The underlying mechanisms responsible for the general increase in species richness from temperate regions to the tropics remain equivocal. Many hypotheses have been proposed to explain this astonishing pattern but additional empirical studies are needed to shed light on the drivers at work. Here we reconstruct the evolutionary history of the cosmopolitan diving beetle subfamily Colymbetinae, the majority of which are found in the Northern hemisphere, hence exhibiting an inversed latitudinal diversity gradient. We reconstructed a dated phylogeny using 12 genes, to investigate the biogeographical history and diversification dynamics in the Colymbetinae. We aimed to identify the role that phylogenetic niche conservatism plays in the inversed diversification pattern seen in this group. Our results suggest that Colymbetinae originated in temperate climates, which supports the hypothesis that their distribution is the result of an ancestral adaptation to temperate environmental conditions rather than tropical origins, and that temperate niche conservatism can generate and/or maintain inverse latitudinal diversity gradients.

The superfamily Dytiscoidea contains six families with an aquatic lifestyle, with most of its extant diversity in two families: the burrowing water beetles (Noteridae) and the diving beetles (Dytiscidae). The other families have few species (up to six) and generally highly disjunct extant distributions. Aspidytidae currently contains one genus with two species, one in China and one in South Africa. Here we provide the first molecular data for the Chinese species, allowing us to explore the phylogenetic relationships and position of both species of this small family for the first time. Based on a matrix of 11 genes we inferred a phylogenetic hypothesis for Dytiscoidea including all extant families. Unexpectedly, Aspidytidae were consistently recovered as paraphyletic relative to Amphizoidae, despite being well characterized by apparently synapomorphic adult features. A re-examination of larval characters in the two aspidytid species revealed that the larva of the Chinese species is strikingly similar to that of Amphizoidae. Both share a series of plesiomorphic features but also some potential synapomorphies, including a dense vestiture of short setae on the head capsule, anteriorly shifted posterior tentorial grooves and widely separated labial palps. Arguably these features may belong to the groundplan of the clade Aspidytidae + Amphizoidae, with far-reaching secondary modifications (including reversals) in the South African Aspidytes niobe. At present we retain the family Aspidytidae, however, due to the strong adult morphological synapomorphies of the two extant species, and the fact that the molecular paraphyly of the family may result from the highly divergent nature of the two extant species. This long evolutionary separation and strong divergence, in terms of gene sequences and Zoological Journal of the Linnean Society, 2016, 176, 537-546. With 11 figures. larval features, is undeniable, substantial levels of saturation in third codon positions of protein-coding genes being present between the two taxa. We address this issue taxonomically by introducing the new genus Sinaspidytes gen. nov. for the Chinese Aspidytes wrasei. The continued contentious relationships amongst Dytiscidae, Hygrobiidae, Aspidytidae and Amphizoidae highlight the need for more data to address dytiscoid phylogenetics, possibly involving a genomic approach.

A comprehensive higher-level phylogeny of diving beetles (Dytiscidae) based on larval characters is presented. Larval morphology and chaetotaxy of a broad range of genera and species was studied, covering all currently recognized subfamilies and tribes except for the small and geographically restricted Hydrodytinae, where the larva is unknown. The results suggest several significant conclusions with respect to the systematics of Dytiscidae including the following: monophyly of all currently recognized subfamilies, although Dytiscinae when considered in a broad context is rendered paraphyletic by Cybistrinae; currently recognized tribes are monophyletic except for Agabini, Hydroporini and Laccornellini; inter-subfamily and inter-tribe relationships generally show weak support, except for a few well supported clades; three distinct clades are recognized within Dytiscinae [Dytiscini sensu lato (i.e. including the genera Dytiscus Linnaeus and Hyderodes Hope), Hydaticini sensu lato, and Cybistrini]; and recognition of Pachydrini as a distinct tribe. Other less robust results include: Methlini sister to the rest of Hydroporinae; relative basal position of Laccornini, Hydrovatini and Laccornellini within Hydroporinae; close relationship of Agabinae and Copelatinae; Matinae nested deep within Dytiscidae, as sister to a large clade including Colymbetinae, Coptotominae, Lancetinae and Dytiscinae sensu lato; the sister-group relationship of Agabetini and Laccophilini is confirmed. The results presented here are discussed and compared with previous phylogenetic hypotheses based on different datasets, and the evolution of some significant morphological features is discussed in light of the proposed phylogeny. All suprageneric taxa are diagnosed, including illustrations of all relevant synapomorphies, and a key to separate subfamilies and tribes is presented, both in traditional (paper) format and as an online Lucid interactive identification key.