
Plant Diversity ›› 2026, Vol. 48 ›› Issue (02): 320-329.DOI: 10.1016/j.pld.2025.12.010
Hong Qiana, Michael Kesslerb, Shenhua Qianc,d
收稿日期:2025-11-04
修回日期:2025-12-13
出版日期:2026-03-25
发布日期:2026-05-19
通讯作者:
Hong Qian,E-mail:hong.qian@illinoisstatemuseum.org
基金资助:Hong Qiana, Michael Kesslerb, Shenhua Qianc,d
Received:2025-11-04
Revised:2025-12-13
Online:2026-03-25
Published:2026-05-19
Contact:
Hong Qian,E-mail:hong.qian@illinoisstatemuseum.org
Supported by:摘要: Liverworts are an important component of terrestrial ecosystems worldwide. They have adapted to and diversified in a wide variety of environments. Investigating variation in net diversification rate is a major goal of biogeographers and ecologists but such investigation is lacking for liverworts at a global scale. Here, we explore global geographic patterns of mean diversification rate (MDR) within genera of liverworts, which are one of the earliest lineages of the extant land plants, and its relationship with latitude, climatic conditions, and regional species richness. We collated species lists of liverworts for each of 390 geographic units (primarily countries, provinces or states) across the world. We related MDR to geographic and current and historical climatic variables, assessed the relative importance of different sets of climatic variables on MDR, and explored the effect of MDR on species richness after accounting for major climatic factors. We analyzed the data with correlation and regression analyses, and structural equation modeling approach. We find that MDR peaks at tropical latitudes and in humid and hot environments, and that at a global scale current climate, temperature-related variables, and climatic seasonality explained more variation in MDR than Quaternary climate change, precipitation-related variables, and climatic extremes, respectively. In addition, we find a positive relationship between MDR and liverwort species richness, with the latter being directly influenced more strongly by climate than by MDR. Most importantly, we find that tropical regions of high liverwort diversity also have high current diversification rates, suggesting ongoing niche occupation. The above-described patterns are similar between the New World and Old World and between the eastern and western parts of the Old World. Our study highlights that tip diversification rates provide a complementary aspect to understand the evolution of liverwort diversity to that recovered by studying phylogenetic diversity and species richness.
Hong Qian, Michael Kessler, Shenhua Qian. Diversification of liverworts across spatial and climatic gradients in the world[J]. Plant Diversity, 2026, 48(02): 320-329.
Hong Qian, Michael Kessler, Shenhua Qian. Diversification of liverworts across spatial and climatic gradients in the world[J]. Plant Diversity, 2026, 48(02): 320-329.
| [1] Adams, D.C., Berns, C.M., Kozak, K.H., et al., 2009. Are rates of species diversification correlated with rates of morphological evolution? Proc. Roy. Soc. B-Biol. Sci. 276, 2729-2738. [2] Alfaro, M.E., Santini, F., Brock, C.D., 2007. Do reefs drive diversification in marine teleosts? Evidence from the pufferfishes and their allies (Order Tetraodontiformes). Evolution 61, 2104-2126. [3] Allen, A.P., Gillooly, J.F., 2006. Assessing latitudinal gradients in speciation rates and biodiversity at the global scale. Ecol. Lett. 9, 947-954. [4] Blomberg, S.P., Garland, T.J., Ives, A.R., 2003. Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution, 57, 717-745. [5] Boucher, F.C., Quatela, A.-S., Ellis, A.G., et al., 2020. Diversification rate vs. diversification density: Decoupled consequences of plant height for diversification of Alooideae in time and space. PLoS One 15, e0233597. [6] Brinda, J.C., Atwood, J.J., 2023. Bryophyte Nomenclator. In: Banki, O., Roskov, Y., Doring, M., et al., Catalogue of Life Checklist (Jan 2023). https://doi.org/10.48580/dfqt-8zm. [7] Cadotte, M.W., Davies, T.J., 2016. Phylogenies in Ecology: A Guide to Concepts and Methods Princeton University Press, Princeton and Oxford. [8] Cai, L., Kreft, H., Taylor, A., et al., 2023. Climatic stability and geological history shape global centers of neo- and paleoendemism in seed plants. Proc. Natl. Acad. Sci. U.S.A. 120, e2300981120. [9] Collart, F., Wang, J., Patino, J., et al., 2021. Macroclimatic structuring of spatial phylogenetic turnover in liverworts. Ecography, 44, 1474-1485. [10] Cooney, C.R., Seddon, N., Tobias, J.A., 2016. Widespread correlations between climatic niche evolution and species diversification in birds. J. Anim. Ecol. 85, 869-878. [11] Dai, Z., Zhang, H.-W., Qian, H., et al., 2025. Effects of microclimates on species richness of epiphytic and non-epiphytic bryophytes along a subtropical elevational gradient in China. J. Biogeogr. 52, e15134. [12] Davies, T.J., Barraclough, T.G., Chase, M.W., et al., 2004. Darwin's abominable mystery: Insights from a supertree of the angiosperms. Proc. Natl Acad. Sci. U.S.A. 101, 1904-1909. [13] Dimitrov, D., Xu, X., Su, X., et al., 2023. Diversification of flowering plants in space and time. Nat. Commun. 14, 7609. [14] Eriksson, O., Bremer, B., 1992. Pollination systems, dispersal modes, life forms, and diversification rates in angiosperm families. Evolution 46, 258-266. [15] Gomez-Rodriguez, C., Baselga, A., Wiens, J.J., 2015. Is diversification rate related to climatic niche width? Glob. Ecol. Biogeogr. 24, 383-395. [16] Guo, Q., Cade, B.S., Dawson, W., et al., 2021. Latitudinal patterns of alien plant invasions. J. Biogeogr. 48, 253-262. [17] Harris, B. J., Clark, J. W., Schrempf, D., et al., 2022. Divergent evolutionary trajectories of bryophytes and tracheophytes from a complex common ancestor of land plants. Nat. Ecol. Evol. 6, 1634-1643. [18] Hughes, C., Eastwood, R., 2006. Island radiation on a continental scale: exceptional rates of plant diversification after uplift of the Andes. Proc. Natl. Acad. Sci. U.S.A. 103, 10334-10339. [19] Igea, J., Tanentzap, A.J., 2020. Angiosperm speciation cools down in the tropics. Ecol. Lett. 23, 692-700. [20] Jetz, W., Thomas, G.H., Joy, J.B., et al., 2012. The global diversity of birds in space and time. Nature 491, 444-448. [21] Kembel, S.W., Cowan, P.D., Helmus, M.R., et al., 2010. Picante: R tools for integrating phylogenies and ecology. Bioinformatics 26, 1463-1464. [22] Laenen, B., Shaw, B., Schneider, H., et al., 2014. Extant diversity of bryophytes emerged from successive post-Mesozoic diversification bursts. Nat. Comm. 5, 5134. [23] Laenen, B., Patino, J., Hagborg, A., et al., 2018. Evolutionary origin of the latitudinal diversity gradient in liverworts. Mol. Phylogenet. Evol. 127, 606-612. [24] Legendre, P., Legendre, L., 2012. Numerical Ecology, 3rd edn. Elsevier, Amsterdam. [25] Lombo-Sanchez, Y.J., Suarez-Contento, K.Y., Silva, M.P.P., et al., 2024. An assessment of liverwort richness, endemicity and conservation in a megadiverse country - Colombia. Biodivers. Conserv. 33, 3781-3797. [26] Magallon, S., Castillo, A., 2009. Angiosperm diversification through time. Am. J. Bot. 96, 349-365. [27] Magallon, S., Sanderson, M.J., 2001. Absolute diversification rates in angiosperm clades. Evolution 55, 1762-1780. [28] Maul, K., Gradstein, S.R., Quandt, D., et al., 2025. Temperature dependence of liverwort diversification reveals a cool origin and hot hotspots. Sci. Rep. 15, 3225. [29] Meyer, A.L.S., Roman-Palacios, C., Wiens, J.J., 2018. BAMM gives misleading rate estimates in simulated and empirical datasets. Evolution, 72, 2257-2266. [30] Meyer, A.L.S., Wiens, J.J., 2018. Estimating diversification rates for higher taxa: BAMM can give problematic estimates of rates and rate shifts. Evolution 72, 39-53. [31] Moore, B.R., Hohna, S., May, M.R., et al., 2016. Critically evaluating the theory and performance of Bayesian analysis of macroevolutionary mixtures. Proc. Natl. Acad. Sci. U.S.A. 113, 9569-9574. [32] Morlon, H., 2014. Phylogenetic approaches for studying diversification. Ecol. Lett. 17, 508-525. [33] Niklas, K.J., Tiffney, B.H., Knoll, A.H., 1983. Patterns in vascular land plant diversification. Nature 303, 614-616. [34] Pagel, M., 1999. Inferring the historical patterns of biological evolution. Nature, 401, 877-884. [35] Qian, H., 2025. Diversification rates in large-scale moss assemblages along latitudinal and climatic gradients across the world. Plant Divers., 47, 833-838. [36] Qian, H., Kessler, M., Zhang, J., et al., 2023. Global patterns and climatic determinants of phylogenetic structure of regional fern floras. New Phytol. 239, 415-428. [37] Qian, H., Kessler, M., 2024. Phylogenetic structure of liverwort assemblages along an elevational gradient in the tropical Andes: Geographic patterns and climatic drivers. Ecography 2024, e07434. [38] Qian, H., Dai, Z., Wang, J. 2024a. Strong evidence for latitudinal diversity gradient in mosses across the world. Plant Divers. 46, 537-541. [39] Qian, H., Kessler, M., Zhang, J., et al., 2024b. Evolutionary causes of global patterns of species richness in regional fern floras across the world. J. Biogeogr. 51, 1429-1437. [40] Qian, H., Qian, S., Zhang, J., et al., 2024c. Effects of climate and environmental heterogeneity on the phylogenetic structure of regional angiosperm floras worldwide. Nat. Comm. 15, 1079. [41] Qian, H., Vanderpoorten, A., Dai, Z., et al., 2024d. Spatial patterns and climatic drivers of phylogenetic structure of regional liverwort assemblages in China. Ann. Bot. 134, 427-436. [42] Qian, H., Kessler, M., Qian, S., 2025a. Global diversification rates of ferns across spatial and climatic gradients. Adv. Sci. 2025, e08106. [43] Qian, H., Qian, S., Kessler, M., et al., 2025b. Latitudinal patterns and macroclimatic drivers of phylogenetic structure in regional liverwort assemblages in North America. J. Biogeogr. 52, 274-283. [44] Quintero, I., Jetz, W., 2018. Global elevational diversity and diversification of birds. Nature 555, 246-250. [45] Rabosky, D.L., 2014. Automatic detection of key innovations, rate shifts, and diversity-dependence on phylogenetic trees. PLoS ONE 9, e89543. [46] Rabosky, D.L., Matute, D.R., 2013. Macroevolutionary speciation rates are decoupled from the evolution of intrinsic reproductive isolation in Drosophila and birds. Proc. Natl Acad. Sci. U.S.A. 110, 15354-15359. [47] Rabosky, D.L., Hurlbert, A.H., 2015. Species richness at continental scales is dominated by ecological limits. Am. Nat. 185, 572-583. [48] Ricklefs, R.E., 2004. A comprehensive framework for global patterns in biodiversity. Ecol. Lett. 7, 1-15. [49] Rohde, K. 1992. Latitudinal gradients in species diversity: the search for the primary cause. Oikos 65, 514-527. [50] Scholl, J.P., Wiens, J.J., 2016. Diversification rates and species richness across the Tree of Life. Proc. Roy. Soc. B-Biol. Sci. 283, 20161335. [51] Stadler, T., 2011. Mammalian phylogeny reveals recent diversification rate shifts. Proc. Natl. Acad. Sci. U.S.A. 108, 6187-6192. [52] Stanley, S.M., 1979. Macroevolution. W. H. Freeman, San Francisco. [53] Stephens, P.R., Farrell, M.J., Davies, T.J., et al., 2025. Global diversity patterns are explained by diversification rates and dispersal at ancient, not shallow, timescales. Syst. Biol. 74, 818-833. [54] Tedesco, P.A., Paradis, E., Leveque, C., et al., 2017. Explaining global-scale diversification patterns in actinopterygian fishes. J. Biogeogr. 44, 773-783. [55] Tietje, M., Antonelli, A., Baker, W.J., et al., 2022. Global variation in diversification rate and species richness are unlinked in plants. Proc. Natl Acad. Sci. U.S.A. 119, e2120662119. [56] Villarreal, J.C., Crandall-Stotler, B.J., Hart, M.L., et al., 2016. Divergence times and the evolution of morphological complexity in an early land plant lineage (Marchantiopsida) with a slow molecular rate. New Phytol. 209, 1734-1746. [57] Wang, J., Vanderpoorten, A., Hagborg, A., et al., 2017. Evidence for a latitudinal diversity gradient in liverworts and hornworts. J. Biogeogr. 44, 487-488. [58] Wang, J., Qian, H., Dai, Z., et al., 2025. Geographic and ecological effects on species richness of liverworts worldwide. Ecography 2025, e07277. [59] Weigand, A., Abrahamczyk, S., Aubin, I., et al., 2020. Global fern and lycophyte richness explained: How regional and local factors shape plot richness. J. Biogeogr. 47, 59-71. [60] Whittaker, R.H., 1975. Communities and Ecosystems. MacMillan Publishing Company, Inc., New York. [61] Wiens, J.J., 2011. The causes of species richness patterns across space, time, and clades and the role of "ecological limits". Q. Rev. Biol. 86, 75-96. [62] Wiens, J.J., 2015a. Explaining large-scale patterns of vertebrate diversity. Biol. Lett. 11, 20150506. [63] Wiens, J.J., 2015b. Faster diversification on land than sea helps explain global biodiversity patterns among habitats and animal phyla. Ecol. Lett. 18, 1234-1241. [64] Wilkinson, L., Hill, M., Welna, J.P., et al., 1992. SYSTAT for Windows: statistics. SYSTAT Inc., Evanston. [65] Wu, G., Wiens, J.J., 2022. The origins of climate-diversity relationships and richness patterns in Chinese plants. Ecol. Evol. 12, e9607. |
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