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Identifying unstable taxa: Efficient implementation of triplet-based measures of stability, and comparison with Phyutility and RogueNaRok
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This paper describes an efficient implementation of triplet-based measures of stability, in the program TNT. The only available implementations of such measures are much slower than the present one, either because of an inefficient implementation (Phyutility, Thor) or because the stability is evaluated with quartets (RogueNaRok, requiring O(t(4)), instead of the O(t(3)) possible for triplets). The method to quickly calculate triplets is applied to solving IterPCR (Pol and Escapa, 2009). It is shown that, in some cases, IterPCR or other algorithms in the program TNT (e.g. commands prunnelsen, prunmajor, or chkmoves) produce more informative results than analysis with RogueNaRok.
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The character information in the rogues can influence the relationships of the other taxa.
“The rogue taxa were excluded from the trees when calculating the consensus; they were never excluded from the matrix in any search for optimal trees (thus, the character information in the rogues can influence the relationships of the other taxa; see [105] for a recent discussion of this problem).”
From Potential for Powered Flight Neared by Most Close Avialan Relatives, but Few Crossed Its Thresholds (2020) · cites this paper as [105]
The rogue taxa were excluded from the trees when calculating the consensus; they were never excluded from the matrix in any search for optimal trees.
“The rogue taxa were excluded from the trees when calculating the consensus; they were never excluded from the matrix in any search for optimal trees (thus, the character information in the rogues can influence the relationships of the other taxa; see [105] for a recent discussion of this problem).”
From Potential for Powered Flight Neared by Most Close Avialan Relatives, but Few Crossed Its Thresholds (2020) · cites this paper as [105]
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TNT version 1.5, including a full implementation of phylogenetic morphometrics
Goloboff, P. A., & Catalano, S. A. (2016) · Cladistics · doi:10.1111/cla.12160 · Cited by 2,254
“…extension of implied weighting to take into account missing entries and average amounts of homoplasy in sets of characters (Goloboff, 2013), new search routines for difficult data sets (Goloboff, 2014, 2015), or the implementation of algorithms to identify wildcard taxa (Goloboff and Szumik, 2015).”
BackgroundMethodSelf-citationMai, U., & Mirarab, S. (2018) · BMC Genomics · doi:10.1186/s12864-018-4620-2 · Cited by 387
Hosner, P. A., Faircloth, B. C., Glenn, T. C., Braun, E. L., & Kimball, R. T. (2015) · Molecular Biology and Evolution · doi:10.1093/molbev/msv347 · Cited by 241
Potential for Powered Flight Neared by Most Close Avialan Relatives, but Few Crossed Its Thresholds
Pei, R., Pittman, M., Goloboff, P. A., Dececchi, T. A., Habib, M. B., Kaye, T. G., Larsson, H. C. E., Norell, M. A., Brusatte, S. L., & Xu, X. (2020) · Current Biology · doi:10.1016/j.cub.2020.06.105 · Cited by 137
“The rogue taxa were excluded from the trees when calculating the consensus; they were never excluded from the matrix in any search for optimal trees (thus, the character information in the rogues can influence the relationships of the other taxa; see [105] for a recent discussion of this problem).”
Self-citationSmith, M. R. (2019) · Biology Letters · doi:10.1098/rsbl.2018.0632 · Cited by 131
(2016) · doi:10.1093/sysbio/syw041 · Cited by 120
“As rogue taxa can frequently have a negative impact on topology or in a bootstrap analysis (see a review in Goloboff and Szumik 2015), we also explored the presence of putative wild card taxa in our data set with RogueNaRok (Aberer et al. 2013), with the aim of identifying a set of taxa that, if…”
MethodPereyra, M. O., Blotto, B. L., Baldo, D., Chaparro, J. C., Ron, S. R., Elias‐Costa, A. J., Iglesias, P. P., Venegas, P. J., Thomé, M. T. C., Ospina‐Sarria, J. J., et al. (2021) · Bulletin of the American Museum of Natural History · doi:10.1206/0003-0090.447.1.1 · Cited by 87
Godoy, P. L., Bronzati, M., Eltink, E., Marsola, J. C. A., Cidade, G. M., Langer, M. C., & Montefeltro, F. C. (2016) · PeerJ · doi:10.7717/peerj.2075 · Cited by 72
“This contribution used TNT v.1.1, a program made freely available thanks to a subsidy by the Willi Hennig Society.”
MethodInfluentialAfrica’s oldest dinosaurs reveal early suppression of dinosaur distribution
Griffin, C. T., Wynd, B. M., Munyikwa, D., Broderick, T. J., Zondo, M., Tolan, S., Langer, M. C., Nesbitt, S. J., & Taruvinga, H. R. (2022) · Nature · doi:10.1038/s41586-022-05133-x · Cited by 73
Herne, M., Nair, J. P., Evans, A. R., & Tait, A. (2019) · Journal of Paleontology · doi:10.1017/jpa.2018.95 · Cited by 68
“Iterative PCR was assessed with the implementation of the original script (‘iterPCR.run’) available in TNT 1.5 (Goloboff and Szumik, 2015).”
MethodInfluentialPlasticity and Convergence in the Evolution of Short-Necked Plesiosaurs
Fischer, V., Benson, R., Zverkov, N. G., Soul, L. C., ARKHANGELSKY, M. S., Lambert, O., Stenshin, I. M., Uspensky, G. N., & Druckenmiller, P. S. (2017) · Current Biology · doi:10.1016/j.cub.2017.04.052 · Cited by 68
Phylogeny of the Caninae (Carnivora): Combining morphology, behaviour, genes and fossils
Zrzavý, J., Duda, P., Robovský, J., Okřinová, I., & Řičánková, V. P. (2018) · Zoologica Scripta · doi:10.1111/zsc.12293 · Cited by 59
Buenaventura, E., Whitmore, D., & Pape, T. (2016) · Cladistics · doi:10.1111/cla.12161 · Cited by 55
“Goloboff and Szumik (2015) recommended against post-pruning analyses, as this will disregard the character information provided by the rogue taxa and may alter the retrieved relationships between the other taxa.”
BackgroundMethodResultInfluentialBuenaventura, E., & Pape, T. (2016) · Molecular Phylogenetics and Evolution · doi:10.1016/j.ympev.2016.12.028 · Cited by 51
Barker, C. T., Hone, D. W. E., Naish, D., Cau, A., Lockwood, J. A. F., Foster, B., Clarkin, C., Schneider, P., & Gostling, N. J. (2021) · Scientific Reports · doi:10.1038/s41598-021-97870-8 · Cited by 49
Torres, A., Goloboff, P. A., & Catalano, S. A. (2021) · Cladistics · doi:10.1111/cla.12477 · Cited by 45
Self-citationGroh, S. S., Upchurch, P., Barrett, P. M., & Day, J. J. (2019) · Zoological Journal of the Linnean Society · doi:10.1093/zoolinnean/zlz117 · Cited by 46
Human population history revealed by a supertree approach
Duda, P., & Zrzavý, J. (2016) · Scientific Reports · doi:10.1038/srep29890 · Cited by 39
Inferring ‘weak spots’ in phylogenetic trees: application to mosasauroid nomenclature
Madzia, D., & Cau, A. (2017) · PeerJ · doi:10.7717/peerj.3782 · Cited by 39
“Therefore, Pol & Escapa (2009) offered to use a TNT script, IterPCR, that provides a list of characters related to the instability of each unstable taxon.”
MethodInfluentialIdentifying and addressing methodological incongruence in phylogenomics: A review
Fleming, J. F., Valero‐Gracia, A., & Struck, T. H. (2023) · Evolutionary Applications · doi:10.1111/eva.13565 · Cited by 41
“Roguenarok's focus on pruning based on tree support to improve a bootstrap score (Felsenstein, 1978; Goloboff & Szumik, 2015) can result in erroneous, but well- supported topologies being favoured by the algorithm, thus removing the uncertainty that would normally be expressed by a low bootstrap…”
BackgroundUsing Information Theory to Detect Rogue Taxa and Improve Consensus Trees
Smith, M. R. (2021) · Systematic Biology · doi:10.1093/sysbio/syab099 · Cited by 31
“…Page: 1093 1088–1094 example by using other quantifications of leaf stability (e.g., Estabrook 1992; Thorley and Wilkinson 1999; Pol and Escapa 2009; Goloboff and Szumik 2015), or by integrating other heuristic methods, such as a “genetic” evolutionary approach (Srivastava et al. 2018).”
MethodRaven, T. J., Barrett, P. M., Joyce, C. B., & Maidment, S. C. R. (2023) · Journal of Systematic Palaeontology · doi:10.1080/14772019.2023.2205433 · Cited by 29
Herrera, Y., Fernández, M., & Vennari, V. V. (2020) · Cretaceous Research · doi:10.1016/j.cretres.2020.104673 · Cited by 29
Pol, D., Baiano, M. A., Černý, D., Novas, F. E., Cerda, I. A., & Pittman, M. (2024) · Cladistics · doi:10.1111/cla.12583 · Cited by 28
“We carried out the IterPCR procedure (Pol and Escapa, 2009; see iterpcr.out in Supplementary Data) as implemented by Goloboff and Szumik (2015) in TNT in order to detect unstable taxa.”
MethodBronzati, M., & Rauhut, O. W. M. (2017) · Zoological Journal of the Linnean Society · doi:10.1093/zoolinnean/zlx029 · Cited by 28
Timilsena, P. R., Barrett, C. F., Piñeyro‐Nelson, A., Wafula, E. K., Ayyampalayam, S., McNeal, J. R., Yukawa, T., Givnish, T. J., Graham, S. W., Pires, J. C., et al. (2022) · Genome Biology and Evolution · doi:10.1093/gbe/evac183 · Cited by 22
“Identifying unstable taxa: efficient implementation of triplet-based measures of stability, and comparison with phyutility and RogueNaRok.”
MethodInfluentialVento, B., & Agraín, F. (2018) · Acta Palaeontologica Polonica · doi:10.4202/app.00493.2018 · Cited by 21
“This methodology was implemented in TNT under the “pcrprune” command (Goloboff and Szumik 2015).”
MethodDíaz, V. D., Mannion, P. D., Csiki‐Sava, Z., & Upchurch, P. (2025) · Journal of Systematic Palaeontology · doi:10.1080/14772019.2024.2441516 · Cited by 21
Blokland, J. C., Reid, C. M., Worthy, T. H., Tennyson, A. J. D., Clarke, J. A., & Scofield, R. P. (2019) · Palaeontologia Electronica · doi:10.26879/1009 · Cited by 20
Sun, J., James, S. W., Jiang, J., Yao, B., Zhang, L., Liu, M., Qiu, J., & Hu, F. (2017) · Molecular Phylogenetics and Evolution · doi:10.1016/j.ympev.2017.07.026 · Cited by 19
Godoy, P. L., Cidade, G. M., Montefeltro, F. C., Langer, M. C., & Norell, M. A. (2020) · Papers in Palaeontology · doi:10.1002/spp2.1339 · Cited by 18
Bronzati, M., Benson, R., & Rauhut, O. W. M. (2017) · Palaeontology · doi:10.1111/pala.12344 · Cited by 18
“The PCRPrune command of TNT (Goloboff & Szumik 2015) was used to generate a reduced strict consensus tree depicting the alternative positions of the unstable OTUs (see Bronzati et al. 2017a).”
MethodCidade, G. M., Fortier, D., & Hsiou, A. S. (2019) · Zoological Journal of the Linnean Society · doi:10.1093/zoolinnean/zlz051 · Cited by 16
Fernández, M., Campos, L., Maxwell, E. E., & Garrido, A. C. (2021) · Journal of Vertebrate Paleontology · doi:10.1080/02724634.2021.1922427 · Cited by 13
“Most of the contributions dealing with missing data agree on the need to include all taxa (although incomplete) that may affect the topology (Wilkinson, 1995; Pol and Escapa, 2009; Goloboff and Szumik, 2015; Siu-Ting et al., 2015).”
BackgroundHolwerda, F. M., & Pol, D. (2018) · Spanish Journal of Palaeontology · doi:10.7203/sjp.33.2.13604 · Cited by 13
Flores, J. R., Bippus, A. C., de Ullivarri, C. F., Suárez, G. M., Hyvönen, J., & Tomescu, A. M. F. (2023) · New Phytologist · doi:10.1111/nph.19254 · Cited by 11
Barker, C. T., Handford, L., Naish, D., Wills, S., Hendrickx, C., Hadland, P. T., Brockhurst, D., & Gostling, N. J. (2024) · Papers in Palaeontology · doi:10.1002/spp2.1604 · Cited by 11
“In the unconstrained analyses, wildcard taxa were identified using the iterPCR procedure (Pol & Escapa 2009; Goloboff & Szumik 2015), and Bremer supports were used to assess the nodal support of the resulting reduced consensus trees.”
Gee, B. M. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.06.22.496729 · Cited by 11
“…by Schoch (2013) are clearly wildcards, either upon reanalysis of the original matrix or upon reanalysis of subsequent derivates, and it is not clear how “problematic” taxa were originally identified (e.g., with an Adams consensus or the IterPCR method of TNT; Goloboff and Szumik, 2015).”
Barker, C. T., Naish, D., & Gostling, N. J. (2023) · PeerJ · doi:10.7717/peerj.15453 · Cited by 10
“In the unconstrained analyses, wildcard OTUs were identified using the iterPCR function ( Goloboff & Szumik, 2015 ; Pol & Escapa, 2009 ), and Bremer support values were calculated as a measure of nodal support in the resulting reduced consensus.”
Herrera, Y., Aiglstorfer, M., & Bronzati, M. (2021) · Journal of Systematic Palaeontology · doi:10.1080/14772019.2021.1883138 · Cited by 9
“The prunnelsen command (Goloboff & Szumik 2015) was used to identify the unstable OTUs, which consisted of Congosaurus bequaerti, Crocodylus acutus, Cr. intermedius, Cr. mindorensis, Cr. moreletti, Cr., rhombifer, Cr. suchus and the ‘Swiss rhacheosaurin’.”
MethodMeasuring inferential importance of taxa using taxon influence indices
Denton, J. S. S., & Goolsby, E. W. (2018) · Ecology and Evolution · doi:10.1002/ece3.3941 · Cited by 9
“Systematic Biology, 62, 162–166. https://doi.org/10.1093/ sysbio/sys078 Aberer, A. J., Pattengale, N. D., & Stamatakis, A. (2010).”
BackgroundMethodInfluentialSong, N., Wei, S., & Wang, M. (2024) · Mitochondrial DNA Part A · doi:10.1080/24701394.2024.2345663 · Cited by 8
Scimitar-crested Spinosaurus species from the Sahara caps stepwise spinosaurid radiation
Sereno, P. C., Vidal, D., Myhrvold, N., Johnson‐Ransom, E., Real, M. C., Baumgart, S. L., Fontela, N. S., Green, T. L., Saitta, E. T., Adamou, B., et al. (2026) · Science · doi:10.1126/science.adx5486 · Cited by 8
TreeShrink: Efficient Detection of Outlier Tree Leaves
Mai, U., & Mirarab, S. (2017) · Lecture notes in computer science · doi:10.1007/978-3-319-67979-2_7 · Cited by 9
Kinateder, T., Drexler, L., Straub, K., Merkl, R., & Sterner, R. (2022) · Protein Science · doi:10.1002/pro.4536 · Cited by 8
“All possible combinations of the three filtering methods ( heatmap approach , rogue taxa and RogueNaRok ; Goloboff & Szumik, 2015) were applied to generate a data set with the best suitable selected sequences.”
MethodDirnberger, M., Peláez‐Campomanes, P., & López‐Antoñanzas, R. (2024) · PeerJ · doi:10.7717/peerj.18440 · Cited by 7
A new large hypercarnivorous crocodyliform from the Maastrichtian of Southern Patagonia, Argentina
Novas, F. E., Pol, D., Agnolín, F. L., Carvalho, I. D. S., Manabe, M., Tsuihiji, T., Rozadilla, S., Lio, G. L., & Isasi, M. P. (2025) · PLoS ONE · doi:10.1371/journal.pone.0328561 · Cited by 7
“Unstable taxa were detected using IterPCR [29,30] and nodal support was evaluated using parsimony jackknife [31] considering the impact of unstable taxa [32].”
A phylogenetic C interpreter for TNT
Goloboff, P. A., & Morales, M. E. (2020) · Bioinformatics · doi:10.1093/bioinformatics/btaa214 · Cited by 7
Self-citationExploring the impact of unstable terminals on branch support values in paleontological data
Flores, J. R., Lehtonen, S., & Hyvönen, J. (2021) · Paleobiology · doi:10.1017/pab.2020.64 · Cited by 6
“Note that taxa that are unstable with minimum differences in fit are known to reduce branch support, and devising methods to efficiently deal with them has been challenging (Aberer and Stamatakis 2011; Aberer et al. 2013; Goloboff and Szumik 2015; Pei et al. 2020; Pol and Goloboff 2020).”
BackgroundMethodFerrari, R. R., & Packer, L. (2020) · Zoological Journal of the Linnean Society · doi:10.1093/zoolinnean/zlaa128 · Cited by 5
A new pachyrukhine (Notoungulata: Typotheria) from the late Early Miocene of south-central Chile
Solórzano, A., Encinas, A., Kramarz, A., Carrasco, G., Núñez‐Flores, M., & Bobe, R. L. (2023) · Historical Biology · doi:10.1080/08912963.2023.2214568 · Cited by 5
“To improve the resolution of the phylogenetic tree, the method ‘Iterative Positional Congruence Reduced’ (IterPCR) of Pol and Escapa (2009), implemented in TNT 1.5 (Goloboff and Szumik 2015), was employed to detect possible unstable taxa.”
MethodInfluentialSánchez-Fenollosa, S., & Cobos, A. (2025) · Vertebrate Zoology · doi:10.3897/vz.75.e146618 · Cited by 4
“The Iter-PCR method (Pol and Escapa 2009) using the command ‘pcrprune’ (Goloboff and Szumik 2015) was employed to identify the rogue taxa.”
Wang, H.-C., Susko, E., & Roger, A. J. (2016) · Molecular Phylogenetics and Evolution · doi:10.1016/j.ympev.2016.08.017 · Cited by 4
“In some cases the whole analysis will have to be re-conducted with the rogue taxa removed (Thomson and Shaffer, 2009; Aberer and Stamatakis, 2011; see also Goloboff and Szumik, 2015 for a discussion of pruning trees or matrices).”
BackgroundTwo new compsognathid-like theropods show diversified predation strategies in theropod dinosaurs
Qiu, R., Wang, X., Jiang, S., Meng, J., & Zhou, Z. (2025) · National Science Review · doi:10.1093/nsr/nwaf068 · Cited by 4
Klawitter, J., Bouckaert, R., & Drummond, A. J. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.09.25.615070 · Cited by 5
“The majority of rogue detection algorithms thus aim to improve both the resolution and the support of the clades (splits) in a consensus tree [1, 2, 13, 28, 31], which however is an NP-hard problem [9].”
BackgroundFernandes, A. E., Pol, D., & Rauhut, O. W. M. (2024) · Royal Society Open Science · doi:10.1098/rsos.241238 · Cited by 2
PickMe : Sample Selection for Species Tree Reconstruction using Coalescent Weighted Quartets
Rusinko, J., Cai, Y., Crysler, A., Thompson, K. L., Boutte, J., Fishbein, M., & Straub, S. C. K. (2025) · Systematic Biology · doi:10.1093/sysbio/syaf017 · Cited by 1
“”Rogue” or ”wildcard” sam‑ ples whose inclusion reduces the resolution of and sup‑ port for splits in gene trees may be identified and re‑ moved (Aberer et al., 2012; Goloboff and Szumik, 2015, and e.g. Foley et al., 2019; Laumer et al., 2019; Li et al., 2021; Salichos and Rokas, 2013).”
Veselić, S., Vujić, A., Radenković, S., Zorić, L. Š., Djan, M., & Ståhls, G. (2025) · Cladistics · doi:10.1111/cla.70009 · Cited by 1
“These kinds of species could be considered as rogue taxa (Wilkinson, 1994; Goloboff and Szumik, 2015; Smith, 2022).”
A new fossil fish sheds light on the rapid evolution of early lungfishes
Qiao, T., Cui, X., Zhao, W., Liu, C., Li, M., Lü, J., Choo, B., & Zhu, M. (2025) · Current Biology · doi:10.1016/j.cub.2025.11.032 · Cited by 1
A comparison of two universal angiosperm bait sets and the phylogenomics of AlismatalesPreprint
Biffin, E., Waycott, M., Hammer, T., & van Dijk, K.-J. (2025) · bioRxiv · doi:10.1101/2025.09.15.676180 · Cited by 1
On the effect of measures for comparing trees on the representation of treespace
Goloboff, P. A., & Morales, M. E. (2025) · Cladistics · doi:10.1111/cla.12614
“Triplets are fewer than quar-tets, and the existing code used by TNT for calculating IterPCR values (Goloboff and Szumik, 2015) can be adapted easily for checking triplets.”
Self-citationGarzón‐Orduña, I. J., Brower, A. V. Z., Joele, F. R., & Torres, A. (2026) · Systematic Entomology · doi:10.1111/syen.70024
“%) and found in all nine datasets, are strong candidates to behave as wild cards or rogue taxa (Goloboff & Szumik, 2015; Sanderson et al., 2011; Wilkinson, 1995) during phylogenetic inference.”
Juárez, M. L., Goloboff, P. A., & Szumik, C. (2026) · American Museum Novitates · doi:10.1206/4048.1
Self-citationreutter, A., Carballido, J. L., Windholz, G. J., Pol, D., & Rauhut, O. W. M. (2026) · PeerJ · doi:10.7717/peerj.20945
Motta, M. J., Rolando, A. M. A., Rozadilla, S., Agnolín, F. L., Egli, F. B., Álvarez-Herrera, G. P., Chimento, N. R., Coco, G. L., Tsuihiji, T., Manabe, M., et al. (2026) · Journal of Vertebrate Paleontology · doi:10.1080/02724634.2026.2656456 · Cited by 1
“Strict consensus and reduced strict consensus were produced, identifying unstable taxa using iterPCR (Pol & Escapa, 2009) as implemented in TNT (Goloboff & Szumik, 2015).”
InfluentialOyabu, S., Kobayashi, Y., Zelenitsky, D. K., Fiorillo, A. R., & Kano, M. (2026) · Cretaceous Research · doi:10.1016/j.cretres.2026.106477
Sualdea, L. R., López‐Antoñanzas, R., Sierra, M., & Peláez‐Campomanes, P. (2026) · Journal of Systematic Palaeontology · doi:10.1080/14772019.2026.2733171
Fernandez, R., Edgecombe, G. D., & Giribet, G. (2016) · Data Archiving and Networked Services (DANS) · doi:10.5061/dryad.8mp17.1
Metadata
- Authors
- Goloboff, P. A., & Szumik, C. A.
- Affiliations
- Consejo Nacional de Investigaciones Científicas y Técnicas (AR)
- Corresponding author
- Goloboff, Pablo A.
- Venue
- Molecular Phylogenetics and Evolution
- Publisher
- Elsevier BV
- Published
- 2015-07
- Volume
- 88
- Pages
- 93-104
- ISSN
- 1055-7903 (ISSN-L)
- OpenAlex
- W1975311257
- Cited by
- 73
- References
- 33
- Citations by year
- 2026: 7 · 2025: 9 · 2024: 6 · 2023: 5 · 2022: 4 · 2021: 7 · 2020: 6 · 2019: 5 · 2018: 5 · 2017: 6 · 2016: 12 · 2015: 1
- FWCI
- 3.47
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Molecular Phylogenetics and Evolution (publishedVersion)
- PubMed (publishedVersion)
- CONICET Digital (CONICET) (submittedVersion, cc-by-nc-nd)
- Funders
- CONICET (112 201101 00687)
- NSF
- NASA
- Fundaçao de Amparo à Pesquisa do Estado de São Paulo
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Genomics and Phylogenetic Studies
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Evolution and Paleontology Studies
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Plant Diversity and Evolution
- Keywords
- phylogenetic stability, TNT, computational efficiency
- MeSH terms
- Algorithms, Phylogeny, Software
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