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Bat Systematics in the Light of Unconstrained Analyses of a Comprehensive Molecular Supermatrix

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Abstract

Bats (Chiroptera) represent the largest diversification of extant mammals after rodents. Here we report the results of a large-scale phylogeny of bats based on unconstrained searches for a data matrix of 804 non-chimeric, taxonomically updated bat terminals (796 species represented by a single terminal plus three species represented by โ‰ฅ2 genetically distinct subspecies), able to preliminary test the systematics of most groups simultaneously. We used nine nuclear and mitochondrial DNA sequence markers fragmentary represented for ingroups (c. 90% and 64% of extant diversity at genus and species level, respectively) and 20 diverse placental outgroups. Maximum Likelihood and Parsimony analyses applied to the concatenated dataset yielded a highly resolved, variously supported phylogeny that recovered the majority of currently recognized clades at all levels of the chiropteran tree. Calibration points based on 44 key fossils allowed the Bayesian dating of bat origins at c. 4 my after the K-Pg boundary, and the determination of stem and crown ages of intraordinal clades. As expected, bats appeared nested in Laurasiatheria and split into Yinpterochiroptera and Yangochiroptera. More remarkable, all polytypic, currently recognized families were monophyletic, including Miniopteridae, Cistugidae, and Rhinonycteridae, as well as most polytypic genera with few expected exceptions (e.g., Hipposideros). The controversial Myzopodidae appeared in a novel position as sister of Emballonuroideaโ€•a result with interesting biogeographic implications. Most recently recognized subfamilies, genera, and species groups were supported or only minor adjustments to the current taxonomy would be required, except Molossidae, which should be revised thoroughly. In light of our analysis, current bat systematics is strongly supported at all levels; the emergent perception of a strong biogeographic imprint on many recovered bat clades is emphasized.

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Acknowledgements

We thank the support from Consejo Nacional de Investigaciones Cientรญficas y Tรฉcnicas (CONICET), Argentina, grant PICT 2008โ€“1798 to NPG, and grants PICT 1930_2011 and PIP 0260 to SAC.

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Authors and Affiliations

  1. Unidad Ejecutora Lillo (UEL: FML-CONICET), Tucumรกn, Argentina

    Lucila I. Amador, R. Leticia Moyers Arรฉvalo, Santiago A. Catalano & Norberto P. Giannini

  2. Genรฉtica y Evoluciรณn de Buenos Aires (IEGEBA: UBA-CONICET), Instituto de Ecologรญa, Buenos Aires, Argentina

    Francisca C. Almeida

  3. Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucumรกn (UNT), Tucumรกn, Argentina

    Santiago A. Catalano & Norberto P. Giannini

  4. American Museum of Natural History (AMNH), Division of Vertebrate Zoology, Department of Mammalogy, NY, USA

    Norberto P. Giannini

Authors
  1. Lucila I. Amador
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  2. R. Leticia Moyers Arรฉvalo
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  3. Francisca C. Almeida
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  4. Santiago A. Catalano
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  5. Norberto P. Giannini
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Corresponding author

Correspondence to Norberto P. Giannini.

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Fig S1.

Pteropodidae. A: Cynopterinae; B: Cynopterini; C: Balionycterini; D: Scotonycterini; E: Rousettinae; F: Eonycterini; G: Rousettini; H: Stenonycterini; I: Myonycterini; J: Epomophorini; K: Macroglossinae; L: Harpyionycterinae; M: Eidolinae; N: Nyctimeninae; O: Pteropodinae (JPEG 3447 kb)

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Fig S2.

Rhinolophoidea. A: African Hipposideros species; B: Asian Hipposideros species; C: Afro-Paleartic Rhinolophus clade; D: Indomalayan Rhinolophus clade (JPEG 3767 kb)

High resolution image (TIFF 6448 kb) (download TIF )

Fig S3.

Yangochiroptera. A: Asian Nycteris clade; B: African Nycteris clade; C: Taphozoinae; D: Emballonurinae; E: Emballonurini; F: Diclidurini; G: Afro-Malagasy emballonurini genera; H: Indo-Pacific emballonurini genus; I: Chaerephon/Mops group; J: Otomops group; K: New World Molossidae group; L: Oriental-Australasian Miniopterus clade; M: Ethiopian Miniopterus clade; N: Indian Ocean Miniopterus clade (JPEG 4114 kb)

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Fig S4.

Relationships within Phyllostomidae. The subfamilies are indicated in bold (JPEG 4357 kb)

High resolution image (TIFF 7445 kb) (download TIF )

Fig S5.

Relationships within Vespertilionidae (part 1: Kerivoulinae, Murininae, Myotinae). A: New World Myotis clade; B: Paleartic lineage โ€œbrandtiiโ€; C: Neartic clade; D: Neotropical clade; E: Old World Myotis clade; F: Ethiopian clade; G: Euroasian clade (JPEG 3654 kb)

High resolution image (TIFF 6354 kb) (download TIF )

Fig S6.

Relationships within Vespertilionidae (part 2: Vespertilioninae). The tribes are indicated in bold (JPEG 3972 kb)

High resolution image (TIFF 6637 kb) (download TIF )

Fig S7.

Pteropodidae. A: Cynopterinae; B: Cynopterini; C: Balionycterini; D: Scotonycterini; E: Rousettinae; F: Eonycterini; G: Rousettini; H: Stenonycterini; I: Myonycterini; J: Epomophorini; K: Macroglossinae; L: Harpyionycterinae; M: Eidolinae; N: Nyctimeninae; O: โ€œPteropodinaeโ€ (JPEG 3509 kb)

High resolution image (TIFF 5707 kb) (download TIF )

Fig S8.

Rhinolophoidea. A: African Hipposideros species; B: Asian Hipposideros species; C: Afro-Paleartic Rhinolophus clade (JPEG 3578 kb)

High resolution image (TIFF 5785 kb) (download TIF )

Fig S9.

Yangochiroptera. A: Asian Nycteris clade; B: African Nycteris clade; C: Taphozoinae; D: Emballonurinae; E: Emballonurini; F: Diclidurini; G: Afro-Malagasy emballonurini genera; H: Indo-Pacific emballonurini genus; I: New World Molossidae group; J: Otomops group; K: Oriental-Australasian Miniopterus clade; L: Ethiopian Miniopterus clade; M: Indian Ocean Miniopterus clade (JPEG 3901 kb)

High resolution image (TIFF 6293 kb) (download TIF )

Fig S10.

Relationships within Phyllostomidae. The subfamilies are indicated in bold (JPEG 3877 kb)

High resolution image (TIFF 6451 kb) (download TIF )

Fig S11.

Relationships within Vespertilionidae (part 1: Kerivoulinae, Murininae, Myotinae). A: New World Myotis clade; B: Paleartic lineage โ€œbrandtiiโ€; C: Neartic clade; D: Neotropical clade; E: Old World Myotis clade; F: Ethiopian clade; G: Euroasian clade; H: Oriental clade (JPEG 3563 kb)

High resolution image (TIFF 5872 kb) (download TIF )

Fig S12.

Relationships within Vespertilionidae (part 2: Vespertilioninae). The tribes are indicated in bold (JPEG 4337 kb)

High resolution image (TIFF 6940 kb) (download TIF )

Fig S13.

Outgroups (JPEG 608 kb)

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Fig S14.

Yinpterochiroptera (JPEG 2716 kb)

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Fig S15.

Yangochiroptera (in part): Noctilionoidea + Emballonuroidea (JPEG 2677 kb)

High resolution image (TIFF 4790 kb) (download TIF )

Fig S16.

Yangochiroptera (in part): Vespertilionoidea (JPEG 3078 kb)

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Fig S17. (download PDF )

Complete molecular dating obtained from a Bayesian analysis (PDF 72 kb)

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Amador, L.I., Moyers Arรฉvalo, R.L., Almeida, F.C. et al. Bat Systematics in the Light of Unconstrained Analyses of a Comprehensive Molecular Supermatrix. J Mammal Evol 25, 37โ€“70 (2018). https://doi.org/10.1007/s10914-016-9363-8

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