Termitophily—the symbiosis of organisms with termite societies—has evolved a disproportionate number of times within the rove beetle subfamily Aleocharinae (Staphylinidae). Among aleocharine termitophiles, defensive (limuloid) and mimetic (physogastric & physothoracic) body forms have evolved convergently, but due to lack of a comprehensive aleocharine phylogeny, the context in which termitophily and associated adaptations evolve is unknown. We present the first example of a robust, morphology-based phylogenetic placement of an exclusively termitophilous tribe, the Termitohospitini. Termitohospitini is recovered to be nested within Myllaenini sensu nov, and sister to Myllaena (new synonymy). Furthermore, we also recovered the small tribe Masuriini nested within Myllaenini sensu nov (new status).Reconstructing ecological transitions within this clade, we present evidence that the stem lineage of Myllaenini sensu nov invaded intertidal marine habitats, the common ancestor for Myllaena + Termitohospitini then transitioned to freshwater riparian habitats, with Termitohospitini alone subsequently shifting to termitophily. We conclude that: (1) Termitohospitini was ancestrally a limuloid-bodied riparian inhabitant; (2) a limuloid form may have been pre-adaptive for defense against host attack during the evolution of termitophily; (3) the strongly tapered abdomen of an ancestral limuloid body was a constraint on the evolution of physogastry, leading to the emergence of the unusual physothoracic body form observed in termitohospitines that likely integrates these obligate termitophiles to life inside termite colonies.“one of the most astonishing spectacles in all natural history.” — Richard Dawkins

Members of the termitophilous subtribe Termitozyrina (Aleocharinae: Lomechusini) associated with Hodotermopsis sjostedti are taxonomically treated. The genera Hodotermophilus Naomi & Terayama and Termophidoholus Naomi & Hirono, and each type species (monotypic) are redescribed. Termophidoholus formosanus, originally described from Taiwan, and its host termite H. sjostedti are newly recorded from Laos. Yakuus iwatai Kanao & Maruyama gen. & sp. nov. and H. gloriosus were collected sympatrically in the same nests in Yaku‐shima, Japan. All of the above species, belonging to the three genera, share the presence of a batch of spurs at the tibial apex of fore and mid legs. Habitus photographs and illustrations of diagnostic features are provided for these species, and their phylogenetic relationships are discussed based on morphological similarities and the extant distribution of the host termite species.

Coptotermocola clavicornis gen. & sp. n. and Neotermitosocius bolivianus gen. & sp. n. of the termite inquilinous tribe Termitohospitini are described from peninsular Malaysia and Bolivia, respectively. The Termitohospitini are most readily diagnosable by the distally migrated anterior tentorial pits that are no longer associated with the antennal fossae, and by the enlarged vertex which obscures the antennal fossae dorsally. Additionally, the Termitohospitini are hypothesized to share a recent common ancestor with the Masuriini and Myllaenini due to shared derived morphologies of the lacinia distal teeth with lateral cuticular processes, presence of a unique maxillary palpomere III sensilla, and anterolateral angles of mentum produced. Habitus photographs and illustrations of diagnostic features are provided for the two new genera in order to facilitate future work.

Madagascar is home to many endemic plant and animal species owing to its ancient isolation from other landmasses. This unique fauna includes several lineages of termites, a group of insects known for their key role in organic matter decomposition in many terrestrial ecosystems. How and when termites colonised Madagascar remains unknown. In this study, we used 601 mitochondrial genomes, 93 of which were generated from Madagascan samples, to infer the global historical biogeography of Neoisoptera, a lineage containing upwards of 80% of described termite species. Our results indicate that Neoisoptera colonised Madagascar between seven to ten times independently during the Miocene, between 8.4-16.6 Ma (95% HPD: 6.1-19.9 Ma). This timing matches that of the colonization of Australia by Neoisoptera. Furthermore, the taxonomic composition of the Neoisopteran fauna of Madagascar and Australia are strikingly similar, with Madagascar harbouring an additional two lineages absent from Australia. Therefore, akin to Australia, Neoisoptera colonised Madagascar during the global expansion of grasslands, possibly helped by the ecological opportunities arising from the spread of this new biome.

Kistnerella malayensis Kanao & Maruyama, gen. et sp.n. (Coleoptera: Staphylinidae: Aleocharinae) is described from Ulu Gombak, Selangor, Peninsular Malaysia. Th is new genus belongs to the termitophilous subtribe Compactopediina in the tribe Aleocharini, and is associated with Longipeditermes longipes (Haviland, 1889) (Isoptera: Termitidae: Nasutitermitinae). A key to the genera in Compactopediina is given. Th e phylogenetic relationships of the genera are investigated, and the host termite relationships and the systematic position of Compactopediina are discussed.