Phylogenetic analyses and their implication on the higher classification of termitesOur phylogenetic reconstructions support the monophyly of all currently recognised termite families (sensu Krishna et al.7, modified by Wang et al.33), except Rhinotermitidae, which is paraphyletic with respect to Serritermitidae and Termitidae, as previously acknowledged3,34,35,36,37,38,39,40,41,42,43,44.The Rhinotermitidae (sensu Krishna et al.7) can be divided into four monophyletic lineages diverging early alongside Serritermitidae and Termitidae (Figs. 1, 2): Heterotermitinae Froggatt + Coptotermitinae Holmgren, Psammotermitinae Holmgren + Prorhinotermitinae Quennedey & Deligne, Rhinotermitinae Froggatt, and Termitogetoninae Holmgren. All our analyses supported the monophyly of these four clades, and our analyses with RogueNaRok found no evidence of rogue taxa within these clades. Assigning each of these four clades a familial rank resolves the challenges embodied in Rhinotermitidae as traditionally conceived. Accordingly, we raise these four lineages to familial rank. We restrict Rhinotermitidae to Acorhinotermes, Dolichorhinotermes, Parrhinotermes, Rhinotermes, and Schedorhinotermes, the five genera composing the Rhinotermitinae (i.e., Rhinotermitidae Froggatt, sensu novo). We synonymise the two largely unused names Arrhinotermitinae and Leucotermitinae with Coptotermitinae and Heterotermitinae, respectively. We further synonymise Coptotermitinae with Heterotermitinae. The synonymy of the monogeneric Coptotermitinae (Coptotermes) with Heterotermitinae is justified by the paraphyly of Heterotermitinae, within which Coptotermitinae are nested. Heterotermitinae, having precedence over the other names, is here elevated to familial rank (i.e., elevated as Heterotermitidae Froggatt, stat. nov.). The monogeneric subfamilies Prorhinotermitinae and Psammotermitinae are together raised to familial rank, the latter having precedence (i.e., Psammotermitidae Holmgren, stat. nov.). Lastly, we elevate Termitogetoninae to Termitogetonidae Holmgren, stat. nov. (see Systematics, infra).Fig. 1: Majority rule consensus summary tree produced from 30 maximum-likelihood phylogenetic reconstructions performed with IQ-TREE on concatenated sequence matrices using various nucleotide, codon, and protein models (for details, see Supplementary Data 5).For readability, we only provided support values (UFBs, ultrafast bootstraps) for nodes representing families and subfamilies. Node labels are site concordance factors (sCFs) calculated on the majority rule consensus tree and the unfiltered UCE supermatrix. Species whose names are in bold were identified as rogue taxa by RogueNaRok (number of times under parentheses; for details, see Supplementary Data 5). The source data used for this figure can be found in the Dryad repository at https://doi.org/10.5061/dryad.02v6wwqbm (trees 1-30 in “File 4: trees.tar”).Fig. 2: Majority rule consensus summary cladogram produced from 21 multi-species coalescence phylogenetic reconstructions with ASTRAL-III using various nucleotide, codon, and protein models (for details, see Supplementary Data 5).The tree presented herein (tree 75 in File 4 on Dryad) was constructed from gene trees for which outliers were pruned with TreeShrink (trees 52–72). The summary topology without pruning outliers is identical for all but one intrasubfamilial node (see tree 74 built from trees 31–51 on Dryad: File 4). For readability, we only provided support values (LPPs, local posterior probabilities) for nodes representing families and subfamilies. Node labels are site concordance factors (sCFs) calculated on the majority rule consensus tree and the unfiltered UCE supermatrix. The source data used for this figure can be found in the Dryad repository at https://doi.org/10.5061/dryad.02v6wwqbm (“File 4: trees.tar”).The phylogenetic relationships of Psammotermitidae and Termitogetonidae, which diverged shortly after the origin of the clade Serritermitidae + Rhinotermitidae + Termitogetonidae + Psammotermitidae + Heterotermitidae + Termitidae, are unresolved and variable among analyses (Fig. 1). Our phylogenetic analyses placed Termitogetonidae in four potential topological conformations, being sister to: Psammotermitidae (analyses on concatenated data: UFBs = 49–93, n = 7/30 analyses; coalescence analyses: LPP = 0.49, n = 1/21); Serritermitidae + Rhinotermitidae (UFBs = 60–99, n = 13/30; LPPs = 0.35–0.54, 6/21); Heterotermitidae + Termitidae (LPPs = 0.49–0.90, n = 6/21); and Psammotermitidae + Heterotermitidae + Termitidae (UFBs = 52–99, n = 10/30; LPPs = 0.38–0.88, n = 8/21). The latter branching was recovered in both majority rule consensus trees, but with sCFs close to 33.3% (32.1–32.5%; Figs. 1, 2), the values obtained for alignments generated at random, indicating the absence of signal for this node in the alignments50. To assess the anomaly zone surrounding the origin of Termitogetonidae and Psammotermitidae, we performed a series of topology tests displacing Termitogetonidae. Topology tests indicated that Termitogetonidae could be sister to non-Stylotermitidae Neoisoptera and Psammotermitidae + Heterotermitidae + Termitidae, as reported in some mitogenome-based Bayesian phylogenies51, but it was never in a direct sister relationship with any of these families (AU tests; Supplementary Data 6).Our sampling of Termitidae included a total of 77 representative genera. Notably, the Syntermitinae were found to be paraphyletic and include the enigmatic monotypic genus Genuotermes Emerson (see Fig. 1). Although sharing key features with the Syntermitinae, Genuotermes was first tentatively assigned to the Termitinae52, based on similar soldier morphologies (Fig. 3 and Supplementary Figs. 17, 18). Genuotermes was identified as a rogue taxon by RogueNaRok in one analysis performed on concatenated sequence data (Fig. 1 and Supplementary Data 5). Genuotermes was also retrieved sister to Onkotermes in one coalescence analysis (Fig. 2; tree 36 in “File 4: trees.tar”, available on Dryad at https://doi.org/10.5061/dryad.02v6wwqbm; LPP = 0.66), both genera forming a clade outside the Syntermitinae and the Microcerotermes clade. The phylogenetic position of Genuotermes is, therefore, not entirely resolved; however, our analyses generally placed Genuotermes within the Syntermitinae53, and we hereby formally transfer this genus to the subfamily Syntermitinae.Fig. 3: Summary cladogram of Isoptera.This cladogram integrates topologies summarised from analyses performed on concatenated data and coalescence analyses. The preferred topology within anomaly zones estimated from approximately unbiased topological tests is indicated in dashed grey. The biogeographic matrix indicates the realms occupied by extant members of all isopteran families and termitid subfamilies (adapted from Krishna et al. 7). Here, we recognise nine realms (sensu Holt et al.54): Afrotropical (including Madagascan), Neotropical (incl. Panamanian), Oriental, Australian, Oceanian, Saharo-Arabian, Sino-Japanese, Nearctic, and Palaearctic. For termitid subfamilies, we report the main diagnostic features based on workers’ digestive tubes and soldiers’ heads. These features are summarised from Supplementary Notes 1 and 3, and Supplementary Figs. 1–18. (i) For workers’ midgut-hindgut junction, we indicate: the presence of a mixed segment (square; stippled section), the presence of Malpighian nodules (circle); the presence of a Malpighian knot (full diamond) or pseudo-knot (full diamond with empty circle); the shape of the junction between the mixed segment and the ileum (P1) either elbowed (empty oval) or arched (full oval). For the hindgut, we indicate the presence of a ventral loop (star) formed under the rectum (P5) by P1; and the position (triangle) of the enteric valve (P2, indicated by an arrow) at the insertion of P1 into the paunch (P3). (ii) For termitid soldiers, we indicate the presence of a frontal projection (hexagon) and whether the fontanelle opens at the tip of the projection (full hexagon with empty circle), or at its base (full hexagon). Greyed shapes indicate that both trait states occur within the considered subfamily. Diagnostic features for all subfamilies are extensively presented in Supplementary Figs. 1–18. Soldier pictures, from left to right: (1) Microcerotermes sp. (Microcerotermitinae); (2) Termes fatalis (Termitinae); (3) Silvestritermes heyeri (Syntermitinae), (4) Genuotermes spinifer (Syntermitinae); (5) Nasutitermes sp. (Nasutitermitinae). Picture credits: M. M. Rocha (4); R. H. Scheffrahn (2,3); J. Šobotník (1,5).Our phylogenetic analyses indicate that all other subfamilies of Termitidae are monophyletic, with the exception of the polyphyletic Termitinae, within which four other subfamilies are nested: Cubitermitinae, Engelitermitinae, Nasutitermitinae, and Syntermitinae. The relationships within the Termitinae + Cubitermitinae + Engelitermitinae + Nasutitermitinae + Syntermitinae clade are largely unresolved. This clade is composed of a dozen monophyletic lineages with uncertain relationships and separated by exceedingly short internodes (see Figs. 1, 2). Based on our phylogenetic reconstructions, we recognise 18 major monophyletic lineages within Termitidae, eight of which already have subfamilial status. Giving subfamilial ranks to the other ten lineages resolves the polyphyly of Termitinae while simultaneously retaining the subfamilial status of four subfamilies supported as monophyletic: Cubitermitinae (UFBs = 100; LPPs = 1; sCFs = 64–64.9%), Engelitermitinae, Nasutitermitinae (UFBs = 100; LPPs = 0.99–1; sCFs = 52.1–53%), and Syntermitinae (UFBs = 100; LPPs = 0.92–1; sCFs = 62.9–64%). Accordingly, we redefine Termitinae Latreille sensu novo and restrict it to Termes and related genera, which unambiguously formed a monophyletic group in all our analyses (UFBs = 100; LPPs = 1; sCFs = 56.5–57%). We advocate the resurrection of three monophyletic subfamilies: Microcerotermitinae Holmgren stat. rev. (UFBs = 100; LPPs = 0.74–1; sCFs = 48.2–49.7%), Mirocapritermitinae Kemner stat. rev. (UFBs = 99–100; LPPs = 1; sCFs = 55–56.1%), and Amitermitinae Kemner stat. rev. (UFBs = 99–100; LPPs = 0.98–1; sCFs = 45.8–46.9%). The aforementioned subfamilies were found monophyletic across all analyses, with the exception of Microcerotermitinae, as Onkotermes was placed sister to Microcerotermes + Syntermitinae in two coalescence analyses (LPPs = 1; trees 34, 50 in “File 4: trees.tar”, available on Dryad at https://doi.org/10.5061/dryad.02v6wwqbm) and sister to Genuotermes in one coalescence analysis (see above). Although not entirely supported by our phylogenetic analyses, the placement of Onkotermes within Microcerotermitinae is also justified by their similar gut morphologies, with a characteristic elbowed junction between the mixed segment and the ileum (Fig. 3). Our topology tests did not reject Microcerotermitinae + Syntermitinae as sisters to other non-Sphaerotermitinae, non-Macrotermitinae, non-Foraminitermitinae and non-Apicotermitinae Termitidae (Supplementary Data 6), while coalescence analyses placed the clade on a polytomy in 8 of 21 coalescence analyses run with diverse sequence matrices and substitution models (Fig. 2).Lastly, we propose to erect six monophyletic lineages to new subfamilies in Termitidae: Crepititermitinae subfam. nov., Cylindrotermitinae subfam. nov. (UFBs = 78–100; LPPs = 0.70–1; sCFs = 44.4–44.7%), Forficulitermitinae subfam. nov., Neocapritermitinae subfam. nov. (UFBs = 100; LPPs = 1; sCFs = 76.6–77%), Protohamitermitinae subfam. nov., and Promirotermitinae subfam. nov. (UFBs = 100; LPPs = 1; sCFs = 77.4–79.5%). These six subfamilies were placed on a polytomy, reflecting their uncertain phylogenetic positions, varying among analyses. The monotypic subfamilies Crepititermitinae and Forficulitermitinae were identified as rogue taxa in, respectively, two and ten analyses performed on concatenated sequence data from edge-trimmed protein-coding loci (Fig. 1 and Supplementary Data 5). Our topology tests did not reject Engelitermitinae and Forficulitermitinae as forming a clade sister to other non-Sphaerotermitinae, non-Macrotermitinae, non-Foraminitermitinae, non-Apicotermitinae, non-Microcerotermitinae and non-Syntermitinae Termitidae (AU tests; Supplementary Data 6). The Crepititermitinae were retrieved as sister to Cylindrotermitinae + Protohamitermitinae in 18 of 30 analyses performed on concatenated sequence data (Fig. 1; UFBs = 51–87), a scenario absent in our coalescence analyses (Fig. 2). The uncertain phylogenetic positions of Crepititermitinae and Forficulitermitinae, together with their unique combinations of morphological and biological characteristics (see the Systematics section below; Fig. 3), justified their placements in two separate subfamilies.SystematicsWe present the revised classification of extant Isoptera in Table 1. The diagnoses and distribution maps of Neoisoptera families and subfamilies presented herein are based on and modified from Krishna et al.7. The biogeographic boundaries follow Holt et al.54. The descriptions of digestive tubes follow Noirot55.Table 1 Revised higher-level classification of the extant Isoptera Brullé, 1832Family-level classification changes for NeoisopteraIn the following, we provide abridged diagnoses for families with changes. Please refer to Supplementary Note 1 for extended diagnoses of all neoisopteran families and Supplementary Note 2 for a revised identification key to all families of Isoptera.Family Rhinotermitidae Froggatt, sensu novoThe subfamily Rhinotermitinae Froggatt21, 1897: 518, is herein elevated to the familial rank.Type genus: Rhinotermes Hagen56, 1858.Included genera (5): Acorhinotermes Emerson, Dolichorhinotermes Snyder & Emerson, Parrhinotermes Holmgren, Rhinotermes Hagen, and Schedorhinotermes Silvestri (Macrorhinotermes Holmgren, syn. nov.).Diagnosis: Imagoes with inflated clypeus, slightly or not at all drawn out over the labrum —reminiscent of a “nose-like” projection; and a narrow groove running from the fontanelle to the clypeus. Imago-worker left mandible with an apical tooth, followed by three distinct (unequal) marginal teeth. Imagoes, workers, and soldiers with anterior margin of postclypeus convex. Soldiers without setae surrounding the fontanelle. Soldier labrum elongated, forming a brush apically and endowed with a groove in the centre along which the chemical secretion of the frontal gland flows from the fontanelle.Remarks: Macrorhinotermes was initially described as a subgenus of Rhinotermes57. Snyder26 considered Macrorhinotermes as a potential synonym of Schedorhinotermes but did not unambiguously synonymize the two names, and Macrorhinotermes has remained a valid genus name. Here, we formally synonymize Macrorhinotermes and Schedorhinotermes, the latter having precedence over the former. There is an interesting diversity of soldier morphologies among Rhinotermitidae. Soldiers can be monomorphic as in Acorhinotermes and Parrhinotermes; dimorphic as in Dolichorhinotermes, Rhinotermes, and most Schedorhinotermes; or trimorphic as in some Schedorhinotermes. Rhinotermitidae occur in the Australian, Afrotropical, Neotropical, Oriental, Palaearctic (Sino-Japanese), and Oceanian realms.Family Termitogetonidae Holmgren, stat. novThe subfamily Termitogetoninae Holmgren23, 1910: 286, is herein elevated to the familial rank.Type genus: Termitogeton Desneux58, 1904Included genus: monogeneric.Diagnosis: Imagoes densely hairy. Head subtriangular with posterior margin sinuate. Pronotum small, with median projections from anterior margins. Soldiers are densely hairy with a heart-shaped head. Soldier mandibles elongated without marginal teeth. Soldier legs with two protibial apical spurs. All castes dorso-ventrally flattened.Remarks: Termitogeton is a wood-feeding termite known from the Oriental and Oceanian realms. It is composed of only two known extant species.Family Psammotermitidae Holmgren, stat. novThe family Psammotermitidae regroups the Psammotermitinae and Prorhinotermitinae subfamilies.Subfamily Psammotermitinae Holmgren24, 1911: 64.Type genus: Psammotermes Desneux59, 1902Subfamily Prorhinotermitinae Quennedey & Deligne60, 1975: 265.Type genus: Prorhinotermes Silvestri61, 1909.Included extant genera (2): Psammotermes Desneux and Prorhinotermes Silvestri.Diagnosis: Imagoes with pronotum flat; tarsi tetramerous. Wings with all veins arising independently from inside the wing scales. Nymphs with fused wing buds forming two shield-looking structures on the mesothorax and metathorax. Imago-worker left mandible with an apical tooth followed by three distinct marginal teeth. Soldiers with tibial spurs formula 3:2:2; presenting a more or less marked groove from the fontanelle to clypeus60.Remarks: The extinct genus †Zophotermes Engel, 2011 is included in the Psammotermitidae. The distribution of Prorhinotermes is mostly insular. This distribution has fuelled speculation that colonies can be transported in driftwood by ocean currents, thus accounting for such a wide distribution on islands of the South Pacific. In sharp contrast, Psammotermes is a desert dweller occurring in the Afrotropical (incl. Madagascan), Saharo-Arabian and Oriental realms.Clade Geoisoptera Engel, Hellemans, & Bourguignon, novThis is an important group of termites as it includes the Termitidae, the most species-rich family of Isoptera, and the Heterotermitidae, which include some of the most widespread and critical pest species. In this clade, the number of chromosomes is fixed to 2n = 42 (reviewed in Jankásek et al.62), and their mitogenomes are characterised by a complex repeat structure in their control region36. Additionally, all species of this clade have a true worker caste (as in Mastotermitidae, Hodotermitidae, and Rhinotermitidae sensu novo) and typically live in large colonies.Etymology: The clade name is a combination of the Ancient Greek prefix geo– (γεω–, from gê / γῆ, meaning “soil,” “land,” or “earth”) and Isoptera. The name references the growing reliance on organically rich soil as a source of nitrogen63.Family Heterotermitidae Froggatt, stat. novThe subfamily Heterotermitinae Froggatt21,1897: 550, is herein elevated to the familial rank.Heterotermitinae Froggatt21, 1897: 550. Type genus: Heterotermes Froggatt21.Leucotermitinae Holmgren23, 1910: 285. Type genus: Leucotermes Silvestri64, 1901 [= Heterotermes Froggatt21, 1897].Coptotermitinae Holmgren23, 1910: 285. Type genus: Coptotermes Wasmann65, 1896. Syn. nov.Arrhinotermitinae Sjöstedt66, 1926: 8. Type genus: Arrhinotermes Wasmann67, 1902 [= Coptotermes Wasmann]. Syn. nov.Type genus: Heterotermes Froggatt21, 1897Included genera (3): Coptotermes Wasmann, Heterotermes Froggatt, and Reticulitermes Holmgren. †Lukotermes Perkovsky & Nel, syn. nov. is an extinct member of Heterotermitidae.Diagnosis: Imagoes with postclypeus devoid of nose-like projection or groove extending from the fontanelle. Head round in Coptotermes and oval with even posterior margin in Heterotermes and Reticulitermes. Pronotum without median projection on the anterior margin; tarsi tetramerous. Workers and soldiers with anterior margin of postclypeus flat or concave. Soldiers with fontanelle surrounded by setae and katana-shaped mandibles devoid of marginal teeth (though with basal serrations); protibia with more than three protibial apical spurs. Soldiers of Coptotermes displaying an egg-shaped head with a large fontanelle directed forward and oozing latex-appearing secretions. Soldiers of Heterotermes and Reticulitermes exhibit a rectangular head with a small indistinct fontanelle.Remarks: Some species exhibit slightly dimorphic soldiers68. The Heterotermitidae now encompass the three genera often referred to as “subterranean termites”, a term historically associated with “Rhinotermitidae” with an extensive pest status in the literature. The Heterotermitidae occur across most temperate and tropical regions.Subfamily-level classification changes for Termitidae LatreilleThe number of subfamilies within the Termitidae is expanded given that the current Termitinae are demonstrably polyphyletic. We describe the new subfamilies of Termitidae erected in this paper and redescribe the Termitinae. We do not address the Syntermitinae, as the inclusion of Genuotermes does not alter the diagnosis of the subfamily. We do not address the other seven subfamilies of Termitinae, as they remain unaltered (i.e., Sphaerotermitinae Engel & Krishna; Macrotermitinae Kemner; Foraminitermitinae Holmgren; Apicotermitinae Grassé & Noirot; Engelitermitinae Romero Arias, Roisin, & Scheffrahn; Nasutitermitinae Hare; and Cubitermitinae Weidner). Diagnoses for these subfamilies can be found in Krishna et al.7 and Romero Arias et al.20. An updated key for all subfamilies of Termitidae is given in Supplementary Note 3. An updated list of all genera of extant Isoptera currently recognised is given in Supplementary Note 4. A list of the top 20 UCEs diagnostic for each subfamily of Termitidae (excluding subfamilies represented by only one sample, i.e., Crepititermitinae, Forficulitermitinae, and Protohamitermitinae) is provided in Supplementary Data 7. An accompanying diagnostic tool is available on the Termite UCE Database (in “File 5: termitidae_diagnosing_database_v1.fasta.gz” available on Dryad at https://doi.org/10.5061/dryad.02v6wwqbm; also available at: https://github.com/sihellem/TER-UCE-DB/). Diagnostic features for each of the 18 subfamilies are given in Supplementary Figs. 1–18.Subfamily Amitermitinae Kemner, stat. novAmitermitinae Kemner69, 1934: 110.Type genus: Amitermes Silvestri64, 1901.Included genera (14): Ahamitermes Mjöberg, Amitermes Silvestri, Dentispicotermes Emerson, Drepanotermes Silvestri, Eremotermes Silvestri, Globitermes Holmgren, Gnathamitermes Light, Hoplotermes Light, Incolitermes Gay, Invasitermes Miller, Orthognathotermes Holmgren, Prohamitermes Holmgren, Pseudhamitermes Holmgren, and Synhamitermes Holmgren.Diagnosis: Imago-worker left mandible with apical tooth (LAt) shorter than marginal teeth (LMt), except in a few genera such as: Ahamitermes and Invasitermes70, or Orthognathotermes (see Supplementary Fig. 1i). Worker gut with ileum (P1) tubular, dilated, or globular, without loop ventrally; each pair of Malpighian tubules united in one base (or nodule), except in Prohamitermes in which each tubule ends in its own nodule. Soldiers generally have a pear-shaped head and are devoid of frontal projection (except in Eremotermes and Dentispicotermes). Diagnostic UCE loci are listed in Supplementary Data 7. Diagnostic features are given in Supplementary Fig. 1.The subfamily is most similar to the Promirotermitinae, whose enteric valve (P2) position, i.e. the insertion of P1 into the paunch (P3), occurs on the right side of the abdomen (rather than the left side, as in Microcerotermitinae, Mirocapritermitinae, and Neocapritermitinae; Fig. 3). The Amitermitinae can be distinguished from the promirotermitines by: P1 tubular to inflated (always inflated in Promirotermitinae); soldier mandibles of various forms, ranging from crushing, piercing, slashing, or symmetrical snapping (always symmetrical snapping in Promirotermitinae); and soldier head pear-shaped without frontal projection (a notable frontal hump or projection is present in Promirotermitinae).Remarks: Amitermitinae are widespread across every tropical and subtropical biogeographic realm. Species feed on various substrates, including mounds, soil, wood (incl. dry grasses), and the soil-wood interface. The genus Invasitermes is reported as soldierless.Subfamily Crepititermitinae Hellemans, Engel, & Bourguignon, subfam. novZoobank registration: https://zoobank.org/urn:lsid:zoobank.org:act:5F7D1841-7543-4914-B552-10318E2224C2.Type genus: Crepititermes Emerson71, 1925.Included genera: monogeneric.Diagnosis: Imago-worker left mandible lacks a third marginal tooth (LMt3). Right mandible lacking second marginal tooth (RMt2). Worker gut with a blind diverticulum or caecum on P3; Malpighian tubules forming two pairs attached individually. Soldiers with subrectangular heads, labrums with concave apical margins and sharp apicolateral points, flattened frons, and symmetrical snapping mandibles. Diagnostic features are given in Supplementary Fig. 3.The presence of a blind diverticulum on P3 and the absence of an indentation between LMt3 and LMt1+2 in Crepititermitinae are reminiscent of the Cubitermitinae, but the former can be distinguished by: the presence of a columnar belt (of the generalised type) in the gizzard (absent in Cubitermitinae); P1 tubular (P1 inflated in Cubitermitinae); the absence of intermediate cushions on the enteric valve armature (intermediate cushions present in Cubitermitinae; see Supplementary Fig. 4j, k); and the blind diverticulum on P3 occurring dorsally (ventrally in Cubitermitinae).Remarks: Crepititermes was previously assigned to the Termes-group. A comprehensive re-description of Crepititermes was provided by Rocha & Cuezzo72. The genus is exclusively Neotropical, with species that feed on humus. Crepititermes workers use an autothysis mechanism that involves the rupture of the upper side of the abdomen, near the junction with the thorax, to expose their sticky gut for colony defence.Subfamily Cylindrotermitinae Hellemans, Engel, & Bourguignon, subfam. novZoobank registration: https://zoobank.org/urn:lsid:zoobank.org:act:DEEDCE51-6696-4C97-BB2E-654EF86C8F9F.Type genus: Cylindrotermes Holmgren73, 1906.Included genera (2): Cephalotermes Silvestri and Cylindrotermes Holmgren.Diagnosis: Imago-worker left mandible with apical tooth (LAt) roughly equal or shorter than LMt1+2 and LMt3 well-developed. Worker gut with long, P1 tubular, forming a loop ventrally under the rectum (P5). Enteric valve bearing six unequal cushions. Soldier head elongated, subrectangular, devoid of frontal projection, covered by short bristles; mandibles short, robust, of biting/slashing type; weak humps on the frons located immediately between the antennae above the basolateral corners of clypeus. Diagnostic UCE loci are listed in Supplementary Data 7. Diagnostic features are given in Supplementary Fig. 5.The Cylindrotermitinae can be differentiated from the Amitermitinae as in the latter, P1 forms a loop ventrally under P5 and P2 occurs on the left side of the abdomen (Fig. 3).Remarks: The two genera of Cylindrotermitinae were previously assigned to the Amitermes-group sensu Krishna et al.7. Cylindrotermitinae are wood-feeding termite genera from separate biogeographic regions: Cephalotermes is Afrotropical, while Cylindrotermes is Neotropical. Cephalotermes builds mounds, and forms populous colonies with few soldiers, headed by one king and one extremely physogastric queen that loses the ability to crawl as the colony matures (J. Šobotník, pers. obs.). Cylindrotermes are found in dead wood and form small colonies.Subfamily Forficulitermitinae Hellemans, Engel, & Bourguignon, subfam. novZoobank registration: https://zoobank.org/urn:lsid:zoobank.org:act:429D8F1D-1D78-44AD-BF91-4AAE9492251F.Type genus: Forficulitermes Emerson74, 1960.Included genera: monogeneric.Diagnosis: Imago-worker right mandible with well-developed second marginal tooth (RMt2). Worker gut with four Malpighian tubules attached in pairs at the junction of the mixed segment. Soldier head subquadrangular with fontanelle located in a small depression, with two flat longitudinal ridges located in front of the opening on each side and forming a faint groove-like structure leading to the base of the clypeus; small, elevated humps on each side between the fontanelle and antennal base; mandibles elongate and slender, without marginal teeth, used to bite and unable to snap; apical margin of the labrum with medioapical convexity and sharply pointed apicolateral angles. Diagnostic features are given in Supplementary Fig. 8.The Forficulitermitinae can be distinguished from termitines in their newly restricted circumscription by the absence of a Malpighian nodule (generally present in Termitinae sensu novo). In addition, the soldier mandibles are elongated and not of snapping type, while in Termitinae the mandibles are generally snapping (except in three genera with biting/slashing mandibles: Divinotermes, Spinitermes, and Tuberculitermes). Lastly, the soldier head has a faint groove running from the fontanelle, which sits in a shallow depression, to the clypeal base and with small humps on either side of the channel between the fontanelle and the antennal toruli, while termitines lack such a groove, depression, or lateral humps and instead have a medial frontal prominence of varying degrees of development.Remarks: Forficulitermes was previously assigned to the Termes-group sensu Krishna et al.7. It is an exclusively Afrotropical genus, including species that feed on soil. A comprehensive re-description of Forficulitermes was provided by Scheffrahn & Křeček75.Subfamily Microcerotermitinae Holmgren, stat. novMicrocerotermitinae Holmgren76, 1910: 145.Type genus: Microcerotermes Silvestri64, 1901.Included genera (2): Microcerotermes Silvestri and Onkotermes Constantino.Diagnosis: Imago-worker left mandible with third marginal tooth (LMt3) well developed, right mandible with posterior margin of LMt2 long and straight. Worker gut with a conspicuous Malpighian knot; junction of the mixed segment with P1 constricted and elbowed, not fully visible in ventral view; P1 tubular, dilated anteriorly, constricted posteriorly at the connection with P2 on the left side of the abdomen, without loop ventrally. P2 is constricted and poorly ornamented. Soldier devoid of frontal projection, head generally long and narrow. Imago head subrectangular. Diagnostic UCE loci are listed in Supplementary Data 7. Diagnostic features are given in Supplementary Fig. 10.Microcerotermitines are similar to Amitermitinae but have P1 inserted on P3 on the left side of the abdomen rather than the right in the latter clade (Fig. 3). In addition, the soldier head of Microcerotermitinae is subrectangular rather than the pear-shaped form of Amitermitinae.Remarks: Species of this group were previously assigned to the Amitermes-group but were found to consistently form a sister relationship with Syntermitinae. The microcerotermitines are pantropical wood feeders (sensu lato: wood and grass).Subfamily Mirocapritermitinae Kemner, stat. novMirocapritermitinae Kemner69, 1934: 166.Type genus: Mirocapritermes Holmgren76, 1914.Included genera (14): Dicuspiditermes Krishna, Homallotermes John, Indocapritermes Chhotani, Kemneritermes Ahmad & Akhtar, Krishnacapritermes Chhotani, Labiocapritermes Krishna, Mirocapritermes Holmgren, Oriencapritermes Ahmad & Akhtar, Pericapritermes Silvestri, Procapritermes Holmgren, Pseudocapritermes Kemner, Rinacapritermes Amina & Rajmohana, Sinocapritermes Ping & Xu, and Syncapritermes Ahmad & Akhtar.Diagnosis: Imago-worker right mandible with anterior margin of RMt2 convex or straight, posterior margin concave. Worker gut with Malpighian tubules attached in two separate pairs fused proximally, devoid of Malpighian nodules. Soldiers with rectangular heads and snapping mandibles are generally asymmetrical (symmetrical in some species of Procapritermes). Soldier head devoid of frontal projection (except in Mirocapritermes). Diagnostic UCE loci are listed in Supplementary Data 7. Diagnostic features are given in Supplementary Fig. 11.Mirocapritermitinae can be distinguished from the termitines in their restricted sense by the absence of a Malpighian nodule (present in Termitinae sensu novo) and the absence of a frontal projection (generally present in Termitinae). In the imago-worker mandible, RMt2 is distinctly present and with a concave posterior margin, while it is absent or vestigial with a straight posterior margin in Termitinae sensu novo.Remarks: Mirocapritermitinae include species previously assigned to the Pericapritermes-group. Their distribution is restricted to the Oriental region, with the exception of Pericapritermes, which is also found in the Australian, Afrotropical, Palaearctic (Sino-Japanese), and Oceanian regions. Mirocapritermitines feed on soil or at the soil-wood interface.Subfamily Neocapritermitinae Hellemans, Engel, & Bourguignon, subfam. novZoobank registration: https://zoobank.org/urn:lsid:zoobank.org:act:B9567537-33EF-4B4D-992F-34E0C16B8701.Type genus: Neocapritermes Holmgren25, 1912.Included genera (3): Neocapritermes Holmgren, Planicapritermes Emerson, and Schievitermes Roisin.Diagnosis: Imago-worker left mandible with apical tooth (LAt) shorter than LMt1+2; right mandible with RMt2 slightly reduced, posterior margin straight. Worker gut with two pairs of Malpighian tubules fused anteriorly and inserted in a single point at the mixed segment junction, devoid of Malpighian nodule; mixed segment long, distally bilobed into mesenteric sacs in Neocapritermes and Schievitermes (see Roisin77); junction of the mixed segment with P1 constricted and elbowed; P3 inflated, with wall bearing numerous spines. Soldiers with asymmetrical snapping mandibles and subrectangular heads devoid of frontal projection. Diagnostic UCE loci are listed in Supplementary Data 7. Diagnostic features are given in Supplementary Fig. 13.Like the Termitinae, the imago-worker mandible has RMt2 reduced and with a straight posterior margin but can be distinguished by the absence of a Malpighian nodule (present in Termitinae sensu novo) and the absence of a frontal prominence on the soldier head (generally present to varying degrees in Termitinae). The Neocapritermitinae differ from the Old World Mirocapritermitinae in three ways: the posterior margin of RMt2 is straight (rather than concave in mirocapritermitines); the molar area has distinct ridges (molar area smooth to faint and reduced striations in mirocapritermitines); the enteric valve armature is composed of three folds (six in mirocapritermitines). The Neocapritermitinae exhibit an elbowed midgut-P1 junction as the Microcerotermitinae but can be distinguished by their soldiers (snapping in Neocapritermitinae, piercing/slashing in Microcerotermitinae).Remarks: The species of Neocapritermitinae were previously assigned to the Termes-group. The subfamily is exclusively Neotropical, with species that feed on decayed wood or at the wood-soil interface.Subfamily Promirotermitinae Hellemans, Engel, & Bourguignon, subfam. novZoobank registration: https://zoobank.org/urn:lsid:zoobank.org:act:1F08A9B3-6F94-498C-A8B4-78ABF508FDA6.Type genus: Promirotermes Silvestri78, 1914.Included genera (2): Angulitermes Sjöstedt and Promirotermes Silvestri.Diagnosis: Imago-worker right mandible with well-developed RMt2. Worker gut with two pairs of Malpighian tubules fused in a common base. Soldier head is subrectangular to egg-shaped (Promirotermes), with symmetrical snapping mandibles and either a pointed frontal projection (Angulitermes) or a blunt frontal hump (Promirotermes). Diagnostic features are given in Supplementary Fig. 14.The subfamily can be distinguished from Termitinae sensu novo by its imago-worker mandible with well-developed RMt2, which is otherwise absent or at least considerably reduced in termitines. The subfamily is most easily distinguished from Amitermitinae by its subrectangular soldier heads with a frontal process (generally absent in Amitermitinae) and bifurcate labrum (rounded, or with slightly pointed tips in Amitermitinae).Remarks: The species of this group were previously assigned to the Termes-group (sensu Krishna et al.7) and are humus-feeders. Promirotermes is exclusively Afrotropical, while Angulitermes is also distributed across the Oriental region, the Middle East, and Africa.Subfamily Protohamitermitinae Hellemans, Engel, & Bourguignon, subfam. novZoobank registration: https://zoobank.org/urn:lsid:zoobank.org:act:2C4DDA82-D86C-40CA-BCB9-F5CD5AA60E7B.Type genus: Protohamitermes Holmgren25, 1912.Included genera (2): Orientotermes Ahmad and Protohamitermes Holmgren.Diagnosis: Imago-worker right mandible with well-developed subsidiary tooth at the base of the first and second marginal teeth (RMt1 and RMt2), with anterior margin angular and posterior margin curved. Worker gut with two pairs of Malpighian tubules connected to the mixed segment by two nodules, one for each pair; with long, tubular ileum (P1), forming a loop ventrally under the rectum (P5) (as in Cylindrotermitinae and part of the Apicotermitinae). Diagnostic features are given in Supplementary Fig. 15.The Protohamitermitinae can be distinguished from the Amitermitinae by the presence of a long P1 that loops ventrally under P5 (ventral loop absent in Amitermitinae) and by the presence of a subsidiary tooth on the right imago-worker mandible (absent in Amitermitinae). The subfamily differs from Cylindrotermitinae by the presence of the subsidiary tooth and the absence of soldiers.Remarks: The Protohamitermitinae include two Oriental soldierless species feeding on wood. They were previously assigned to the Amitermes-group sensu Krishna et al.7. Although the type-genus Protohamitermes was not included in our analyses, its relationship with Orientotermes is known from previous phylogenetic analyses3.Subfamily Termitinae Latreille, sensu novoTermitina Latreille80, 1802: 293. Type genus: Termes Linnaeus81.Mirotermitini Weidner82, 1956: 99. Type genus: Mirotermes Wasmann83, 1897 [= Termes Linnaeus81, 1758].Capritermitini Weidner82, 1956: 100. Type genus: Capritermes Wasmann83, 1897.Type genus: Termes Linnaeus81, 1758Included genera (23): Apsenterotermes Miller, Capritermes Wasmann, Cavitermes Emerson, Cornicapritermes Emerson, Cristatitermes Miller, Dihoplotermes Araujo, Divinotermes Carrijo & Cancello, Ekphysotermes Miller, Ephelotermes Miller, Hapsidotermes Miller, Hesperotermes Gay, Inquilinitermes Mathews, Lophotermes Miller, Macrognathotermes Silvestri, Palmitermes Hellemans & Roisin, Paracapritermes Hill, Protocapritermes Holmgren, Quasitermes Emerson, Saxatilitermes Miller, Spinitermes Wasmann, Termes Linnaeus, Tuberculitermes Holmgren, and Xylochomitermes Miller.Diagnosis: Imago-worker right mandible with RMt2 reduced or absent and posterior margin straight. Left mandible with apical tooth (LAt) longer or slightly shorter than LMt1+2. Worker gut with two pairs of Malpighian tubules attached on a nodule connected to the mixed segment. P2 is on the left side of the abdomen (except in Cornicapritermes, whose P1 forms a ventral loop and P2 is on the right; M.M. Rocha and T.F. Carrijo, pers. obs.). Soldiers with head subrectangular, a frontal prominence varying in size among genera, and snapping mandibles, except in Divinotermes and Spinitermes, which exhibit biting/slashing mandibles; and Tuberculitermes, which has snapping/slashing mandibles (A. Buček, pers. obs.). Diagnostic UCE loci are listed in Supplementary Data 7. Diagnostic features are given in Supplementary Fig. 18. Additional diagnoses are provided in the systematic sections of Promirotermitinae, Neocapritermitinae, Mirocapritermitinae, and Forficulitermitinae (above).Remarks: The Termitinae have a pantropical distribution, found in the Afrotropical, Australian, Neotropical, Oceanian and Oriental regions. Termitinae comprises species feeding on diverse substrates, including soil, mounds, wood, and the soil-wood interface.
