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Mixture models unveil the origin of the enigmatic Satyrinae butterfly genera Calisto and Llorenteana (Lepidoptera, Nymphalidae, Satyrinae)

Núñez, Rayner ; Bodenheim, Anja ; Álvarez, Yosiel ; Wahlberg, Niklas LU and Espeland, Marianne (2026) In Systematic Entomology 51(2).
Abstract

We provide the first comprehensive analysis of the origin of two enigmatic Satyrinae genera of uncertain affinities. Calisto Hübner, the only Satyrinae genus from the West Indies and endemic to these islands, has resisted numerous attempts at phylogenetic placement, regardless of the data type or methods used. Llorenteana Viloria & Luis-Martínez, a monotypic genus from northwestern Mexico, has never been included in a molecular phylogenetic study, and past authors have placed it in five different genera and subtribes. We used not only mostly published genomic data but also newly sequenced whole genome data from museum specimens and old DNA extracts, extracted BUSCO genes and prepared several datasets. These datasets differed in the... (More)

We provide the first comprehensive analysis of the origin of two enigmatic Satyrinae genera of uncertain affinities. Calisto Hübner, the only Satyrinae genus from the West Indies and endemic to these islands, has resisted numerous attempts at phylogenetic placement, regardless of the data type or methods used. Llorenteana Viloria & Luis-Martínez, a monotypic genus from northwestern Mexico, has never been included in a molecular phylogenetic study, and past authors have placed it in five different genera and subtribes. We used not only mostly published genomic data but also newly sequenced whole genome data from museum specimens and old DNA extracts, extracted BUSCO genes and prepared several datasets. These datasets differed in the degree of heterogeneity and saturation, the number of nucleotide positions used (all positions or only the first two), and were analysed as nucleotides or as amino acids. We employed maximum likelihood methods, using both partitioned and mixture models, as well as a multispecies coalescent method, ASTRAL, to infer the phylogenetic relationships. Additionally, we estimated divergence times and the ancestral areas of origin for Calisto and Llorenteana. The phylogenetic placement of Calisto varied among datasets when we used partitioned models and ASTRAL; however, most datasets resulted in the same topology under mixture models despite incomplete lineage sorting and gene flow affecting our data. Our results suggest that Calisto is part of a clade of Old World origin that colonized the New World from north to south, thus sharing ancestry with Nearctic taxa. The placement of Llorenteana remained stable in all analyses and datasets. It constitutes one of the earliest splits within the Euptychiina, a subtribe of Neotropical origin, but descending together with the Pronophilina from Nearctic ancestors. We propose the recognition of Erebiina stat. rev. as the only subtribe comprising the former Calistina syn. nov., Callerebiina syn. nov, Maniolina syn. nov. and Ypthimina syn. nov.

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author
; ; ; and
organization
publishing date
type
Contribution to journal
publication status
published
subject
keywords
biogeography, divergence times, gene flow, incomplete lineage sorting, phylogenetic relationships, Satyrini, systematics
in
Systematic Entomology
volume
51
issue
2
article number
e70057
publisher
Wiley-Blackwell
external identifiers
  • scopus:105040697003
ISSN
0307-6970
DOI
10.1111/syen.70057
language
English
LU publication?
yes
additional info
Publisher Copyright: © 2026 The Author(s). Systematic Entomology published by John Wiley & Sons Ltd on behalf of Royal Entomological Society.
id
58d5b3d8-94d4-4574-86fb-b9e284c581b4
date added to LUP
2026-08-18 16:14:40
date last changed
2026-08-21 10:37:53
@article{58d5b3d8-94d4-4574-86fb-b9e284c581b4,
  abstract     = {{<p>We provide the first comprehensive analysis of the origin of two enigmatic Satyrinae genera of uncertain affinities. Calisto Hübner, the only Satyrinae genus from the West Indies and endemic to these islands, has resisted numerous attempts at phylogenetic placement, regardless of the data type or methods used. Llorenteana Viloria &amp; Luis-Martínez, a monotypic genus from northwestern Mexico, has never been included in a molecular phylogenetic study, and past authors have placed it in five different genera and subtribes. We used not only mostly published genomic data but also newly sequenced whole genome data from museum specimens and old DNA extracts, extracted BUSCO genes and prepared several datasets. These datasets differed in the degree of heterogeneity and saturation, the number of nucleotide positions used (all positions or only the first two), and were analysed as nucleotides or as amino acids. We employed maximum likelihood methods, using both partitioned and mixture models, as well as a multispecies coalescent method, ASTRAL, to infer the phylogenetic relationships. Additionally, we estimated divergence times and the ancestral areas of origin for Calisto and Llorenteana. The phylogenetic placement of Calisto varied among datasets when we used partitioned models and ASTRAL; however, most datasets resulted in the same topology under mixture models despite incomplete lineage sorting and gene flow affecting our data. Our results suggest that Calisto is part of a clade of Old World origin that colonized the New World from north to south, thus sharing ancestry with Nearctic taxa. The placement of Llorenteana remained stable in all analyses and datasets. It constitutes one of the earliest splits within the Euptychiina, a subtribe of Neotropical origin, but descending together with the Pronophilina from Nearctic ancestors. We propose the recognition of Erebiina stat. rev. as the only subtribe comprising the former Calistina syn. nov., Callerebiina syn. nov, Maniolina syn. nov. and Ypthimina syn. nov.</p>}},
  author       = {{Núñez, Rayner and Bodenheim, Anja and Álvarez, Yosiel and Wahlberg, Niklas and Espeland, Marianne}},
  issn         = {{0307-6970}},
  keywords     = {{biogeography; divergence times; gene flow; incomplete lineage sorting; phylogenetic relationships; Satyrini; systematics}},
  language     = {{eng}},
  month        = {{04}},
  number       = {{2}},
  publisher    = {{Wiley-Blackwell}},
  series       = {{Systematic Entomology}},
  title        = {{Mixture models unveil the origin of the enigmatic Satyrinae butterfly genera Calisto and Llorenteana (Lepidoptera, Nymphalidae, Satyrinae)}},
  url          = {{http://dx.doi.org/10.1111/syen.70057}},
  doi          = {{10.1111/syen.70057}},
  volume       = {{51}},
  year         = {{2026}},
}