Abstract
Fossils recorded information of the past phenotype and geographic distribution directly. Incorporating fossils information into phylogenies of living taxa would deepen our understanding of their macroevolution. Accurate taxonomy of these fossils is the first step before employing them. However, the taxonomy of some fossils has not been evaluated carefully. Here we take Artemisia as an example. We investigated the morphology of all the available Artemisia macrofossils based on the latest phylogeny and evaluated their taxonomic identities by comparing them with extant representative species. Our result showed that the taxonomic identities of these macrofossils are doubted. They would better not be treated as members of Artemisia for subsequent phylogenetic or biogeographic studies. This study improves our knowledge of the morphology of Artemisia, highlights the importance of careful morphological comparisons between fossils and living taxa, and reveals that attention should be given to the uncertainty of the fossil taxonomy.
References
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span style="color: #090911;"><span lang="en-GB">Barreda, V.D., Palazzesi, L., Teller</span></span>ía, M.C., Katinas, L., Crisci, J.V., Bremer, K., Passalia, M.G, Corsolini, R, Rodríguez Brizuela, R. & Bechis, F. (2010) Eocene Patagonia fossils of the daisy family. Science 329 (5999): 1621. https://doi.org/10.1126/science.1193108</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Bremer, K. & Humphries, C. (1993) Generic monograph of the Asteraceae-Anthemideae. Bulletin of the Natural History Museum 23: 1–177.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Chen, S.B. & Zhang, J.T. (1991) A Study on Pollen Morphology of Some Chinese Genera in Tribe Anthemideae. Journal of Systematics and Evolution 29 (3): 246–251.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Crepet, W.L. & Stuessy, T.F. (1978) A reinvestigation of the fossil Viguiera cronquistii (Compositae). Brittonia 30: 483–491. https://doi.org/10.2307/2806356</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Donoghue, M.J., Doyle, J.A., Gauthier, J., Kluge, A.G. & Rowe<span lang="en-GB">,</span> T. (1989) The importance of fossils in phylogeny reconstruction. Annual Review of Ecology, Evolution, and Systematics 20: 431–460. https://doi.org/10.1146/annurev.es.20.110189.002243</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Dos Reis, M. & Yang, Z. (2011) Approximate likelihood calculation on a phylogeny for Bayesian estimation of divergence times. Molecular Biology and Evolution 28: 2161–2172. https://doi.org/10.1093/molbev/msr045</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Graham, A. (1996) A contribution to the geologic history of the Compositae. In: Hind, D.N.N. & Beentje, H. (Eds.) Proceedings of the International Compositae Conference, Kew, 1994, vol. 1, Compositae: Systematics. Royal Botanical Gardens, Kew, pp. 123–140.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Hayat, M.Q., Ashraf, M., Khan, M.A., Yasmin, G., Shaheen, N. & Jabeen, S. (2009) Phylogenetic analysis of Artemisia L. (Asteraceae) based on micromorphological traits of pollen grains. African Journal of Biotechnology 8: 6561–6568.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Hermsen, E.J. & Hendricks, J.R. (2008) W(h)ither Fossils? Studying Morphological Character Evolution in the Age of Molecular Sequences. Annals of the Missouri Botanical Garden 95 (1): 72–100. https://doi.org/10.3417/2006206</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Iles, W.J.D., Smith, S.Y., Gandolfo, M.A. & Graham, S.W. (2015) Monocot fossils suitable for molecular dating analyses. Botanical Journal of the Linnean Society 178 (3): 346–374. https://doi.org/10.1111/boj.12233</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Jiang, L., Wang, Q., Ye, L.Z. & Ling, Y.R. (2005) Pollen morphology of Artemisia L. and its systematic significance. Wuhan University Journal of Natural Sciences 10: 448–454. https://doi.org/10.1007/BF02830685</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Landis, M.J., Eaton, D.A.R., Clement, W.L., Park, B., Spriggs, E.L., Sweeney, P.W., Edwards, E.J. & Donoghue, M.J. (2020) Joint phylogenetic estimation of geographic movements and biome shifts during the global diversification of Viburnum. Systematic Biology 70: 67–85. https://doi.org/10.1093/sysbio/syaa027</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Ling, Y.R., Humphries, C.J. & Gilbert, M.G. (2011) Artemisia Linnaeus. In: Wu, Z.Y., Raven, P.H. & Hing, D.Y. (Eds.) Flora of China, vol. 20. Science Press, Beijing & Missouri Botanical Garden Press, Saint-Louis, pp. 1151–1259.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Linnaeus<span lang="en-GB">,</span> C. (1754) Genera Plantarum. L. Salvius, Stockholm, 1200 pp.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Linnaeus<span lang="en-GB">,</span> C. (1767) Systerna Naturae, vol. 2. Salvius, Stockholm.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Malik, S., Vitales, D., Hayat, M., Korobkov, A., Garnatje, T. & Vallès<span lang="en-GB">,</span> J. (2017) Phylogeny and biogeography of Artemisia subgenus Seriphidium (Asteraceae: Anthemideae). Taxon 66: 934–952. https://doi.org/10.12705/664.8</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martín, J., Torrell, M. & Valles, J. (2001) Palynological features as a systematic marker in Artemisia L. and related genera (Asteraceae, Anthemideae). Plant Biology 3 (4): 372–378. https://doi.org/10.1055/s-2001-16462</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martín, J., Torrell, M., Korobkov<span lang="en-GB">,</span> A.A. & Valles, J. (2003) Palynological features as a systematic marker in Artemisia L. and related genera (Asteraceae, Anthemideae) – II: Implications for subtribe Artemisiinae delimitation. Plant Biology 5 (1): 85–93. https://doi.org/10.1055/s-2003-37979</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Matsunaga, K.K.S. & Smith, S.Y. (2021) Fossil palm reading: using fruits to reveal the deep roots of palm diversity. American Journal of Botany 108: 472–494. https://doi.org/10.1002/ajb2.1616</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Mei, Q., Chen, X., Xiang, L., Liu, Y., Su, Y., Gao, Y., Dai, W., Dong, P. & Chen, S. (2016) DNA Barcode for Identifying Folium Artemisiae argyi from Counterfeits. Biological and Pharmaceutical Bulletin 39 (9): 1531–1537. https://doi.org/10.1248/bpb.b16-00336</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miao, Y., Meng, Q., Fang, X., Yan, X., Wu, F. & Song<span lang="en-GB">,</span> C. (2011) Origin and development of Artemisia (Asteraceae) in Asia and its implications for the uplift history of the Tibetan Plateau: A review. Quaternary International 236: 3–12. https://doi.org/10.1016/j.quaint.2010.08.014</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Oberprieler, C., Himmelreich, S., Källersjö, M., Vallès, J., Watson, L.E. & Vogt, R. (2009) Anthemideae. In: Funk, V.A., Susanna, A., Stuessy, T.F. & Bayer, R.J. (Eds.) Systematics, Evolution and Biogeography of the Compositae. International Association Plant Taxonomy, Vienna. pp. 632–666.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Parham, J.F., Donoghue, P.C.J., Bell, C.J., Calway, T.D., Head, J.J., Holroyd, P.A., Inoue, J.G., Irmis, R.B., Joyce, W.G., Ksepka, D.T., Patané, J.S.L., Smith, N.D., Tarver, J.E., Van Tuinen, M., Yang, Z., Angielczyk, K.D., Greenwood, J., Hipsley, C.A., Jacobs, L., Makovicky, P.J., Müller, J., Krister, T.S., Jessica, M.T., Rachel, C.M.W. & Michael, J.B. (2012) Best practices for justifying fossil calibrations. Systematic Biology 61 (2): 346–359. https://doi.org/10.1093/sysbio/syr107</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sanderson, M.J. (2003) r8s: inferring absolute rates of molecular evolution and divergence times in the absence of a molecular clock. Bioinformatics 19: 301–302. https://doi.org/10.1093/bioinformatics/19.2.301</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sanz, M., Vilatersana, R., Hidalgo, O., Garcia-Jacas, N., Susanna, A., Schneeweiss, G. & Vallès, J. (2008) Molecular phylogeny and evolution of floral characters of Artemisia and allies (Anthemideae, Asteraceae): Evidence from nrDNA ETS and ITS sequences. Taxon 57: 1–13.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sauquet, H., Ho, S.Y.W., Gandolfo, M.A., Jordan, G.J., Wilf, P., Cantrill, D.J., Bayly, M.J., Bromham, L., Brown, G.K., Carpenter, R.J., Lee, D.M., Murphy, D.J., Sniderman, J.M.K. & Udovicic, F. (2012) Testing the impact of calibration on molecular divergence times using a fossil-rich group: the case of Nothofagus (Fagales). Systematic Biology 61: 289–313. https://doi.org/10.1093/sysbio/syr116</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sauquet, H. & Magallon, S. (2018) Key questions and challenges in angiosperm macroevolution. New Phytologist 219 (4): 1170–1187. https://doi.org/10.1111/nph.15104</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Shultz, L.M. (2006) Genus Artemisia (Asteraceae: Anthemideae). In: Flora of North America Editorial Committee (Ed.) The Flora of North America north of Mexico, vol. 19: Asterales. Oxford University Press, New York, pp. 503–534.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Smith, S.Y. (2013) The fossil record of non-commelinid monocots. In: Wilkin, P. & Mayo, S.J. (Eds.) Early events in monocot evolution. Cambridge University Press, Cambridge, pp. 29–59. https://doi.org/10.1017/CBO9781139002950.003</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Tkach, N.V., Hoffmann, M.H., Röser, M. & von Hagen, K.B. (2008) Temporal patterns of evolution in the Arctic explored in Artemisia L. (Asteraceae) lineages of different age. Plant Ecology & Diversity 1: 161–169. https://doi.org/10.1080/17550870802331912</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Tomescu, A.M.F. & Rothwell, G.W. (2022) Fossils and plant evolution: structural fingerprints and modularity in the evo-devo paradigm. EvoDevo 13: 8. https://doi.org/10.1186/s13227-022-00192-7</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vallès, J., Garcia, S., Hidalgo, O., Martín, J., Pellicer, J., Sanz, M. & Garnatje, T. (2011) Biology, genome evolution, biotechnological issues, and research including applied perspectives in Artemisia (Asteraceae). Advances in Botanical Research 60: 349–419. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Wood, H.M., Matzke, N.J., Gillespie, R.G. & Griswold, C.E. (2013) Treating fossils as terminal taxa in divergence time estimation reveals ancient vicariance patterns in the palpimanoid spiders. Systematic Biology 62: 264–284. https://doi.org/10.1093/sysbio/sys092</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Yang, Z. & Rannala, B. (2006) Bayesian estimation of species divergence times under a molecular clock using multiple fossil calibrations with soft bounds. Molecular Biology and Evolution 23: 212–226. https://doi.org/10.1093/molbev/msj024</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Zazula, G.D., Froese, D.G., Schweger, R.W., Mathewes, A.B., Beaudoin, A.M., Telka, C.R. & Harington, J.A. (2003) Westgate Palaeobotany: ice-age steppe vegetation in east Beringia. Nature 423. https://doi.org/10.1038/423603a</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Zazula, G.D., Froese, D.G., Elias, S.A., Kuzmina, S. & Mathewes, R.W. (2007) Arctic ground squirrels of the mammoth-steppe: paleoecology of middens from the last glaciation, Yukon Territory, Canada. Quaternary Science Reviews 26: 979–1003. https://doi.org/10.1016/j.quascirev.2006.12.006</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Zhang, C., Huang, C.H., Liu, M., Hu, Y., Panero, J.L., Luebert, F., Gao, T. & Ma, H. (2021) Phylotranscriptomic insights into Asteraceae diversity, polyploidy, and morphological innovation. Journal of Integrative Plant Biology 63 (7): 1273–1293. https://doi.org/10.1111/jipb.13078</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Bremer, K. & Humphries, C. (1993) Generic monograph of the Asteraceae-Anthemideae. Bulletin of the Natural History Museum 23: 1–177.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Chen, S.B. & Zhang, J.T. (1991) A Study on Pollen Morphology of Some Chinese Genera in Tribe Anthemideae. Journal of Systematics and Evolution 29 (3): 246–251.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Crepet, W.L. & Stuessy, T.F. (1978) A reinvestigation of the fossil Viguiera cronquistii (Compositae). Brittonia 30: 483–491. https://doi.org/10.2307/2806356</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Donoghue, M.J., Doyle, J.A., Gauthier, J., Kluge, A.G. & Rowe<span lang="en-GB">,</span> T. (1989) The importance of fossils in phylogeny reconstruction. Annual Review of Ecology, Evolution, and Systematics 20: 431–460. https://doi.org/10.1146/annurev.es.20.110189.002243</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Dos Reis, M. & Yang, Z. (2011) Approximate likelihood calculation on a phylogeny for Bayesian estimation of divergence times. Molecular Biology and Evolution 28: 2161–2172. https://doi.org/10.1093/molbev/msr045</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Graham, A. (1996) A contribution to the geologic history of the Compositae. In: Hind, D.N.N. & Beentje, H. (Eds.) Proceedings of the International Compositae Conference, Kew, 1994, vol. 1, Compositae: Systematics. Royal Botanical Gardens, Kew, pp. 123–140.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Hayat, M.Q., Ashraf, M., Khan, M.A., Yasmin, G., Shaheen, N. & Jabeen, S. (2009) Phylogenetic analysis of Artemisia L. (Asteraceae) based on micromorphological traits of pollen grains. African Journal of Biotechnology 8: 6561–6568.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Hermsen, E.J. & Hendricks, J.R. (2008) W(h)ither Fossils? Studying Morphological Character Evolution in the Age of Molecular Sequences. Annals of the Missouri Botanical Garden 95 (1): 72–100. https://doi.org/10.3417/2006206</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Iles, W.J.D., Smith, S.Y., Gandolfo, M.A. & Graham, S.W. (2015) Monocot fossils suitable for molecular dating analyses. Botanical Journal of the Linnean Society 178 (3): 346–374. https://doi.org/10.1111/boj.12233</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Jiang, L., Wang, Q., Ye, L.Z. & Ling, Y.R. (2005) Pollen morphology of Artemisia L. and its systematic significance. Wuhan University Journal of Natural Sciences 10: 448–454. https://doi.org/10.1007/BF02830685</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Landis, M.J., Eaton, D.A.R., Clement, W.L., Park, B., Spriggs, E.L., Sweeney, P.W., Edwards, E.J. & Donoghue, M.J. (2020) Joint phylogenetic estimation of geographic movements and biome shifts during the global diversification of Viburnum. Systematic Biology 70: 67–85. https://doi.org/10.1093/sysbio/syaa027</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Ling, Y.R., Humphries, C.J. & Gilbert, M.G. (2011) Artemisia Linnaeus. In: Wu, Z.Y., Raven, P.H. & Hing, D.Y. (Eds.) Flora of China, vol. 20. Science Press, Beijing & Missouri Botanical Garden Press, Saint-Louis, pp. 1151–1259.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Linnaeus<span lang="en-GB">,</span> C. (1754) Genera Plantarum. L. Salvius, Stockholm, 1200 pp.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Linnaeus<span lang="en-GB">,</span> C. (1767) Systerna Naturae, vol. 2. Salvius, Stockholm.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Malik, S., Vitales, D., Hayat, M., Korobkov, A., Garnatje, T. & Vallès<span lang="en-GB">,</span> J. (2017) Phylogeny and biogeography of Artemisia subgenus Seriphidium (Asteraceae: Anthemideae). Taxon 66: 934–952. https://doi.org/10.12705/664.8</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martín, J., Torrell, M. & Valles, J. (2001) Palynological features as a systematic marker in Artemisia L. and related genera (Asteraceae, Anthemideae). Plant Biology 3 (4): 372–378. https://doi.org/10.1055/s-2001-16462</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martín, J., Torrell, M., Korobkov<span lang="en-GB">,</span> A.A. & Valles, J. (2003) Palynological features as a systematic marker in Artemisia L. and related genera (Asteraceae, Anthemideae) – II: Implications for subtribe Artemisiinae delimitation. Plant Biology 5 (1): 85–93. https://doi.org/10.1055/s-2003-37979</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Matsunaga, K.K.S. & Smith, S.Y. (2021) Fossil palm reading: using fruits to reveal the deep roots of palm diversity. American Journal of Botany 108: 472–494. https://doi.org/10.1002/ajb2.1616</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Mei, Q., Chen, X., Xiang, L., Liu, Y., Su, Y., Gao, Y., Dai, W., Dong, P. & Chen, S. (2016) DNA Barcode for Identifying Folium Artemisiae argyi from Counterfeits. Biological and Pharmaceutical Bulletin 39 (9): 1531–1537. https://doi.org/10.1248/bpb.b16-00336</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miao, Y., Meng, Q., Fang, X., Yan, X., Wu, F. & Song<span lang="en-GB">,</span> C. (2011) Origin and development of Artemisia (Asteraceae) in Asia and its implications for the uplift history of the Tibetan Plateau: A review. Quaternary International 236: 3–12. https://doi.org/10.1016/j.quaint.2010.08.014</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Oberprieler, C., Himmelreich, S., Källersjö, M., Vallès, J., Watson, L.E. & Vogt, R. (2009) Anthemideae. In: Funk, V.A., Susanna, A., Stuessy, T.F. & Bayer, R.J. (Eds.) Systematics, Evolution and Biogeography of the Compositae. International Association Plant Taxonomy, Vienna. pp. 632–666.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Parham, J.F., Donoghue, P.C.J., Bell, C.J., Calway, T.D., Head, J.J., Holroyd, P.A., Inoue, J.G., Irmis, R.B., Joyce, W.G., Ksepka, D.T., Patané, J.S.L., Smith, N.D., Tarver, J.E., Van Tuinen, M., Yang, Z., Angielczyk, K.D., Greenwood, J., Hipsley, C.A., Jacobs, L., Makovicky, P.J., Müller, J., Krister, T.S., Jessica, M.T., Rachel, C.M.W. & Michael, J.B. (2012) Best practices for justifying fossil calibrations. Systematic Biology 61 (2): 346–359. https://doi.org/10.1093/sysbio/syr107</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sanderson, M.J. (2003) r8s: inferring absolute rates of molecular evolution and divergence times in the absence of a molecular clock. Bioinformatics 19: 301–302. https://doi.org/10.1093/bioinformatics/19.2.301</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sanz, M., Vilatersana, R., Hidalgo, O., Garcia-Jacas, N., Susanna, A., Schneeweiss, G. & Vallès, J. (2008) Molecular phylogeny and evolution of floral characters of Artemisia and allies (Anthemideae, Asteraceae): Evidence from nrDNA ETS and ITS sequences. Taxon 57: 1–13.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sauquet, H., Ho, S.Y.W., Gandolfo, M.A., Jordan, G.J., Wilf, P., Cantrill, D.J., Bayly, M.J., Bromham, L., Brown, G.K., Carpenter, R.J., Lee, D.M., Murphy, D.J., Sniderman, J.M.K. & Udovicic, F. (2012) Testing the impact of calibration on molecular divergence times using a fossil-rich group: the case of Nothofagus (Fagales). Systematic Biology 61: 289–313. https://doi.org/10.1093/sysbio/syr116</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sauquet, H. & Magallon, S. (2018) Key questions and challenges in angiosperm macroevolution. New Phytologist 219 (4): 1170–1187. https://doi.org/10.1111/nph.15104</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Shultz, L.M. (2006) Genus Artemisia (Asteraceae: Anthemideae). In: Flora of North America Editorial Committee (Ed.) The Flora of North America north of Mexico, vol. 19: Asterales. Oxford University Press, New York, pp. 503–534.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Smith, S.Y. (2013) The fossil record of non-commelinid monocots. In: Wilkin, P. & Mayo, S.J. (Eds.) Early events in monocot evolution. Cambridge University Press, Cambridge, pp. 29–59. https://doi.org/10.1017/CBO9781139002950.003</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Tkach, N.V., Hoffmann, M.H., Röser, M. & von Hagen, K.B. (2008) Temporal patterns of evolution in the Arctic explored in Artemisia L. (Asteraceae) lineages of different age. Plant Ecology & Diversity 1: 161–169. https://doi.org/10.1080/17550870802331912</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Tomescu, A.M.F. & Rothwell, G.W. (2022) Fossils and plant evolution: structural fingerprints and modularity in the evo-devo paradigm. EvoDevo 13: 8. https://doi.org/10.1186/s13227-022-00192-7</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vallès, J., Garcia, S., Hidalgo, O., Martín, J., Pellicer, J., Sanz, M. & Garnatje, T. (2011) Biology, genome evolution, biotechnological issues, and research including applied perspectives in Artemisia (Asteraceae). Advances in Botanical Research 60: 349–419. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Wood, H.M., Matzke, N.J., Gillespie, R.G. & Griswold, C.E. (2013) Treating fossils as terminal taxa in divergence time estimation reveals ancient vicariance patterns in the palpimanoid spiders. Systematic Biology 62: 264–284. https://doi.org/10.1093/sysbio/sys092</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Yang, Z. & Rannala, B. (2006) Bayesian estimation of species divergence times under a molecular clock using multiple fossil calibrations with soft bounds. Molecular Biology and Evolution 23: 212–226. https://doi.org/10.1093/molbev/msj024</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Zazula, G.D., Froese, D.G., Schweger, R.W., Mathewes, A.B., Beaudoin, A.M., Telka, C.R. & Harington, J.A. (2003) Westgate Palaeobotany: ice-age steppe vegetation in east Beringia. Nature 423. https://doi.org/10.1038/423603a</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Zazula, G.D., Froese, D.G., Elias, S.A., Kuzmina, S. & Mathewes, R.W. (2007) Arctic ground squirrels of the mammoth-steppe: paleoecology of middens from the last glaciation, Yukon Territory, Canada. Quaternary Science Reviews 26: 979–1003. https://doi.org/10.1016/j.quascirev.2006.12.006</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Zhang, C., Huang, C.H., Liu, M., Hu, Y., Panero, J.L., Luebert, F., Gao, T. & Ma, H. (2021) Phylotranscriptomic insights into Asteraceae diversity, polyploidy, and morphological innovation. Journal of Integrative Plant Biology 63 (7): 1273–1293. https://doi.org/10.1111/jipb.13078</span></span></span>