A remarkable troglomorphic ant, Yavnella laventa sp. nov. (Hymenoptera: Formicidae: Leptanillinae), identified as the first known worker of Yavnella Kugler by phylogenomic inference
Zachary Hayes Griebenow
A * , Marco Isaia B and Majid Moradmand C
+ Author Affiliations
- Author Affiliations
A Department of Entomology and Nematology, University of California, Davis, CA, USA.
B Laboratorio di Ecologia, Ecosistemi terrestri, Dipartimento di Scienze della Vita e Biologia dei Sistemi, Università di Torino, Torino, Italy.
C Department of Plant and Animal Biology, Faculty of Biological Science and Technology, University of Isfahan, Isfahan, Iran.
Invertebrate Systematics 36(12) 1118-1138 https://doi.org/10.1071/IS22035
Submitted: 21 July 2022 Accepted: 27 October 2022 Published: 5 December 2022
© 2022 The Author(s) (or their employer(s)). Published by CSIRO Publishing. This is an open access article distributed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License (CC BY-NC-ND)
Abstract
The ant subfamily Leptanillinae (Hymenoptera: Formicidae) consists of minute soil-dwelling species, with several genera within this clade being based solely upon males, including Yavnella Kugler. The dissociation of males and workers has resulted in taxonomic confusion for the Leptanillinae. We here describe the worker caste of Yavnella, facilitated by maximum-likelihood and Bayesian inference from 473 partitioned ultra-conserved element loci, this dataset including 49 other leptanilline species, both described and undescribed. Yavnella laventa sp. nov. is described from seven worker specimens collected in south-western Iran from the Milieu Souterrain Superficiel, a subterranean microhabitat consisting of air-filled cavities among rock and soil fragments, which is subject to similar environmental conditions as caves. This species has bizarrely elongated appendages, which suggests that it is confined to cavities, in contrast with the soil-dwelling behaviour observed in other leptanilline ants. Based on its gracile phenotype relative to other Leptanillinae, Y. laventa shows remarkable adaptations for subterranean life, making it one of a very few examples of this syndrome among the ants. Moreover, the discovery of the worker caste of Yavnella expands our morphological knowledge of the leptanilline ants. We provide worker- and male-based diagnoses of Yavnella, along with a key to the genera of the Leptanillinae for which workers are known. The worker caste of Yavnella as known from this species is immediately recognisable, but the possibility must be noted that described workers of Leptanilla may in fact belong to Yavnella. Further molecular sampling is required to test this hypothesis.
ZooBank: urn:lsid:zoobank.org:pub:A54A5766-F35A-4074-9353-1C70FE3955D3
Keywords: Insecta, Iran, Milieu Souterrain Superficiel, molecular systematics, Palaearctic, phylogenomics, subterranean biology, troglomorphism.
References
Abbassi, HR, Aleni, MR, Feiznia, S, Darvish, M, Ahmadian, M, and Shahbazi, A (2015). Effects of geological formation on desertification in the Mond watershed. Iranian Journal of Range & Desert Research 22, 583–593.Aberer, AJ, Kobert, K, and Stamatakis, A (2014). ExaBayes: massively parallel Bayesian tree inference for the whole-genome era. Molecular Biology and Evolution 31, 2553–2556.
| ExaBayes: massively parallel Bayesian tree inference for the whole-genome era.Crossref | GoogleScholarGoogle Scholar |
Aswaj, P, Anoop, K, and Priyadarsanan, DR (2020). First record of the rarely collected ant Protanilla gengma Xu, 2012 (Hymenoptera, Formicidae, Leptanillinae) from the Indian subcontinent. Check List 16, 1621–1625.
| First record of the rarely collected ant Protanilla gengma Xu, 2012 (Hymenoptera, Formicidae, Leptanillinae) from the Indian subcontinent.Crossref | GoogleScholarGoogle Scholar |
Badino, G (2010). Underground meteorology – “what’s the weather underground?” Acta Carsologica 39, 427–448.
| Underground meteorology – “what’s the weather underground?”Crossref | GoogleScholarGoogle Scholar |
Baidya, P, and Bagchi, S (2020). A new species of Protanilla Taylor 1990 (Hymenoptera: Formicidae: Leptanillinae) from India. Halteres 11, 19–24.
| A new species of Protanilla Taylor 1990 (Hymenoptera: Formicidae: Leptanillinae) from India.Crossref | GoogleScholarGoogle Scholar |
Bankevich, A, Nurk, S, Antipov, D, Gurevich, AA, Dvorkin, M, Kulikov, AS, Lesin, VM, Nikolenko, SI, Pham, S, Prjibelski, AD, Pyshkin, AV, Sirotkin, AV, Vyahhi, N, Tesler, G, Alekseyev, MA, and Pevzner, PA (2012). SPAdes: a new genome assembly algorithm and its applications to single-cell sequencing. Journal of Computational Biology 19, 455–477.
| SPAdes: a new genome assembly algorithm and its applications to single-cell sequencing.Crossref | GoogleScholarGoogle Scholar |
Barden, P, and Grimaldi, D (2012). Rediscovery of the bizarre Cretaceous ant Haidomyrmex Dlussky (Hymenoptera: Formicidae), with two new species. American Museum Novitates 3755, 1–16.
| Rediscovery of the bizarre Cretaceous ant Haidomyrmex Dlussky (Hymenoptera: Formicidae), with two new species.Crossref | GoogleScholarGoogle Scholar |
Baroni Urbani, C (1977). Materiali per una revisione della sottofamiglia Leptanillinae Emery (Hymenoptera: Formicidae). Entomologica Basiliensia 2, 427–488.
Baroni Urbani, C, and de Andrade, ML (2006). A new Protanilla Taylor, 1990 (Hymenoptera: Formicidae: Leptanillinae) from Sri Lanka. Myrmecologische Nachrichten 8, 45–47.
Barranco, P, Gilgado, JD, and Ortuño, VM (2013). A new mute species of the genus Nemobius Serville (Orthoptera, Gryllidae, Nemobiinae) discovered in colluvial, stony debris in the Iberian Peninsula: a biological, phenological and biometric study. Zootaxa 3691, 201–219.
| A new mute species of the genus Nemobius Serville (Orthoptera, Gryllidae, Nemobiinae) discovered in colluvial, stony debris in the Iberian Peninsula: a biological, phenological and biometric study.Crossref | GoogleScholarGoogle Scholar |
Bichuette, ME, Rantin, B, Hingst-Zaher, E, and Trajano, E (2015). Geometric morphometrics throws light on evolution of the subterranean catfish Rhamdiopsis krugi (Teleostei: Siluriformes: Heptapteridae) in eastern Brazil. Biological Journal of the Linnean Society 114, 136–151.
| Geometric morphometrics throws light on evolution of the subterranean catfish Rhamdiopsis krugi (Teleostei: Siluriformes: Heptapteridae) in eastern Brazil.Crossref | GoogleScholarGoogle Scholar |
Billen, J, Bauweleers, E, Hashim, R, and Ito, F (2013). Survey of the exocrine system in Protanilla wallacei (Hymenoptera, Formicidae). Arthropod Structure & Development 42, 173–183.
| Survey of the exocrine system in Protanilla wallacei (Hymenoptera, Formicidae).Crossref | GoogleScholarGoogle Scholar |
Bolton, B (1990). The higher classification of the ant subfamily Leptanillinae (Hymenoptera: Formicidae). Systematic Entomology 15, 267–282.
| The higher classification of the ant subfamily Leptanillinae (Hymenoptera: Formicidae).Crossref | GoogleScholarGoogle Scholar |
Bolton, B (2003). Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71, 1–370.
Borowiec, ML (2016). AMAS: a fast tool for alignment manipulation and computing of summary statistics. PeerJ 4, e1660.
| AMAS: a fast tool for alignment manipulation and computing of summary statistics.Crossref | GoogleScholarGoogle Scholar |
Borowiec, ML, Schulz, A, Alpert, GD, and Baňař, P (2011). Discovery of the worker caste and descriptions of two new species of Anomalomyrma (Hymenoptera: Formicidae: Leptanillinae) with unique abdominal morphology. Zootaxa 2810, 1–14.
| Discovery of the worker caste and descriptions of two new species of Anomalomyrma (Hymenoptera: Formicidae: Leptanillinae) with unique abdominal morphology.Crossref | GoogleScholarGoogle Scholar |
Borowiec, ML, Rabeling, C, Brady, SG, Fisher, BL, Schultz, TR, and Ward, PS (2019). Compositional heterogeneity and outgroup choice influence the internal phylogeny of the ants. Molecular Phylogenetics and Evolution 134, 111–121.
| Compositional heterogeneity and outgroup choice influence the internal phylogeny of the ants.Crossref | GoogleScholarGoogle Scholar |
Boudinot, BE (2018). A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota. Arthropod Structure & Development 47, 563–613.
| A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota.Crossref | GoogleScholarGoogle Scholar |
Boudinot, BE, Perrichot, V, and Chaul, JCM (2020). †Camelosphecia gen. nov., lost ant-wasp intermediates from the mid-Cretaceous (Hymenoptera, Formicoidea). ZooKeys 1005, 21–55.
| †Camelosphecia gen. nov., lost ant-wasp intermediates from the mid-Cretaceous (Hymenoptera, Formicoidea).Crossref | GoogleScholarGoogle Scholar |
Boudinot, BE, Moosdorf, OTD, Beutel, RG, and Richter, A (2021). Anatomy and evolution of the head of Dorylus helvolus (Formicidae: Dorylinae): patterns of sex- and caste-limited traits in the sausagefly and the driver ant. Journal of Morphology 282, 1616–1658.
| Anatomy and evolution of the head of Dorylus helvolus (Formicidae: Dorylinae): patterns of sex- and caste-limited traits in the sausagefly and the driver ant.Crossref | GoogleScholarGoogle Scholar |
Branstetter, MG, Longino, JT, Ward, PS, and Faircloth, BC (2017). Enriching the ant tree of life: enhanced UCE bait set for genome-scale phylogenetics of ants and other Hymenoptera. Methods in Ecology & Evolution 8, 768–776.
| Enriching the ant tree of life: enhanced UCE bait set for genome-scale phylogenetics of ants and other Hymenoptera.Crossref | GoogleScholarGoogle Scholar |
Castresana, J (2000). Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17, 540–552.
| Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis.Crossref | GoogleScholarGoogle Scholar |
Chernomor, O, von Haeseler, A, and Minh, BQ (2016). Terrace aware data structure for phylogenomic inference from supermatrices. Systematic Biology 65, 997–1008.
| Terrace aware data structure for phylogenomic inference from supermatrices.Crossref | GoogleScholarGoogle Scholar |
Christiansen KA (2005) Morphological adaptations. In ‘Encyclopedia of Caves’. (Eds D Culver, WB White) pp. 517–527. (Elsevier: Amsterdam, Netherlands)
Cigna, AA (2002). Modern trend [sic] in cave monitoring. Acta Carsologica 31, 35–54.
Cokendolpher JC, Reddell JR, Taylor SJ, Krejca JK, Suarez AV, Pekins CE (2009) Further ants (Hymenoptera: Formicidae) from caves of Texas. In ‘Studies on the Cave & Endogean Fauna of North America. Vol. 7’. (Eds JC Cokendolpher, JR Reddell) pp. 151–168. (University of Texas at Austin: Austin, TX, USA)
Collingwood, CA, and Agosti, D (1996). Formicidae (Insecta: Hymenoptera) of Saudi Arabia (part 2). Fauna of Saudi Arabia 15, 300–385.
Creighton, WS (1930). A review of the genus Myrmoteras (Hymenoptera: Formicidae). Journal of the New York Entomological Society 38, 177–192.
Cruaud, A, Nidelet, S, Arnal, P, Weber, A, Fusu, L, Gumovsky, A, Huber, J, Polaszek, A, and Rasplus, J-Y (2019). Optimized DNA extraction and library preparation for minute arthropods: application to target enrichment in chalcid wasps used for biocontrol. Molecular Ecology Resources 19, 702–710.
| Optimized DNA extraction and library preparation for minute arthropods: application to target enrichment in chalcid wasps used for biocontrol.Crossref | GoogleScholarGoogle Scholar |
Culver DC, Pipan T (2014) ‘Shallow Subterranean Habitats: Ecology, Evolution, & Conservation.’ (Oxford University Press: New York, NY, USA)
Eisenbeis G, Wichard W (1987) ‘Atlas on the Biology of Soil Arthropods.’ (Springer: Berlin, Germany)
Emery, C (1910). Hymenoptera. Fam. Formicidae. Subfam. Dorylinae. Genera Insectorum 102, 1–34.
Faircloth, BC (2016). PHYLUCE is a software package for the analysis of conserved genomic loci. Bioinformatics 32, 786–788.
| PHYLUCE is a software package for the analysis of conserved genomic loci.Crossref | GoogleScholarGoogle Scholar |
Griebenow, ZH (2020). Delimitation of tribes in the subfamily Leptanillinae (Hymenoptera: Formicidae), with a description of the male of Protanilla lini Terayama, 2009. Myrmecological News 30, 229–250.
| Delimitation of tribes in the subfamily Leptanillinae (Hymenoptera: Formicidae), with a description of the male of Protanilla lini Terayama, 2009.Crossref | GoogleScholarGoogle Scholar |
Griebenow, ZH (2021). Synonymisation of the male-based ant genus Phaulomyrma (Hymenoptera : Formicidae) with Leptanilla based upon Bayesian total-evidence phylogenetic inference. Invertebrate Systematics 35, 603–636.
| Synonymisation of the male-based ant genus Phaulomyrma (Hymenoptera : Formicidae) with Leptanilla based upon Bayesian total-evidence phylogenetic inference.Crossref | GoogleScholarGoogle Scholar |
Guindon, S, Dufayard, JF, Lefort, V, Anisimova, M, Hordijk, W, and Gascuel, O (2010). New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology 59, 307–321.
| New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0.Crossref | GoogleScholarGoogle Scholar |
Harris, RA (1979). A glossary of surface sculpturing. Occasional Papers in Entomology 28, 1–31.
Hoang, DT, Chernomor, O, von Haeseler, A, Minh, BQ, and Vinh, LS (2018). UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35, 518–522.
| UFBoot2: improving the ultrafast bootstrap approximation.Crossref | GoogleScholarGoogle Scholar |
Hölldobler B, Wilson EO (1990) ‘The Ants.’ (Harvard University Press: Cambridge, MA, USA)
Hsu, P-W, Hsu, F-C, Hsiao, Y, and Lin, CC (2017). Taxonomic notes on the genus Protanilla (Hymenoptera: Formicidae: Leptanillinae) from Taiwan. Zootaxa 4268, 117–130.
| Taxonomic notes on the genus Protanilla (Hymenoptera: Formicidae: Leptanillinae) from Taiwan.Crossref | GoogleScholarGoogle Scholar |
Ito, F, and Yamane, S (2020). Behavior of the queen of Leptanilla clypeata Yamane et Ito collected in the Bogor Botanical Gardens, West Java, Indonesia (Hymenoptera; Formicidae), with a note on colony composition and a description of the ergatoid queen. Asian Myrmecology 12, e012004.
| Behavior of the queen of Leptanilla clypeata Yamane et Ito collected in the Bogor Botanical Gardens, West Java, Indonesia (Hymenoptera; Formicidae), with a note on colony composition and a description of the ergatoid queen.Crossref | GoogleScholarGoogle Scholar |
Ito, F, Hashim, R, Mizuno, R, and Billen, J (2022). Notes on the biology of Protanilla sp. (Hymenoptera, Formicidae) collected in Ulu Gombak, Peninsular Malaysia. Insectes Sociaux 69, 13–18.
| Notes on the biology of Protanilla sp. (Hymenoptera, Formicidae) collected in Ulu Gombak, Peninsular Malaysia.Crossref | GoogleScholarGoogle Scholar |
Juberthie C, Decu V (2006) Interstitial habitat (terrestrial). In ‘Encyclopedia of Caves & Karst Science’. (Ed J Gunn) pp. 984–987. (Taylor and Francis Group: New York, NY, USA)
Kalyaanamoorthy, S, Minh, BQ, Wong, TKF, von Haeseler, A, and Jermiin, LS (2017). ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods 14, 587–589.
| ModelFinder: fast model selection for accurate phylogenetic estimates.Crossref | GoogleScholarGoogle Scholar |
Katoh, K, and Toh, H (2010). Parallelization of the MAFFT multiple sequence alignment program. Bioinformatics 26, 1899–1900.
| Parallelization of the MAFFT multiple sequence alignment program.Crossref | GoogleScholarGoogle Scholar |
Kugler, J (1986). The Leptanillinae (Hymenoptera: Formicidae) of Israel and a description of a new species from India. Israel Journal of Entomology 20, 45–47.
Lanfear, R, Calcott, B, Kainer, D, Mayer, C, and Stamatakis, A (2014). Selecting optimal partitioning schemes for phylogenomic datasets. BMC Evolutionary Biology 14, 82.
| Selecting optimal partitioning schemes for phylogenomic datasets.Crossref | GoogleScholarGoogle Scholar |
Leong, C-M, Yamane, S, and Guénard, B (2018). Lost in the city: discovery of the rare ant genus Leptanilla (Hymenoptera: Formicidae) in Macau with description of Leptanilla macauensis sp. nov. Asian Myrmecology 10, e010001.
| Lost in the city: discovery of the rare ant genus Leptanilla (Hymenoptera: Formicidae) in Macau with description of Leptanilla macauensis sp. nov.Crossref | GoogleScholarGoogle Scholar |
Lieberman, ZE, Billen, J, van de Kamp, T, and Boudinot, BE (2022). The ant abdomen: the skeletomuscular and soft tissue anatomy of Amblyopone australis workers (Hymenoptera: Formicidae). Journal of Morphology 283, 693–770.
| The ant abdomen: the skeletomuscular and soft tissue anatomy of Amblyopone australis workers (Hymenoptera: Formicidae).Crossref | GoogleScholarGoogle Scholar |
Liu, SP, Richter, A, Stoessel, A, and Beutel, RG (2019). The mesosomal anatomy of Myrmecia nigrocincta workers and evolutionary transformations in Formicidae (Hymenoptera). Arthropod Systematics & Phylogeny 77, 1–19.
Mammola, S, and Isaia, M (2016). The ecological niche of a specialized subterranean spider. Invertebrate Biology 135, 20–30.
| The ecological niche of a specialized subterranean spider.Crossref | GoogleScholarGoogle Scholar |
Mammola, S, Giachino, PM, Piano, E, Jones, A, Barberis, M, Badino, G, and Isaia, M (2016). Ecology and sampling techniques of an understudied subterranean habitat: the Milieu Souterrain Superficiel (MSS). The Science of Nature 103, 88.
| Ecology and sampling techniques of an understudied subterranean habitat: the Milieu Souterrain Superficiel (MSS).Crossref | GoogleScholarGoogle Scholar |
Man, P, Ran, H, Chen, Z, and Xu, Z (2017). The northernmost record of Leptanillinae in China with description of Protanilla beijingensis sp. nov. (Hymenoptera: Formicidae). Asian Myrmecology 9, e009008.
| The northernmost record of Leptanillinae in China with description of Protanilla beijingensis sp. nov. (Hymenoptera: Formicidae).Crossref | GoogleScholarGoogle Scholar |
Masuko, K (1990). Behavior and ecology of the enigmatic ant Leptanilla japonica Baroni Urbani (Hymenoptera: Formicidae: Leptanillinae). Insectes Sociaux 37, 31–57.
Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In ‘Proceedings of the Gateway Computing Environments Workshop (GCE)’, 14 November 2010, New Orleans, LA, USA. INSPEC Accession Number 11705685. (IEEE)
| Crossref |
Minh, BQ, Schmidt, HA, Chernomor, O, Schrempf, D, Woodhams, MD, von Haeseler, A, and Lanfear, R (2020). IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution 37, 1530–1534.
| IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era.Crossref | GoogleScholarGoogle Scholar |
Naka, T, and Maruyama, M (2018). Aphaenogaster gamagumayaa sp. nov.: the first troglobiotic ant from Japan (Hymenoptera: Formicidae: Myrmicinae). Zootaxa 4450, 135–141.
| Aphaenogaster gamagumayaa sp. nov.: the first troglobiotic ant from Japan (Hymenoptera: Formicidae: Myrmicinae).Crossref | GoogleScholarGoogle Scholar |
Nguyen, L-T, von Haeseler, A, and Minh, BQ (2018). Complex models of sequence evolution require accurate estimators as exemplified with the invariable site plus gamma model. Systematic Biology 67, 552–558.
| Complex models of sequence evolution require accurate estimators as exemplified with the invariable site plus gamma model.Crossref | GoogleScholarGoogle Scholar |
Ogata, K, Terayama, M, and Masuko, K (1995). The ant genus Leptanilla: discovery of the worker-associated male of L. japonica, and a description of a new species from Taiwan (Hymenoptera: Formicidae: Leptanillinae). Systematic Entomology 20, 27–34.
| The ant genus Leptanilla: discovery of the worker-associated male of L. japonica, and a description of a new species from Taiwan (Hymenoptera: Formicidae: Leptanillinae).Crossref | GoogleScholarGoogle Scholar |
Pape, R (2016). The importance of ants in cave ecology, with new records and behavioral observations of ants in Arizona caves. International Journal of Speleology 45, 185–205.
| The importance of ants in cave ecology, with new records and behavioral observations of ants in Arizona caves.Crossref | GoogleScholarGoogle Scholar |
Pérez-González, S, Gómez-Durán, JM, and Martínez-Ibáñez, MD (2020). Highlighting the elusive: new findings and a redescription of the rare ant Leptanilla plutonia López, Martínez et Barandica, 1994, presenting morphological novelties for the genus. Annales Zoologici 70, 289–304.
| Highlighting the elusive: new findings and a redescription of the rare ant Leptanilla plutonia López, Martínez et Barandica, 1994, presenting morphological novelties for the genus.Crossref | GoogleScholarGoogle Scholar |
Petersen, B (1968). Some novelties in presumed males of Leptanillinae (Hym., Formicidae). Entomologiske Meddelelser 36, 577–598.
| Some novelties in presumed males of Leptanillinae (Hym., Formicidae).Crossref | GoogleScholarGoogle Scholar |
Rabeling, C, Brown, JM, and Verhaagh, M (2008). Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences 105, 14913–14917.
| Newly discovered sister lineage sheds light on early ant evolution.Crossref | GoogleScholarGoogle Scholar |
Rambaut, A, Drummond, AJ, Xie, D, Baele, G, and Suchard, MA (2018). Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67, 901–904.
| Posterior summarization in Bayesian phylogenetics using Tracer 1.7.Crossref | GoogleScholarGoogle Scholar |
Reddell JR (1977) A preliminary survey of the caves of the Yucatan peninsula. In ‘Studies of the Caves & Cave Fauna of the Yucatan Peninsula’. (Ed. JR Reddell) Association for Mexican Cave Studies Bulletin, Vol. 6, pp. 215–296. (Speleo Press: Austin, TX, USA)
Richter, A, Garcia, FH, Keller, RA, Billen, J, Katzke, J, Boudinot, BE, Economo, EP, and Beutel, RG (2021). The head anatomy of Protanilla lini (Hymenoptera: Formicidae: Leptanillinae), with a hypothesis of their mandibular movement. Myrmecological News 31, 81–114.
| The head anatomy of Protanilla lini (Hymenoptera: Formicidae: Leptanillinae), with a hypothesis of their mandibular movement.Crossref | GoogleScholarGoogle Scholar |
Roncin, E, and Deharveng, L (2003). Leptogenys khammouanensis sp. nov. (Hymenoptera: Formicidae). A possible troglobitic species of Laos, with a discussion on cave ants. Zoological Science 20, 919–924.
| Leptogenys khammouanensis sp. nov. (Hymenoptera: Formicidae). A possible troglobitic species of Laos, with a discussion on cave ants.Crossref | GoogleScholarGoogle Scholar |
Saroj, S, Mandi, A, and Dubey, A (2022). A new species of the rare ant genus, Leptanilla Emery (Hymenoptera: Formicidae) from Eastern Himalaya, India. Asian Myrmecology 15, e015005.
| A new species of the rare ant genus, Leptanilla Emery (Hymenoptera: Formicidae) from Eastern Himalaya, India.Crossref | GoogleScholarGoogle Scholar |
Sendra, A, Komerički, A, Lips, J, Luan, Y, Selfa, J, and Jiménez-Valverde, A (2021). Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha). European Journal of Taxonomy 731, 1–46.
| Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha).Crossref | GoogleScholarGoogle Scholar |
Tagliacollo, VA, and Lanfear, R (2018). Estimating improved partitioning schemes for ultraconserved elements. Molecular Biology and Evolution 35, 1798–1811.
| Estimating improved partitioning schemes for ultraconserved elements.Crossref | GoogleScholarGoogle Scholar |
Talbot, CJ, and Alavi, M (1996). The past of a future syntaxis across the Zagros. Geological Society, London, Special Publications 100, 89–109.
| The past of a future syntaxis across the Zagros.Crossref | GoogleScholarGoogle Scholar |
Uéno, SI (1980). The anophthalmic trechine beetles of the group of Trechiama oshimai. Bulletin of the National Science Museum of Tokyo, Series A 6, 195–274.
Ward, PS, and Blaimer, BB (2022). Taxonomy in the phylogenomic era: species boundaries and phylogenetic relationships among North American ants of the Crematogaster scutellaris group. Zoological Journal of the Linnean Society 194, 893–937.
| Taxonomy in the phylogenomic era: species boundaries and phylogenetic relationships among North American ants of the Crematogaster scutellaris group.Crossref | GoogleScholarGoogle Scholar |
Ward, PS, and Brady, SG (2009). Rediscovery of the ant genus Amyrmex Kusnezov (Hymenoptera: Formicidae) and its transfer from Dolichoderinae to Leptanilloidinae. Zootaxa 2063, 46–54.
| Rediscovery of the ant genus Amyrmex Kusnezov (Hymenoptera: Formicidae) and its transfer from Dolichoderinae to Leptanilloidinae.Crossref | GoogleScholarGoogle Scholar |
Ward, PS, and Fisher, BL (2016). Tales of dracula ants: the evolutionary history of the ant subfamily Amblyoponinae (Hymenoptera: Formicidae). Systematic Entomology 41, 683–693.
| Tales of dracula ants: the evolutionary history of the ant subfamily Amblyoponinae (Hymenoptera: Formicidae).Crossref | GoogleScholarGoogle Scholar |
Wheeler WM (1938) Ants from the caves of Yucatan. In ‘Fauna of the Caves of Yucatan’. (Ed. AS Pearse) pp. 251–255. (Carnegie Institution of Washington: Washington, DC, USA)
Wilson, EO (1955). A monographic revision of the ant genus Lasius. Bulletin of the Museum of Comparative Zoology 113, 1–201.
Wong, MKL, and Guénard, B (2016). Leptanilla hypodracos sp. n., a new species of the cryptic ant genus Leptanilla (Hymenoptera, Formicidae) from Singapore, with new distribution data and an updated key to Oriental Leptanilla species. ZooKeys 551, 129–144.
| Leptanilla hypodracos sp. n., a new species of the cryptic ant genus Leptanilla (Hymenoptera, Formicidae) from Singapore, with new distribution data and an updated key to Oriental Leptanilla species.Crossref | GoogleScholarGoogle Scholar |
Xu, Z-H (2012). Furcotanilla, a new genus of the ant subfamily Leptanillinae from China with descriptions of two new species of Protanilla and P. rafflesi Taylor (Hymenoptera: Formicidae). Sociobiology 59, 477–491.
| Furcotanilla, a new genus of the ant subfamily Leptanillinae from China with descriptions of two new species of Protanilla and P. rafflesi Taylor (Hymenoptera: Formicidae).Crossref | GoogleScholarGoogle Scholar |
Yamada, A, Nguyen, DD, and Eguchi, K (2020). Unveiling the morphology of the Oriental rare monotypic ant genus Opamyrma Yamane, Bui & Eguchi, 2008 (Hymenoptera: Formicidae: Leptanillinae) and its evolutionary implications, with first descriptions of the male, larva, tentorium, and sting apparatus. Myrmecological News 30, 27–52.
| Unveiling the morphology of the Oriental rare monotypic ant genus Opamyrma Yamane, Bui & Eguchi, 2008 (Hymenoptera: Formicidae: Leptanillinae) and its evolutionary implications, with first descriptions of the male, larva, tentorium, and sting apparatus.Crossref | GoogleScholarGoogle Scholar |
Zarei M (2010) Hydrogeology of salt diapirs in the south of Iran. PhD dissertation, Shiraz University, Shiraz, Iran.
