher
taxonomic levels (referred to as suprageneric records herein).
Records containing indications of uncertainty in the genus or
species identifications, or taxa in open nomenclature (“sp.”,
“spp.”,“cf.”,“aff.”,“?”, etc.) were included in the database and
subsequent analyses where they were informative of floral diversity.
There are 191 plant fossil records where the number of species
discussed was unclear and thus were not included in analyses of
counts of species reports from each locality (Supplementary Table
S3): 129 suprageneric records (including three records where two
genera were indicated as equally probable), and 62 records only
specified to genus level, 11 of which were reported as “spp.”
indicating the presence of at least one distinct fossil-species from
a given genus.
2.2 PAI case study
To determine whether the currently reported levels of
arthropod herbivore damage are representative of Triassic
assemblages across Gondwana, we also conducted a case study of
the Middle Triassic Benolong Flora of the Napperby Formation in
the Gunnedah Basin, Australia (Supplementary Table S4). The
Benolong Flora is from a site at Ugothery, near Benolong, New
South Wales. All specimens investigated in this study (N= 591) are
housed in the Australian Museum Palaeontological Collection,
Sydney, Australia. For data collection, all hand specimens c. ≤40
cm long were photographed using a Canon 80D camera with a 50
mm Canon macro lens. Dust and loose particles were removed from
the specimens prior to imaging using an air blower brush tool.
Photography of each sample was conducted under three light
conditions: one ‘ambient’high-angle light condition, and two
conditions of low-angle light with orthogonal light directions. For
each light condition, a series of photos were taken at slightly
different focal lengths. Each photo series was then merged into a
composite focused image using Helicon Focus v.8.2.2. Individual
hand specimens were typically imaged with a consistent field of
view and, hence, focal length (notwithstanding the minor length
changes for the composite image-stitching outlined above).
However, average focal length varied between samples (c. 30–100
cm), thus resulting in an inconsistent spatial resolution.
All fossils identifiable as individual plant fragments were
included in the analyses. For each rock surface, the number of
distinct plant fragments was counted, and assessed for PAIs using
the DT/FFG categories described by Labandeira et al. (2007) and
later unpublished additions (C. Labandeira, pers. comm.). Physical
herbivore defenses, such as hairs or spines, were also documented.
Artefacts that could be misidentified as PAIs include abiotic or
non-herbivore biotic damage to the plant prior to fossilization, the
effects of taphonomic processes, extraction marks, overlying
sediment, and adhering dust. The presence of reaction tissue—
produced by the living plant around the damaged area to limit tissue
infection—was utilized to more confidently identify biotic damage
inflicted on living plant tissue. In this way, traces of herbivory were
distinguishable from detritivore damage on dead plant material and
subsequent taphonomic artefacts. Furthermore, stereotypy, the
repeated pattern of size, shape and position of damage, was used
to distinguish PAI evidence from abiotic environmental damage to
the plant (Labandeira et al., 2016;Currano et al., 2021). Imaging the
fossil using low-angle light from orthogonal directions highlighted
low-relief, positive and negative topographic features on the rock
surfaces, allowing for areas where fossil layers were absent or
covered by overlying sediment to be distinguished from plant
fossil areas removed by herbivory.
Additional potential confounding variables include the
difficulty in accurately counting specimens on rock surfaces with
multiple overlapping plant fragments. Moreover, over the course of
examining 591 plant fragments, it is possible that the accuracy of
the assessment improved due to increased familiarity with the DT
guide and assessors’skills in recognizing more subtle PAIs and in
excluding pseudotraces. As noted by Thomas and Hill (2023), none
of the images used to assign a PAI to the DT categories of
Labandeira et al. (2007) is of an Australian specimen. Moreover,
most of the specimens illustrated by Labandeira et al. (2007) are:
1, of much younger material than the Middle Triassic Benolong
Flora; and 2, consist almost exclusively of angiosperms, a group of
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plants for which there is no unequivocal Middle Triassic evidence.
Hence, some variation in the expression of these FFGs is expected
and a significant proportion of PAIs identified were not confidently
assignable to a DT, but were referred only to an FFG.
3 Results
3.1 Database: localities and
stratigraphic units
Eighty-nine stratigraphic units yielding 1559 plant records were
included in the Gondwanan Early–Middle Triassic fossil database
(Figure 2;Supplementary Tables S1,S2): six from Africa, three from
Antarctica, two from Jordan, fifty from Oceania, twenty-three from
South America, and five from South Asia.
3.1.1 Late Permian to Early Triassic assemblages
Fossil leaves of latest Permian to earliest Triassic age (i.e., close
in age to the EPE) are represented in South Asia from the Mianwali
Formation (Schneebeli-Hermann et al., 2015) and the Nidpur beds
and adjacent strata (Bose and Srivastava, 1971;Banerji et al., 1976;
Srivastava, 1988;Bhowmik and Parveen, 2014). Stratigraphic units
of other Gondwanan regions that span this interval are: the Puesto
Tscherig Formation (Argentina) and the Sakamena Formation
(Madagascar) (Carpentier, 1935,1936;Artabe, 1985a,b;Spalletti
et al., 1999;Falco et al., 2020). This time bin contains 96 records
(including seven genus-only/“spp.”records and eight
suprageneric records).
The Jordanian Umm Irna Formation, of putative latest Permian
age (LP; Supplementary Table S2), hosts abundant Dicroidium and the
floral records and PAI reports were included in the plant fossil database
for comparison (57 plant records including one suprageneric record)
(Abu Hamad et al., 2008,2017;Blomenkemper et al., 2018,2019,2020,
2021;Kerp et al., 2021,2024). However, there are no strata that extend
into the specified Early to Middle Triassic age range and so the floral
records were not included in any further analyses.
3.1.2 Early Triassic assemblages
The majority of records of Early Triassic leaves are from Australian
strata: the Bald Hill Claystone, Bulgo Sandstone, Munmorah
Conglomerate, Tuggerah Formation, a single species from the Banks
Wall Sandstone, and two plant assemblages with imprecise stratigraphic
provenance, are all from the Sydney Basin (Raggatt, 1969;Helby, 1970;
Balme and Helby, 1973;Retallack, 1980a). In addition to the Sydney
Basin localities, there is also the Camden Head Claystone and Grants
FIGURE 2
Distribution maps of Early and Middle Triassic Gondwanan basins indicating the numbers of localities (n) included in the plant fossil database. Green
circles indicate >10 localities, orange circles indicate 3–10 localities, red circles indicate <3 localities. Regions with high concentrations of basins
have been combined for simplicity and are represented by larger circles. Central eastern Australian basins: the Bowen, Gunnedah, Lorne, Nymboida
and Sydney basins and the Esk and Warialda troughs. Southern South American basins: the Cuyo/Cuyana, El Tranquilo, Los Menucos and San Rafael
basins of Argentina and the Antofagasta, Atacama, Coquimbo and Valparaıso regions of Chile. Southeastern African basins: the Luangwa and Ruhuhu
basins. Basins in bold have relevant localities across both the Early and Middle Triassic. Fossiliferous units spanning the Early to Middle Triassic have
been duplicated into both age categories as they likely represent transitional floras and may contain assemblages from either, or both the Early and
Middle Triassic.
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Head Formation (one reported species) from the Lorne Basin, the
Arcadia Formation from the Bowen Basin, the Blina Shale and Millyit
Sandstone from the Canning Basin, the Kockatea Shale from the Perth
Basin, and the Knocklofty Formation from Tasmania (White and
Yeates, 1976;Ash, 1979;Holmes and Ash, 1979;Chaloner and
Turner, 1987;Retallack, 1995,1997;Cantrill and Webb, 1998;Haig
et al., 2015). Non-Australian Early Triassic assemblages include those
from the Flagstone Bench Formation (Ritchie Member) in Antarctica
and the Katberg and Verkykerskop formations from South Africa
(McLoughlin et al., 1997;Gastaldo et al., 2005;Gastaldo and
Bamford, 2023). This investigation did not identify any records of
plant macrofossil assemblages of definitive Early Triassic age from the
Arabian Peninsula, South America or South Asia.
This time bin contains 139 records (including 16 genus-only
records and 27 suprageneric records).
3.1.3 Late Early to early Middle
Triassic assemblages
The fossil-bearing stratigraphic units that may span the Lower to
Middle Triassic boundary in Gondwana are largely from Australia:
the Burralow, Garie, Gosford and Terrigal formations of the Sydney
Basin, the Clematis Sandstone and Glenidal Formation of the Bowen
Basin, and the Culvida and Erskine sandstones from the Canning
Basin (White, 1965;Goldbery, 1969;Raggatt, 1969;White, 1971;
White and Yeates, 1976;Retallack, 1977;Ash, 1979;Retallack, 1980a;
Chaloner and Turner, 1987;Holmes, 2001b;McLoughlin, 2011a).
The Murihiku Supergroup from New Zealand includes the Potiki
Siltstone and the Wairuna Peak Beds of this age, along with some leaf
fossil occurrences not allocated to a stratigraphic unit (Retallack,
1985). The palaeoflora-bearing Burgersdorp Formation (South
Africa) and the Puesto Vera Formation (Argentina) have also been
attributed to the Early to Middle Triassic (Anderson and Anderson,
1985;Artabe, 1985b,a;Bamford, 1999;Spalletti et al., 1999;Hancox
et al., 2002;Bamford, 2004;Gastaldo et al., 2005;Falco et al., 2020;
Hancox et al., 2020).
This time bin contains 159 records (including seven genus-
only/”spp.”records and 15 suprageneric records).
3.1.4 Middle Triassic assemblages
Fossil leaves confidently ascribed to the Middle Triassic are also
described predominantly from Australian sedimentary basins.
Fossiliferous units of this interval include the Gragin Conglomerate
and Gunnee Beds from the Warialda Trough, the Napperby
Formation from the Gunnedah Basin, the Ashfield Shale, Bringelly
Shale, Brookvale Shale, Hawkesbury Sandstone and the Newport
Formation from the Sydney Basin, the Moolayember and Teviot
formations from the Bowen Basin, the Basin Creek Formation from
the Nymboida Basin, and the Bryden and Esk formations, together
with a single record from the Neara Volcanics in the Esk Trough
(Walkom, 1924,1928;White, 1965;Branagan, 1969;Jell, 1969;
Lovering and McElroy, 1969;White, 1969,1971;Bourke et al.,
1977;Rigby, 1977;Retallack, 1980a;Holmes, 1982;Playford et al.,
1982;Webb, 1982;Webb and Holmes, 1982;Webb, 2001). Also from
the Sydney Basin, another small assemblage reported as “Mittagong
Formation or basal Ashfield Shale”by Retallack (1980a, p. 426), and
one unit yielding only one species, the Minchinbury Sandstone, were
included (Lovering and McElroy, 1969). From New Zealand, the
Black Jacks Formation and the Tank Gully Coal Measures are
assigned to the Middle Triassic (Retallack, 1980b,1983), along with
the remaining Murihiku Supergroup units and unspecified
assemblages (Retallack, 1985): The North Etal Group, the North
Peak (Jacobs River Member) and Pears formations, an assemblage
“southeast of Mautaura island”(p. 11), an assemblage “above the
Franklin Conglomerate”(p. 9) and the ‘Tauringatura Group’(p. 13),
the latter two with only one species reported each. The remaining
Middle Triassic strata are: 1, the Mukheiris Formation from Jordan
(Abu Hamad et al., 2019), and; 2, the Barreal, Cerro de Las Cabras,
Corral de Piedra, Monina, Montaña, Paramillos, and Quebrada de los
Fosiles formations from Argentina (Brea et al., 2008;Cariglino et al.,
2016,2018;Bodnar et al., 2019;Pedernera et al., 2019;Drovandi et al.,
2020;Pedernera et al., 2022).
This time bin contains 582 records (including 19 genus-only/
“spp.”records and 39 suprageneric records).
3.1.5 Middle to Late Triassic assemblages
Formations with strata indicated to extend from the Middle
Triassic into the Upper Triassic are located predominantly in South
America, the vast majority of which have uncertain age controls:
1, the Cañadon Largo Formation from the El Tranquilo Basin
(Herbst, 1988;Jalfin and Herbst, 1995;Gnaedinger and Herbst,
1998a,b,1999;Troncoso et al., 2000;Herbst et al., 2001;Herbst
and Gnaedinger, 2002;Adami-Rodrigues et al., 2008;Villalva, 2017;
Gnaedinger et al., 2023;Villalva et al., 2023); 2, the Agua de la Zorra,
Cortaderita (Don Raul Member), Agua de Los Pajaritos and
Potrerillos formations from the Cuyo/Cuyana Basin in Argentina
(Ottone et al., 2011;Cariglino et al., 2016;Lara et al., 2017;Bodnar
et al., 2019;Pedernera et al., 2019); 3, the El Bordo and El Mono beds,
and the Pular, Tuina, Guanaco Sonso, Las Breas and El Puquen
formations of Chile (Azcarate and Fasola, 1970;Breitkreuz et al.,
1992;Lutz et al., 1999;Troncoso and Herbst, 1999;Herbst, 2000;
Herbst and Troncoso, 2000;Melchor and Herbst, 2000;Troncoso
et al., 2000;Gnaedinger and Herbst, 2001;Herbst and Gnaedinger,
2002;Gnaedinger and Herbst, 2004a,b;Niemeyer et al., 2008;
Moisan, 2024); and 4, the Santa Maria Formation of Brazil
(Guerra-Sommer and Cazzulo-Klepzig, 2000;Cenci, 2013;Barboni
and Dutra, 2015;Barboni et al., 2016;Cenci and Adami-Rodrigues,
2017;Cenci et al., 2019). Very few plant species have been identified
from the Tuina Formation and El Mono beds in this review.
Similarly, only four species were reported from the Ntawere
Formation in Zambia, and only one plant species was identifiable
from the Manda Formation of Tanzania, which is poorly dated but
suggested to be within this age range (Dixey, 1937;Lacey and Smith,
1970). The rich plant assemblages from the permineralized peat (and
associated beds) of the upper Fremouw Formation of the
Transantarctic Mountains (Schopf, 1978;Stubblefield and Taylor,
1986;Perovich and Taylor, 1989;Taylor et al., 1989;Millay and
Taylor, 1990;Pigg, 1990;Delevoryas et al., 1992;Meyer-Berthaud
et al., 1993;Taylor et al., 1994;Del Fueyo et al., 1995;Yao et al., 1995,
1997;Axsmith et al., 1998;Phipps et al., 2000;Klavins et al., 2002;
Rothwell et al., 2002;Cuneo et al., 2003;Klavins et al., 2003;Kellogg
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and Taylor, 2004;Klavins et al., 2004;Retallack et al., 2005;Hermsen
et al., 2006,2007a,b;Decombeix et al., 2010;Schwendemann et al.,
2010;Bergene et al., 2013;Decombeix et al., 2014) have commonly
been assigned to the Middle (or even Lower) Triassic (Taylor et al.,
1986,1993). However, palynoassemblages from these beds are
preferable to the Alisporites Assemblage Zone C or Australian
APT4 zone, indicating a possible Carnian (Late Triassic) age
(Farabee et al., 1990). The Antarctic Lashly Formation and the
Long Gully Formation of New Zealand have also been assigned to
this geochronological interval (Retallack, 1981;Escapa et al., 2011;
Bomfleur et al., 2013b).
This time bin contains 444 records (including ten genus-only/
“spp.”records and 33 suprageneric records).
3.1.6 Assemblages of Early to Late Triassic or
uncertain age
There are three remaining formations in the database, indicated
to be in the range of Lower to Upper Triassic without more precise
stratigraphic resolution: the Chilean San Felix Formation (Mohr
and Schöner, 1985;Bell and Suarez, 1994;Troncoso et al., 2000;
Salazar et al., 2013) and the Indian Panchet and upper Kamthi
formations (Bose and Banerji, 1974;Banerji et al., 1976;Banerji and
Bose, 1977;Pal and Ghosh, 1997;Pal et al., 2010,2014;Saxena et al.,
2019;Ghosh et al., 2021). As plant fossil records from these
formations (82 records, including three genus-only/“spp.”records
and six suprageneric records) cannot be grouped into any particular
time bin or used for comparison across the Early to Middle Triassic,
they were not included in the analyses.
Additional units in India are possibly referable to the Middle
Triassic (e.g., the Pali, Parsora, and Panchmarhi formations) but age
constraints are currently insufficient for confident assignment of
fossil assemblages from these units to any particular time bin (Joshi
et al., 2014).
3.2 Floral composition
Gondwanan terrestrial ecosystems were dominated by glossopterid
gymnosperms throughout most of the Permian (McLoughlin, 2011b).
The end-Permian event in Gondwana was marked by the demise of
glossopterids (Figure 1) and many associated free-sporing and
gymnospermous plants (Anderson et al., 1998;Retallack et al., 2005;
Lindström and McLoughlin, 2007;Fielding et al., 2019;Gastaldo and
Bamford, 2023). Reports of glossopterids surviving the end-Permian
extinction event and persisting into the earliest Triassic are equivocal.
Records of glossopterids in units close to or spanning the Permian-
Triassic boundary collated in the present database derive from: 1, the
Umm Irna Formation (LP; Jordan; Kerp et al., 2021); 2, the Sakamena
Formation (LP–ET; Madagascar; Carpentier, 1935,1936); 3, the
Munmorah Conglomerate, Blina Shale, Millyit Sandstone (ET;
Australia) the Erskine Sandstone (ET–MT; Australia) (Raggatt, 1969;
White and Yeates, 1976); and 4, several units in India, including the
Nidpur beds and associated strata, the Mianwali Formation (all LP–
ET), and the Panchet and Kamthi formations (ET–LT)
(Supplementary Table S1;Banerji et al., 1976;Banerji and Bose,
1977;Srivastava, 1988;Pal and Ghosh, 1997;Pal et al., 2010;
Schneebeli-Hermann et al., 2015;Saxena et al., 2019). Additional
Indian strata hosting putative Glossopteris of possible Triassic age are
the Pali, Hinjir, Maitur, Parsora, and Hirapur/“Upper Kamthi”
formations (Maheshwari, 1992;Shah, 2000;Ghosh et al., 2016;Shah,
2021); however, these records may represent misidentifications of
typical Triassic taxa, such as Gontriglossa. With the exception of the
Munmorah Conglomerate and Blina Shale, Dicroidium has also been
reported from these same strata (Carpentier, 1935,1936;Raggatt, 1969;
Bose and Srivastava, 1971;Bose and Banerji, 1974;Banerji et al., 1976;
White and Yeates, 1976;Srivastava, 1988;Maheshwari, 1992;Retallack,
1995;Pal and Ghosh, 1997;Shah, 2000;Pal et al., 2014;Schneebeli-
Hermann et al., 2015;Saxena et al., 2019;Kerp et al., 2021). Srivastava
et al. (2010) also reported Dicroidium associated with Glossopteris from
Lower Permian strata of the Barakar Formation, however, the putative
Dicroidium specimens were later reinterpreted as peltaspermalean
foliage (Srivastava et al., 2011). Other putative co-occurrences of
Dicroidium and Glossopteris recorded in this database may include
conflations of pre-EPE Permian and post-EPE Permian or Triassic
plant assemblages that have been grouped into the same formation or
have become stratigraphically misplaced by faulted juxtapositioning of
beds of different ages (Tiwari and Ram-Awatar, 1990). Although we
cannot exclude the possibility that some glossopterids persisted as post-
EPE survivors, this conclusion awaits verification. No Glossopteris has
yet been identified in any of the Antarctic or South American Triassic
units included in this database, although Rigby and Schopf (1969)
report the co-occurrence of thesetaxainanAntarcticassemblage
they determined to be equivalent to the Panchet Stage (no
formation indicated).
Later in the Early Triassic, phases of alternating dominance by
pleuromeian lycopsids, voltzialean conifers and Dicroidium
(Umkomasiales), together with spikes in charcoal abundance,
indicate repeated fluctuations in environmental conditions
(Retallack, 1977;Helby et al., 1987;Mays et al., 2020;Mays and
McLoughlin, 2022). By the end of the Early Triassic, Umkomasiales
had become established as the dominant gymnosperm group across
southern Gondwana (Figures 1,3;Retallack, 1977;Anderson and
Anderson, 2003).
Through the Middle Triassic, Umkomasiales diversified and
retained dominance of the Gondwanan temperate vegetation
(Retallack, 1977;Anderson and Anderson, 1985;Taylor et al., 1993).
They were associated with a broad range of sphenophytes, ferns, broad-
leafed conifers, cycads, bennettitopsids, peltasperms, voltzialean
conifers and various gymnosperms of uncertain affinity (Figures 3,4;
Walkom, 1928;Anderson and Anderson, 1985;Taylor et al., 1993;
McLoughlin et al., 2018b). Overall land-plant diversity continued to
increase in the austral temperate zone (>35°S) through the Middle and
Late Triassic (Anderson and Anderson, 1989,2003), before suffering
depletion and replacement of key taxa during the end-Triassic
extinction event (Anderson et al., 1998;Kustatscher et al., 2018).
Afloral transition is evidenced in the present database by the
higher proportion of bryophytes, lycophytes, sphenophytes and
conifers in the Early Triassic compared to the Middle Triassic,
whereas ferns, bennettitaleans, cycads, ginkgophytes, gnetopsids
and gymnosperms of uncertain affinity constitute a greater
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FIGURE 3
Gondwanan Early and Middle Triassic seed fern diversity. The number of records of each order in the plant fossil database is reported. The
“uncertain”group consists of fossil records where authors have indicated affinity to seed ferns, but for which designations of order were not
specified. Seed fern records from assemblages spanning Early to Middle Triassic age have been included in both age categories: Early Triassic
(n=123); Middle Triassic (n=407).
FIGURE 4
Comparison of Gondwanan Early and Middle Triassic floral composition. The number of records of each higher taxonomic group in the plant fossil
database is reported. Uncoloured bars represent Early to Middle Triassic plant fossil records, with equal values above the data for both the Early
Triassic and Middle Triassic age categories. Higher taxonomic groups follow the same sequence for all charts: bryophytes; lycophytes; ferns;
sphenophytes; bennettitaleans; conifers; cordaitaleans; cycads; ginkgophytes; gnetopsids; seed ferns; gymnosperm incertae sedis;incertae sedis;“?”.
The “?”group consists of plant fossil genera for which higher taxonomic designations were unable to be identified in this review, in contrast to the
incertae sedis group defined by uncertainty in higher taxonomic affinity of the genera. Pie chart values represent proportional contributions of each
higher taxonomic group to the total floral records in that age category (in %). Plant fossil records from assemblages spanning Early to Middle Triassic
age have been included in the pie charts of both age categories; total Early Triassic records (n=394); total Middle Triassic records (n=1185).
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proportion of Middle Triassic diversity than in the Early Triassic
assemblages (Figure 4). Seed ferns, including umkomasialeans and
peltasperms, although dominant in both Early and Middle Triassic
assemblages, constitute a greater proportion of Middle Triassic total
species richness, demonstrating their continued diversification
(Figures 3–5). The total number of plant species reported
increases in the Middle Triassic, particularly in Oceania and
South America (Figures 3–5,Supplementary Figure S1;
Supplementary Table S3). This likely represents a combination of
recovery in species diversity after the EPE, and increased collection
and taxonomic identification efforts for Middle Triassic
assemblages compared to those of the Early Triassic.
3.3 PAI record of Gondwana
Of the 85 stratigraphic units included in the Gondwanan Early–
Middle Triassic analyses (Supplementary Table S1), only twelve
(14.1%) yielded PAIs reported in the surveyed literature (Table 1).
3.3.1 Africa
We could find no reports of insect damage from Early to Middle
Triassic African floral assemblages. Labandeira et al. (2018) noted
the conspicuous absence of galls from a collection of 1386 plant
specimens of the Burgersdorp Formation (ET–MT).
3.3.2 Antarctica
The majority of PAI records from the late Middle to early Late
Triassic of Antarctica derive from the Fremouw Formation, with
several organ types affected, although no PAIs have yet been found on
leaves. These trace fossils include evidence of boring—likely caused
by oribatid mites—in 220 specimens of stems, petioles and roots
including a petiole of the fern Antarctipteris (Kellogg and Taylor,
2004). This assemblage also yielded insect coprolites containing
pollen located in the cycad pollen cone Delemaya spinulosa
(Klavins et al., 2005). Additionally, Hermsen et al. (2006,2009)
identified feeding galleries in stems of the cycad Antarcticycas
schopfiiof the Fremouw assemblage, which was likely caused by
wood-boring arthropod larvae. Fungal damage has also been
observed in almost eighty specimens of Araucarioxylon-type wood
and one specimen of the coniferous ovulate organ Parasciadopitys
from this formation (Stubblefield and Taylor, 1986;Bomfleur et al.,
2013a). Bomfleur et al. (2013a) also reported fungal hyphae on one
coniferous leaf of Heidiphyllum from the Lashly Formation of the
same age. However, as noted above, the ages of the upper Fremouw
and Lashly formations may be Late Triassic.
FIGURE 5
Gondwanan Early and Middle Triassic floral composition by region. The number of records of each higher taxonomic group in the plant fossil
database is reported. Higher taxonomic groups follow the same sequence for both charts: bryophytes, lycophytes, ferns, and sphenophytes (spore
producers); bennettitaleans, conifers, ‘cordaitaleans’, cycads, ginkgophytes, gnetopsids, seed ferns, and gymnosperm incertae sedis (seed producers);
incertae sedis;“?”. The “?”group consists of plant fossil genera for which higher taxonomic designations were unable to be identified in this review,
in contrast to the incertae sedis group defined by uncertainty in higher taxonomic affinity of the genera. Plant fossil records from assemblages
spanning Early to Middle Triassic age have been included in both age categories. Early Triassic taxon records: Africa (n=57); Antarctica (n=54);
Arabian Peninsula (n=0); Oceania (n=241); South America (n=28); South Asia (n=65). Middle Triassic taxon records: Africa (n=29); Antarctica (n=54);
Arabian Peninsula (n=2); Oceania (n=631); South America (n=469); South Asia (n=0). Functional feeding groups of reported plant–arthropod
interactions are indicated beneath the chart for each region.
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TABLE 1 Records of plant–arthropod interactions (PAIs) from formations in the Gondwanan plant fossil database.
Gondwanan
region
Age
group
Formation Margin
Feeding
Hole
Feeding
Skeleton-
isation
Surface
Feeding
Piercing
& Sucking
Mining Galling Oviposition Boring Pollinivory Pathogen Uncertain
PAI
References
Antarctica MT-LT Fremouw >220 ≥1 ~80 a, b, c,
d, e, f
Antarctica MT-LT Lashly 1f
Oceania MT Basin Creek 1 g
Oceania MT Esk 11h
Oceania MT Napperby 23 2 3 6 10 11 *
Oceania MT Newport 1 h
South America MT Quebrada de
los Fosiles ≥1≥1i
South America MT-LT Cañadon Largo (X) (X) (X) (X) (X) 1 (X) i, j, k
South America MT-LT Las Breas 2l
South America MT-LT Potrerillos 6 2 1 1 2 1 1 m, n
South America MT-LT Santa Maria >1 >1 >7 >164 >4 3 o, p, q
South Asia LP-ET Nidpur Beds 1r
PAI traces have been identified on fossil plants from formations in the Late Permian-Early Triassic (LP-ET), Middle Triassic (MT), and Middle Triassic-Late Triassic (MT-LT) age categories. Reported numbers of affected plant fossils are indicated. (X) indicates records
from a combination of the Cañadon Largo and Laguna Colorada (LT) formations. References: a, Stubblefield and Taylor (1986);b,Hermsen et al. (2006);c,Hermsen et al. (2009);d,Klavins et al. (2005);e,Kellogg and Taylor (2004);f,Bomfleuret al. (2013a);g,Holmes and
Anderson (2007);h,McLoughlin (2011a);i,Cariglino et al. (2021);j,Adami-Rodrigues et al. (2008);k,Gnaedinger et al. (2023);l,Gnaedinger et al. (2014);m,Cariglino et al. (2022);n,Lara et al. (2017);o,Cenci(2013) ;p,Cenci and Adami-Rodrigues (2017);q,Cenci et al.
(2019); and r, Bhowmik and Parveen (2014); *Benolong Flora case study presented herein (bold text).
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3.3.3 Arabian peninsula
Hole feeding on three Dicroidium irnensis cuticles from the late
Permian Umm Irna Formation (Jordan) was identified by
McLoughlin et al. (2021a). However, no definitive Triassic PAIs
have yet been reported.
3.3.4 Oceania
This review identified three Middle Triassic Australian
formations hosting examples of PAI traces on fossil plants.
Holmes and Anderson (2007) identified margin feeding on a leaf
apex of the ginkgoalean Sphenobaiera densinerva from the Basin
Creek Formation (MT). McLoughlin (2011a) recognized
oviposition scars, possible margin feeding and either hole feeding
or surface feeding on a leaf of Taeniopteris parvilocus from the Esk
Formation (MT) and galling on a Dicroidium odontopteroides leaf
from the Newport Formation (MT). The present study describes
several newly identified PAIs from the Benolong Flora (Napperby
Fm, MT); see section 3.4 below for more details.
3.3.5 South America
Permian and Triassic PAI records from South America were
extensively compiled and reviewed by Gnaedinger et al. (2014),
Cariglino et al. (2021) and Romero-Lebron et al. (2022).An
updated summary of these occurrences is provided below (in
stratigraphic order).
The Argentinian Quebrada de los Fosiles Formation (MT)
yielded gall-bearing leaves of the seed fern Ptilozamites longifolia
(possibly = Kurtziana sp.) and oviposition traces on the frond of
another seed fern of uncertain affinity (Cariglino et al., 2020,2021).
Adami-Rodrigues et al. (2008) identified PAIs from the
Argentinian El Tranquilo Group, including the Cañadon Largo
Formation (MT–LT), and the Late Triassic Laguna Colorada
Formation. That study listed several types of external foliage
feeding, together with piercing and sucking, mining, galling and
oviposition. PAIs were identified on sphenophytes, ginkgoaleans,
conifers, gymnosperms of uncertain affinity and umkomasialean
seed ferns, with fossils of the latter group hosting the majority of
documented PAIs (Adami-Rodrigues et al., 2008;Cariglino et al.,
2021). However, it was not clear which floral remains and their
associated PAIs were recovered from strata of definitively Middle
versus Late Triassic age. Gnaedinger et al. (2023) identified
oviposition scars on a stem of the sphenopsid Equisetites lateralis,
also from the Cañadon Largo Formation.
Two occurrences of oviposition traces were reported from two
Taeniopteris sp. leaves of the Las Breas Formation, Chile (MT–LT;
Gnaedinger et al., 2014).
The flora of the Potrerillos Formation from Argentina (MT–LT)
hosts several PAIs. Lara et al. (2017) assessed 56 plant specimens,
reporting 11 specimens hosting PAI traces: margin feeding on leaves
of the umkomasialeans Dicroidium argenteum, D. odontopteroides,
Johnstonia stelzneriana and the matatiellalean Kurtziana brandmayri,
hole-feeding traces on leaves of Johnstonia sp. and Yabeiella
wielandii, surface feeding and piercing-and-sucking traces on leaves
of uncertain affinity, and oviposition traces on an indeterminate
sphenophyte stem. Cariglino et al. (2022) identified mines on leaves
of the conifer Heidiphyllum cacheutense and the gnetalean Yabeiella
wielandii, with possible oviposition or a fungal spot identified
alongside the mine in one Y. wielandii leaf.
The leaf floras of the Santa Maria Formation of Brazil (MT–LT)
have been the most extensively investigated assemblages of any
Gondwanan locality, with the most PAI records in the database
herein (Cenci, 2013;Cenci and Adami-Rodrigues, 2017;Cenci et al.,
2019). Cenci (2013) noted that, of a total of 414 plant samples from
the Santa Maria Formation, 149 (36%) hosted evidence of PAIs: 135
umkomasialean leaves (52 external foliage feeding (EFF) traces, seven
mines, 578 galls), 11 ginkgoalean leaves (three mines, 49 galls), one
voltzialean leaf (one mine), eight equisetalean stems (one mine, 22
galls) and nine incertae sedis (one EFF, two mines, 10 galls). Cenci
and Adami-Rodrigues (2017) also found galls to be especially
common in this unit, with 164 gall-bearing specimens from their
sample of 414 (39.6%), apparently representing the same dataset as
that employed by Cenci (2013). The vast majority of the galls were
identified on umkomasialean leaves. Six-hundred and fifty-nine
individual galls were identified across umkomasialean, ginkgoalean,
equisetalean and incertae sedis taxa, including leaves of Dicroidium
odontopteroides, Dicroidium sp., Sphenobaiera sp. and Taeniopteris
sp., an umkomasialean seed, and a stem of the sphenophyte
Neocalamites.Cenci et al. (2019) assessed 550 plant specimens, and
reported oviposition (six DTs), surface feeding (three DTs), piercing
and sucking (four DTs), mining (two DTs), margin feeding (one DT)
and hole feeding (one DT). They found that the majority of PAI
traces identified were hosted by the dominant umkomasialeans of the
Santa Maria Formation, though the ginkgoaleans displayed the
greatest proportion of affected specimens.
3.3.6 South Asia
To our knowledge, records of PAIs from this region are almost
entirely absent for the Early to Middle Triassic. Bhowmik and
Parveen (2014) reported evidence of potential plant-insect
interactions from the Indian Nidpur beds (LP–ET). They
suggested that clumped pollen grains inside the pollen chamber
of the pteridospermous seed Gopadispermum papillatus, indicate
the possibility of fluid feeding, palynophagy and insect pollination
in the seeds, although such pollen characters are also common in
anemophilous plants (Benson, 1908;McLoughlin et al., 2018a).
Ghosh et al. (2015) reported galls on Dicroidium leaves from the
Parsora Formation (of unspecified Triassic age and, therefore, not
included in the database).
3.4 Case study: Benolong Flora
Five hundred and ninety-one fossil leaf fragments were assessed
from the Benolong Flora (Supplementary Table S4). Fifty-five
potential traces of plant–arthropod interaction were identified on
forty-nine plant fragments (8.29% of fragments assessed;
Figures 6A,7). Forty-four of these PAI traces (78.6%), hosted by
40 plant fragments (1.1 PAIs per affected fragment), matched the
DT categories described by Labandeira et al. (2007). Margin feeding
was the most common FFG, with 23 examples referable to
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Frontiers in Ecology and Evolution frontiersin.org13
established DTs (identified on 3.72% of plant fragments; Figures 6B,
7A,B;Table 1); this was more than twice as common as oviposition
(identified on 1.69% of plant fragments) and all other recognized
(non-ambiguous) FFGs combined (Figures 6B,7C–H;Table 1).
However, margin-feeding traces may be more identifiable on
compression fossils than other FFGs as they do not require relief
to be observed and are especially visible where there is a high degree
of contrast between the fossil and the host sedimentary matrix.
Eleven traces were not consistent with any established DTs or had
characters in common with DTs of multiple FFGs (e.g., resembling
features of various hole feeding, oviposition or piercing and sucking
DTs). In such cases, these were recorded as ‘ambiguous’.
Preservational artefacts resembling leaf mine traces
(Supplementary Figure S2A) and leaf features with atypical
textures and surficial patterns (Supplementary Figures S2B–D),
potentially of biotic origin but not able to be categorized into any
FFG using the guide images, were not included in the analysis. One
fossilized structure resembled a gall with possible radiating
partitions (Supplementary Figure S2E); however, this feature was
not located on any observable plant remains and had the same color
as the host sediment and so was also excluded from analyses. Other
leaf features that highlighted the potential for the inflation of
identified PAIs included a semicircular line of discoloration on
the margin of another leaf fossil, suggestive of pathogenic, possibly
fungal, damage (Supplementary Figure S2F resembles DT058) and
the presence of pseudotraces, such as fern sori resembling the
texture of a gall (Supplementary Figure S2G resembles DT106).
Twenty-seven fossil fragmentsofleavesandstemsdisplayed
textural evidence of hairs or spines.
The data collection process in this case study, which involved
systematic photography of an entire floral assemblage, is inherently
limited by the resolution of the images. Even in cases where small
specimens were imaged, evidence of smaller PAIs, such as piercing
and sucking, was difficult to distinguish from preservational
artefacts and adhering dust on the rock surface. Another potential
source of error of this approach is the inconsistent focal length,
hence spatial resolution, of images taken of separate specimens. At
the maximum focal length, as used for the largest hand specimens,
much less detail is captured on the image, resulting in lower levels of
confidence in distinguishing PAIs from artefacts where these are
not verified by microscopic inspection. For increased validity of
inter-sample comparisons, we recommend a consistent, high-
resolution field-of-view and stereomicroscopic evaluation of each
specimen. Despite the limitations, we report abundant PAIs from
this case study of a relatively small Middle Triassic fossil
assemblage. These findings clearly demonstrate marked under-
reporting of arthropod trace fossils from the post-EPE recovery
phase, as discussed below.
4 Discussion
4.1 Patterns of Early to Middle Triassic
plant–arthropod interactions in Gondwana
4.1.1 Temporal trends and gaps
With the inclusion of the 13 Early to Middle Triassic localities
into the data of both age groups, only one of 35 localities from the
FIGURE 6
(A) Percentage of Benolong Flora fossil fragments hosting plant–arthropod interaction (PAI) traces relative to the total sample (n=591);
(B) Proportions of functional feeding groups from identified PAI trace fossils (n=55). Values indicate percentages of plant fragments hosting each
functional feeding group relative to the total sample.
Turner et al. 10.3389/fevo.2024.1419254
Frontiers in Ecology and Evolution frontiersin.org14
Early Triassic group reported any PAIs (2.9% of localities), and
eleven of 63 localities from the Middle Triassic group (17.5% of
localities) yielded PAIs (Figure 2;Table 1;Supplementary Table S2).
Our survey highlights the distinct paucity of arthropod trace fossil
records from Early Triassic floras across Gondwana, with only one
pteridospermous seed hosting putative arthropod traces reported
from the Nidpur beds of India (Table 1;Supplementary Table S1).
In contrast, at least 544 Middle and Middle-to-Late Triassic plant
fragments from eleven localities have been confirmed to host PAI
traces, from ten of eleven categories of FFG: margin feeding (5.70%
of quantifiable reports); hole feeding (0.74%); surface feeding
(0.37%); piercing and sucking (0.74%); mining (1.65%); galling
(31.62%); oviposition (3.68%); boring (40.44%); palynophagy
(0.18%); pathogenic/fungal damage (14.89%). The remaining
FFG, skeletonization, has been recognized but is possibly of Late
Triassic age (Table 1;Supplementary Table S1).
Austral temperate floras were fundamentally restructured after
the end-Permian extinction event. Earliest Triassic vegetation
consisted of low-diversity open forests dominated by lepidopterid
peltasperms and scale-leafed voltzialean conifers in the upper story,
FIGURE 7
Plant–arthropod interaction traces on Benolong Flora fossil fragments. (A) gall (top arrow) and margin feeding on two fragments of Dicroidium sp.
(F61131); (B) apical margin feeding on two fragments of Dicroidium odontopteroides (F61138); (C) oviposition scars on two fragments of
Heidiphyllum elongatum (F61501); (D) oviposition scars on Heidiphyllum elongatum (F50237); (E) possible hole feeding on Dicroidium
odontopteroides (F61140); (F) piercing and sucking scars on Dicroidium sp. leaf (F61143); (G) gall (top arrow) and an uncertain feature (F61129); and
(H) gall on seed fern rachis (F61130). Scale bars: 1 cm.
Turner et al. 10.3389/fevo.2024.1419254
Frontiers in Ecology and Evolution frontiersin.org15
and pleuromeian lycopsids, sphenopsids, and ferns in the
understory (Retallack, 1980a;Mays et al., 2020;Vajda et al., 2020;
McLoughlin et al., 2021a). There are 247 plant species across 118
genera reported from the Gondwanan Early Triassic, and 608
species from 200 genera identified from the Middle Triassic
(Supplementary Table S1). Whereas the latest Permian to Early
Triassic PAI was identified on a seed fern taxon, the Middle to Late
Triassic PAIs are distributed across ferns, sphenophytes, conifers,
cycads, ginkgophytes, seed ferns, and plants of uncertain taxonomic
affinity (Supplementary Table S1). This disparity may be due to the
relatively low productivity and slow recovery of the immediate post-
EPE flora compared to the Middle Triassic. Low productivity and
low diversity of vegetation would have contributed to fewer Early
Triassic PAIs by: 1, producing fewer leaves for potential burial and
2, sustaining a smaller and less diverse population of arthropods.
The difficulty in confidently identifying feeding traces on small
needle- and scale-shaped leaves (e.g., voltzialean leaves and
pleuromeian microphylls), owing to their diminutive size and
commonly dense occurrence, may have implications for assessing
damage frequency in the Early Triassic, when leaves of this type
were proportionately more common (Figure 4). The absence of
coals immediately post-EPE (Retallack et al., 1996) may provide
another, economic, bias for the observed trends, as historic coal-
mining efforts have produced a significant proportion of known
plant fossil collections.
4.1.2 Geographic trends and gaps
Strata from South America and Oceania contain the vast
majority of Lower and Middle Triassic palaeobotanical
assemblages (Figure 2;Supplementary Table S2), floral diversity
(Figure 5;Supplementary Figure S1;Supplementary Table S3) and
PAIs (Figure 5;Table 1), constituting 75% of Gondwanan PAI-
reporting localities and suggesting a major sampling bias towards
these regions. Of the 23 South American localities included in the
plant fossil database, five (21.7%) contain PAI traces attributable to
at least seven FFG categories, whereas in Oceania only four of 50
localities (8%) host reported PAIs, in at least five FFGs. As both
regions report sufficiently high numbers of plant species
(Supplementary Table S3), this observed difference in the number
of recorded FFGs from localities in Oceania versus South America
likely represents a bias in effort towards South American arthropod
trace fossil investigations. The PAI investigations of South America
constitute the majority of quantitative PAI assessments included in
the database (Potrerillos and Santa Maria formations in particular;
all MT–LT; Table 1), however, there is a notable lack of Lower
Triassic fossiliferous strata reported from South America
(Supplementary Tables S2,S3). As discussed in section 4.1.1, the
disparity in floral productivity between the Early and Middle
Triassic is likely to impact the probability of finding PAIs over
these two time intervals, and the greater proportion of Lower
Triassic strata in Oceania suggests another potential reason for
the observed South American PAI report abundance.
The Lashly Formation (Antarctica; MT–LT) yielded one leaf
hosting fungal hyphae, the Fremouw Formation (Antarctica; MT–
LT) and the Nidpur beds (India; LP-ET) do not report any PAIs on
leaves at all, and the Umm Irna Formation (Jordan; late Permian)
reports only hole-feeding on one species of Dicroidium to make a
combined total of four FFGs across these three Gondwanan regions
(possibly five, as it was not clear what type of interaction was
occurring on the Nidpur seed). There is clearly great potential in
advancing research on the Early Triassic floras and PAIs of these
broad regions.
To the authors’knowledge, the Benolong Flora investigation
presented herein represents the first quantitative PAI assessment of
an Early or Middle Triassic macrofloral assemblage from Oceania.
More PAI traces were identified in the Benolong Flora than the
majority of other assemblages in the database, except for the well-
studied Santa Maria Formation fossil suites, and the quantitative
investigations undertaken on wood specimens of the Fremouw
Formation (Table 1); both of these assemblages are possibly Late
Triassic in age. Furthermore, higher FFG diversity was identified in
the Benolong Flora than any Antarctic, Arabian, Oceanian, and
Indian localities, and three out of five of the South American
stratigraphic units (Table 1). These findings indicate a high
likelihood of under-reporting and under-investigation of PAIs for
Triassic strata across the majority of Gondwana and clearly
demonstrates how a concerted investigation can greatly increase
the reported number of PAIs.
4.2 Future work and recommendations
Although there has been increased interest in PAI investigations
in recent years, this compilation of data indicates large temporal
and spatial gaps in our knowledge of plant–arthropod interactions
during this interval of Gondwanan history. The pattern of gaps in
the database can be informative, since they are not equally
distributed in time or space, and our understanding of ancient
plant–arthropod interactions is almost entirely dependent on the
accuracy of identification and completeness of the plant
fossil record.
The variation in floral composition between the Early to Middle
Triassic resulted in major changes in the representation of shapes,
sizes, and densities of leaves and other organs. The arthropod
populations surviving in these evolving plant communities may
have resulted in taxonomically specialized plant–arthropod
interactions and spatiotemporal variations in FFG abundances.
Taxonomic trends in FFG distribution on plants are difficult to
discern from the current post-EPE PAI record, since specific plant–
arthropod associations have not yet been reported in sufficient
detail in most cases. Although some affected taxa and species were
described in the reports included in this review, many PAIs/FFGs
identified were not linked to specific plant taxa outside of individual
instances in the figured specimens, and the numbers of affected
leaves/organs per FFG were not clearly stated in many cases.
Thomas and Hill (2023) identified significant under-reporting
of insect damage from plant fossils illustrated in the figures of
Cenozoic publications, owing, in part, to the tendency for
palaeobotanical researchers to preferentially illustrate undamaged
leaves in their papers. This highlights the necessity for greater
awareness of, and emphasis on, the reporting of PAIs by researchers
conducting these plant taxonomic studies. Therefore, increased
Turner et al. 10.3389/fevo.2024.1419254
Frontiers in Ecology and Evolution frontiersin.org16
efforts in describing new localities and floral assemblages, revisiting
existing palaeobotanical collections and publications, and
conducting PAI assessments would greatly increase available data
and the reliability of any trends observed.
More extensive Early Triassic Gondwanan floral assemblage
identification, collection, taxonomic description and PAI
assessment would enable more accurate comparisons to Middle
Triassic PAI records, and the determination of ecological recovery
trends from this interval. Description of more floral assemblages
from Middle Triassic strata of Africa, Antarctica, South Asia and
the Arabian Peninsula would also provide a more representative
pan-Gondwanan assessment of PAI spatiotemporal trends. Oceania
hosts the majority of described palaeobotanical records in this
database; however, PAIs from the Australasian flora are relatively
understudied compared to South American assemblages. Eastern
Australia in particular, hosts a thick succession of continental upper
Permian to Middle Triassic stratigraphic units (Herbert, 1997;
Fielding et al., 2021;McLoughlin et al., 2021a;Fielding et al.,
2022) and an abundance of fossil plant-bearing strata (e.g., in the
Bowen, Nymboida and Sydney basins and the Esk Trough;
Walkom, 1924;White, 1965;Raggatt, 1969;White, 1969,1971;
Rigby, 1977;Retallack, 1980a;Playford et al., 1982;Cantrill and
Webb, 1998;Holmes and Anderson, 2013b) that represent useful
targets for tracking the evolution of Gondwanan Permian–Triassic
plant and arthropod interactions.
Detailed quantification of arthropod damage (e.g., DT/FFG
intensity and diversity; Schachat et al., 2022) is not currently
practical for this time interval of Gondwana. With larger datasets
resulting from detailed taxonomic assessment of more floral
assemblages, and increased reporting of insect damage types,
more quantitative analyses can then be conducted. A more
accurate understanding of the changes in relative abundances of
plant groups over time will contextualize implications for the PAI
record (e.g., preferential or specialized arthropod interactions with
specific plant groups, DT/FFG preservation biases between
leaf types).
To this end, it is recommended that a standardized approach for
the assessment and reporting of PAIs is implemented, necessitating
approaches that accurately identify, record and report PAI data.
Variation in preservational conditions between floral assemblages
(e.g., impressions vs coalified compressions) may influence the
detectability of various DTs and FFGs, potentially leading to
reporting biases in DT frequency. Similarly, different PAI
assessment techniques may be better for highlighting particular
FFGs (e.g., analyzing images vs hand specimens, using microscopy,
low-angle lighting, UV light, and X-ray tomography). Incorporating
the DT code of Labandeira et al. (2007; unpublished additions) into
PAI assessments will standardize records and increase confidence in
DT identification. The ‘sampling unit’per individual PAI recorded
should be clearly defined as either one rock specimen, one plant
fossil, or one plant organ (e.g., one leaf). Furthermore, a standard of
quantitative reporting of findings should be established: for each
DT, the average number of traces per plant sampling unit, the total
number of sampling units involved, and the affected plant taxa
should be reported; for each FFG, the DT diversity and distribution
across plant taxa could then be compared (see Labandeira
et al., 2018).
Addressing these concerns will also enable researchers to
contrast trends in plant–arthropod interactions from Gondwana
to those of the Laurasian and Tethyan regions for a global
perspective on the terrestrial ecological response to late Permian
to Middle Triassic climatic instability.
5 Conclusions
This review summarizes our present understanding of an
essential component of terrestrial ecosystem function following
themostsevereextinctioneventinEarth’shistory.The
compilation of Early and Middle Triassic fossil leaf floras of
Gondwana and their associated arthropod traces presented
herein demonstrates:
1. The present state of the published fossil record is
temporally and spatially biased, providing an unreliable
gauge of terrestrial ecosystem changes during the end-
Permian event recovery in Gondwana. These spatial and
temporal biases, particularly the patchiness and under-
representation of PAI reporting, need to be addressed
before the available data can be meaningfully assessed for
quantitative ecological trends.
2. The reliability of the Gondwanan PAI record is
compromised by two main biases. Firstly, observational
biases, which include a geographical skew against
Gondwanan collections in favor of those from Laurasia
and the Tethys regions, and within Gondwana, a greater
representation of assemblages from South America and
Oceania. There is also a stratigraphic observational bias,
particularly against Lower Triassic strata, which have
historically received less attention owing to their scarcity
of fossils and coal. Secondly, preservational biases will tend
to facilitate the identification of PAIs in certain styles of
fossilization and in specific types of leaves or plant organs
over others (e.g., broad vs scale-like leaves). In addition to
these biases, an absence of standardized data collection
currently limits meaningful comparisons across the region.
3. By mitigating the limiting factors above, it may in the future
be possible to utilize the PAI record of Gondwana to
construct distinctive ecological and evolutionary trends
during the recovery period following the end-Permian event.
4. In contrast to existing PAI reports from the Australian
Early to Middle Triassic (that documented three fossil plant
specimens hosting one PAI trace each), our systematic
investigation of the Benolong Flora revealed 44 PAI
traces consistent with published DTs, hosted by 40 fossil
plant fragments. This demonstrates that a concerted
quantitative investigation can greatly increase the number
of PAIs identified on an assemblage and reaffirms the
likelihood of under-reporting of PAIs for similarly
overlooked Gondwanan sites.
Turner et al. 10.3389/fevo.2024.1419254
Frontiers in Ecology and Evolution frontiersin.org17
5. We propose several key next steps to address the
demonstrable gaps in our collective knowledge of Early to
Middle Triassic Gondwanan floras and arthropod
interactions. Specifically, we advocate for establishment of
a standardized approach for assessing and reporting PAIs
that will facilitate accurate identification, recording and
reporting of PAI data. Finally, we highlight that the
excellent Early and Middle Triassic macrofloral
assemblages from age-controlled, and readily accessible
localities in eastern Australia offer a remarkable
opportunity to track the long-term patterns of recovery in
plant and arthropod herbivores from before the end-
Permian extinction through the environmental crises of
the Early to Middle Triassic.
Data availability statement
The original contributions presented in the study are included
in the article/Supplementary Material. Further inquiries can be
directed to the corresponding author/s.
Author contributions
H-AT: Conceptualization, Funding acquisition, Investigation,
Methodology, Writing –original draft, Writing –review & editing.
SM: Conceptualization, Funding acquisition, Methodology,
Supervision, Writing –original draft, Writing –review & editing.
CM: Conceptualization, Funding acquisition, Methodology,
Software, Supervision, Writing –original draft, Writing –review
& editing.
Funding
The author(s) declare financial support was received for the
research, authorship, and/or publication of this article. This project
was supported by an Irish Research Council Government of Ireland
Postgraduate Scholarship awarded to H-AT (GOIPG/2022/2071), a
Swedish Research Council (VR) grant (2022-03920) awarded to SM,
and research grants from Science Foundation Ireland’s Research
Centre in Applied Geosciences (13/RC/2092_P2) and Science
Foundation Ireland’s Frontiers for the Future Project (22/FFP-P/
11448) awarded to CM.
Acknowledgments
We would like to thank Matthew McCurry, Patrick Smith,
Graham McLean and Ailie MacKenzie for access to and assistance
with the Benolong specimens in the Australian Museum
Palaeontological Collection. Special thanks are given to Keith
Holmes and Heidi Anderson for their input and insights
regarding the Middle Triassic plant records of eastern Australia,
including the Benolong Flora. We thank Editor Sandra Schachat,
Zhou Feng, Barbara Cariglino and one anonymous reviewer for
their helpful insights that have improved the manuscript.
Conflict of interest
The authors declare that the research was conducted in the
absence of any commercial or financial relationships that could be
construed as a potential conflict of interest.
Publisher’s note
All claims expressed in this article are solely those of the authors
and do not necessarily represent those of their affiliated organizations,
or those of the publisher, the editors and the reviewers. Any product
that may be evaluated in this article, or claim that may be made by its
manufacturer, is not guaranteed or endorsed by the publisher.
Supplementary material
The Supplementary Material for this article can be found online
at: https://www.frontiersin.org/articles/10.3389/fevo.2024.1419254/
full#supplementary-material
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