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Dirini
Dirini is one of the tribes in the butterfly subfamily Satyrinae of the family Nymphalidae. Consisting of 29 species[a] in 6 genera,[b] the group's members are exclusively found in southern Africa. Initially distinguished by having a forewing cell length shorter than half the length of the forewing, the morphological synapomorphy that links the members of the Dirini together is the presence of the scaphium on the male genitalia. The group is remarkable for its restricted distribution within South Africa and Lesotho, with a single species whose distribution who extends into Zimbabwe. It is closely affiliated with the tribe Melanitini, of whom the Neotropical Manataria hercyna is closest related to the Dirini as a whole. DescriptionMembers of the tribe are endemic to southern Africa,[3] with the group consisting of 29 species[a] in 6 genera.[b][4] The larvae of the members feed on grasses.[5] First characterized by Miller in 1968 as exhibiting a forewing cell length shorter than half of the length of the forewing, this feature united the genera of Dira, Dingana, Serradinga, Torynesis and Tarsocera.[5] This circumscription excluded Aeropetes or Paralethe, which were alternatively placed into Lethini by Miller in 1968, Elymniini by Ackery et al. in 1995, and Melanitini by Peña et al. in 2005.[6][5] Phylogenetic analysis in 2010 by Price et al. confirmed their continued placement within the Dirini.[4] The presence of a scaphium on the male genitalia was identified as the sole morphological synapomorphy which united the group.[5] Systematics and taxonomyPer Peña & Wahlberg, 2008, the tribe is around 25 million years old.[4] The divergence of Dirini and Melanitini occurred between 23 and 47 Ma.[7] The initial point of radiation for the tribe was just before or at the Oligocene-Miocene transition around 29-24 Ma.[4] The seven genera of the tribe were described as "convincingly monophyletic" by Price et al., reflected in the morphological and biological homogeneity of the genera that contain more than a single species, with the relative diversity in the morphological and biological sense between the various genera.[4] ,_Tambopata.jpg) The most stable group within the Satyrinae is the sister group relationship between Dirini and Melanitini, having been recovered in studies by Peña et al. 2006, Peña and Wahlberg 2008, Wahlberg et al. 2009, and Price et al. 2010.[8] Manataria hercyna was confirmed to be a sister taxon of the tribe, with that relationship supported by previous analyses Peña et al. in 2006, and Peña & Wahlberg in 2008.[6][4] Studies showed that Manataria would be placed into the Melanitini, but its placement would render Melanitini paraphyletic, which suggests that it would be best to either place Manataria into its own tribe or within the Dirini.[4] The relationship between Manataria and the remainder of the Dirini is strange, due to the former's Neotropical distribution compared to the tribe's Afrotropical distribution, and its estimated divergence from the remainder of the tribe 40 Ma from the present.[4] This would mean that divergence had occurred when South America was several thousand kilometers from the African coast, which is a process that has occurred similarly for the amphisbaenid lizards, platyrrine monkeys and caviomorph rodents, suggesting an underlying process in a westward direction.[4] The following phylogenetic trees are adapted from Wahlberg and Brower, 2011:[5][9]
DistributionMembers of the tribe are noted for their remarkable degree of endemism, being found almost exclusively in South Africa and Lesotho, with a single member whose distribution extends into eastern Zimbabwe.[c][4] They are almost all found on high-lying ground, but are described by Price et al. as not a truly montane group, occurring on slopes from the south and east and from elevations at 2,000 metres (6,600 ft) in the north and west.[4] Members of the tribe are found in South Africa's Succulent Karoo region and grasslands, which are dominated by C3 producing grasses, with the exception of Paralethe dendrophilus, which inhabits coastal and Afromontane forests.[4] GeneraGenus, species, and subspecies list derived from Price et al. 2010,[4] and Williams 2018.[11] Price et al. recognizes Torynesis hawequas as a subspecies of Torynesis mintha (as Torynesis mintha hawequas),[4] while Williams recognizes it at full species status.[11][12] Genomic testing conducted in 2023 by Zhang et al. revealed that Serradinga was better suited as a subgenus of Dingana.[1]
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