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Abstract The classification of ants (Hymenoptera: Formicidae) has progressed in waves since the first 17 species were described by Linnaeus in the 1758 edition of Systema Naturae. Since then, over 18,000 species-rank names have accumulated for the global myrmecofauna, of which ~14,260 living and ~810 fossil species are valid. Here, we provide a synopsis of ant biodiversity and review the history and classification of the family, while highlighting the massive growth of the field in the new millennium. We observe that major transformation has occurred for ant classification due to advances in DNA sequencing technologies, model-based hypothesis testing, and imaging technologies. We therefore provide a revised and illustrated list of diagnostic character states for the higher clades of Formicidae, recognizing that vastly more work is to be done. To facilitate discussion and the systematic accumulation of evolutionary knowledge for the early evolution of the ants, we suggest an informal nomenclatural system for the higher clades of ants, based on names currently in use and a set of names that have been democratically selected by the authors. To guide future work on ant systematics, we summarize currently available databases and present perspectives on regions in need of biodiversity exploration, challenges facing the field, and the future of ant taxonomy. 

Theta and gamma oscillations in the hippocampus have been hypothesized to play a role in the encoding and retrieval of memories. Recently, it was shown that an intrinsic fast gamma mechanism in medial entorhinal cortex can be recruited by optogenetic stimulation at theta frequencies, which can persist with fast excitatory synaptic transmission blocked, suggesting a contribution of interneuronal network gamma (ING). We calibrated the passive and active properties of a 100-neuron model network to capture the range of passive properties and frequency/current relationships of experimentally recorded PV+ neurons in the medial entorhinal cortex (mEC). The strength and probabilities of chemical and electrical synapses were also calibrated using paired recordings, as were the kinetics and short-term depression (STD) of the chemical synapses. Gap junctions that contribute a noticeable fraction of the input resistance were required for synchrony with hyperpolarizing inhibition; these networks exhibited theta-nested high frequency oscillations similar to the putative ING observed experimentally in the optogenetically-driven PV-ChR2 mice. With STD included in the model, the network desynchronized at frequencies above ~200 Hz, so for sufficiently strong drive, fast oscillations were only observed before the peak of the theta. Because hyperpolarizing synapses provide a synchronizing drive that contributes to robustness in the presence of heterogeneity, synchronization decreases as the hyperpolarizing inhibition becomes weaker. In contrast, networks with shunting inhibition required non-physiological levels of gap junctions to synchronize using conduction delays within the measured range.