Dirini is one of the tribes in the butterfly subfamily Satyrinae of the family Nymphalidae. Consisting of 29 species in 6 genera, 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 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.
| Dirini | |
|---|---|
 | |
| Dira clytus | |
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | Insecta |
| Order: | Lepidoptera |
| Family: | Nymphalidae |
| Subfamily: | Satyrinae |
| Tribe: | Dirini Verity, 1953 |
| Genera | |
| See text | |
Description
Members of the tribe are endemic to southern Africa, with the group consisting of 29 species in 6 genera. The larvae of the members feed on grasses. 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. 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. Phylogenetic analysis in 2010 by Price et al. confirmed their continued placement within the Dirini. The presence of a scaphium on the male genitalia was identified as the sole morphological synapomorphy which united the group.
Systematics and taxonomy
Per Peña & Wahlberg, 2008, the tribe is around 25 million years old. The divergence of Dirini and Melanitini occurred between 23 and 47 million years ago (Ma). The initial point of radiation for the tribe was just before or at the Oligocene-Miocene transition around 29-24 Ma. 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.
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.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. 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. The relationship between Manataria and the remainder of the Dirini was described by Price et al. as "odd", 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. 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.
The following phylogenetic trees are adapted from Wahlberg and Brower, 2011:
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| Dirini |
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Distribution
Members 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. 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. 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.
Genera
Genus, species, and subspecies list derived from Price et al. 2010, and Williams 2018. Price et al. recognizes Torynesis hawequas as a subspecies of Torynesis mintha (as Torynesis mintha hawequas), while Williams recognizes it at full species status. Genomic testing conducted in 2023 by Zhang et al. revealed that Serradinga was better suited as a subgenus of Dingana.
| Photograph | Scientific name | Species |
|---|---|---|
 | Aeropetes Billberg, 1820 |
|
 | Dingana van Son, 1955 |
|
 | Dira Hübner, 1819 |
|
 | Paralethe van Son, 1955 |
|
 | Tarsocera Butler, 1899 |
|
 | Torynesis Butler, 1899 |
|
Footnotes
- Price et al. recognizes 28 species in 7 genera, while Williams' Afrotropical Butterflies and Skippers (2016) and Wahlberg (2018) elevated Torynesis mintha hawequas to full species status as Torynesis hawequas.
- Genomic analysis by Zhang et al. conducted in 2023 revealed that Serradinga was better suited as a subspecies of Dingana.
- Price et al. does not specify which member of the Dirini has the distribution that extends into eastern Zimbabwe, but per Van Son in 1955 and Williams in 2019, Aeropetes tulbaghia has a distribution that extends into eastern Zimbabwe.
References
- Zhang, Jing; Cong, Qian; Shen, Jinhui; Song, Leina; Grishin, Nick V. (13 December 2023). "Genomic analysis reveals new species and subspecies of butterflies". The Taxonomic Report of the International Lepidoptera Survey. 11 (6): 9. doi:10.5281/zenodo.10372378. ISSN 2643-4806. PMC 11759507. PMID 39867645.
- Williams, Mark C. (29 January 2024). "Genus Dingana van Son, 1955 Widows". AFROTROPICAL BUTTERFLIES (PDF). Lepidopterists' Society of Africa. p. 17. Retrieved 2 January 2025.
- Price, Benjamin Wills; Walton, Shaun M.; Barker, Nigel P.; Villet, Martin H. (January 2011). "Price et al - Dirini (supporting information)". Systematic Entomology. Retrieved 29 December 2024.
- Price, Ben W.; Villet, Martin H.; Walton, Shaun M.; Barker, Nigel P. (April 2011). "Using molecules and morphology to infer the phylogenetic relationships and evolutionary history of the Dirini (Nymphalidae: Satyrinae), a tribe of butterflies endemic to Southern Africa". Systematic Entomology. 36 (2): 300–316. Bibcode:2011SysEn..36..300P. doi:10.1111/j.1365-3113.2010.00560.x. Retrieved 29 December 2024.
- Wahlberg, Niklas; Brower, Andrew V. Z. (9 December 2011). "Dirini". Tree of Life Web Project. Retrieved 29 December 2024.
- Peña, Carlos; Wahlberg, Niklas; Weingartner, Elisabet; Kodandaramaiah, Ullasa; Nylin, Sören; Freitas, André V.L.; Brower, Andrew V.Z. (July 2006). "Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data" (PDF). Molecular Phylogenetics and Evolution. 40 (1): 29–49. Bibcode:2006MolPE..40...29P. doi:10.1016/J.YMPEV.2006.02.007. PMID 16563805. Retrieved 30 December 2024.
- Matos-Maraví, Pável; Wahlberg, Niklas; Antonelli, Alexandre; Penz, Carla M. (October 2019). "Species limits in butterflies (Lepidoptera: Nymphalidae): reconciling classical taxonomy with the multispecies coalescent" (PDF). Systematic Entomology. 44 (4): 745–756. Bibcode:2019SysEn..44..745M. doi:10.1111/syen.12352. Retrieved 30 December 2024.
- Marín, M. A.; Peña, C.; Freitas, A. V. L.; Wahlberg, N.; Uribe, S. I. (February 2011). "From the Phylogeny of the Satyrinae Butterflies to the Systematics of Euptychiina (Lepidoptera: Nymphalidae): History, Progress and Prospects". Neotropical Entomology. 40 (1): 1–13. doi:10.1590/S1519-566X2011000100001. ISSN 1519-566X. PMID 21437476. Retrieved 29 December 2024.
- Wahlberg, Niklas; Brower, Andrew V. Z. (2006). "Satyrinae". Tree of Life Web Project. Retrieved 30 December 2024.
- Williams, Mark C. (18 January 2019). "Genus Aeropetes Billberg, 1820 Mountain Pride". AFROTROPICAL BUTTERFLIES (PDF). Lepidopterists' Society of Africa. p. 2. Retrieved 2 January 2025.
- Wahlberg, Niklas (12 February 2018). "Dirini Classification". Nymphalidae.net. Retrieved 30 December 2024.
- Williams, Mark C. (2016-01-09). "Afrotropical Butterflies: Nymphalidae, Satyrinae, Genus Torynesis" (PDF). Metamorphosis. Retrieved 28 August 2017.
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 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 Dirini Dira clytus Scientific classification Domain Eukaryota Kingdom Animalia Phylum Arthropoda Class Insecta Order Lepidoptera Family Nymphalidae Subfamily Satyrinae Tribe Dirini Verity 1953 Genera See text Contents 1 Description 2 Systematics and taxonomy 3 Distribution 4 Genera 5 Footnotes 6 ReferencesDescriptioneditMembers 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 Pena 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 taxonomyeditPer Pena amp Wahlberg 2008 the tribe is around 25 million years old 4 The divergence of Dirini and Melanitini occurred between 23 and 47 million years ago 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 nbsp Manataria hercyna is supported by several studies to be sister to the Dirini 4 The most stable group within the Satyrinae is the sister group relationship between Dirini and Melanitini having been recovered in studies by Pena et al 2006 Pena 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 Pena et al in 2006 and Pena amp 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 was described by Price et al as odd 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 Elymniini Zetherini Amathusiini Brassolini Morphini Melanitini Dirini Haeterini Satyrini Dirini Dingana Serradinga b Dira Torynesis Tarsocera Aeropetes ParaletheDistributioneditMembers 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 GeneraeditGenus 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 Photograph Scientific name Species nbsp Aeropetes Billberg 1820 Aeropetes tulbaghia Linnaeus 1764 nbsp Dingana van Son 1955 Dingana alaedeus Henning amp Henning 1996 Dingana alticola Henning amp Henning 1996 Dingana angusta Henning amp Henning 1996 Dingana Serradinga bowkeri Trimen 1870 Dingana bowkeri bella van Son 1955 Dingana bowkeri bowkeri Trimen 1870 Dingana clara van Son 1940 Dingana Serradinga clarki van Son 1955 Dingana clarki amissivallis Henning amp Henning 1996 Dingana clarki clarki van Son 1955 Dingana clarki dracomontana Henning amp Henning 1996 Dingana clarki ocra Henning amp Henning 1996 Dingana dingana Trimen 1873 Dingana fraterna Henning amp Henning 1996 Dingana jerinae Henning amp Henning 1996 Dingana Serradinga kammanassiensis Henning amp Henning 1994 nbsp Dira Hubner 1819 Dira clytus Linnaeus 1764 Dira clytus clytus Linnaeus 1764 Dira clytus eurina Quickelberge 1978 Dira jansei Swierstra 1909 Dira oxylus Trimen 1881 Dira swanepoeli van Son 1939 Dira swanepoeli isolata van Son 1955 Dira swanepoeli swanepoeli van Son 1939 nbsp Paralethe van Son 1955 Paralethe dendrophilus Trimen 1862 Paralethe dendrophilus albina van Son 1955 Paralethe dendrophilus dendrophilus Trimen 1862 Paralethe dendrophilus indosa Trimen 1879 Paralethe dendrophilus junodi van Son 1935 nbsp Tarsocera Butler 1899 Tarsocera cassina Butler 1868 Tarsocera cassus Linnaeus 1764 Tarsocera cassus cassus Linnaeus 1764 Tarsocera cassus outeniqua Vari 1971 Tarsocera dicksoni van Son 1962 Tarsocera fulvina Vari 1971 Tarsocera imitator Vari 1971 Tarsocera namaquensis Vari 1971 Tarsocera southeyae Dickson 1969 nbsp Torynesis Butler 1899 Torynesis hawequas Dickson 1973 Torynesis magna van Son 1941 Torynesis mintha Geyer 1837 Torynesis mintha mintha Geyer 1837 Torynesis mintha piquetbergensis Dickson 1967 Torynesis orangica Vari 1971 Torynesis pringlei Dickson 1979Footnotesedit a b Price et al recognizes 28 species in 7 genera while Williams Afrotropical Butterflies and Skippers 2016 and Wahlberg 2018 elevated Torynesis mintha hawequas to full species status as Torynesis hawequas a b c Genomic analysis by Zhang et al conducted in 2023 revealed that Serradinga was better suited as a subspecies of Dingana 1 2 Price et al does not specify which member of the Dirini has the distribution that extends into eastern Zimbabwe but per Van Son in 1955 and Williams in 2019 Aeropetes tulbaghia has a distribution that extends into eastern Zimbabwe 10 Referencesedit a b Zhang Jing Cong Qian Shen Jinhui Song Leina Grishin Nick V 13 December 2023 Genomic analysis reveals new species and subspecies of butterflies The Taxonomic Report of the International Lepidoptera Survey 11 6 9 doi 10 5281 zenodo 10372378 ISSN 2643 4806 PMC 11759507 PMID 39867645 Williams Mark C 29 January 2024 Genus Dingana van Son 1955 Widows AFROTROPICAL BUTTERFLIES PDF Lepidopterists Society of Africa p 17 Retrieved 2 January 2025 Price Benjamin Wills Walton Shaun M Barker Nigel P Villet Martin H January 2011 Price et al Dirini supporting information Systematic Entomology Retrieved 29 December 2024 a b c d e f g h i j k l m n o Price Ben W Villet Martin H Walton Shaun M Barker Nigel P April 2011 Using molecules and morphology to infer the phylogenetic relationships and evolutionary history of the Dirini Nymphalidae Satyrinae a tribe of butterflies endemic to Southern Africa Systematic Entomology 36 2 300 316 Bibcode 2011SysEn 36 300P doi 10 1111 j 1365 3113 2010 00560 x Retrieved 29 December 2024 a b c d e Wahlberg Niklas Brower Andrew V Z 9 December 2011 Dirini Tree of Life Web Project Retrieved 29 December 2024 a b Pena Carlos Wahlberg Niklas Weingartner Elisabet Kodandaramaiah Ullasa Nylin Soren Freitas Andre V L Brower Andrew V Z July 2006 Higher level phylogeny of Satyrinae butterflies Lepidoptera Nymphalidae based on DNA sequence data PDF Molecular Phylogenetics and Evolution 40 1 29 49 Bibcode 2006MolPE 40 29P doi 10 1016 J YMPEV 2006 02 007 PMID 16563805 Retrieved 30 December 2024 Matos Maravi Pavel Wahlberg Niklas Antonelli Alexandre Penz Carla M October 2019 Species limits in butterflies Lepidoptera Nymphalidae reconciling classical taxonomy with the multispecies coalescent PDF Systematic Entomology 44 4 745 756 Bibcode 2019SysEn 44 745M doi 10 1111 syen 12352 Retrieved 30 December 2024 Marin M A Pena C Freitas A V L Wahlberg N Uribe S I February 2011 From the Phylogeny of the Satyrinae Butterflies to the Systematics of Euptychiina Lepidoptera Nymphalidae History Progress and Prospects Neotropical Entomology 40 1 1 13 doi 10 1590 S1519 566X2011000100001 ISSN 1519 566X PMID 21437476 Retrieved 29 December 2024 Wahlberg Niklas Brower Andrew V Z 2006 Satyrinae Tree of Life Web Project Retrieved 30 December 2024 Williams Mark C 18 January 2019 Genus Aeropetes Billberg 1820 Mountain Pride AFROTROPICAL BUTTERFLIES PDF Lepidopterists Society of Africa p 2 Retrieved 2 January 2025 a b Wahlberg Niklas 12 February 2018 Dirini Classification Nymphalidae net Retrieved 30 December 2024 Williams Mark C 2016 01 09 Afrotropical Butterflies Nymphalidae Satyrinae Genus Torynesis PDF Metamorphosis Retrieved 28 August 2017 Retrieved from https en wikipedia org w index php title Dirini amp oldid 1274481323
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