Morphotropism in fumarolic mineral

  -related anhydrous sulfates: novel representatives in A + 2 M 2+(SO4)2 and A + 2 M 2+ 2(SO4)3 series Oleg I Siidra, Diana Nekrasova, Olga Blatova, Marie Colmont, Olivier Mentré, Dmitri Charkin To cite this version: Oleg I Siidra, Diana Nekrasova, Olga Blatova, Marie Colmont, Olivier Mentré, et al.. Morphotropism in fumarolic mineral-related anhydrous sulfates: novel representatives in A + 2 M 2+(SO4)2 and A + 2 M 2+ 2(SO4)3 series. Acta Crystallographica Section B : Structural Science, Crystal Engineering and Materials [2014-..], 2022, 78 (2), pp.153 -161. 10.1107/s2052520622000919. hal-03870080


Corentin Jouault
email: jouaultc0@gmail.com

Andre Nel

Frederic Legendre

Fabien L Con- Damine

Estimating the drivers of diversification of stoneflies through time and the limits of their fossil record

Keywords: Crises, extinctions, fossil record, Plecoptera, PyRate !

Deciphering the timing of lineage diversification and extinction has greatly benefited in the last decade from methodological developments in fossil-based analyses. If these advances are increasingly used to study the past dynamics of vertebrates, other taxa such as insects remain relatively neglected. Our understanding of how insect clades waxed and waned or of the impact of major paleoenvironmental changes during their periods of diversification and extinction (mass extinction) are rarely investigated. Here, we compile and analyze the fossil record of Plecoptera (1,742 vetted occurrences) to investigate their genus-level diversification and diversity dynamics using a Bayesian process-based model that incorporates temporal preservation biases. We found that the Permian-Triassic mass extinction has drastically impacted Plecoptera, while the Cretaceous Terrestrial Revolution corresponds with a turnover of plecopteran fauna.

We also unveiled three major gaps in the plecopteran fossil record: the Carboniferous-Permian transition, the late Early Cretaceous, and the late Cenomanian to Bartonian, which will need to be further investigated. Based on the life history of extant Plecoptera, we investigate the correlations between their past dynamic and a series of biotic (Red Queen hypothesis) and abiotic (Court Jester hypothesis) factors. These analyses highlight the major role of continental fragmentation in the evolutionary history of stoneflies, which is in line with phylogeny-based biogeographic analyses showing how vicariance drove their diversification. Our study advocates analyzing the fossil record with caution, while attempting to unveil the diversification and extinction periods plus the likely triggers of this past dynamics of diversification.

3 Documenting the biodiversity dynamics in the deep time of various organisms is among the major goals of evolutionary biologists [START_REF] Raup | Mass Extinctions in the Marine Fossil Record[END_REF], Alroy et al. 2008, Close et al. 2020). However, despite their huge diversity (Stork 2018), the diversification dynamics of insects remains relatively poorly studied with fossil-based analyses (e.g., Smith and Marcot 2015, Clapham et al. 2016[START_REF] Condamine | Global patterns of insect diversification: towards a reconciliation of fossil and molecular evidence?[END_REF][START_REF] Fochetti | Diversity, Threats, Decline and Conservation of European Stoneflies (Plecoptera, Insecta)[END_REF]). Yet, the fossil record of several insect lineages is relatively continuous (Schachat and Labandeira 2021) so that it could be integrated into diversification analyses to better understand both the evolutionary history of these organisms and the impact of major environmental changes in the deep time. Among those drivers of diversification, mass extinctions as well as biotic (Red Queen hypothesis) and abiotic (Court Jester hypothesis) factors are regularly invoked [START_REF] Grimaldi | Evolution of the Insects[END_REF]Engel 2005, Benton 2009, Shachat and Labandeira 2021), and their relative impact can be disentangled with the latest methods used to analyze the fossil record [START_REF] Silvestro | Improved estimation of macroevolutionary rates from fossil data using a Bayesian framework[END_REF]. Those methods take advantage of the full fossil record and can account for its well-known biases [START_REF] Foote | Principles of Paleontology[END_REF]Miller 2007, Alroy 2016). Benefiting from these methods, one can study a group of insects whose origin goes back to the Permian and address questions related to past diversification dynamics and the likely drivers affecting this diversification.

The order Plecoptera (stoneflies) is of particular interest because today they are considered as bio-indicators of water quality due to their need for well oxygenated and unpolluted freshwater (e.g., Törnblom et al. 2011). The latest molecular dated phylogeny estimated that the order originated ≈265 million years ago (Ma, 95% credibility interval: 236-294 Ma, [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF]. Investigating the diversification dynamics of Plecoptera can provide insights into the impact of three of the Big Five mass extinctions-the Permian-Triassic, Triassic-Jurassic and Cretaceous-Paleogene boundaries [START_REF] Raup | Biological extinction in earth history[END_REF]Sepkoski 1982, Hoyal Cuthill et al. 2020)and during the Carnian Humid Episode [START_REF] Ruffell | The Carnian Humid Episode of the late Triassic: A review[END_REF]). These events have strongly impacted the diversity dynamics of various organisms (e.g., [START_REF] Hull | Life in the Aftermath of Mass Extinctions[END_REF][START_REF] Chen | The timing and pattern of biotic recovery following the end-Permian mass extinction[END_REF], Benton et al. 2021). For instance, the Permian-Triassic boundary is known as the most important crisis for insects (e.g., Labandeira andSepkoski 1993, Labandeira 2005), while the Carnian Humid Episode corresponds to a period of climate changes, extinctions and radiations [START_REF] Dal Corso | Extinction and dawn of the modern world in the Carnian (Late Triassic)[END_REF]. We might expect that these events have also impacted the evolutionary history of Plecoptera. Studying Plecoptera can also contribute to better understand the impact of past environmental changes on continental aquatic communities.

In addition to these major events, the diversification of Plecoptera has likely been linked with a series of abiotic (i.e. climate and/or geological changes: see [START_REF] Zwick | (0); Angiosperms (1); continental fragmentation (2); Gymnosperms (3); temperatures (4). Figure 1. Net diversification rate for the entire evolutionary history of Plecoptera. A. From Carboniferous to mid-Eocene; B. Focus on the Carboniferous to Middle Triassic period; C. Focus on the late Middle Triassic to the late Lower Cretaceous period. The rates are deduced through the analysis of the fossil register at the genus-level, via the RJMCMC model implemented in PyRate. Abbreviation: Pen = Pennsylvanian, Cis= Cisuralian, Gua= Guadalupian, L= Lopingian, ET= Lower Triassic, MT = Middle Triassic, LP= Upper Triassic, EJ= Lower Jurassic, MJ= Middle Jurassic, LJ= Upper Jurassic, EC= Lower Cretaceous, LC= Upper Cretaceous, Pa= Paleocene, Eoc= Eocene[END_REF] for a discussion on the putative effect of the break-up of Gondwana) and biotic factors (e.g. ecological changes and/or diversity dependence). Among the most notable environmental changes, the Cretaceous Terrestrial Revolution (KTR: [START_REF] Lloyd | Dinosaurs and the Cretaceous Terrestrial Revolution[END_REF], Benton et al. 2021), along with the rapid diversification of angiosperms, has radically altered ecosystems [START_REF] Chaboureau | Tectonic-driven climate change and the diversification of angiosperms[END_REF]) and the diversification of other plant groups (Condamine et al. 2020). Plecoptera have likely been affected by these changes, including some specialized herbivores [START_REF] Monakov | Feeding of freshwater invertebrates[END_REF] or predators, whose diets or abilities to detect or pursue preys have likely been disturbed [START_REF] Fuller | Influence of substrate type on vulnerability of prey to predacious aquatic insects[END_REF]. The replacement of gymnosperms by angiosperms as dominant plant groups in terrestrial environments is indeed often seen as the trigger for major faunal changes in past freshwater ecosystems (e.g., replacement of subfamilies of Chironomidae, extinction of certain groups of Odonata and Plecoptera: Sinitschenkova 2003). Angiosperms tend to produce more biomass from their leaves than gymnosperms, and this biomass is recycled into the soil and freshwater by microorganisms that consume dissolved oxygen in the water, causing eutrophication of water, especially lakes. Such eutrophication is highly deleterious for Plecoptera, adapted to welloxygenated waters [START_REF] Gaufin | Studies on the tolerance of aquatic insects to low oxygen concentrations[END_REF], Sinitschenkova 2003[START_REF] Malison | Remarkable anoxia tolerance by stoneflies from a floodplain aquifer[END_REF]). However, these hypotheses have never been formally tested, questioning the impact of the angiosperm diversification or of the Cretaceous Terrestrial Revolution on Plecoptera and, more generally, on freshwater communities.

The fossil record of Plecoptera represents currently 322 species distributed in 25 families and two suborders: Antarctoperlaria and Arctoperlaria [START_REF] Zwick | Insecta: Plecoptera. Phylogenetisches System und Katalog. Das Tierreich[END_REF]. But, as for extant species (more than 3,500), the number of fossil representatives of each sub-order is largely unequal, with about 10% of extinct species for Antarctoperlaria, while the Arctoperlaria represents 90% of the past diversity (data compiled from http://fossilworks.org; last accessed March 27, 2021; [START_REF] Fochetti | Global diversity of stoneflies (Plecoptera; Insecta) in freshwater[END_REF]. The Perlopseina record-a Permian stem group [START_REF] Martynov | Permian fossil insects from Chekarda[END_REF][START_REF] Zwick | (0); Angiosperms (1); continental fragmentation (2); Gymnosperms (3); temperatures (4). Figure 1. Net diversification rate for the entire evolutionary history of Plecoptera. A. From Carboniferous to mid-Eocene; B. Focus on the Carboniferous to Middle Triassic period; C. Focus on the late Middle Triassic to the late Lower Cretaceous period. The rates are deduced through the analysis of the fossil register at the genus-level, via the RJMCMC model implemented in PyRate. Abbreviation: Pen = Pennsylvanian, Cis= Cisuralian, Gua= Guadalupian, L= Lopingian, ET= Lower Triassic, MT = Middle Triassic, LP= Upper Triassic, EJ= Lower Jurassic, MJ= Middle Jurassic, LJ= Upper Jurassic, EC= Lower Cretaceous, LC= Upper Cretaceous, Pa= Paleocene, Eoc= Eocene[END_REF]-is limited in number, with only three species. Such unbalanced species richness brings the question of the evolutionary drivers underlying the diversification of the two suborders.

As in many insect groups, the number of fossil plecopteran species is likely underestimated because there are less than a handful of people studying them. Moreover, their long evolutionary history and the difficulty to get a rich and continuous fossil record over such a long time inevitably led to a knowledge bias towards certain periods and the virtual absence of fossil for others. Also, the phylogenetic placement of many fossil species at the family level needs to be revised; a task made even more complicated because of the high plasticity of the wing veins of stoneflies. This situation is often ignored, most fossil plecopteran species being described from a single-sometimes partial-specimen (Jouault et al. 2021a). Altogether, these features make any inference drawn from the fossil record sensitive to several paleontological biases, which need to be considered.

Here we study how Plecoptera have waxed and waned through time, analyzing all stoneflies fossil occurrences and modeling preservation rates through time and across taxa. Our analyses not only allow highlighting the periods of radiation and those of decline, but also discuss-ing gaps in the plecopteran fossil record throughout geological times. We further perform analyses to tease apart the factors that may have impacted the diversification or extinction of Plecoptera by studying correlations between their diversification dynamics and different biotic and abiotic variables.

Material and methods

Fossil occurrences and dataset

The fossil record of Plecoptera was compiled from the Paleobiology Database (http://fossilworks.org/, https://paleobiodb.org: last accessed July 10, 2021). The original dataset contained about 430 occurrences and was then corrected and emended. The dataset was cleaned of synonyms, outdated combinations, nomina dubia and other erroneous and doubtful records. After correction including data addition from the literature, it is composed of 322 species (130 genera) for 1,742 occurrences resulting from an in-depth study of the entire bibliography of fossil Plecoptera (Supp Table S1 [online only]). The differences between genus level or sub-order occurrence numbers are explained by the systematic placement of some genera that can only be placed confidently in a sub-order but not in a family (Supp Table S2 [online only]). Note that some occurrences of Plutopteryx beata [START_REF] Sinitshenkova | New Jurassic stoneflies of the family Baleyopterygidae[END_REF] are not included since the species is known from more than 1,400 specimens but 1,000 of them are not labelled as indicated by Sinichenkova (1985: p. 133). Therefore, we only include the labelled specimens, viz. 467 specimens (Sinitshenkova 1985: p. 133) to avoid overrepresentation of a genus while not being informative for the whole analysis. We removed the species Marciperla curta Lin, 1986 from our dataset. Initially considered as Plecoptera incertae sedis, it cannot be placed with certainty in this order [START_REF] Lin | Early Mesozoic fossil insects from South China[END_REF]). This taxon is possibly a Plecoptera because of its two cercomeres, but the partial preservation challenges a clear attribution of this species (Lin 1986: fig. 62). It has for instance been considered as a possible Grylloblattodea (Storozhenko 1998: figs. 82-87). To avoid unsupported placement, authors are encouraged to keep some specimens in open nomenclature until additional material is discovered. Similarly, the systematic position of numerous fossil Plecoptera at the subfamily or family level being poorly supported, we refrain from conducting analyses at these taxonomic levels. The complete datasets used for each analysis are available as supplementary data.

Temporal dynamics of origination and extinction rates

Traditionally the past dynamic of lineages was inferred using the fossil record (e.g., [START_REF] Stanley | Macroevolution: Pattern and Process[END_REF], although recent progress have led to estimate diversification rates from dated phylogenies (e.g., [START_REF] Harvey | Phylogenies without fossils[END_REF][START_REF] Morlon | Reconciling molecular phylogenies with the fossil record[END_REF][START_REF] Rabosky | Rates of speciation and morphological evolution are correlated across the largest vertebrate radiation[END_REF]. Numerous issues have arisen regarding the analyses of the fossil record: (1) using discrete time bins, (2) analyzing only first and last appearances of taxa, thus ignoring other occurrences if available while first and last occurrences cannot be assumed to reflect the true times of speciation/origination and extinction (Strauss andSadler 1989, Marshall 1990), (3) the inability to perform model testing against over-parameterization, which limits models' robustness [START_REF] Burnham | Model selection and multimodel inference: a practical information-theoretic approach[END_REF], [START_REF] Fochetti | Global diversity of stoneflies (Plecoptera; Insecta) in freshwater[END_REF] the poor or sole incorporation of extant taxa in the analyses, which in both cases represents only a small fraction of the total diversity that has existed since their origination (e.g., [START_REF] Raup | Biological extinction in earth history[END_REF]).

To overcome these issues, new methodologies were developed and implemented in PyRate (Silvestro et al. 2014a[START_REF] Silvestro | Improved estimation of macroevolutionary rates from fossil data using a Bayesian framework[END_REF]. PyRate estimates speciation and extinction rates and their temporal dynamics from fossil occurrence data and has proven to be robust thanks to simulations under a range of potential biases, including violations of the sampling assumptions, variable preservation rates, and incomplete taxon sampling (Silvestro et al. 2014b(Silvestro et al. , 2015a[START_REF] Silvestro | Improved estimation of macroevolutionary rates from fossil data using a Bayesian framework[END_REF].

Simulation results showed that the dynamics of speciation and extinction rates, including sudden rate changes and mass extinctions, are correctly estimated under a wide range of conditions, such as low levels of preservation (down to 1-3 fossil occurrences per species on average), severely incomplete taxon sampling (up to 80% missing), and high proportion of singletons (exceeding 30% of the taxa in some cases).

We analyzed the plecopteran fossil record using a Bayesian approach, as implemented in PyRate 3.0 [START_REF] Silvestro | Improved estimation of macroevolutionary rates from fossil data using a Bayesian framework[END_REF], to simultaneously infer (1) the times of speciation (Ts) and of extinction (Te) for each taxon in the dataset, (2) the temporal dynamics of origination and extinction rates, (3) the preservation rate across taxa, and (4) the preservation rate through time.

To model the processes of diversification and preservation, this approach uses as input all fossil occurrences that can be assigned to taxa, in our case fossil genera. All analyses were set with the best-fit preservation process after comparing (-PPmodeltest option) the homogeneous Poisson process (-mHPP option), the non-homogeneous Poisson process (default option), and the time-variable Poisson process (-qShift option). The time-variable Poisson process assumes that preservation (expected number of occurrences per lineage per Ma) rates are constant within a predefined time frame but may vary over time. In the present study we used a predefined time frame of bins of 10 million years since the Carboniferous. This model is thus appropriate when rates over time are heterogeneous. We also accounted for varying preservation rates across taxa using the Gamma model (-mG option), which is with gamma-distributed rate heterogeneity with four rate categories.

We jointly estimated the number of rate shifts as determined by log Bayes factors and the times at which origination and extinction undergo a shift using the reversible jump Markov Chain Monte Carlo (RJMCMC, -A 4 option; [START_REF] Silvestro | Improved estimation of macroevolutionary rates from fossil data using a Bayesian framework[END_REF] or a birth-death model with constrained shifts (BDCS, Silvestro et al. 2015b). This allowed us to obtain an overview of the general trends of rate variation across a long timescale. Both the preservation and the birthdeath processes are modeled in continuous time. Thus, the origination and extinction rates are measured as the number of origination and extinction events per lineage per Ma, while the preservation rate is expressed as expected occurrences per taxon per Ma. We performed two BDCS analyses: one to estimate diversification rates per bin of 10 million years, and the second to estimate diversification rates per geological epochs. The BCDS model is one of the first to be implemented in PyRate, but recent progress suggests that the RJMCMC model provides more robust results [START_REF] Silvestro | Improved estimation of macroevolutionary rates from fossil data using a Bayesian framework[END_REF]. To cross-validate our results we have opted for a multi-model approach. To look for differences in the diversification patterns of the two suborders of Plecoptera, we also analyzed two sub-datasets corresponding to the fossil record of the Antarctoperlaria composed of 13 genera for 398 occurrences (Supp Table S3 [online only]) and to the Arctoperlaria composed of 108 genera and 1,242 occurrences (Supp Table S4 [online only]).

For each analysis, we ran PyRate for 10 million MCMC iterations with sampling frequency of 1,000 and combined the posterior samples of the parameters from the 10 randomly replicated datasets after excluding the first 10% of the samples as burnin. These 10 datasets were generated using the extract.ages function to randomly resample the age of fossil occurrences within their respective temporal ranges (i.e., resampled ages are randomly drawn between the minimum and the maximum ages of the geological stratum). We monitored chain mixing and effective sample sizes by examining the log files in Tracer 1.7.1 [START_REF] Rambaut | Posterior summarisation in Bayesian phylogenetics using Tracer 1.7[END_REF]. The parameters are considered convergent when the effective sample size (ESS) is greater than 200. We combined the posterior estimates of the origination and extinction rates across all replicates to generate rate-through-time plots. Thus, we obtained ten posterior estimates of the Ts and Te for all genera and we estimated the past diversity dynamics by calculating the number of living taxa at every point in time based on the Ts and Te (-ltt 1 option). We also looked at the marginal posterior distributions of origination and extinction rates during the largest extinction events documented in geological history. We particularly examined the di-versification dynamics at three of the Big Five events: the Permian-Triassic, the Triassic-Jurassic and the Cretaceous-Paleogene boundaries (sensu [START_REF] Raup | Biological extinction in earth history[END_REF]Sepkoski 1982, Hoyal Cuthill et al. 2020). Shifts of diversification were considered significant when log Bayes factors > 6 with the RJMCMC model, while we considered shifts to be significant with the BDCS model when mean rates in a time bin do not overlap with the 95% CI of the rates of adjacent time bins.

Correlations with biotic and abiotic variables

To propose a global scenario of the evolution of Plecoptera, we focused on biotic (e.g. relative diversity of angiosperms or gymnosperms) and abiotic (e.g. temperature, continental fragmentation) factors, potentially responsible for the phenomena of apparitions or extinctions. Analyses were performed with PyRate 3.0 [START_REF] Silvestro | Improved estimation of macroevolutionary rates from fossil data using a Bayesian framework[END_REF]) using the multivariate birth-death (MBD) model. This model estimates speciation and extinction rates based on several continuous variables over time assuming linear or exponential functions linking temporal variations in birth-death rates to change as a function of one or more variables [START_REF] Lehtonen | Environmentally driven extinction and opportunistic origination explain fern diversification patterns[END_REF]). In the MBD model, a correlation parameter is estimated to quantify independently the role of each variable on speciation and extinction. A horseshoe prior (explained in [START_REF] Lehtonen | Environmentally driven extinction and opportunistic origination explain fern diversification patterns[END_REF]) is used to reduce the correlation parameters around zero, thereby reducing the risk of over-parameterization and the need to test explicit models. The data used for the analyses (Ts and Te data) are derived from our previous RJMCMC inferences. In this study, we tested five variables that we assume, because of the group's biology, having an impact on their diversification or extinction, namely: (0) intra-clade competition; (1) fluctuation in relative diversity of angiosperms (Silvestro et al. 2015b); (2) fragmentation of continents, which is an index with low values reflecting a continental agglomeration (e.g., Pangea) and high values a continental splitting [START_REF] Zaffos | Plate tectonic regulation of global marine animal diversity[END_REF]); (3) fluctuation in relative diversity of gymnosperms (Silvestro et al. 2015b); and (4) average global temperatures over geological time, which was obtained combining the δ 18 O data from [START_REF] Prokoph | Compilation and time-series analysis of a marine carbonate δ18O, δ13C, 87Sr/86Sr and δ34S database through Earth history[END_REF] and [START_REF] Zachos | An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics[END_REF] to encompass the period ranging from the Holocene to the lowermost Carboniferous [START_REF] Condamine | Assessing the causes of diversification slowdowns: temperature-dependent and diversity-dependent models receive equivalent support[END_REF].

All the datasets for these five variables are available in the publications cited aside each variable. The MBD analyses were performed over 10 million generations, and visualization of results and convergence parameters was performed on Tracer 1.7.1 [START_REF] Rambaut | Posterior summarisation in Bayesian phylogenetics using Tracer 1.7[END_REF]. We considered correlation parameters to be significant when the shrinkage weights (ω) > 0.5 and the 95% CI does not overlap with 0.

Results

Global diversification pattern

The net diversification rate (origination minus extinction) corresponds to the number of events per million years per lineage. The phases of diversification are visible when the net diversification rate is superior to 0; oppositely when it is below 0 it indicates periods of decline.

Heterogeneous diversification between the Antarctoperlaria and Arctoperlaria

The fossil record of Antarctoperlaria is very fragmentary while that of Arctoperlaria is much more diverse and somewhat continuous (Supp Table S2 [online only]). Fossil representatives of Antarctoperlaria range only from the Guadalupian to the Aptian-Albian, resulting in a partial modeling of their past dynamic (Figs. 1B-C). Despite their current diversity, there is no fossil Antarctoperlaria in the Late Cretaceous and the Cenozoic. Antarctoperlaria feebly diversified from the Guadalupian to Aptian-Albian without experimenting peaks of strong extinction or diversification (net diversification between 0.01 and 0.05) (Figs. 1B-C).

The oldest Arctoperlaria are nearly contemporaneous with the oldest Antarctoperlaria but their fossil record, more abundant, allows modeling their past dynamic showing a pattern strongly resembling that of the order. They are impacted by the Permo-Triassic crisis (net diversification ≈ -1.5) and suffer a strong decline during the Early Cretaceous (net diversification ≈ -1.25) (Fig. 1C). They diversified slightly during the Middle Triassic (max. net diversification ≈ 0.25) and strongly diversified from the mid-Cretaceous (net diversification between 0.2 and 1.75) until the Priabonian (Fig. 1C). The Carnian Humid Episode, the Triassic-Jurassic events or the subsequent mass extinction events have no real impact on the dynamic of each group since we found no change in net diversification (Figs. 1B-C).

Disparities in preservation rates and deposits

Three gaps are recorded during the evolutionary history of the Plecoptera. To better illustrate these gaps, we used the preservation rates and the number of lineages from the Carboniferous to present. The preservation rate indicates four time-intervals in which the preservation is low (ca. or inferior to 1, Fig. 4A). The first is around the Carboniferous-Permian boundary (preservation rate ≈ 1), the second during the Early and early Middle Triassic (preservation rate ≈ 0), the third extends from the middle of the Late Triassic to the Early Jurassic (preservation rate ≈ 1), and the fourth encompasses the period ranging from the late Early Cretaceous to nowadays (preservation rate between 1.3 and 0) (Fig. 4A). Plotting the diversity dynamic through time, we found three gaps (Fig. 4B), corresponding to the periods where there is no lineage recorded over time. The first encompasses the Carboniferous-Permian transition (Fig. 4B) and ranges from the Kasimovian (ca. 306 Ma) to the Asselian (ca. 295 Ma). The second gap occurs during the lower Cretaceous (ca. 111 Ma) and the mid-Cretaceous (ca. 100 Ma) (Fig. 4B). The third and largest gap extends from the uppermost Cenomanian (ca. 94 Ma) to the Bartonian (ca. 39 Ma) (Fig. 4B).

Correlations with abiotic and biotic variables

The results of the MBD analyses are summarized in Table 1. Shrinkage weights indicate good support (ω > 0.8) for all correlation parameters (G) of origination and extinction rates, except for the ω of extinction depending on the clade's diversity and the ω of the extinction depending on the relative abundance of gymnosperms (ω < 0.5). The effect of a variable is assessed as significant when ω > 0.5 and 95% HPD does not encompass 0.

The variables positively correlated with the origination of Plecoptera are the temperature (ω = 0.74, G ≈ 0.03) and the fragmentation of continents (ω = 0.9, G ≈ 12.15) (Table 1; Fig. 5C). However, only the continental fragmentation is significantly correlated with origination (indicated by a star in Fig. 5C). The variables negatively correlated with the origination of Plecoptera are the relative abundance of gymnosperms (ω = 0.63, G ≈ -1.2), angiosperms (ω = 0.82, G ≈ -1.6) and the intra-clade competition (ω = 0.9, G ≈ -2.8). Only the intra-clade competition is significantly correlated with origination (Table 1; Fig. 5C).

The variables positively correlated with the extinction of Plecoptera are the temperature (ω = 0.91, G ≈ 0.08) and the fragmentation of continent (ω = 0.96, G ≈ 20.2), with a significant correlation only for the latter (Table 1; Fig. 5D). The relative abundance of gymnosperms (ω = 0.44, G ≈ -0.09), angiosperms (ω = 0.88, G ≈ -2.3) and the intra-clade competition (ω = 0.50, G ≈ -0.39) are all negatively correlated with the extinction, and none is significantly correlated with extinction (Table 1; Fig. 5D).

Discussion

We analyzed the plecopteran fossil record, comprising >300 species and >1,700 specimen occurrences, using a Bayesian framework to investigate the impact of major crises, biotic and abiotic factors on the diversification dynamics of these insects that now thrive in well-oxygenated waters. We have unveiled different periods of diversification, extinction and turnovers 5). The first critical period corresponds to the early dynamics of Plecoptera, at the transition between stem-Plecoptera and crown-Plecoptera. The second corresponds to the 'mid'-Cretaceous, a period characterized by the Cretaceous Terrestrial Revolution and the outcomes of Gondwana fragmentation. We then focus on the results of the MBD analyses to contrast the support for the Red Queen and Court Jester hypotheses, i.e. for biotic and abiotic factors, respectively (Fig. 5). Finally, we discuss a few critical gaps in the plecopteran fossil record, leveraging known gaps for insects and new analyses of preservation rate (Fig. 4). Pinpointing these gaps could help guide future efforts that would benefit analyses of plecopteran diversification and, more broadly, aquatic insect communities.

Early dynamics of Plecoptera derived from the fossil record

Chronologically, the first transitional events (from diversification to extinction) occur between the Carboniferous and middle Permian. The stem Plecoptera is uniquely recorded in the Carboniferous [START_REF] Béthoux | At last, a Pennsylvanian stem-stonefly (Plecoptera) discovered[END_REF][START_REF] Schubnel | Two new stem-stoneflies discovered in the Pennsylvanian Avion locality, Pas-de-Calais, France (Insecta: 'Exopterygota')[END_REF]). These pre-Pangean faunas, as shown by the fossil record in China and France during the Carboniferous [START_REF] Béthoux | At last, a Pennsylvanian stem-stonefly (Plecoptera) discovered[END_REF][START_REF] Schubnel | Two new stem-stoneflies discovered in the Pennsylvanian Avion locality, Pas-de-Calais, France (Insecta: 'Exopterygota')[END_REF], have likely led to new Pangean faunas developing on the northern continental block, dividing during the Permian [START_REF] Blakey | Gondwana paleogeography from assembly to breakup -a 500 m.y. odyssey[END_REF][START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF]. Our analyses indicate an increase in origination rate around 300 Ma (Figs. 1-2, 5A-B), which can be attributed to this continental re-organization. Because of the relative scarcity of the plecopteran fossil record at this period, a fine understanding of their early diversification is out of reach, but it is certain that this period was critical in their evolution. It is also important to associate this diversification period with the Carboniferous-Permian glaciation and deglaciation events that have substantially reset terrestrial ecosystems [START_REF] Montañez | CO2-forced climate and vegetation instability during late Paleozoic deglaciation[END_REF]). These events likely created suitable habitats for Plecoptera, which are known to live in cold waters (particularly in glacier streams).

Additionally, this period also coincides with a period of continental agglomeration, while the distribution of Plecoptera would have been restricted [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF]) because of arid areas in the center of the Pangea [START_REF] Pardo | Carboniferous-Permian climate change constrained early land vertebrate radiations[END_REF].

The second remarkable event-and likely the most important of the early dynamics of Plecoptera-is the Permian-Triassic boundary (Figs. 1-2, 5A-B). It corresponds to a general mass extinction event (e.g., Labandeira andSepkoski 1993, Labandeira 2005;Benton andNewell 2014, Zhao et al. 2021) that we record for Plecoptera as well with a sudden decrease in the net diversification (Fig. 1A). This extinction event is visible when confronting the numerous stoneflies from the Lopingian (Supp Table S2 [online only]) with the lack of stonefly from the Early and the Middle Triassic (Fig. 4) despite several fossil insect deposits, e.g., Brookvale in Australia, Vosges in France, or Tongchuan in China [START_REF] Zheng | Middle-Late Triassic insect radiation revealed by diverse fossils and isotopic ages from China[END_REF]. This period would also correspond to the divergence of the two infraorders of the Arctoperlaria (Euholognatha and Systellognatha, [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF]) but we have recorded only a massive extinction and not any increase in origination before the Middle Triassic (Fig. 1; Supp Figs. S1-2). The end-Permian extinction is supposed to be linked to massive volcanic eruptions in Siberia causing drastic changes in ecosystems that have affected most organisms (e.g., [START_REF] Benton | How to kill (almost) all life: the end-Permian extinction event[END_REF][START_REF] Ogden | Explosive eruption of coal and basalt and the end-Permian mass extinction[END_REF][START_REF] Burgess | High-precision geochronology confirms voluminous magmatism before, during, and after Earth's most severe extinction[END_REF][START_REF] Fielding | Age and pattern of the southern high-latitude continental end-Permian extinction constrained by multiproxy analysis[END_REF]. Temperatures rose by ca. 15°C, a global warming that, combined with volcanic gases resulted in acid rains that have destroyed trees and led to a loss of vegetation cover [START_REF] Benton | The Triassic[END_REF]. This crisis was unfavorable to life at equatorial latitudes and prevented ecosystem recovery after greenhouse events (Sun et al. 2012). Our results indicate that this dramatic event has strongly affected Plecoptera (Figs. 1A,5A,B) and most likely the whole aquatic insect communities.

During the Late Triassic, another period has recently gained attention: the Humid Episode of the Carnian, which has mainly affected marine diversity (e.g., Dal Corso et al. 2020).

Given the biology of Plecoptera (life in torrent waters), an effect of the paleoenvironmental conditions of the Carnian on their diversification could have been expected. However, we have not detected any change in net diversification during the Carnian (Fig. 1A).

Similarly, we have not detected the Triassic-Jurassic extinction in our analyses (Figs. 1A, 2C-D, 5A-B), suggesting that this crisis may have not been so severe for insects (e.g., [START_REF] Labandeira | The Fossil Record of Insect Extinction: New Approaches and Future Directions[END_REF][START_REF] Condamine | Global patterns of insect diversification: towards a reconciliation of fossil and molecular evidence?[END_REF][START_REF] Nel | Diversification of insects since the Devonian: a new approach based on morphological disparity of mouthparts[END_REF]. Instead, it corresponds with the beginning of the radiation of many extant insect orders that have their oldest known representatives during the Triassic [START_REF] Montagna | Recalibration of the insect evolutionary time scale using Monte San Giorgio fossils suggests survival of key lineages through the End-Permian Extinction[END_REF]. For Plecoptera, this period would correspond to an increase in diversification in Perloidea [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF] and to the colonization of the southern hemisphere by Notonemouridae and Antarctoperlaria after a long dispersal on lands, while the continents were still connected (Cui et al. 2019[START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF]). However, these phylogeny-derived hypotheses are not confirmed with our analyses of the fossil record (e.g., stable origination, extinction and net diversification before and after the Triassic-Jurassic boundary) (Figs. 1A,2). Given that the number of lineages in this period is relatively high (Fig. 4), we would have expected to record these events if they had been impactful. After this period, no significant change in the diversification of the Plecoptera was recorded before the Early Cretaceous (Figs. 1A,2).

The Cretaceous Terrestrial Revolution and the subsequent diversification of Plecoptera

For Plecoptera, the most dramatic extinction event occurred from the early Aptian (ca. 125 Ma) to the early Cenomanian (ca. 100 Ma; Fig. 1A). This brutal extinction is sometimes recorded at a broader taxonomic scale when studying the complete fossil record of insects [START_REF] Labandeira | The Fossil Record of Insect Extinction: New Approaches and Future Directions[END_REF]) but also tempered in other studies [START_REF] Labandeira | Insect diversity in the fossil record[END_REF][START_REF] Labandeira | Why did terrestrial insect diversity not increase during the angiosperm radiation? Mid-Mesozoic, plant-associated insect lineages harbor clues[END_REF][START_REF] Condamine | Global patterns of insect diversification: towards a reconciliation of fossil and molecular evidence?[END_REF]; see also our section on fossil gaps below and how they interfere with diversification analyses of Plecoptera). A few hypotheses have been proposed to explain this major event and, for Plecoptera, both the importance of the KTR and of the fragmentation of Gondwana can be defended. The KTR is characterized by the radiation of the angiosperms, a drastic ecological change that reshuffled the ecosystems [START_REF] Lloyd | Dinosaurs and the Cretaceous Terrestrial Revolution[END_REF][START_REF] Meredith | Impacts of the Cretaceous Terrestrial Revolution and KPg extinction on mammal diversification[END_REF], Benton et al. 2021) and that translated into biotic turnovers (sensu Schachat and Labandeira 2021) with a high extinction and origination in different lineages. We recorded such a turnover at this period for Plecoptera (Figs. 1A,2). Alternatively or concomitantly, major geological events might have played a strong role in the diversification of Plecoptera, e.g., the fragmentation of the Gondwana or the opening of the Atlantic, which continued into the Late Jurassic and Cretaceous [START_REF] Daly | Rift basin evolution in Africa: the influence of reactivated steep basement shear zones[END_REF][START_REF] Seton | Global continental and ocean basin reconstructions since 200 Ma[END_REF]. The fragmentation of continents is the only variable tested strongly positively correlated with origination and extinction rates (Figs. 5C,D); we discuss it further in the next section. The fragmentation of continents could have favored the diversification of Antarctoperlaria in the southern hemisphere (Letsch et al. 2021: fig. 4). However, several land masses such as India, Africa or Burmese Block lack evidence for a colonization by Antarctoperlaria at that time (e.g., [START_REF] Jouault | A new mid-Cretaceous fossil genus of stonefly (Plecoptera: Perlidae) from the Burmese amber[END_REF]. This is rather surprising given that the biome homogeneity in central Gondwana could have facilitated wide dispersion between South America and Africa during the Early Cretaceous (Jouault et al. 2021b, Nel andJouault 2021) and by extension up to the Burma Terrane (Jouault et al. 2021c). Similarly, Antarctica may have served as an exchange route between Australia and South America when a large expanse of temperate rainforests was present on the continent [START_REF] Klages | Temperate rainforests near the South Pole during peak Cretaceous warmth[END_REF] and until the opening of the Drake Passage around 41 Ma [START_REF] Scher | Timing and climatic consequences of the opening of Drake Passage[END_REF]Martin 2006, Letsch et al. 2021). The availability of new habitats (e.g., in Antarctica) and the colonization of new continental land masses [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF]) would suggest a phase of plecopteran diversification but we record an extinction during the Early Cretaceous, rather indicating that these opportunities fail to counterbalance extinction events (Figs. 1A,2,).

Interestingly, two shifts in diversification rate, estimated from a molecular phylogeny and located in Nemouridae and Leuctrinae, occurred during the KTR in the late Early Cretaceous [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF], further supporting our results derived from the fossil record. These inferred diversification rate shifts suggest that the KTR has generated an increase in the diversification of Nemouridae and Leuctrinae, and additional studies must be conducted to decipher its causes.

The second major event of the Cretaceous period is the Cretaceous-Paleogene (K/Pg) extinction, which corresponds to a series of drastic events leading to the most famous mass extinction with the final demise of non-avian dinosaurs (e.g., [START_REF] Brusatte | The extinction of the dinosaurs[END_REF]. The origin of this mass extinction event is far from being consensual, but a combination of biotic and abiotic factors likely led to these dramatic events (e.g., [START_REF] Mitchell | Late Cretaceous restructuring of terrestrial communities facilitated the end-Cretaceous mass extinction in North America[END_REF][START_REF] Chiarenza | Asteroid impact, not volcanism, caused the end-Cretaceous dinosaur extinction[END_REF], Condamine et al. 2021). Like for the early Aptian extinction, recorded in our analyses, the impact of K/Pg events does not seem to be strong when analyses are conducted on the insect fossil record at the family or order level [START_REF] Labandeira | Insect diversity in the fossil record[END_REF][START_REF] Labandeira | Why did terrestrial insect diversity not increase during the angiosperm radiation? Mid-Mesozoic, plant-associated insect lineages harbor clues[END_REF][START_REF] Condamine | Global patterns of insect diversification: towards a reconciliation of fossil and molecular evidence?[END_REF]. However, given that the main trigger of the K/Pg extinction is supposed to be, most of the time simplified to, a cataclysm related to the impact of an asteroid [START_REF] Alvarez | Extraterrestrial Cause for the Cretaceous-Tertiary Extinction[END_REF], we wonder if this crisis has also affected the Plecoptera. Based on the results detailed above, and the elevated net diversification rate recorded before and after the K/Pg boundary, it appears that this event did not have a significant impact on Plecoptera (Fig. 1A). Like recent results for vertebrate clades, the decline or faunal turnover appears to begin before the impact of the asteroid [START_REF] Condamine | Dinosaur biodiversity declined well before the asteroid impact, influenced by ecological and environmental pressures[END_REF]. For Plecoptera, these diversification rate changes occur earlier in the Early Cretaceous (Fig. 2), suggesting that the K/Pg events had a low impact on the clade dynamics. This is corroborated by the moderately high preservation rate of this period, suggesting that drastic events should have been recorded in the fossil record.

In addition, a significant shift in diversification rate is recorded at the K/Pg boundary for the Capniidae (Letsch et al. 2021: fig. 2), suggesting that at least this family flourished during this period. According to all these elements, the K/Pg severity on the diversification of Plecoptera appears to be reduced.

How Plecoptera diversified: Red Queen or Court Jester hypotheses?

The diversification dynamics of organisms is often investigated by contrasting the importance of biotic and abiotic drivers, what is commonly known as the Red Queen and Court Jester hypotheses, respectively [START_REF] Van Valen | A new evolutionary law[END_REF][START_REF] Barnosky | Distinguishing the effects of the Red queen and Court Jester on Miocene mammal evolution in the northern Rocky Mountains[END_REF]. Using the plecopteran fossil record, we aimed at deciphering the relative importance of those hypotheses, looking at correlations between a few factors (intra-clade competition, relative abundance of angiosperms and gymnosperms vs. temperature variation, continental fragmentation) and the dynamics of Plecoptera over geological time (Figs. 5C-D).

One biotic factor-intra-clade competition-and one abiotic factor-fragmentation of continents-were significantly correlated with the diversification of Plecoptera (Figs. 5C-D).

Intra-clade competition was negatively correlated with origination, whereas continental fragmentation was positively correlated with both origination and extinction. The first result indicates that origination decreases when the diversity of Plecoptera increases, suggesting intraclade competition or diversity dependence (Fig. 5C). This is in line with the life history traits of Plecoptera, whose adults are relatively poor flyers and whose nymphs have specialized ecological requirements (fresh and well-oxygenated waters), resulting in low dispersal rates [START_REF] Marden | Surface-skimming stoneflies: a possible intermediate stage in insect flight evolution[END_REF], Thomas et al. 2000[START_REF] Zwick | (0); Angiosperms (1); continental fragmentation (2); Gymnosperms (3); temperatures (4). Figure 1. Net diversification rate for the entire evolutionary history of Plecoptera. A. From Carboniferous to mid-Eocene; B. Focus on the Carboniferous to Middle Triassic period; C. Focus on the late Middle Triassic to the late Lower Cretaceous period. The rates are deduced through the analysis of the fossil register at the genus-level, via the RJMCMC model implemented in PyRate. Abbreviation: Pen = Pennsylvanian, Cis= Cisuralian, Gua= Guadalupian, L= Lopingian, ET= Lower Triassic, MT = Middle Triassic, LP= Upper Triassic, EJ= Lower Jurassic, MJ= Middle Jurassic, LJ= Upper Jurassic, EC= Lower Cretaceous, LC= Upper Cretaceous, Pa= Paleocene, Eoc= Eocene[END_REF][START_REF] Fochetti | Global diversity of stoneflies (Plecoptera; Insecta) in freshwater[END_REF], McCulloch et al. 2009). Based on these considerations, it is accepted that long-range dispersal capabilities of Plecoptera are reduced. Therefore, when a population or diversity increases locally, competition for resources and particularly for food reaches its climax, limiting speciation by intra-clade competition. This is especially relevant since plecopteran nymphs are represented in all functional feeding groups (e.g., predator, phytophagous, detritivores, etc.) (Tierno de Figueroa and López-Rodríguez 2019) and sometimes change diet according to their developmental stage [START_REF] Yule | The life cycle and dietary habits of Illiesoperla mayi Perkins (Plecoptera: Gripopterygidae) in Victoria, Australia[END_REF]. Competition between species may increase depending on the stage of development of the different local populations. Similarly, cases of cannibalism are reported [START_REF] Yule | The life cycle and dietary habits of Illiesoperla mayi Perkins (Plecoptera: Gripopterygidae) in Victoria, Australia[END_REF]) and we can imagine a similar behavior in deep time under strong selection and competition pressure. Therefore, reaching suitable ecological niches might be challenging for these insects and competition very disadvantageous.

Continental fragmentation has likely driven both origination and extinction of Plecoptera, likely by promoting allopatric speciation but also by altering previous ecosystems and increasing extinctions (Figs. 5C-D). Our results are congruent with the main distribution of extant stonefly species, which postulates that continental fragmentation is a driver of their diversification and distribution. In fact, previous authors named the two sub-orders after the Antarctica (Antarctoperlaria) and the Arctic (Arctoperlaria) because of the restricted distribution of their constitutive families [START_REF] Zwick | (0); Angiosperms (1); continental fragmentation (2); Gymnosperms (3); temperatures (4). Figure 1. Net diversification rate for the entire evolutionary history of Plecoptera. A. From Carboniferous to mid-Eocene; B. Focus on the Carboniferous to Middle Triassic period; C. Focus on the late Middle Triassic to the late Lower Cretaceous period. The rates are deduced through the analysis of the fossil register at the genus-level, via the RJMCMC model implemented in PyRate. Abbreviation: Pen = Pennsylvanian, Cis= Cisuralian, Gua= Guadalupian, L= Lopingian, ET= Lower Triassic, MT = Middle Triassic, LP= Upper Triassic, EJ= Lower Jurassic, MJ= Middle Jurassic, LJ= Upper Jurassic, EC= Lower Cretaceous, LC= Upper Cretaceous, Pa= Paleocene, Eoc= Eocene[END_REF]. The only exceptions are the arctoperlarian families, Notonemouridae and Perlidae, which are present and diversified in the southern hemisphere.

Therefore, it is considered that Antarctoperlaria originated in Gondwana, while Arctoperlaria originated in Laurasia [START_REF] Zwick | (0); Angiosperms (1); continental fragmentation (2); Gymnosperms (3); temperatures (4). Figure 1. Net diversification rate for the entire evolutionary history of Plecoptera. A. From Carboniferous to mid-Eocene; B. Focus on the Carboniferous to Middle Triassic period; C. Focus on the late Middle Triassic to the late Lower Cretaceous period. The rates are deduced through the analysis of the fossil register at the genus-level, via the RJMCMC model implemented in PyRate. Abbreviation: Pen = Pennsylvanian, Cis= Cisuralian, Gua= Guadalupian, L= Lopingian, ET= Lower Triassic, MT = Middle Triassic, LP= Upper Triassic, EJ= Lower Jurassic, MJ= Middle Jurassic, LJ= Upper Jurassic, EC= Lower Cretaceous, LC= Upper Cretaceous, Pa= Paleocene, Eoc= Eocene[END_REF], but a recent biogeographic analysis suggests a more complicated scenario [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF]). An alternative hypothesis states that the current restricted distribution of Plecoptera suborders could be explained by the disappearance of Antarctoperlaria on Laurasia and of Arctoperlaria on Gondwana [START_REF] Zwick | (0); Angiosperms (1); continental fragmentation (2); Gymnosperms (3); temperatures (4). Figure 1. Net diversification rate for the entire evolutionary history of Plecoptera. A. From Carboniferous to mid-Eocene; B. Focus on the Carboniferous to Middle Triassic period; C. Focus on the late Middle Triassic to the late Lower Cretaceous period. The rates are deduced through the analysis of the fossil register at the genus-level, via the RJMCMC model implemented in PyRate. Abbreviation: Pen = Pennsylvanian, Cis= Cisuralian, Gua= Guadalupian, L= Lopingian, ET= Lower Triassic, MT = Middle Triassic, LP= Upper Triassic, EJ= Lower Jurassic, MJ= Middle Jurassic, LJ= Upper Jurassic, EC= Lower Cretaceous, LC= Upper Cretaceous, Pa= Paleocene, Eoc= Eocene[END_REF]. In any case, the ancestral plecopteran stock on the Pangea was divided during major geological events of continental fragmentation [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF]. The link between diversification and continental fragmentation is also strengthened by the limited dispersal abilities of stoneflies letting us assume that vicariance (i.e. the isolation of previously connected ancestral groups as a result of plate tectonics or other geographic barriers) shaped their biogeography and diversification [START_REF] Zwick | (0); Angiosperms (1); continental fragmentation (2); Gymnosperms (3); temperatures (4). Figure 1. Net diversification rate for the entire evolutionary history of Plecoptera. A. From Carboniferous to mid-Eocene; B. Focus on the Carboniferous to Middle Triassic period; C. Focus on the late Middle Triassic to the late Lower Cretaceous period. The rates are deduced through the analysis of the fossil register at the genus-level, via the RJMCMC model implemented in PyRate. Abbreviation: Pen = Pennsylvanian, Cis= Cisuralian, Gua= Guadalupian, L= Lopingian, ET= Lower Triassic, MT = Middle Triassic, LP= Upper Triassic, EJ= Lower Jurassic, MJ= Middle Jurassic, LJ= Upper Jurassic, EC= Lower Cretaceous, LC= Upper Cretaceous, Pa= Paleocene, Eoc= Eocene[END_REF]. Vicariance and allopatric speciation are driven by historical glaciation, orogenesis, and small-scale population fragmentation [START_REF] Zwick | (0); Angiosperms (1); continental fragmentation (2); Gymnosperms (3); temperatures (4). Figure 1. Net diversification rate for the entire evolutionary history of Plecoptera. A. From Carboniferous to mid-Eocene; B. Focus on the Carboniferous to Middle Triassic period; C. Focus on the late Middle Triassic to the late Lower Cretaceous period. The rates are deduced through the analysis of the fossil register at the genus-level, via the RJMCMC model implemented in PyRate. Abbreviation: Pen = Pennsylvanian, Cis= Cisuralian, Gua= Guadalupian, L= Lopingian, ET= Lower Triassic, MT = Middle Triassic, LP= Upper Triassic, EJ= Lower Jurassic, MJ= Middle Jurassic, LJ= Upper Jurassic, EC= Lower Cretaceous, LC= Upper Cretaceous, Pa= Paleocene, Eoc= Eocene[END_REF][START_REF] Fochetti | Global diversity of stoneflies (Plecoptera; Insecta) in freshwater[END_REF][START_REF] Fochetti | Molecular systematics and biogeography of the western Mediterranean stonefly genus Tyrrhenoleuctra (Insecta, Plecoptera)[END_REF], 2011[START_REF] Weiss | Molecular systematics, evolution and zoogeography of the stonefly genus Siphonoperla (Insecta: Plecoptera, Chloroperlidae)[END_REF], Theissinger et al. 2013). Continental fragmentation can also affect mountain orogeny and thus habitats for Plecoptera, which often occur in mountain streams (similar observations are made for other freshwater lineages: [START_REF] Hrivniak | The impact of Miocene orogeny for the diversification of Caucasian Epeorus (Caucasiron) mayflies (Ephemeroptera: Heptageniidae)[END_REF].

Examples of clade radiations in mountains across the globe can testify for the impact of orogenic events in Plecoptera (e.g., Stark andNelson 1994, Sproul et al. 2015). For instance, the endemic radiation of the genus Chilenoperla in the Andes exemplifies the link between mountain building and speciation. Chilenoperla diverged in the Eocene [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF] when the separation of South America, Antarctica and Australia led to a cooling of the circum-Antarctic zone and to the formation of mountains, like the Andes [START_REF] Barrett | Cooling a continent[END_REF]. This synchronicity between the Andean orogeny, which took place during the late Eocene-early Miocene and led to the growth of the Western Cordillera (e.g., [START_REF] Armijo | Coupled tectonic evolution of Andean orogeny and global climate[END_REF], Riesner et al. 2018[START_REF] Boschman | Andean mountain building since the Late Cretaceous: A paleoelevation reconstruction[END_REF], and the diversification of some extant genera like Chilenoperla in the region might reflect the strong influence of continental fragmentation on the diversification of Plecoptera. This influence needs to be further examined when significant gaps in the fossil record are filled (see below) or using phylogenetic approaches [START_REF] Boschman | Mountain radiations are not only rapid and recent: ancient diversification of South American frog and lizard families related to Paleogene Andean orogeny and Cenozoic climate variations[END_REF]. As evidenced by the strong correlation recorded in our MBD analysis, it appears that continental fragmentation, a component of the Court Jester hypothesis, is a major driver of the diversification dynamics of Plecoptera (Figs. 5C-D).

Gaps in Plecoptera fossil record: Where should we look?

As for many extant insect orders, the order Plecoptera has a long evolutionary history that can be traced from the fossil record. Nonetheless, the fossil record has a few gaps that challenge the estimation of evolutionary history of Plecoptera and that should guide future efforts (Fig. 4).

Beyond known gaps in the paleo-entomological record [START_REF] Nel | Diversification of insects since the Devonian: a new approach based on morphological disparity of mouthparts[END_REF], our analyses inferred preservation rate through time, which allows us to expand our critical analysis of those gaps for Plecoptera (Fig. 4).

The first gap (Fig. 4B), from the Kasimovian (ca. 306 Ma) to the Asselian (ca. 295 Ma), is flanked by the appearance of stem-Plecoptera [START_REF] Béthoux | At last, a Pennsylvanian stem-stonefly (Plecoptera) discovered[END_REF][START_REF] Schubnel | Two new stem-stoneflies discovered in the Pennsylvanian Avion locality, Pas-de-Calais, France (Insecta: 'Exopterygota')[END_REF]) and the early diversification of crown-Plecoptera. It makes the transition between stem-and crown-Plecoptera opaque and prevents us from understanding the morphological changes (wing venation, genitalia, etc.) that took place between these lineages. As a result, the early evolutionary history of Plecoptera remains poorly understood. Several fossil deposits could inform us (e.g., Mazon Creek: USA, Wettin: Germany, Avion: France) on this pivotal period for Plecoptera.

A second gap is recorded during the late Early Cretaceous (Fig. 4B), but it may simply reflect the lack of deposits covering this period. Filling this gap might thus be more challenging than the first one. It is nonetheless crucial because it corresponds to the angiosperm diversification. If not filled, this gap may weaken our ability to quantify the impact of angiosperm diversification on Plecoptera. This period also corresponds to a sudden diminution of the net diversification (Fig. 1A) rate and to two major shifts in the plecopteran diversification [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF], for the Leuctrinae and Nemouridae, and to some major events in continental fragmentation. Disentangling the relationships between these results and putative driving forces would require a better documentation of the late Early Cretaceous gap.

The third and largest gap extends from the mid-Cretaceous to the end of the Eocene (Fig. 4B) and thus encompasses the K/Pg event. It prevents from a robust quantifying of the impact of this crisis and of the warming events of the Cenozoic (e.g., Paleocene-Eocene Thermal Maximum, Middle Miocene Climatic Optimum). This gap is bewildering because many fossil entomofaunas are known for this time interval from amber deposits, compressions or imprints (Schachat and Labandeira 2020). If the fossil record of Plecoptera trapped in amber is relatively scarce, the imprints or compressions of Plecoptera are generally more abundant.

Therefore, the absence of described Plecoptera from the Paleocene of Menat (France) or from the Eocene of the Green River Formation or Florissant (USA) is puzzling, especially since a few fossils are known from these localities (for example in the collection of the Museum of Natural History, University of Colorado). This period also includes a major shift in diversification rate for the Capniidae [START_REF] Letsch | Combining molecular datasets with stronglyheterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera)[END_REF], supposedly around the K/Pg boundary. However, the lack of fossils of this family or other Plecoptera impedes further study.

  Using the RJMCMC model, Plecoptera seem to diversify (net diversification between 0.3 and 0.6) between ca. 310 Ma and ca. 275 Ma, which corresponds with the period ranging from the middle Pennsylvanian to the middle Cisuralian (Figs. 1-3, 5A,B). Then the speciation and extinction rates drop to reach zero until the Permo-Triassic boundary (between 259-247 Ma) (Figs. 1A), where Plecoptera undergo a strong decline (max. net diversification ≈ -0.25) (Figs. 1-2, 5A,B). The origination (Figs. 2A, B) and the extinction (Figs. 2C, D) seem to increase before the Permo-Triassic boundary, during the Cisuralian and Guadalupian, with the origination stronger than the extinction (Fig. 2) but the dramatic event occurs at the Permo-Triassic boundary with an impressive burst of extinction and no origination (= mass extinction) (Figs. 1-3, 5A,B). After this event, the net diversification, the origination and extinction rates remain relatively constant (≈ 0 or at least <0.2) from the Triassic (ca. 247 Ma) to the Aptian (ca. 125 Ma) (Figs. 1-3, 5A,B). Following this plateau, the fossil record of Plecoptera indicates a period of major decline (max. net diversification ≈ -1.4) extending from the lower Aptian (ca. 125 Ma) to the lower Cenomanian (ca. 100 Ma). This period is the outcome of a feeble speciation and a massive extinction peak (= turnover) (Figs. 1-3, 5A,B). This extinction peak is followed by a period of diversification but differences in diversification rates are recorded between the RJMCMC and the BDCS models (Figs. 1-3, 5A,B; Supp Figs. 1-2 [online only]).

Figure 2 .Figure 3 .

 23 Figure 2. Plecoptera diversification dynamics. A. Origination rate; B. Probability of changes in

Figure 4 .

 4 Figure 4. Preservation and gaps throughout Plecoptera fossil record. A. Mean preservation rates

Figure 5 .

 5 Figure 5. A. Dynamics of speciation and extinction rates through time as estimated with the

Acknowledgements

We are grateful to the three anonymous reviewers for the insightful comments and corrections. This work is part of CJ's Ph.D. project. This project benefited from an "Investissements d'Avenir" program managed by the Agence Nationale de la Recherche (CEBA, ref. ANR-10-LABX-25-01). correlation parameters on speciation and extinction with abiotic factors like global temperature, and global continental fragmentation; with biotic factors like the relative diversity through time of gymnosperms, and angiosperms; and diversity-dependence factors with the diversity through time of all fossil Plecoptera. The asterisk (*) indicates significant correlation parameter for a given variable (shrinkage weights (ω) > 0.5). Thermometer from CJ; Angiosperms, Gymnosperms and Plecoptera silhouettes from http://phylopic.org/; Triassic world maps pictures of courtesy of Eikeskog1225 (© Wikimedia Commons): https://creativecommons.org/licenses/by-sa/4.0/; the colors of the geological time scale correspond to the same periods as in Fig. 1A.

Overall, our results show the crucial interest in documenting fossil taxa to analyze the evolutionary history of a clade and infer scenarios about periods of diversification or decline.

In this study, we analyzed, for the first time, the fossil record of Plecoptera to study their diversification and diversity dynamics in the deep time. Our analysis suggests that the Permian-Triassic crisis had a notable impact on the diversification of the Plecoptera, with a drastic decrease in the number of lineages, whereas the rainy episodes of the Carnian and the Triassic-Jurassic transition did not. A second, major extinction event is also recorded at the end of the Early Cretaceous and may be interpreted as a turnover between 'pre-angiosperm' Plecoptera and 'new' Plecoptera developing in novel ecosystems dominated by angiosperms. We also unveiled three major gaps (i.e. the Carboniferous-Permian transition; the late Early Cretaceous; uppermost Cenomanian to Bartonian) to be filled for a better understanding of the evolutionary history of this insect clade. These gaps in the fossil record are even more troublesome when they overlap with a crisis or when major diversification events are expected to occur.

Supplementary Data

Supplementary data are available at Insect Systematics and Diversity online. baseline rates; Gλ correlation parameters to origination; Gμ correlation parameters to extinctions. ωλ and ωμ statistical support (significant between 0.8-1). Parameter end numbers:

Supp Table 1. Fossil occurrences of Plecoptera at the genus-level with their status and life span.

Supp Table 2. Fossil record of Plecoptera with their current systematic placement (the systematic proposed here is widely accepted but does not reflect the opinion of the authors).

Supp Table 3. Fossil occurrences of Antarctoperlaria at the genus-level with their status and life span.

Supp Table 4. Fossil occurrences of Arctoperlaria at the genus-level with their status and life span.