Corentin Jouault
email: jouaultc0@gmail.com

Frédéric Legendre

Philippe Grandcolas

Frederic Legendre
email: frederic.legendre@mnhn.fr@orcid.org

Andre Nel
email: anel@mnhn.fr

André Nel
email: anel@mnhn.fr@orcid.org

Frederic Legendre

Andre Nel

Revising dating

Keywords: Time divergence, FBD, tip-dating, morphological phylogeny, Cretaceous, calibration, ghost lineages

teaching and research institutions in France or abroad, or from public or private research centers.

Introduction

Deciphering the timing and tempo of lineage diversification of organisms has greatly benefited from advances in Bayesian phylogenetic analyses using morphological data, allowing to include fossil taxa in tip-dating or total-evidence analyses [START_REF] Dos Reis | Bayesian molecular clock dating of species divergences in the genomics era[END_REF][START_REF] Wright | A systematist's guide to estimating Bayesian phylogenies from morphological data[END_REF]. Even if some methodological refinements are still under development (e.g. [START_REF] Keating | Morphological phylogenetics evaluated using novel evolutionary simulations[END_REF], those analyses have shed insights on the evolution of organisms, sometimes resulting in a better fit with stratigraphic evidence [START_REF] King | Tip dating supports novel resolutions of controversial relationships among early mammals[END_REF], sometimes clarifying the tempo of diversification of several lineages with a robust fossil record such as mammals [START_REF] King | Tip dating supports novel resolutions of controversial relationships among early mammals[END_REF], Mesozoic birds (Zhang & Wang, 2019), Lissamphibia [START_REF] Pyron | Divergence time estimation using fossils as terminal taxa and the origins of Lissamphibia[END_REF], and Hymenoptera (Ronquist et al., 2012). Despite several remarkable fossils-including two new species based on soldier specimens that we describe here-the timing of the evolution of termites and their eusocial system has never been investigated with those latest developments.

The spectacular evolutionary and ecological success of termites is mostly attributed to their highly integrated eusocial system, defined by cooperative brood care, overlapping generations within a colony, and the division of labor into reproductive and non-reproductive castes [START_REF] Wilson | The Insect societies[END_REF][START_REF] Wilson | Sociobiology: the new synthesis[END_REF][START_REF] Lin | Evolution of sociality in insects[END_REF][START_REF] Wilson | Eusociality: origin and consequences[END_REF]. Termites are regarded as the first animals that have evolved eusociality [START_REF] Grimaldi | Evolution of the insects[END_REF]Legendre & Condamine, 2018;[START_REF] Barden | Fossil social insects[END_REF]Zhao et al. 2020b) because they show the oldest record of morphologically specialized soldiers [START_REF] Engel | Morphologically specialized termite castes and advanced sociality in the Early Cretaceous[END_REF][START_REF] Zhao | The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites[END_REF]Zhao et al., , 2020a)), although the discovery of Cretaceous ant fossils has questioned this assertion [START_REF] Perrichot | New Cretaceous records and the diversification of crown-group ants (Hymenoptera: Formicidae)[END_REF]. Clarifying the timing of diversification of termites is also critical to understand the underlying roles of environmental factors in the evolution of highly integrated social systems [START_REF] Grandcolas | The origin of a 'true' worker caste in termites: phylogenetic evidence is not decisive[END_REF]Legendre et al., 2008Legendre et al., , 2013Legendre et al., , 2015)).

Most molecular phylogenies have agreed on a termite crown-age dating back to ca. 150 Ma (e.g. [START_REF] Bourguignon | The evolutionary history of termites as inferred from 66 mitochondrial genomes[END_REF]Legendre et al., 2015). Nevertheless, discrepancies exist among dating estimates for this group and its families [START_REF] Evangelista | An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites (Blattodea)[END_REF], whether estimated from molecular or morphological evidence. Some of the most recent phylogenetic analyses have relied on molecular data, using fossils as node calibrations only [START_REF] Bourguignon | The evolutionary history of termites as inferred from 66 mitochondrial genomes[END_REF]Legendre et al., 2015;[START_REF] Wu | Phylogenetic position of the enigmatic termite family Stylotermitidae (Insecta: Blattodea)[END_REF][START_REF] Bucek | Evolution of termite symbiosis informed by transcriptome-based phylogenies[END_REF][START_REF] Condamine | Fossil and phylogenetic analyses reveal recurrent periods of diversification and extinction in dictyopteran insects[END_REF]. Others have used morphological data with fossils as terminals, but the phylogenies were built in a parsimony framework, wherein crown-and stem-ages of termite families were educated guesses [START_REF] Engel | Termites (Isoptera): their phylogeny, classification, and rise to ecological dominance[END_REF][START_REF] Krishna | Treatise on the Isoptera of the world[END_REF][START_REF] Zhao | The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites[END_REF]Zhao et al., , 2020a)). A single study [START_REF] Wappler | The effects of fossil placement and calibration on divergence times and rates: an example from the termites (Insecta: Isoptera)[END_REF] used extant and fossil taxa as terminals in a Bayesian framework; it was conducted, however, before significant methodological improvements in tip-dating analyses were developed (e.g. [START_REF] Heath | The fossilized birth-death process for coherent calibration of divergence-time estimates[END_REF]. Those different practices led to different results, sometimes tens of Myrs apart in their mean estimates. To obfuscate things even further, some ichnofossils, namely Triassic, Early and Late Jurassic putative nests, are controversial. Sometimes regarded as evidence of a Triassic, or even earlier origin of termites [START_REF] Hasiotis | Termite (Insecta: Isoptera) nest ichnofossils from the upper triassic chinle formation, petrified forest national park, Arizona[END_REF][START_REF] Bordy | Advanced Early Jurassic termite (Insecta: Isoptera) nests: evidence from the Clarens Formation in the Tuli Basin, Southern Africa[END_REF][START_REF] Bordy | Advanced Early Jurassic termite (Insecta: Isoptera) nests: evidence from the Clarens Formation in the Tuli Basin, Southern Africa[END_REF][START_REF] Bordy | New data on the palaeontology and sedimentology of the Lower Jurassic Lisbon Formation (Karoo Supergroup), Ellisras Basin, South Africa[END_REF][START_REF] Smith | New social insect nests from the Upper Jurassic Morrison Formation of Utah[END_REF], the oldest ones are judged by others as lacking proper analyses to form conclusive evidence of Triassic antiquity of termites [START_REF] Genise | Comment -advanced Early Jurassic Termite (Insecta: Isoptera) nests: evidence from the Clarens Formation in the Tuli Basin, Southern Africa[END_REF][START_REF] Genise | Ichnoentomology: insect traces in soils and paleosols[END_REF]. To our knowledge, however, they have never been used to calibrate phylogenetic trees.

In order to clarify the timing of the origin of termites and their constitutive extant and extinct families, we first expand the termite fossil record by describing two new species from soldier specimens. Next, we take advantage of the fossilized birth-death (FBD) model [START_REF] Heath | The fossilized birth-death process for coherent calibration of divergence-time estimates[END_REF] and of a previous morphological dataset [START_REF] Engel | Termites (Isoptera): their phylogeny, classification, and rise to ecological dominance[END_REF][START_REF] Engel | Morphologically specialized termite castes and advanced sociality in the Early Cretaceous[END_REF][START_REF] Zhao | The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites[END_REF] to estimate datings that we contrast with estimates from a node-dating analysis and from previous works. We then discuss those estimates and their implications, including the origin of termite eusociality.

Materials and methods

Origin, preparation and examination of amber pieces

The amber pieces containing the specimens studied herein come from the two main amber deposits of the world [START_REF] Grimaldi | Fossiliferous Cretaceous amber from Myanmar (Burma): its rediscovery, biotic diversity, and paleontological significance[END_REF][START_REF] Weitschat | Baltic amber[END_REF], respectively Burmese and Baltic amber.

One is embedded in a piece of yellow Burmese amber with numerous debris. This amber comes from Noije Bum in the Hukawng Valley (26° 29′ N, 96° 35′ E), Kachin State, northern Myanmar (see detailed map in Grimaldi and Ross, 2017: fig. 2). Taphonomic analyses of pholadids and radiometric data, based on zircons from volcanic clasts found within the amber-bearing sediments, established an early Cenomanian age (98.79 ± 0.62 Ma) for Kachin amber (Shi et al., 2012;Smith & Ross, 2017). Some ammonites found in the amber-bearing bed and within amber corroborate a Late Albian / Early Cenomanian age (Cruickshank & Ko, 2003;Yu et al., 2019).

The other is embedded in a piece of clear yellow Baltic amber with at least three Diptera and two Collembola as syninclusions. This amber comes from the Blue Earth Formation (or Blue Earth member of the Prussian Formation) from the Kaliningrad Russian enclave (54°41'19.5"N, 20°20'50.6"E). Blue Earth Formation was dated as late Bartonian to Priabonian (Upper Eocene, based on palynological data (Kosmoswska-Ceranowicz et al., 1997;Aleksandrova & Zaporozhets, 2008). Discussions on the age and paleobiota of Baltic amber can be found in [START_REF] Weitschat | Baltic amber[END_REF], [START_REF] Sadowski | Conifers of the "Baltic amber forest" and their palaeoecological significance[END_REF][START_REF] Alekseev | The revision of gymnosperm species from Eocene Baltic amber[END_REF].

The amber pieces were polished with a grinder polisher (Buehler EcoMet 30) by using a very thin silicon carbide sanding paper (grit size = 7000). Then, the fragment was embedded with cedar oil between microscopic slides to minimize light scattering during focus stacking.

Both specimens are housed in the amber collection of the Geological Department and Museum (IGR) of the University of Rennes (France), respectively under the collection number IGR.BU-012 and IGR.BA-020. Amber pieces and specimens were examined with a Leica MZ APO stereomicroscope, and a Nikon SMZ25 stereomicroscope. Photographs were taken using a Canon EOS 5D mark II mounted on a Leica MZ APO stereomicroscope. All images are digitally stacked photomicrographic composites of several individual focal planes, obtained using HeliconFocus 6.7 software. Figures were composed with Adobe Illustrator CC 2019 and Photoshop CC 2019 software.

We follow the morphological terminology and the classification of termites as presented in [START_REF] Krishna | Treatise on the Isoptera of the world[END_REF], and thus retain the rank Isoptera.

Morphological dataset

Phylogenetic analyses were conducted to i) clarify the phylogenetic positions of the two new genera described herein, and ii) estimate divergence times for termites and their subgroups. The morphological matrix (Appendix 1) was modified from [START_REF] Engel | Morphologically specialized termite castes and advanced sociality in the Early Cretaceous[END_REF] with insightful corrections from [START_REF] Zhao | The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites[END_REF]. The data set is composed of 111 morphological characters (Appendix 2) coded for 89 extant and extinct taxa. The outgroup comprises four cockroaches and one mantid (Chaeteessa sp., Panchlora sp., Periplaneta americana, Cryptocercus sp. 1, and Cryptocercus sp. 2).

The ingroup has representatives for all extant and extinct families. The morphological matrix was established using Mesquite software ver. 3.61 [START_REF] Maddison | Mesquite: a modular system for evolutionary analysis[END_REF], with all characters unordered, non-polarized, and of equal weight. Characters coded as inapplicable were treated as missing data and coded '-', while unknown characters were coded '?'. Missing data and unknown characters represent respectively 8.57 % and 22.13 % of the dataset. All consensus trees were visualized and drawn using FigTree ver. 1.4.4 [START_REF] Rambaut | FigTree v1.4.4[END_REF], and modified with Adobe Illustrator CC2019.

Tree reconstruction and dating estimates in Bayesian inference

We performed two kinds of analyses using relaxed clock models in MrBayes 3.2.7a [START_REF] Huelsenbeck | MRBAYES: Bayesian inference of phylogeny[END_REF][START_REF] Ronquist | MrBayes 3: Bayesian phylogenetic inference under mixed models[END_REF]Ronquist et al., 2012). First, we computed node-dating analyses with only extant taxa, fossils being used to calibrate nodes (see below for justifications of fossil calibrations). Second, we computed tip-dating analyses with a fossilized birth-death model (FBD; [START_REF] Heath | The fossilized birth-death process for coherent calibration of divergence-time estimates[END_REF] wherein fossil taxa are terminals. Because not all the expected inter-familial relationships were found in unconstrained analyses-an unsurprising outcome for a relatively small dataset comprising only morphological characters (see Supporting Information)-and because our aim is not to reassess phylogenetic relationships but rather age estimates, we added four topology constraints (i.e. monophyly of Blattodea, (Cryptocercus + Isoptera), Isoptera, and Neoisoptera but Stylotermitidae). Those constraints fit the current understanding in termite phylogenetics (notably [START_REF] Wu | Phylogenetic position of the enigmatic termite family Stylotermitidae (Insecta: Blattodea)[END_REF] for Stylotermitidae), thus ensuring better estimates of the parameters under investigation (i.e. ages; Zhang & Wang, 2019;[START_REF] Luo | A simulation-based evaluation of tip-dating under the fossilized birth-death process[END_REF].

All but one analyses were computed with a Mkv model (Lewis, 2001) with a gamma rate variation across characters and with the independent gamma relaxed clock model (Lepage et al., 2007;lset rates = invgamma, prset clockvarpr = igr, prset igrvarpr = exp(10); the exception was computed with no rate variation across characters: lset rates = equal). The prior used for the mean clock rate was gamma (2,200), except in one analysis wherein we tested a normal but very flat prior (prset clockratepr = normal(0.0025,1) and a flat uniform speciation prior (prset speciationpr = uniform(0,10); Matzke & Wright, 2016). The proportion of extant taxa was set to 0.012 (37 termite terminals for ca. 3000 species; prset sampleprob=0.012). Sampling strategy of taxa was either set to diversity (prset samplestrat=diversity wherein fossils are sampled randomly and can be tips or ancestors), random (prset samplestrat=random wherein fossils can be tips or ancestors) or fossiltip (prset samplestrat=fossiltip wherein fossil and extant taxa are assumed to be sampled randomly and fossil ancestor is not allowed). An exponential prior and a beta prior were used for the net speciation rate and the relative extinction rate, respectively [prset speciationpr = exp(100); prset extinctionpr = beta(1,1)], while the node age prior was set to 'calibrated'. All analyses comprised two runs and four chains, and were launched for 50 million generations. Chains were sampled every 5000 generations and a burn-in fraction of 0.25 was used. Convergence diagnostics were checked for each analysis [i.e. average standard deviation of split frequencies < 0.01, PRSF close to 1.0, and ESS > 200 in Tracer [START_REF] Rambaut | Posterior summarisation in Bayesian phylogenetics using Tracer 1.7[END_REF]; see also Supporting Information].

Node-dating analyses -These four analyses have a reduced taxonomic sampling comprising only extant taxa (Ntax = 42). The prior probability distribution on branch lengths was either set to clock:birthdeath (two analyses) or clock:uniform (two analyses). Three nodes were assigned offset exponential priors: the root of the tree (minimum age: 225 Ma, mean age: 270 Ma), Blattodea (150,225 Ma) and Isoptera (137,150 Ma). Those ages were set according to the latest studies [START_REF] Bourguignon | The evolutionary history of termites as inferred from 66 mitochondrial genomes[END_REF][START_REF] Bourguignon | Transoceanic dispersal and plate tectonics shaped global cockroach distributions: evidence from mitochondrial phylogenomics[END_REF]Legendre et al., 2015;[START_REF] Engel | Morphologically specialized termite castes and advanced sociality in the Early Cretaceous[END_REF][START_REF] Bucek | Evolution of termite symbiosis informed by transcriptome-based phylogenies[END_REF][START_REF] Evangelista | An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites (Blattodea)[END_REF]Zhao et al., 2020a; more details in Appendix 3).

Tip-dating analyses -For the 13 tip-dating analyses, extinct taxa were calibrated with uniform distributions (Appendix 3) bounded according to the minimum and maximum ages of their deposits [START_REF] Barido-Sottani | Ignoring stratigraphic age uncertainty leads to erroneous estimates of species divergence times under the fossilized birth-death process[END_REF] ten analyses) or with fixed distributions bounded according to the minimum ages of their deposits (three analyses). For the fossilized birth-death process, sampling fossil ancestors [START_REF] Gavryushkina | Bayesian inference of sampled ancestor trees for epidemiology and fossil calibration[END_REF][START_REF] Zhang | Total-evidence dating under the fossilized birth-death process[END_REF] was allowed in four analyses, whereas all fossils were set as tips in the others (prset samplestrat = fossiltip). The prior probability distribution on branch lengths was set to clock:fossilization, except in a single analysis (prset brlenspr = clock:uniform). Because the dataset comprises characters of alates, workers, soldiers, and life history traits, we conducted analyses with and without partitioning of the data (i.e. a single or three partitions; in the latter, alate and worker characters belong to the same partition). Our aim was to check whether data partitioning would affect age estimates, as well as evolutionary rates.

Finally, some analyses were computed, modifying one prior at a time relative to the one partition analysis, to assess the robustness of our results. The modified priors were the mean age of termites MRCA (difference with the minimum age halved or doubled; i.e. 144 or 163 Ma) and a variable partition specific rate prior (i.e. prset ratepr = variable).

Overall, 17 tip-dating and node-dating analyses were performed on CIPRES portal with MrBayes 3.2.7a on XSEDE [START_REF] Miller | Creating the CIPRES Science Gateway for inference of large phylogenetic trees[END_REF]; all scripts are provided as Supporting Information.

Tree reconstruction in parsimony

We performed a tree reconstruction using maximum of parsimony (MP) in PAUP ver. 4.0a166 [START_REF] Swofford | PAUP: Phylogenetic Analysis Using Parsimony (and Other Methods)[END_REF] to compare the topology with the Bayesian inference. The outgroup was treated as paraphyletic with respect to the ingroup, and the tree search was performed using a heuristic search method with the following options: maximum number of trees saved equal to 10000, only optimal trees retain, collapse of zero-length branches, and a TBR swapping algorithm. The search produced more than one equally parsimonious cladogram so that a strict consensus cladogram was built to summarize the results. Bremer indices were computed [START_REF] Bremer | Branch support and tree stability[END_REF], as well as Bootstrap support values (bs) using the heuristic search option for 100 replicates [START_REF] Felsenstein | Confidence limits on phylogenies: an approach using the bootstrap[END_REF][START_REF] Hillis | An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis[END_REF] [START_REF] Zhao | The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites[END_REF]. In termites, workers/soldiers body length is about 0.8 times that of alates [START_REF] Engel | Morphologically specialized termite castes and advanced sociality in the Early Cretaceous[END_REF] so that Milesitermes gen. nov. (body length 6.4 mm) does not belong to

Anisotermes xiai (body length 12.9 mm).

Milesitermes gen. nov. differs from the soldiers of the extant Mastotermes darwiniensis and of the Miocene Mastotermes electromexicanus in its thinner mandible without a strong basal tooth and its pronotum trapezoidal.

An epicranial scar (plesiomorphy) could be present on the head of Milesitermes gen. nov. but the discovery of new specimens is necessary to confirm this hypothesis. Numerous wood fragments are embedded in the amber piece and may suggest a wood feeding habit. Note that [START_REF] Zhao | The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites[END_REF] attributed a soldier to Anisotermes xiai (whose type is an alate specimen) on the basis of the unique presence of this species in the Burmese amber when they described it.

The peculiar shape of the mandible of Milesitermes gen. nov. is unknown because it has been damaged on this specimen. We consider unwise to derive any systematic affiliation based on the mere shape of this mandible. Nonetheless, note that soldiers of Hodotermitidae may have similar mandibles but they do not have five tarsomeres. Also, Hodotermitidae soldiers have their pronota narrower than their heads and do not possess additional spines on tibia. Furthermore, it is unlikely that Milesitermes gen. nov. belong to the Hodotermitidae whose soldiers have heavily toothed mandibles, a feature we did not observe on Milesitermes gen. nov.

Family Rhinotermitidae Froggatt, 1897

Subfamily Heterotermitinae Froggatt, 1897

Genus Reticulitermes [START_REF] Holmgren | Termitenstudien. 4. Versuch einer systematischen Monographie der Termiten der Orientalischen Region[END_REF] Reticulitermes grimaldii sp. nov. urn:lsid:zoobank.org:act:A25EC97F-73EA-4EF8-A30F-DF1B6F3F1516

Holotype: Number IGR.BA-020 (a nearly complete and well-preserved soldier).

Etymology:

The specific epithet is a patronym honoring our colleague Pr. David A. Grimaldi, eminent American paleoentomologist, and specialist of Isoptera. The species epithet is to be treated as a noun in the genitive case. Color. Head clearly orange, except black mandibles and brown labrum; rest of body grayish white.

Locality and horizon:

Remarks: Based on the presence of four tarsomeres, head with fontanelle (hole partially visible), flat pronotum, mandible without serrations along the inner margins (maybe some at the base, but not visible), we can undoubtedly assign Reticulitermes grimaldii sp. nov. to the Rhinotermitidae. Within this family, Reticulitermes grimaldii sp. nov. keys out in the Heterotermitinae because of the eye spot absent or inconspicuous, the head with a small fontanelle, the absence of a groove from the fontanelle to the base of the clypeus, and the head with subparallel sides [START_REF] Emerson | Tertiary fossil species of the Rhinotermitidae (Isoptera), phylogeny of genera, and reciprocal phylogeny of associated Flagellata (Protozoa) and the Staphylinidae (Coleoptera)[END_REF]. The presence of setulae surrounding the fontanelle is also distinctive of the Heterotermitinae [START_REF] Engel | Termites (Isoptera): their phylogeny, classification, and rise to ecological dominance[END_REF].

The Heterotermitinae comprise the genera Heterotermes Froggatt, 1897 and Reticulitermes. Despite the fact that the mandibles cross, the flat postclypeus exclude any affinity with Heterotermes [START_REF] Scheffrahn | A new subterranean termite introduced to Florida: Heterotermes Froggatt (Rhinotermitidae: Heterotermitinae) established in Miami[END_REF]: Table 1). We assigned specimen IGR.BA-020 to the genus Reticulitermes based on its flat postclypeus, its mandibles distinctly hooked at tip, and its tongue-shaped labrum [START_REF] Krishna | Treatise on the Isoptera of the world[END_REF]. [START_REF] Carrijo | New World Heterotermes (Isoptera, Rhinotermitidae): molecular phylogeny, biogeography and description of a new species[END_REF] proposed a phylogeny of the New World Heterotermes and have highlighted the hyaline labrum of soldiers as a generic character (Carrijo et al., 2020: figs 1B,C), while the labrum in the genus Reticulitermes is fully sclerotized. This strengthens our taxonomic attribution. Interestingly, fossils of the genus Reticulitermes are rather frequent in the Baltic amber [START_REF] Nel | Les Isoptera fossiles (Insecta, Dictyoptera)[END_REF], with two species described from alate specimens, viz. R.

antiquus [START_REF] Germar | Insecten in Bernstein eingeschlossen, beschrieben aus dem academischen Mineralien-Cabinet zu Halle[END_REF] and R. minimus [START_REF] Snyder | A new Reticulitermes from Baltic Sea amber (Insecta Isoptera)[END_REF], making comparisons with the newly described soldier specimen impossible. Engel et al. (2007a) 2008. In terms of topologies, all BI analyses resulted in very similar tree reconstructions, except for the poorly supported (PP < 0.5) relationships within the 'Meiatermes-grade'.

In addition to defining the relationships of the two new fossil taxa, our main aim was to refine time divergence estimates in Isoptera (Fig. Partitioning the data reveals that each partition (alates/workers, soldiers, and life-history traits)

has its own history, as shown with their relative rates of changes. While higher rates were found along the branch leading to termites without any partitioning (Figure 6), higher rates were found either in most of the deepest nodes (alates/workers partition), for Nasutitermitinae (soldiers partition), or for the outgroups (life-history traits partition).

Discussion

A possible Triassic/Jurassic origin of eusociality

If we assume that tip-dating results better reflect the timing of evolution of termites than nodedating analyses (see below The disagreements between node-and tip-dating estimates underline that further investigations are needed to clarify the timing of termite diversification and of the origin of their eusociality. Several reasons may explain those differences (Matzke & Wright, 2016), including recent fossil discoveries that could not be included in earlier studies [START_REF] Bourguignon | The evolutionary history of termites as inferred from 66 mitochondrial genomes[END_REF][START_REF] Bourguignon | Transoceanic dispersal and plate tectonics shaped global cockroach distributions: evidence from mitochondrial phylogenomics[END_REF]Legendre et al., 2015;[START_REF] Engel | Morphologically specialized termite castes and advanced sociality in the Early Cretaceous[END_REF][START_REF] Bucek | Evolution of termite symbiosis informed by transcriptome-based phylogenies[END_REF][START_REF] Evangelista | An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites (Blattodea)[END_REF][START_REF] Zhao | The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites[END_REF]Zhao et al., , 2020a; but see Ware et al., 2010 for older estimates). In many taxonomic groups, recent fossil discoveries have impacted our understanding of their timing of evolution. Termites are not an exception (e.g. [START_REF] Selden | A golden orb-weaver spider (Araneae: Nephilidae: Nephila) from the Middle Jurassic of China[END_REF][START_REF] Huang | Diverse transitional giant fleas from the Mesozoic era of China[END_REF][START_REF] Huang | Amphibious flies and paedomorphism in the Jurassic period[END_REF][START_REF] Toussaint | To what extent do new fossil discoveries change our understanding of clade evolution? A cautionary tale from burying beetles (Coleoptera: Nicrophorus)[END_REF][START_REF] Nicholson | Changes to the fossil record of insects through fifteen years of discovery[END_REF].

Our dating increases the discrepancy between phylogenetic age estimates and the termite fossil record. Such a mismatch is not uncommon and is often explained by fossilization biases. Even at uncontroversial times-i.e. when termites were undoubtedly present-their presence in the fossil record might be scarce. Less than 1% of all insect specimens from the Early Cretaceous are termites (Engel et al., 2009: Table 2) and there is currently no Triassic or Jurassic record of termites. Putative termite nests have been suggested from the Triassic and Early Jurassic [START_REF] Hasiotis | Termite (Insecta: Isoptera) nest ichnofossils from the upper triassic chinle formation, petrified forest national park, Arizona[END_REF][START_REF] Bordy | Advanced Early Jurassic termite (Insecta: Isoptera) nests: evidence from the Clarens Formation in the Tuli Basin, Southern Africa[END_REF] but those ichnofossils remain controversial [START_REF] Genise | Ichnoentomology: insect traces in soils and paleosols[END_REF]. Our datings of crown-Isoptera, concomitant to those putative nests, could fuel the debate around those nests and their significance for the origin of eusociality, although the construction of large mounds is a derived trait, acquired more recently, within termites. Irrespectively of those dubious fossils (Legendre et al. 2015), nest structures in termites result from advanced cooperation among colony members, revealing highly integrated social systems [START_REF] Grassé | Comportement -socialité -écologie -évolution -systématique[END_REF][START_REF] Invernizzi | Deconstructing collective building in social insects: implications for ecological adaptation and evolution[END_REF]. The capacity to build a nest was probably inherited by the common ancestor of the crown-Isoptera [START_REF] Mizumoto | Modern termites inherited the potential of collective construction from their common ancestor[END_REF]. Although nest building does not require a soldier caste, it does require a worker caste ('true' workers or pseudergates), which has been found in Early Cretaceous deposits for the latest [START_REF] Martínez-Delclòs | The oldest known record of social insects[END_REF]. Recent descriptions of workers and soldiers [START_REF] Engel | Morphologically specialized termite castes and advanced sociality in the Early Cretaceous[END_REF][START_REF] Zhao | The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites[END_REF]Zhao et al., , 2020a) ) have confirmed that eusociality was already established in the Early to 'mid'-Cretaceous.

The oldest termite fossils correspond to alates, which are indecisive for deciphering the origin of eusociality. Those fossils have wing sutures, like living alate termites, suggesting that these ancient termites had adult swarms and lived the rest of their life in hidden, cryptic environments. But it does not mean that they were eusocial insects. In other words, although it is reasonable to think that fossil taxa of crown-Isoptera were eusocial, alates from stem-Isoptera do not inform about the putative eusociality of those stem-lineages.

Living in cryptic environments during the Late Triassic-Early Jurassic was undoubtedly advantageous because dinosaurs, small lizards, and mammals diversified at that time, with at least some of those mammals that were insectivorous [START_REF] Luo | Transformation and diversification in early mammal evolution[END_REF]Melo et al., 2019). The Late Jurassic mammal Fruitafossor convergently developed the main morphological characters that allow extant armadillos and aardvarks to dig the ground and feed on termite nests [START_REF] Luo | A Late Jurassic digging mammal and early mammalian diversification[END_REF][START_REF] Luo | Transformation and diversification in early mammal evolution[END_REF].

Interestingly, Fruitafossor was found in the Late Jurassic Morrison Formation (Colorado, USA), in layers where some putative termite nests were also described [START_REF] Engelmann | Stratigraphic and geographic distribution of fossils in the upper part of the Upper Jurassic Morrison Formation of the Rocky Mountain region[END_REF][START_REF] Koch | Paleontological discoveries at Curecanti National Recreation Area and Black Canyon of the Gunnison National Park, Upper Jurassic Morrison Formation, Colorado[END_REF][START_REF] Smith | New social insect nests from the Upper Jurassic Morrison Formation of Utah[END_REF]. Its presence may provide additional support to the hypothesis that termite nests and eusociality were already established at that time, sufficient to provide a food source for a highly specialized mammal. But, alternatively, Fruitafossor might also have fed on other unknown insects, even if there is no obvious candidate group in the rather well-known Late Jurassic entomofauna; those hypotheses need to be investigated further. As indicated by the fossil record and the estimates of isopteran stem and crown ages, it is highly probable that the cryptic termites became eusocial during the period between the Early and Late Jurassic.

It is however unclear whether first termites (i.e. stem-Isoptera but also †Cratomastotermes, †Baissatermes, etc.) were deprived of a soldier caste like the first diverging lineages of corbiculates, some halictine bees, and ants ( †Haidomyrmecinae, †Sphecomyrminae, and poneromorphs).

Similarly, it is unclear whether stem-lineages of termites had a worker caste. But if they did, it is often believed that the polymorphism between workers and reproducers was reduced (Engel et al., 2007b;[START_REF] Michener | The social behavior of the bees: a comparative study[END_REF][START_REF] Hölldobler | Caste and division of labor[END_REF]Barden & Grimaldi, 2016).

Refining stem and crown-ages within Isoptera and identifying ghost lineages

Differentiating stem-from crown-groups is pivotal to our understanding of how lineages waxed and waned [START_REF] Budd | The dynamics of stem and crown groups[END_REF]. In Isoptera, the difference has sometimes been elusive for three reasons. First, the termite fossil record is full of gaps, for instance between the 'mid'-Cretaceous and the Paleocene-Middle Eocene. Second, some families (e.g. Mastotermitidae) are only represented by one or two species. Third, the assignment of certain fossil species to extant lineages is questionable due to their poor preservation. All these issues obfuscate the delineation between stem-and crowngroups, and lead to misestimating their ages. It might, however, be clarified with tip-dating analyses wherein fossils are included in the phylogenetic reconstructions.

With Mastotermes darwiniensis as its single living species and several extinct species sharing plesiomorphies, the Mastotermitidae is one of the families for which delineating stem-and crowngroups is the most complicated [START_REF] Zhao | The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites[END_REF]. Although its phylogenetic position as the firstdiverging lineage within Isoptera is well-established (e.g. [START_REF] Kambhampati | Phylogenetic relationship among termite families based on DNA sequence of mitochodrial 16S rRNA gene[END_REF][START_REF] Donovan | Morphological phylogenetics of termites (Isoptera)[END_REF][START_REF] Thompson | Phylogenetic evidence for a single, ancestral origin of a 'true' worker caste in termites[END_REF][START_REF] Inward | A comprehensive phylogenetic analysis of termites (Isoptera) illuminates key aspects of their evolutionary biology[END_REF]Legendre et al., 2008), its synapomorphy that could be discernable on fossils has been clarified only recently: a thick and sclerotized M vein, stronger than CuA [START_REF] Zhao | The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites[END_REF]. On the basis of this synapomorphy, the family includes now six genera, i.e., Anisotermes, Blattotermes, Garmitermes Mastotermes, Milesitermes gen. nov., and Miotermes. Zhao et al. (2019: 621) excluded the genera Valditermes, Idanotermes, and Khanitermes from the Mastotermitidae. The relationships of Spargotermes remain uncertain [START_REF] Wappler | The effects of fossil placement and calibration on divergence times and rates: an example from the termites (Insecta: Isoptera)[END_REF][START_REF] Bezerra | New record of Mastotermitidae from Fonseca Basin, Eocene-Oligocene boundary of southeastern Brazil[END_REF]. The genus Mastotermes is possibly not monophyletic but its definition is out of the scope of this paper. We dated this highly-supported clade from the Early Cretaceous ( 130 Mylacrotermes and some Meiatermes species, and may have originated in the Early Jurassic ( 188Ma, 166-209 95% HPD). The composition of stem-Mastotermitidae is, however, not clearly settled because deeper nodes in the phylogeny of termites are weakly supported. This is notably due to the quality of the fossil record, with several species of the so-called Meiatermes-grade based on incomplete specimens.

Similarly, we have clarified the delineation between stem-and crown-groups of the following families: Archotermopsidae, Stolotermitidae, Hodotermitidae, and Kalotermitidae. Our age estimates agree with those previously found for the Archotermopsidae but not for the Hodotermitidae [START_REF] Bourguignon | The evolutionary history of termites as inferred from 66 mitochondrial genomes[END_REF][START_REF] Bucek | Evolution of termite symbiosis informed by transcriptome-based phylogenies[END_REF]. The split between those two families would have occurred in the Early Cretaceous (116 Ma, 73-160 mean 95% HPD), revealing a ghost lineage of ca.

90 Ma for Hodotermitidae. However, in all recent molecular works (e.g. [START_REF] Bourguignon | The evolutionary history of termites as inferred from 66 mitochondrial genomes[END_REF][START_REF] Bourguignon | Transoceanic dispersal and plate tectonics shaped global cockroach distributions: evidence from mitochondrial phylogenomics[END_REF]Legendre et al., 2015), Hodotermitidae are nested within Archotermopsidae, contrary to what we found here. This disagreement, which might be reduced with future works on Hodotermitid fossils (CJ in prep.), unavoidably affects dating estimates. Another ghost lineage, the one leading to Stolotermitidae, has been recently investigated (Zhao et al., 2020a). This study and ours rely on the same dataset and their results concur; we nonetheless suggest a putative stem-Stolotermitidae (i.e.

Tanytermes) that would require further investigation as its phylogenetic position is poorly supported.

Ghost lineages have always challenged the understanding of organism diversification. They emphasize how rocks and clocks disagree, how the fossil record and phylogenetic dating estimates might diverge [START_REF] Ronquist | Closing the gap between rocks and clocks using total-evidence dating[END_REF]. Gaps in the insect fossil record are notorious [START_REF] Nel | Diversification of insects since the Devonian: a new approach based on morphological disparity of mouthparts[END_REF] and are more prone to occur for some lineages than others. For termites, the putative cryptic way of life of stem-lineages was an unfavorable condition for their fossilization, which might explain the absence of a fossil record in the Early and Middle Jurassic. But even some lineages with important mating swarms, like the Hodotermitidae, might show important gaps in their fossil record.

Underlining those gaps and refining their temporal delimitation contribute to clarify which deposits should be further investigated, and hopefully make termite rocks and clocks converge.

Better models for better dating estimates

Because of the well-identified limitations of node-dating analyses, tip-dating analyses are becoming popular [START_REF] Parham | Best practices for justifying fossil calibrations[END_REF]Ronquist et al., 2012;[START_REF] Heath | The fossilized birth-death process for coherent calibration of divergence-time estimates[END_REF]Matzke & Wright, 2016;[START_REF] Simões | Sphenodontian phylogeny and the impact of model choice in Bayesian morphological clock estimates of divergence times and evolutionary rates[END_REF]. They are, however, not devoid of their own limitations, which has occasionally triggered some doubts in their resulting age estimates, which might have seemed unreasonably old (e.g. [START_REF] Arcila | Mass extinction in tetraodontiform fishes linked to the Palaeocene-Eocene thermal maximum[END_REF]. Several reasons have been suggested from the lack of a FBD process, to inaccurate topologies and inadequate models of evolution [START_REF] Arcila | Mass extinction in tetraodontiform fishes linked to the Palaeocene-Eocene thermal maximum[END_REF][START_REF] Luo | A simulation-based evaluation of tip-dating under the fossilized birth-death process[END_REF][START_REF] Simões | Sphenodontian phylogeny and the impact of model choice in Bayesian morphological clock estimates of divergence times and evolutionary rates[END_REF]. Here, tip-dating analyses all provide older estimates than nodedating analyses. However, to both test and circumvent the aforementioned potential issues, we have implemented several FBD-based analyses and constrained a few nodes in the topology. Using uniform or fixed calibrations for fossil tips resulted in similar datings (Supplementary Information).

Fixed calibrations slightly pulled down time divergence estimates, but using uniform priors seems more appealing and relevant because the exact age of each fossil is unknown. Likewise, we have tested different priors (i.e. clockrate and speciation priors, mean age for the ingroup) but it did not significantly change our results. One parameter-allowing fossil ancestor to be sampled or not-had a deeper impact on dating estimates but none of the analyses resulted in ages as young as in the node-dating analyses. Analyses wherein fossil ancestor were allowed to be sampled even produced the oldest estimates (if we omit the analysis with a uniform clock), but some parameter estimates are dubious (e.g. estimated mean of branch rate variance) so that ages estimated from those analyses should be interpreted with caution. A recent study on sphenodontian reptiles came to the same conclusion regarding the negative impact of sampling for ancestors in dating estimates relying only on morphological datasets [START_REF] Simões | Sphenodontian phylogeny and the impact of model choice in Bayesian morphological clock estimates of divergence times and evolutionary rates[END_REF].

Overall, we consider the dating estimates produced with the birth-death-serial-sampling (BDSS) models as reasonable up-to-date hypotheses, while the estimates resulting from sampledancestor BDSS are dubious. Note that for some of the shallowest nodes, the topology-and thus the associated datings-is inaccurate, which is a typical consequence of a reduced morphological dataset [START_REF] Luo | A simulation-based evaluation of tip-dating under the fossilized birth-death process[END_REF]. It has arguably a very limited effect on dating estimates of the deepest nodes, those we mostly focused on in this work, especially knowing that we include numerous fossil taxa in our analyses [START_REF] Luo | A simulation-based evaluation of tip-dating under the fossilized birth-death process[END_REF].

Beyond including more fossil taxa into tip-dating or total-evidence phylogenetic analyses to produce more robust age estimates, improving the models for morphological data is a complementary direction [START_REF] Wright | Bayesian analysis using a simple likelihood model outperforms parsimony for estimation of phylogeny from discrete morphological data[END_REF][START_REF] Simões | Sphenodontian phylogeny and the impact of model choice in Bayesian morphological clock estimates of divergence times and evolutionary rates[END_REF]. Herein, we have tested a Mkv and Mkv + G models, as well as the impact of morphological data partitioning. It had an insignificant effect on dating estimates but it unsurprisingly revealed that each partition had its own evolutionary trajectory.

In other words, evolutionary rates for soldier-related characters differ from the rates of worker/alaterelated characters. The differences are striking for some lineages (e.g. within Termitidae; Figure 6) and a partitioning approach should be more often used for morphological data, like it is now routinely done with molecular data. Partitioning morphological data might affect tree reconstructions and dating estimates, or be the foundation for studies aiming at deciphering morphological evolution (e.g. [START_REF] Zhang | Bayesian tip dating reveals heterogeneous morphological clocks in Mesozoic birds[END_REF]. Whatever the aim, the ongoing developments in models for morphological data must go on (e.g. [START_REF] O'reilly | Bayesian methods outperform parsimony but at the expense of precision in the estimation of phylogeny from discrete morphological data[END_REF][START_REF] Puttick | Uncertain-tree: discriminating among competing approaches to the phylogenetic analysis of phenotype data[END_REF][START_REF] Goloboff | Weighted parsimony outperforms other methods of phylogenetic inference under models appropriate for morphology[END_REF].

Conclusion

Time divergence estimates of the Isoptera and of some constitutive families may be older than previously stated. If our results are correct, the crown Isoptera may have arisen ca. 205 , contrasting with most molecular-based node-dating analyses (between 134-165 Ma) [START_REF] Bourguignon | The evolutionary history of termites as inferred from 66 mitochondrial genomes[END_REF]Legendre et al., 2015;[START_REF] Evangelista | An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites (Blattodea)[END_REF][START_REF] Bucek | Evolution of termite symbiosis informed by transcriptome-based phylogenies[END_REF]. We infer those results from the termite fossil record that we have enriched with two new species from the Burmese mid-Cretaceous amber and Baltic Eocene amber. This suggests that termites have developed a 'highly specialized' eusociality with a tripartite caste system before ants and other eusocial taxa.

This social organization might have influenced their ability to survive mass extinctions (e.g. [START_REF] Bakker | Dinosaur mid-life crisis: the Jurassic-Cretaceous transition in Wyoming and Colorado[END_REF] or favored the appearance of other modern eusocial insects such as Hymenoptera (particularly wasps and ants) by providing an abundant food source. Additionally, acquiring a eusocial system has certainly enabled termites to become a group with a major ecological role in various palaeoenvironments. By refining and providing new time divergence estimates of the crown-groups of termite families, our study might allow better palaeoecological inferences from their extant representatives. Clarifying the distinction between stem-and crown-groups among termite families will also contribute to a better understanding of termite diversification.

Although we have proposed a new hypothesis for the tempo of diversification of termites, further analyses and material are needed, especially for the 'Meiatermes-grade', whose lineages probably belong to the stem-Mastotermitidae or to a transition stage between Mastotermitidae and Euisoptera. Further advances might also be achieved from improvements in evolutionary models for morphological data like our partitioned analyses suggest.

Data availability statement

The data that support the findings of this study are available from the corresponding authors upon reasonable request. The phylogeny of termites (Dictyoptera: Isoptera) based on mitochondrial and nuclear markers:

implications for the evolution of the worker and pseudergate castes, and foraging behaviors. (2,200) 0.001103 0.000568 -0.001722 IGR 0.003359 -1015. 27 -1061.35 4333.46 gamma (2,200) 0.000750 0.000426 -0.001114 IGR 0.002953 -1015. 97 -1060.75 4730.44 gamma (2,200) 0.000748 0.000425 -0.001110 IGR 0. 005948 -1652.89 -1726.05 3041.98 gamma (2,200) 0.001468 0.000968 -0.002038 IGR 0.004800 -1655. 68 -1735.25 3351.94 gamma (2,200) 0.001526 0.000985 -0.002088 IGR 0. 005522 -1655.21 -1730.83 3611.75 gamma (2,200) 0.001434 0.000924 -0.001979 IGR 0.007870 -1638. 53 -1729.58 2052.92 gamma (2,200) 0.001447 0.000959 -0.001965 IGR 0.006890 -1643. 08 -1726.95 2436.13 gamma (2,200) 0.001524 0.000984 -0.002120 IGR 0.006027 -1695. 03 -1769.22 2352.46 gamma (2,200) 0.001215 0.000938 -0.001500 IGR 0.005028 -1654.29 -1725.98 3540.81 normal(0.0025,1) 0.001427 0.000944 -0.001975 IGR 0. 005025 -1781.23 -1838.72 2280.86 gamma (2,200) 0.003034 0.002167 -0.003995 IGR 0.004831 -1648.80 -1734.16 3462.06 gamma (2,200) 0.001498 0.000980 -0.002054 IGR 0.005033 -1740. 29 -1801.50 2012.46 gamma (2,200) 0.001586 0.001092 -0.002088 IGR 0.004945 -1750.22 -1807.29 1330.00 gamma (2,200) 0.001564 0.001099 -0.002059 IGR 0.004066 -1744. 75 -1798.99 1619.54 gamma (2,200) 0.001508 0.001046 -0.001991 IGR 0.005301 -1650. 89 -1728.98 2773.18 gamma (2,200) 0.000921 0.000571 -0.

Molecular

  Ma,. Stem-Mastotermitidae-containing by definition all extinct organisms more closely related to crown-Mastotermitidae than to any other crown-group[START_REF] Budd | The dynamics of stem and crown groups[END_REF])would include Idanotermes,Krishnatermes, Valditermes, Cratomastotermes, Carinatermes,

  Latreille, P.A. (1802) Histoire naturelle, générale et particulière des Crustacés et des Insectes. Ouvrage faisant suite aux oeuvres de Leclerc de Buffon et partie du cours complet d'Histoire naturelle rédigé par C.S. Sonnini. T. 3-4, an X. Familles naturelles et genres. Paris, Dufart, 1-467 and 1-387. Latreille, P.A. (1810) Histoire naturelle, générale et particulière des Crustacés et des Insectes. Ouvrage faisant suite aux oeuvres de Leclerc de Buffon et partie du cours complet d'Histoire naturelle rédigé par C.S. Sonnini. T. 13, an XIII. Familles naturelles et genres. Paris, Dufart, 1-432.Legendre, F., Whiting, M. F., Bordereau, C., Cancello, E. M., Evans, T. A.& Grandcolas, P. (2008)

Fig. 1 .Fig. 2 .

 12 Fig. 1. Milesitermes engeli gen. et sp. nov., holotype IGR.BU-012. Photographs. (A) habitus in dorsal

Fig. 3 .

 3 Fig. 3. Reticulitermes grimaldii sp. nov., holotype IGR.BA-020 (A) photographs. Habitus in ventro-

Fig. 4 .Fig. 5 .

 45 Fig. 4. Reticulitermes grimaldii sp. nov., holotype IGR.BA-020. Reconstruction drawings. (A)

Fig. 6 .

 6 Fig. 6. Dated-trees recovered from Bayesian analyses. (A) node-dating (Script_node-dating), only

Fig. 7 .

 7 Fig. 7. Dated-trees recovered from Bayesian analyses. FBD model with three partitions (script_FBD-

1904 Milesitermes gen. nov. Jouault & Nel

  . The consensus cladogram with bs and Bremer indices is provided as a Supplementary file Even if this fossil is partly damaged, we can undoubtedly assign it to the Mastotermitidae based on the five tarsomeres, head without fontanelle (even if partially destroyed, we did not notice any external groove or internal gland or structures suggesting the presence of a fontanelle), large pronotum as wide as head, slightly saddle-shaped, and cerci four-segmented. Additionally, Milesitermes gen. nov. differs from the only other Mastotermitidae soldier (Anisotermes xiai) from the Cretaceous period in having thin hooked mandibles without a basal tooth, pronotum trapezoidal-

	Diagnosis: Head capsule extended, cylindrical-rectangular shaped. Antenna moniliform. Epicranial
	suture maybe present. Mandibles stout slightly elongated, and hooked apically, right mandible with
	one marginal tooth. Pronotum remarkable, trapezoidal, large, as wide as head capsule, anterior margin
	concave, slightly saddle shape, lateral margin convex, posterior margin nearly straight. Procoxal
	carina present. Legs with tarsi pentamerous, tibial spur formula 3-4-4, all tibiae with extra spines
	(Suppl. fig. 1). and rows of macrosetae. Abdomen plump and slightly elongate, and cerci four-segmented.
	Published work and nomenclatural acts are registered in ZooBank (http://www.zoobank.org/, Milesitermes engeli sp. nov. Jouault & Nel (Figs 1-2)
	last urn:lsid:zoobank.org:act:70D5C4CA-3CA2-4E09-98E7-D943E2C909B1 access: 20 February 2021), with the following	LSID	(reference):
	urn:lsid:zoobank.org:pub:D3EBBA6B-F488-4B56-AC98-0418A6B353FC Holotype: soldier, accession number IGR.BU-012 (a nearly complete specimen in a rounded piece of
	amber measuring 5 × 10 × 3 mm with numerous debris and wood fiber).
	Results Etymology: The specific epithet is a patronym honoring our colleague Pr. Michael S. Engel, eminent
	Systematic Palaeontology American paleoentomologist, and specialist of Isoptera. The species epithet is to be treated as a noun
	Order Blattodea, Latreille, 1810 in the genitive case.
	Infraorder Isoptera Brullé, 1832 Locality and horizon: Noije Bum Hill, Hukawng Valley, Kachin State, Myanmar; upper Albian to
	Epifamily Termitoidae Latreille, 1802 lower Cenomanian, mid-Cretaceous.
	Diagnosis: As for the genus. Description: Large soldier termite, body length ca. 6.40 mm from tip of mandible to apex of abdomen. Head massive, width ca. 1.65 mm, length ca. 2.35 mm from buccal aperture to posterior Family Mastotermitidae Desneux, (Figs 1-2) margin of head, ca. 2.7 mm total (including mandibles); head capsule longer than wide; lateral
	urn:lsid:zoobank.org:act:5711364A-9468-4315-A380-23CB17AF96BB margins nearly parallel; posterior border touching anterior margin of pronotum. Mandibles strongly
	Etymology: The generic epithet is a combination of the Latin word Miles meaning 'soldier' and sclerotized, clearly prognathous, dentate; left mandible destroyed; right mandible partially preserved,
	Termes commonly used to designate Isoptera. The generic epithet refers to the description based on ca. 0.82 mm long, greatest exposed width ca. 0.28 mm; dentition visible on right mandible only; apex
	a soldier specimen. Gender masculine.
	Type species: Milesitermes engeli sp. nov.

of right mandible slender, pointed and hooked, with one shallow triangular median tooth. Labrum, anteclypeus, postclypeus, and clypeus not preserved. Antennae partially destroyed, with more than six antennomeres (six preserved on left antenna). Compound eye not visible. Posterior part of head capsule with maybe remains of a faint epicranial ecdysial cleavage scar. Maxillary palps with five palpomeres, basal palpomere shortest, ultimate one longest; maxillary palpomeres length (when measurable) in mm: P5 0.26, P4 0.21, P3 0.2; labial palps with three(?) visible palpomeres, basal palpomere shortest, apical one longest. Paraglossa three(?)-segmented with apical segment the longest and apically rounded. Lacinial incisor conspicuous with one apical tooth and a basal one shorter. Galea visible but poorly preserved, other mouthparts hardly visible. Cervical sclerites massive, two-segmented (an inner cervical sclerite and lateral cervical sclerite on each side).

Pronotum massive in contact with head posterior margin, widest than head, trapezoidal shaped with anterior part the widest, anterior margin concave, antero-lateral corners rounded; lateral margins gently convex; posterior margin nearly straight with a small median concavity, postero-lateral corner broadly rounded; pronotal dimensions: median length 0.71 mm; greatest length 0.8 mm; greatest width 1.78 mm (across anterior portion); median portion of pronotum slightly raised longitudinally, two reduced depressions lateral to this raising. Mesonotum and metanotum largely transverse, mesonotal greatest length 0.66 mm, width 1.32 mm; mesonotum trapezoidal shaped with antero-and postero-lateral corners rounded, mesonotum overlapping metanotum. Legs with tarsi wholly pentamerous (i.e., no cryptic tarsomere), tibiae spinose, tibial spines asymmetrical, with finely serrate margin; arolium present between pretarsal claws. Foreleg smallest, procoxa small with carina present, profemora ca. 1 mm long, protibia slender, ca. 0.7 mm long, protarsus ca. 0.3 mm, pretarsal claws 0.1 mm. Mesofemur ca 1.1 mm long; mesotibia ca. 0.9 mm, mesotarsus ca. 0.45 mm, mesotarsal claws stout, ca. 0.14 mm. Hind leg larger than others, metafemur stout and medially enlarged, maximal width ca. 0.36 mm, 1.2 mm long; metatibia 1.15 mm long, metatarsus long, ca. 0.5 mm long, metapretarsal claw 0.16 mm long. Tibial spines: protibia with f1, f2, f3, plus one more proximal (x); mesotibia with four spines m1, m2, m3, m4 plus one ventral spine (ma); metatibia with four spines h1, h2, h3, h4 (h1 and h4 small), plus two ventral spines (hb, hc) and a row of stout macrosetae.

Eleven visible abdominal tergites, last one the smallest, lengths for TIV to apical terga measured dorsally (in mm): 0.31, 0.25, 0.31, 0.35, 0.22, 0.14, 0.11, 0.06. Total length of abdomen ca. 2.5 mm.

Styli not observed; cerci slender, short (0.2 mm), and with four cercomeres, basal cercomeres the widest and apical most the longest.

Remarks:

shaped, peculiar tibial spurs arrangement, and almost no pilosity (but possible problem of preservation)

  Soldier. Body length around ca. 5.25 mm from tip of mandible to apex of abdomen, but body is slightly twisted so total length slightly greater. Head massive, width ca. 1.05 mm, length ca.2.25 mm from anterior margin of labrum to posterior margin of head, ca. 2.5 mm total (including

	during preparation. Hindleg apparently the largest, metacoxa 0.3 mm long (L), 0.2 mm wide (W);
	metatrochanter 0.16 mm L, 0.08 mm W; metafemur: 0.75 mm L, 0.23 mm W; metatibia swollen
	mandibles); head capsule rectangular, tapering posteriorly, longer than wide; lateral margins apically 0.83 mm L, 0.11 W, with two spines, metatarsus long, 0.33 mm L, metapretarsal claw ca.
	subparallel to slightly convex; posterior border virtually separated from anterior margin of pronotum. 0.07 mm L. Tibial spines: protibia with three spines f1, f2, f3, without a raw of stout ventral setae;
	Mandibles very heavily sclerotized, strongly prognathous; left mandible overlapping right mandible; mesotibiae with two spines m1, m2; metatibia with two spines: h2, h3 and a raw of stout ventral setae.
	mandible bases rather broad; right mandibles slightly shorter than left one but partly hidden; dentition At least eight abdominal sternites visible ventro-laterally, lengths along midline, from SII-SVIII (in
	not visible due to amber fractures, apparently absent, but, if present, only composed of minutes teeth mm): 0.22, 0.18, 0.18, 0.18, 0.07, 0.15, 0.11. Total length of abdomen ca. 1.75 mm. Styli ca. 0.07
	mm long; cerci slender, short (ca. 0.12 mm), and two-segmented.
	to preservation), with concave anterior margin, median portion raised longitudinally, lateral parts
	narrower than head, apparently slightly saddle-shaped with concave anterior margin; lateral margins depressed; lateral margins nearly parallel (very slightly convex), posterior margin slightly concave;
	nearly parallel, posterior margin slightly concave. Legs with tarsi tetramerous, tibial spur formula 3-pronotal dimensions: medial length 0.35 mm; greatest length 0.48 mm; greatest width 0.75 mm.
	2-2. Pro-and metatibia with a row of macrosetae. Abdomen plump; styli slender; cerci two-Meso-and metanotum slightly convex in lateral view but not describable. Legs with tarsi tetramerous,
	segmented.

In Baltic amber, Kaliningrad Oblast, Russia, Late Eocene (likely

Priabonian,

.

Diagnosis:

The new species can be readily distinguished from the two other Reticulitermes in Baltic amber, because it is the only soldier. Body and head medium of sized. Head capsule rectangular slightly tapering posteriorly, sides slightly convex. Labrum with four(?) apical setae. Mandibles elongated and strongly hooked in their apical third, right mandibles apparently slightly shorter than left one; left mandible overlapping right mandible. Fontanelle small situated close. Antennae moniliform with 14 antennomeres; antennal insertion surrounded by a small rim. Pronotum large,

Description:

at mandible bases; apex of left mandible slender, very pointed and hooked; left mandible 0.78 mm long, greatest exposed width ca. 0.15 mm. Maxillary palps with at least four palpomeres, basal palpomere shortest, ultimate one apparently the longest; labial palps not visible. Galea, cardo, stipes, and other mouthparts not visible (covered by a white veil). Cervical sclerites not visibles. Labrum long, triangular with convex sides, 0.45 mm wide x 0.35 mm long, covered with sparse short appressed setae, apically with four(?) distinct long setae on margin. Anteclypeus shallow with anterior margin convex and posterior margin slightly concave, 0.44 mm wide, 0.08 mm long; postclypeus somewhat rectangular, 0.55 mm wide, 0.11 mm long. Clypeus without any projection; postclypeus flat. Fontanelle preserved as a small concavity situated posterior to clypeus; setulae surrounding fontanelle present. Compound eye spot not discernable, absent or inconspicuous.

Antenna with 14 preserved antennomeres, antenna insertion delimited by a small rim. Pronotum large, narrower than head, partially bend due to preservation, apparently slightly saddle-shaped (maybe due tibiae spinose, tibial spines slightly symmetrical and smooth; arolium absent between pretarsal claws. Foreleg smallest, procoxa small, protibia slender, ca. 0.46 mm long, protarsus 0.22 mm long, pretarsal claws slightly smaller than metapretarsal claws; profemur thick, stout. Mid legs partly destroyed

  indicated that R. antiquus is the most common termite in Baltic amber. Additionally, these authors have proposed to distinguished R. antiquus from R. minimus by its larger body size (length of forewing from basal suture 6.5-8.8 mm in R. antiquus vs. 3.94 mm in R. minimus) and more numerous antennomeres (17-20 in R. antiquus vous. 13-14 in R. minimus)(Engel et al., 2007a). The size of Reticulitermes grimaldii sp. nov. could match with R. antiquus since termites soldiers are rarely larger than alates. However, the number of antennal segments cannot be used to assign Reticulitermes grimaldii sp. nov. to one of the previously described species since it varies between soldier and alate castes. In termites, a general trend established that the body length of workers and soldiers relative to alates is close to 0.8(Engel et al., 2016 page 6 and fig.S3). Therefore, Reticulitermes grimaldii sp. nov. cannot be attributed to R.

minimus because its body is nearly twice as long. Similarly, Reticulitermes grimaldii sp. nov. cannot be attributed to R. antiquus since the soldier is slightly longer than the alates of R. antiquus. Phylogenetic analyses Milesitermes engeli gen. et sp. nov. and Reticulitermes grimaldii sp. nov. are nested, respectively, within Mastotermitidae and Rhinotermitidae in BI and MP (Figs 5-6; Supp. fig. 1), confirming our taxonomic attributions. The 'mid'-Cretaceous genus Krishnatermes Engel et al., 2016 is closely related to the Mastotermitidae as the sister taxon of the Middle Eocene Idanotermes desioculus Engel,

  5,. All tip-dating analyses (run with the FBD model, except one analysis with a uniform strict clock) push back the time divergence estimates of stem-and crown-Isoptera of several tens of Myrs: 232 Ma (172-362 Ma min and max values of 95% HPD) and 205 for stem-and crown Isoptera across all FBD analyses, respectively, vs 189 and 144 in the nodedating analyses (Fig.6A, Script_node-dating). In the tip-dating analysis with one partition (Fig.5), the most recent common ancestor (MRCA) of Euisoptera is estimated to have arisen 185. The MRCA of Neoisoptera is estimated to have arisen 109 Ma (99-126 Ma 95% HPD), while the MRCA of Termopsidae is estimated to have arisen 54 Ma (37-83 Ma 95% HPD).Archotermopsidae and Hodotermitidae are sister-taxa and their crown-groups would date back from the K/T and the Oligocene/Miocene boundaries, respectively (68 Ma, 42-101 95% HPD and 28 Ma, 6-56 95% HPD). Extant Stolotermitidae have their MRCA from the Eocene (38 Ma, 9-75 95% HPD), while their oldest relatives (from the genus Cosmotermes) would date back to the Lower Cretaceous (115 Ma, 101-137 95% HPD). As for Kalotermitidae, the crown-group would have originated during the Paleocene (58 Ma, 36-86 95% HPD), while the stem-Kalotermitidae would date back to the Lower Cretaceous (126 Ma, 110-146 95% HPD).

Overall, the oldest age estimates are found, for all nodes, when a uniform clock is used. The tip-dating analysis that gives the youngest age (viz. with fixed calibrations and a diversity sampling), still produces older mean estimates than node-dating analyses, except for the shallowest nodes. Time divergence estimates for the main isopteran lineages are summarized across all analyses in Table

1

, whereas several priors, statistics and parameter values are summarized in Supporting Information.

  ), crown-Isoptera would have arisen around the Late Triassic / Early Jurassic boundary (ca.205 Ma,, 50-60 Ma earlier than currently thought (as reflected with node-dating results: 144 Ma, 137-166 mean 95% HPD). This suggests an even older origin of stem-Isoptera, dating back to the Late Triassic. Those age estimates were consistently found across all tip-dating analyses, even with very flat priors or when priors were centered on much younger ages (e.g. mean age for crown Isoptera set to 144 Ma); some tip-dating analyses even produced older age estimates but those might be biased (e.g. random sampling, see below).

Table 1 .

 1 Date estimates (in Ma) for Isoptera and subclades. Age estimates for stem-and crowngroups are distinguished when possible. Estimates are older for Fossilized Birth-Death analyses that include extinct species when compared to the node dating analyses (with only extant species); analyses wherein sampled ancestors are allowed give old estimates and several dubious values for other parameters (see Supporting Information); 144 and 163 Ma correspond to the mean age for Isoptera (150 Ma for other analyses).

	Figure 1

Systematic Entomology

Acknowledgments

CJ is grateful to Dr. Phillip Barden (NJIT, Newark; AMNH, New York, USA) who provided literature on phylogenetic analyses. We thank three anonymous reviewers for their constructive comments.

Author contributions

C.J. and F.L. are first authors with equal rank and performed the phylogenetic analyses; C.J. and A.N. coordinated the manuscript, wrote the descriptions, and prepared the figures; all authors drafted the manuscript.

Conflicts of interest statement

The authors declare that there are no conflicts of interest.

Supplementary files:

Appendix 2. List of character states. [START_REF] Iturralde-Vincent | Age and palaeogeographical origin of Dominican amber[END_REF][START_REF] Seyfullah | Production and preservation of resins -past and present[END_REF]. Due to the lack of information concerning the exact provenance of the amber we do not constrain to precisely the age of taxa derivating from this deposit. Radiometric data established an early Cenomanian age (98.79 ± 0.62 Ma) while some ammonites found in the amber-bearing bed and within amber corroborate a late Albian / early Cenomanian age (Cruickshank & Ko, 2003;Yu et al., 2019). We use the minimum age recovered from zircon and a mid-Albian age allowing errors due to the older stratigraphic levels currently mined and where the amber piece might have come from. Blue Earth Formation was dated as late Bartonian to Priabonian (upper Eocene, ca.

33.9-37.2 Ma) based on palynological data (Kosmoswska-Ceranowicz et al., 1997;Aleksandrova & Zaporozhets, 2008).

Most paleozoologists assign the Zaza Formation to the Neocomian, most probably Berriasian [START_REF] Rasnitsyn | Wasps (Insecta: Vespida = Hymenoptera) from the Purbeck and Wealden (Lower Cretaceous) of southern England and their biostratigraphical and palaeoenvironmental significance[END_REF]. However, the presence of Asteropollenites lead palynologists and paleobotanists to believe that the Baissa deposits are younger, possibly Aptian (Vakhrameev & Kotova, 1997). In any case, the faunistic data indicate pre-Aptian. We therefore us the stratigraphic age of former Aptian (when the taxa was described) and a upper most Beriasian age fitting with the controversial age of the deposit.

The amber deposit is Turonian age [START_REF] Doyle | Angiosperm pollen donation of the continental Cretaceous of the Atlantic Coastal Plain and its application to deep wells in the Salisbury Embayment[END_REF][START_REF] Christopher | Normapolles and triporate pollen assemblages from the Raritan and Magothy Formations (Upper Cretaceous) of New Jersey[END_REF] while the base of the Raritan formation is Cenomanian [START_REF] Brenner | Early angiosperm pollen differentiation in the Albian to Cenomanian deposits of Delaware (U.S.A.)[END_REF]Cooban & Kennedy, 1990) Dominican amber is mined from several deposit dated from the Langhian-Burdigalian [START_REF] Seyfullah | Production and preservation of resins -past and present[END_REF][START_REF] Iturralde-Vincent | Age and palaeogeographical origin of Dominican amber[END_REF]. Due to the lack of information concerning the exact provenance of the amber we do not constrain to precisely the age of taxa derivating from this deposit. Dominican amber is mined from several deposit dated from the Langhian-Burdigalian [START_REF] Seyfullah | Production and preservation of resins -past and present[END_REF][START_REF] Iturralde-Vincent | Age and palaeogeographical origin of Dominican amber[END_REF]. Due to the lack of information concerning the exact provenance of the amber we do not constrain to precisely the age of taxa derivating from this deposit.

The fossiliferous unit has been dated as Aptian or perhaps early Albian on the basis of its palynology [START_REF] Pons | Quelques observations sur la palynologie de l'Aptien supeŕieur et de l'Albien du bassin d'Araripe (N.E. du Breśil)[END_REF] and therefore, we follow the age proposed by http://fossilworks.org/ The fossiliferous unit has been dated as Aptian or perhaps early Albian on the basis of its palynology [START_REF] Pons | Quelques observations sur la palynologie de l'Aptien supeŕieur et de l'Albien du bassin d'Araripe (N.E. du Breśil)[END_REF] and therefore, we follow the age proposed by http://fossilworks.org/ Dominican amber is mined from several deposit dated from the Langhian-Burdigalian [START_REF] Seyfullah | Production and preservation of resins -past and present[END_REF][START_REF] Iturralde-Vincent | Age and palaeogeographical origin of Dominican amber[END_REF]. Due to the lack of information concerning the exact provenance of the amber we do not constrain to precisely the age of taxa derivating from this deposit. Dominican amber is mined from several deposit dated from the Langhian-Burdigalian (Seyfullah et al., 2018; Iturralde-Vincent & MacPhee, 1996). Due to the lack of information concerning the exact provenance of the amber we do not constrain to precisely the age of taxa derivating from this deposit. Radiometric data established an Early Cenomanian age (98.79 ± 0.62 Ma) while some ammonites found in the amber-bearing bed and within amber corroborate a late Albian / early Cenomanian age (Cruickshank & Ko, 2003;Yu et al., 2019). We use the minimum age recovered from zircon and a mid-Albian age allowing errors due to the older stratigraphic levels currently mined and where the amber piece might have come from. Mexican amber-bearing strata extend from the Balumtun Sandstone of the lower Miocene to the La Quinta Formation of the upper Oligocene [START_REF] Poinar | Hymenaea mexicana sp. nov. (Leguminosae: Caesalpinioideae) from Mexican amber indicates Old World connections[END_REF].

A lower Miocene mid-Oligocene age has been proposed by Berggren & Van Couvering, (1974) while a recent publication proposed a Serravallian-Burdigalian age [START_REF] Seyfullah | Production and preservation of resins -past and present[END_REF]. Therefore we chose a mean age corresponding to the Burdigalian-Aquitinian and fitting with most of previous studies (as proposed in http://fossilworks.org/) Blue Earth Formation was dated as late Bartonian to Priabonian (upper Eocene, ca.

33.9-37.2 Ma) based on palynological data (Kosmoswska-Ceranowicz et al., 1997;Aleksandrova & Zaporozhets, 2008). Blue Earth Formation was dated as late Bartonian to Priabonian (upper Eocene, ca.

33.9-37.2 Ma) based on palynological data (Kosmoswska-Ceranowicz et al., 1997;Aleksandrova & Zaporozhets, 2008). Dominican amber is mined from several deposit dated from the Langhian-Burdigalian [START_REF] Seyfullah | Production and preservation of resins -past and present[END_REF][START_REF] Iturralde-Vincent | Age and palaeogeographical origin of Dominican amber[END_REF]. Due to the lack of information concerning the exact provenance of the amber we do not constrain to precisely the age of taxa derivating from this deposit. Radiometric data established an early Cenomanian age (98.79 ± 0.62 Ma) while some ammonites found in the amber-bearing bed and within amber corroborate a late Albian / early Cenomanian age (Cruickshank & Ko, 2003;Yu et al., 2019). We use the minimum age recovered from zircon and a mid-Albian age allowing errors due to the older stratigraphic levels currently mined and where the amber piece might have come from.

The fossiliferous unit has been dated as Aptian or perhaps early Albian on the basis of its palynology [START_REF] Pons | Quelques observations sur la palynologie de l'Aptien supeŕieur et de l'Albien du bassin d'Araripe (N.E. du Breśil)[END_REF] and therefore, we follow the age proposed by http://fossilworks.org/ Dominican amber is mined from several deposit dated from the Langhian-Burdigalian (Seyfullah et al., 2018; Iturralde-Vincent & MacPhee, 1996). Due to the lack of information concerning the exact provenance of the amber we do not constrain to precisely the age of taxa derivating from this deposit. Mexican amber-bearing strata extend from the Balumtun Sandstone of the lower Miocene to the La Quinta Formation of the upper Oligocene [START_REF] Poinar | Hymenaea mexicana sp. nov. (Leguminosae: Caesalpinioideae) from Mexican amber indicates Old World connections[END_REF].

A lower Miocene mid-Oligocene age has been proposed by Berggren & Van Couvering, (1974) while a recent publication proposed a Serravallian-Burdigalian age [START_REF] Seyfullah | Production and preservation of resins -past and present[END_REF]. Therefore we chose a mean age corresponding to the Burdigalian-Aquitinian and fitting with most of previous studies (as proposed in http://fossilworks.org/) Oise amber is dated back from Sparnacian (early Eocene) continental facies [START_REF] Franceschi | Origine de l'ambre des faciès sparnaciens (Éocène inférieur) du Bassin de Paris : le bois de l'arbre producteur[END_REF] The fossiliferous unit has been dated as Aptian or perhaps early Albian on the basis of its palynology [START_REF] Pons | Quelques observations sur la palynologie de l'Aptien supeŕieur et de l'Albien du bassin d'Araripe (N.E. du Breśil)[END_REF] and therefore, we follow the age proposed by http://fossilworks.org/ The fossiliferous unit has been dated as Aptian or perhaps early Albian on the basis of its palynology [START_REF] Pons | Quelques observations sur la palynologie de l'Aptien supeŕieur et de l'Albien du bassin d'Araripe (N.E. du Breśil)[END_REF] and therefore, we follow the age proposed by http://fossilworks.org/ The fossiliferous unit has been dated as Aptian or perhaps early Albian on the basis of its palynology [START_REF] Pons | Quelques observations sur la palynologie de l'Aptien supeŕieur et de l'Albien du bassin d'Araripe (N.E. du Breśil)[END_REF] and therefore, we follow the age proposed by http://fossilworks.org/ Amber found in three intervals in the upper part of the Grès du Liban. Hammana is found in the upper interval, located between the Jezzinian above and the "Banc de Mrejatt" below. The "Banc de Mrejatt" includes one biostratigraphically significant benthic foraminifer: Eopalorbitolina transiens [START_REF] Cherchi | Montseciella, a new orbitolinid genus (Foraminiferida) from the Uppermost Hauterivian -Early Barremian of SW Europe[END_REF], which is a zonal marker of the lower to upper Barremian transition according to [START_REF] Schroeder | Revised orbitolinid biostratigraphic zonation for the Barremian -Aptian of the eastern Arabian Plate and implications for regional stratigraphic correlations[END_REF]. Accordingly the "Banc de Mrejatt" is correlated to the transgression of sequence Ba3 of [START_REF] Clavel | Dating and progradation of the Urgonian limestone from the Swiss Jura to South-East France[END_REF] and ascribed a late early Barremian age. The upper subunit is assigned a late Barremian age, although the amber may be reworked [START_REF] Maksoud | New data on the age of the Lower Cretaceous amber outcrops of Lebanon[END_REF]. we follow the age proposed by http://fossilworks.org/ Dominican amber is mined from several deposit dated from the Langhian-Burdigalian [START_REF] Iturralde-Vincent | Age and palaeogeographical origin of Dominican amber[END_REF][START_REF] Seyfullah | Production and preservation of resins -past and present[END_REF]. Due to the lack of information concerning the exact provenance of the amber we do not constrain to precisely the age of taxa derivating from this deposit. Radiometric data established an early Cenomanian age (98.79 ± 0.62 Ma) while some ammonites found in the amber-bearing bed and within amber corroborate a late Albian / early Cenomanian age (Cruickshank & Ko, 2003;Yu et al., 2019). We use the minimum age recovered from zircon and a mid-Albian age allowing errors due to the older stratigraphic levels currently mined and where the amber piece might have come from. Dominican amber is mined from several deposit dated from the Langhian-Burdigalian [START_REF] Iturralde-Vincent | Age and palaeogeographical origin of Dominican amber[END_REF][START_REF] Seyfullah | Production and preservation of resins -past and present[END_REF]. Due to the lack of information concerning the exact provenance of the amber we do not constrain to precisely the age of taxa derivating from this deposit. Dominican amber is mined from several deposit dated from the Langhian-Burdigalian [START_REF] Iturralde-Vincent | Age and palaeogeographical origin of Dominican amber[END_REF][START_REF] Seyfullah | Production and preservation of resins -past and present[END_REF]. Due to the lack of information concerning the exact provenance of the amber we do not constrain to precisely the age of taxa derivating from this deposit. Blue Earth Formation was dated as late Bartonian to Priabonian (upper Eocene, ca. List of the 17 analyses computed in 'Revising dating estimates and the antiquity of eusociality in termites using the fossilized birth-death process'

Run

ND : node dating analysis with a birth-death tree prior; one partition; diversity sampling ND-BD-random: same as ND but with a random sampling ND-uni-diversity: node dating analysis with a uniform tree prior; one partition; diversity sampling ND-uni-random: same as ND-uni-diversity but with a random sampling 1p-clock_uni: tip-dating analysis with a uniform clock tree prior; one partition, uniform fossil calibrations 1p: tip-dating analysis with a birth-death serial sampling (i.e. no sampled ancestor); one partition; uniform fossil calibrations 1p halved: same as 1p but with a more restricted age prior for the ingroup 1p twice: same as 1p but with an extended age prior for the ingroup 3p: same as 1p but with three partitions 3p rate variable: same as 3p but with a variable relative rate setting for partitions 1p-equal: same as 1p but with a Mkv model (and not a Mkv + G) 1p-uni-diversity: same as 1p but with a diversity sampling (sampled ancestor allowed) 1p-uni-random: same as 1p but with a random sampling (sampled ancestor allowed) 1p-flatpriors: same as 1p but with flat clock rate and speciation priors 1p-fixed-fossiltip: same as 1p but with fixed fossil calibrations 1p-fixed-diversity: same as 1p but with fixed fossil calibrations and a diversity sampling (sampled ancestor allowed) 1p-fixed-random: same as 1p but with fixed fossil calibrations and a random sampling (sampled ancestor allowed)