Middle Stone Age humans in high-altitude Africa: Recent archaeological research has produced evidence of the earliest human occupation of high-altitude habitats in the Andes and the Tibetan Plateau. Ossendorf et al. now present the oldest evidence of human settlement and adaptation to areas above 4000-meter elevation in Africa (see the Perspective by Aldenderfer). Their excavations at a rock shelter in the Bale Mountains of Ethiopia reveal obsidian artifacts and faunal remains, including abundant burnt bones, mostly of giant mole-rats. The findings reveal the environmental conditions and show how Late Pleistocene humans adapted to the harsh environments of these glaciated high-altitude African landscapes. Science , this issue p. 583 ; see also p. 541

The Himalayan mountain arc is one of the hotspots of biodiversity on earth, and species diversity is expected to be especially high among insects in this region. Little is known about the origin of the Himalayan insect fauna. With respect to the fauna of high altitude cloud forests, it has generally been accepted that Himalayan lineages are derived from ancestors that immigrated from Western Asia and from adjacent mountainous regions of East and Southeast Asia (immigration hypothesis). In this study, we sought to test a Tibetan Origin as an alternative hypothesis for groups with a poor dispersal ability through a phylogeographic analysis of the Ethira clade of the genus Pterostichus. We sequenced COI mtDNA and the 18S and 28S rDNA genes in 168 Pterostichini specimens, including 46 species and subspecies of the Ethira clade. In our analysis, we were able to show that the Ethira clade is monophyletic and, thus, represents a Himalayan endemic clade, supporting endemism of two of the basal lineages to the Central Himalaya and documenting large distributional gaps within the phylogeographic structure of the Ethira clade. Furthermore, the molecular data strongly indicate very limited dispersal abilities of species and subspecies of these primary wingless ground beetles. These results are consistent with the hypothesis of a Tibetan Origin, which explains the evolution, diversity and distribution of the Himalayan ground beetle Ethira clade much more parsimoniously than the original immigration hypothesis.

Balticeler kerneggeri gen. nov., sp. nov., is described based on six fossil specimens preserved in Eocene Baltic amber and imaged using light microscopy and X-ray micro-computed tomography. Based on certain characters observed in the fossil species it is considered a “middle grade” Carabidae, outside of the large family Harpalinae (as it possesses a scrobal seta, the lack of which is a synapomorphy of that subfamily), but possessing four synapomorphies that indicate Balticeler belongs to a large clade of carabids including Harpalinae (anisochaetous Grade B antennal cleaner, conjunct mesocoxae, closed procoxal cavities, and a well-developed external lobe of the metepimeron). This remarkable beetle has several striking features, including lack of externally-visible sexually dimorphic characters, lack of lateral borders on the pronotum, and very long and thin mandibles and maxillae. In combination, these states are unique within Carabidae. We consider the presence of a dorsally completely open aedeagal median lobe as a synapomorphy of the fossil species with the subfamily Trechinae, a pubescent and relatively long second antennomere and a 4+2+2 pattern of umbilicate setae as synapomorphies of the supertribe Trechitae, and a quadrisetose clypeus as a synapomorphy with the Trechitae clade Bembidarenini + Trechini sensu Maddison et al. (2019). As it lacks a synapomorphy of Bembidarenini + Trechini, we propose that it is a member of the stem group of that clade.

The trechine beetle fauna (Coleoptera, Carabidae) of the Ethiopian Highlands is known to be highly diverse in species, and many species groups were recognized to be characterized by unusual character states of external and genital morphology. Earlier authors described several genera and subgenera of Ethiopian Trechina endemic to certain high mountains of the country. However, the relationships of these species groups and their evolutionary history are unknown so far. Here, we present the first molecular phylogenetic analysis of Ethiopian Trechina, detect several synonymic names under Trechus sensu lato, and introduce two new species groups to the country’s fauna: the monotypic genus Baehria Schmidt & Faille, gen. nov., with the type species B. separatasp. nov. from Mt. Choke in northern Ethiopia, and the Trechus subgenus Abunetrechus Schmidt & Faille, subgen. nov., with the type species T. bipartitus Raffray, 1885; this subgenus includes three species of northern Ethiopia. We show that the composition of the Ethiopian fauna is based on multiple events of immigration, which started simultaneously with or some million years after the Oligocene-Early Miocene orogenic events north and south of the Rift Valley. Our results support the habitat island hypothesis for the evolution of the Ethiopian highland fauna. We found no evidence for an alternative hypothesis assuming a close connection of the Trechina immigration to Ethiopia and Pleistocene cooling. We, thus, conclude that the geomorphological development rather than the climatic changes are the main drivers of the diversification of the high-altitude Trechina fauna in Ethiopia.

The first platynine ground beetle, Limodromus hoffeinsorum sp. n., is described from Baltic amber at species level. The fossil species is considered the sister taxon of the modern species group of L. assimilis (Paykull, 1790). However, it was adapted to warmer climate conditions. The habitat of the new species is assumed similar to that of the modern L. klausnitzeri Schmidt, 2005, which represents a more basal branch of Limodromus and which occurs in the mountains of northern Vietnam.

The phylogeny of the carabid beetle supertribe Nebriitae is inferred from analyses of DNA sequence data from eight gene fragments including one nuclear ribosomal gene (28S), four nuclear-protein coding genes (CAD, topoisomerase 1, PEPCK, andwingless), and three mitochondrial gene fragments (16S + tRNA-Leu + ND1, COI (“barcode” region) and COI (“Pat/Jer” region)). Our taxon sample included 264 exemplars representing 241 species and subspecies (25% of the known nebriite fauna), 39 of 41 currently accepted genera and subgenera (all exceptNotiokasisandArchileistobrius), and eight outgroup taxa. Separate maximum likelihood (ML) analyses of individual genes, combined ML analyses of nuclear, nuclear protein-coding, and mitochondrial genes, and combined ML and Bayesian analyses of the eight-gene-fragment matrix resulted in a well-resolved phylogeny of the supertribe, with most nodes in the tree strongly supported. Within Nebriitae, 167 internal nodes of the tree (out of the maximum possible 255) are supported by maximum-likelihood bootstrap values of 90% or more. The tribes Notiophilini, Opisthiini, Pelophilini, and Nebriini are well supported as monophyletic but relationships among these are not well resolved.Nippononebriais a distinct genus more closely related toLeistusthanNebria.Archastes,Oreonebria,Spelaeonebria, andEurynebria, previously treated as distinct genera by some authors, are all nested within a monophyletic genusNebria.WithinNebria, four major clades are recognized: (1) theOreonebriaSeries, including eight subgenera arrayed in two subgeneric complexes (theEonebriaandOreonebriaComplexes); (2) the Nebriola Series, including only subgenus Nebriola; (3) theNebriaSeries, including ten subgenera arrayed in two subgeneric complexes, theBoreonebriaandNebriaComplexes, with the latter further subdivided into three subgeneric subcomplexes (theNebria,Epinebriola, andEunebriaSubcomplexes)); and (4) theCatonebriaSeries, including seven subgenera arrayed in two subgeneric complexes (theReductonebriaandCatonebriaComplexes). A strong concordance of biogeography with the inferred phylogeny is noted and some evident vicariance patterns are highlighted. A revised classification, mainly within the Nebriini, is proposed to reflect the inferred phylogeny. Three genus-group taxa (Nippononebria,VancouveriaandArchastes) are given revised status and seven are recognized as new synonymies (NebrioritesJeannel, 1941 andMarggiaHuber, 2014 =OreonebriaDaniel, 1903;PseudonebriolaLedoux & Roux, 1989 =BoreonebriaJeannel, 1937;PatrobonebriaBänninger, 1923,ParanebriaJeannel, 1937 andBarbonebriolaHuber & Schmidt, 2017 =EpinebriolaDaniel & Daniel, 1904; andAsionebriaShilenkov, 1982 =PsilonebriaAndrewes, 1923). Six new subgenera are proposed and described for newly recognized clades: Parepinebriola Kavanaugh subgen. nov. (type species:Nebria delicataHuber & Schmidt, 2017), Insulanebria Kavanaugh subgen. nov. (type species:Nebria carbonariaEschscholtz, 1829), Erwinebria Kavanaugh subgen. nov. (type speciesNebria sahlbergiiFischer von Waldheim, 1828),NivalonebriaKavanaughsubgen. nov.(type species:Nebria paradisiDarlington, 1931),NeaptenonebriaKavanaughsubgen. nov.(type species:Nebria ovipennisLeConte, 1878), andPalaptenonebriaKavanaughsubgen. nov.(type species:Nebria mellyiGebler, 1847). Future efforts to better understand relationships within the supertribe should aim to expand the taxon sampling of DNA sequence data, particularly within subgeneraLeistusandEvanoleistus of genusLeistusand theNebriaComplex of genusNebria.