Lionepha, Casey, 1918

Lionepha View in CoL specimens sequenced 143 143 140

Legend

143 70 47 52 21

Black cells: bootstrap support for the clade 90 or above

Gray cells: bootstrap support for the clade 50-89

White cells: ML tree has clade, with bootstrap support 0-49

Pink cells: ML tree has contradictory clade, with bootstrap support 0-49

Red cells: bootstrap support against clade 50-100

INDELS IN 28S

There are data in the DNA sequences not considered in any of the phylogenetic analyses, in the form of indels.The 28S gene, in particular, shows an amount of insertions and deletions in the evolutionary history of Lionepha that is unusual within bembidiines; every species of Lionepha has a unique set of indels. Many of these indels are in a part of the D2 expansion region (Supporting Information, Fig. S2). There is variation within species as well. Lionepha disjuncta , in particular, has striking variation: among the ten specimens sequenced there are eight unique patterns of insertions and deletions over three regions of the gene. There was less variation observed in other species: four bases missing from the northernmost specimen of L. erasa and an extra two bases in the Mt. Hood specimen; an extra two bases in two Oregon and one Washington specimens of L. casta ; an extra base in the Utah specimens of L. probata ; four indel differences between the California and Oregon specimens of L. tuulukwa .

MORPHOLOGICAL RESULTS

The morphological diversity evident among specimens of Lionepha (depicted in Figs 9–19, and described in ‘Taxonomic treatment’, below) corresponds to the patterns evident in the gene trees, with the male genitalic structures in particular ( Figs 11–14) being consistent within a proposed species and showing disparities among species.

CYTOGENETIC RESULTS

Eight of the species examined have 12 pairs of autosomes, and an X0/XX sex-chromosome system, for a complement of 25 chromosomes in males ( Table 5). Preparations of the ninth species examined, Lionepha pseudoerasa ( Lindroth, 1963) , were insufficient to determine the sex chromosomes,but the25 chromosomes seen in the single male studied were consistent with results from other species. Re-examination of notes taken about the single male of L. casta examined by Maddison (1985) suggest that the tentative male count of 22+XY reported was in error.