Ochlodes (Ochluma) sylvanoides (Boisduval, 1852)

Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina & Grishin, Nick V., 2025, Update to: Advancing butterfly systematics through genomic analysis, The Taxonomic Report of the International Lepidoptera Survey 12 (6), pp. 1-36 : 26

Ochlodes (Ochluma) sylvanoides group

Previously treated as a single species Ochlodes sylvanoides (Boisduval, 1852) (type locality in USA: California, Plumas Co.), this species group is currently placed in the subgenus Ochluma Grishin, 2025 (type species Hesperia yuma W. H. Edwards, 1873 ) and, as here defined, consists of three genetically differentiated species: Ochlodes (Ochluma) santacruza J. Scott, 1981 (type locality USA: California, Santa Barbara Co., Santa Cruz Island), O. sylvanoides , and Ochlodes (Ochluma) napa (W. H. Edwards, 1865) (type locality in USA: Colorado, Clear Creek Co. ) ( Zhang et al. 2023b). We expanded genomic sequencing of this group by sampling additional specimens across the range from southwestern Canada through all 13 western states of the USA to Baja California, Mexico ( Fig. 19 ). The results confirm the phylogenetic separation of the group into three species and reveal additional insights, some of which are unexpected.

First, the nominate subspecies of O. napa ( Fig. 19 yellow circles) is restricted to the mountainous region of Colorado and its immediate neighborhood in southeastern Wyoming, eastern Utah, and northern New Mexico. Second, Ochlodes (Ochluma) napa kaibab Grishin, 2023 (type locality in USA: Arizona, Coconino Co.) ( Fig. 19 , blue circles) has a much wider distribution than anticipated and spreads as a narrow strip north from the type locality through central Utah, reaching northwestern Wyoming. This subspecies of O. napa , genetically differentiated from the nominate, represents populations between it and neighboring O. sylvanoides while being phylogenetically associated with the former. It is also possible that the populations of O. napa in Wyoming, Utah, and

Arizona are best partitioned into several subspecies. They have different, albeit more closely related, mitogenome haplotypes ( Fig. 20b ) and show different levels of introgression with other taxa. For instance, specimens of O. napa kaibab from Emery County, Utah, introgress stronger with O. napa napa and thus are placed near the base of the nuclear genome tree ( Fig. 20a ) and possess mitochondrial genomes of O. napa napa . Nevertheless, specimens shown as blue circles in Fig. 19 are in the clade with O. napa kaibab , and we presently treat them as this subspecies. Furthermore, the populations of O. napa kaibab in Utah are coming close to and may even be sympatric with O. sylvanoides , a question to be addressed in future studies.

Third, paler-colored populations to the west of the main Rocky Mountains chain north of Central Wyoming ( Fig. 19 magenta and violet squares) traditionally associated with O. napa do not belong to this species and are O. sylvanoides instead. Due to their wing pattern differences that resulted in this misidentification, these populations are described below as two new subspecies that are somewhat differentiated genetically from other O. sylvanoides populations. These populations of O. sylvanoides are geographically close to O. napa kaibab in northwestern Wyoming and south-central Montana, another region to study interactions between the two species O. napa and O. sylvanoides .

Fourth, phylogenetic analysis did not reveal prominent genetic differences of previously described O. sylvanoides subspecies: Ochlodes sylvanoides orecoasta J. Scott, 1981 (type locality in USA: Oregon, Clatsop Co.), Ochlodes sylvanoides bonnevilla J. Scott, 1981 (type locality in USA: Nevada, Elko Co.), and Ochlodes sylvanoides omnigena Austin, 1998 (type locality in USA: Nevada, Lander Co.). While we do not propose synonymizing these subspecies, studies of their distinction are beyond the scope of this work and will be addressed in future research. Here , all these populations combined are shown as green squares in Fig. 19 .

Figures

Fig. 19. A map of sequenced specimens from the Ochlodes sylvanoides group. Species are shown as symbols of different shapes and subspecies are denoted by color: O. santacruza (stars), O. sylvanoides (squares), and O. napa (circles); O. santacruza santacruza (red), O. santacruza catalina (orange), O. sylvanoides sylvanoides including orecoasta, bonnevilla, and omnigena (green), O. sylvanoides dempwolfi ssp. n. (violet), O. sylvanoides paranapa ssp. n. (magenta), O. napa kaibab (blue), and O. napa napa (yellow). The type localities are marked with tiny white circles inside symbols, and names of corresponding subspecies are placed nearby.

Fig. 20 (see the previous two pages). Phylogenetic trees of the Ochlodes sylvanoides group species constructed from proteincoding regions in: a) the nuclear genome (autosomes), based on 312,678 positions, and b) the mitochondrial genome. Different