Entheus Hübner

Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina & Grishin, Nick V., 2025, Advancing butterfly systematics through genomic analysis, The Taxonomic Report of the International Lepidoptera Survey 12 (5), pp. 1-201 : 61-64

A preliminary taxonomic list of Entheus Hübner , [1819] species

Here, we integrate our new results into the previous taxonomic arrangement of Entheus Hübner, [1819] (Evans 1952; Steinhauser 1989; Austin 1997; Austin et al. 1997; Mielke 2005; Grishin 2013) to compile a preliminary taxonomic list for the genus and suggest a phylogenetic order of the species. The phylogeny of Entheus ( Fig. 50 ) generally agrees with phenotypic considerations and division into species groups. The eumelus , gentius , priassus , and matho groups agree with the Evans’s (1952) identification key. However, we find that E. warreni and E. bogoteus sp. n., both of which would be identifiable as members of the matho group by the presence of a separate orange spot between the forewing orange bands in males, are not monophyletic with E. matho and form a clade sister to the priassus group. Therefore, we define a separate species group for them. Furthermore, in agreement with discussions by Austin (1997) and Austin et al. (1997) based on phenotypic differences and similarities, we partition the priassus group into three subgroups, distinguishing the telemus and pralina subgroups, which are closely related to each other and fully separate only in the nuclear genome tree ( Fig. 50a ).

We attempt to order species to maximize the phenotypic similarity and geographic proximity of the list neighbors but without disrupting phylogenetic orders given in genomic trees ( Fig. 50 ): i.e., a strongly supported clade in the trees is a continuous segment in the list. The evolution and diversification of Entheus are riddled with complexities due to the incongruence of the genomic trees ( Fig. 50 ). Most notably, Entheus latifascius M. Hering, 1925 , stat. rest. ( type locality in Colombia: Rio Micay) is confidently placed deep within the matho group in the nuclear genome tree inferred from autosomes ( Fig. 50a ), but is sister to the priassus group in both the Z chromosome and the mitochondrial genome trees ( Fig. 50b, c ). Until such discrepancies are explained, we side with phenotypic similarity in choosing between the conflicting phylogenetic hypotheses in different phylogenetic trees. Here, we chose to order species groups to maintain the traditional order of Evans (1952), which agrees with phylogenetic trees. Within each group, species are arranged to maximize wing pattern resemblance (within phylogenetic constraints) between neighboring species from different groups, i.e., the brightest member of the priassus group (e.g., E. telemus ) is placed after the gentius group species consisting of brightly colored species, and E. dius with the dark anal fold similar to the priassus group is near the beginning of the matho group.

This arrangement results in the placement of E. crux and E. guato sp. n., large species with dark females without a white area on the hindwing, last. Further refinements of the list order are encouraged.

In the resulting arrangement below, we also include two species discovered by Burns and coauthors ( Janzen et al. 2011) but still unnamed, shown in gray font. Based on COI barcode comparison, we hypothesize that the third species, Entheus Burns 03, may be conspecific with E. matho . Type localities (general area only: state, region, department, or county) are in gray font. New taxa described in this study and the category of taxonomic change are in red font. Taxonomic treatment before this work (for valid names) or the category of synonym (for synonyms) and comments (phenotypic characters for species groups and subgroups refer to males) are shown in smaller font following a vertical bar | after the type locality; an equal sign = precedes synonyms given in their original genus combination; and a double dagger ‡ marks permanently invalid synonyms (e.g., objective synonyms) and unavailable names. The list covers 36 valid taxa, all at the species level, with 12 newly proposed here (and 2 yet undescribed) and 6 elevated (from subspecies or synonyms) to the species status: i.e., 16 previously listed as species with 5 additional subspecies and 1 synonym: 22 known and 14 undescribed before this work. Our study follows the trend to reveal approximately as many new Hesperiidae taxa as previously described ones in nearly every genus under revision (Austin and Mielke 1998; Austin and Mielke 2008; Medeiros et al. 2019; Siewert et al. 2020). Because our work on Entheus has not been completed yet, the list below is preliminary, given as a guide to the published knowledge, and with additions to follow.

Figures

Fig. 50. Phylogenetic trees of Entheus inferred from protein-coding regions in: a) the nuclear genome (autosomes), based on 3,486,282 positions, b) the Z chromosome, based on 279,462 positions, and c) the mitochondrial genome. Ultrafast bootstrap (Minh et al. 2013) values are shown at nodes. Different species groups and subgroups are colored: eumelus group (olive), gentius group (blue), priassus group (cyan, with telemus and pralina subgroups in purple and red, respectively), warreni group (magenta), and matho group (green). New taxa proposed in this work are labeled in red, and those with taxonomic changes, such as subspecies-to-species or synonym-to-species status (changes indicated in brackets), are labeled in blue.