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Thermophilization and reshuffling of montane leaf beetle communities over a two-decade period
Alba Rodríguez-Pacheco, Victoria Formoso-Freire, M. Olalla Lorenzo-Carballa, Andrés Baselga, and Carola Gómez-Rodríguez
J Syst Evol 2024, 62 (2): 321-331.  
doi: 10.1111/jse.13054
Given the sensitivity of mountain biodiversity to human pressure, it is essential to quantify changes in montane biological communities and contrast them with expectations based on potential drivers of change. This need is particularly pressing for biological groups representing important but little-studied fractions of biodiversity, such as insects. We analyze the temporal changes (between 1998 and 2015) of leaf beetle communities in an altitudinal gradient in the Sierra de Ancares (NW Spain). Our results show temporal changes in the composition of local communities, with a tendency to assemblage thermophilization, as well as a homogenization of the spatial turnover pattern, mostly driven by an increased similarity between communities at the lower and intermediate altitudes. These temporal changes in community composition and in the spatial structure of biodiversity were associated with upward shifts of the upper altitudinal limit of warm-adapted species and with downward shifts of the lower altitudinal limit of cold-adapted species. While this upward shift is consistent with expectations of climate change effects, the observed downward shift suggests a land-use change effect. Our results point to the joint effect of multiple factors (climate and land-use change) behind temporal changes of these leaf beetle communities, which result in compositional reorganization and biotic homogenization, rather than a mere coherent displacement toward higher altitudes. More generally, we show that understanding temporal change of biodiversity requires assessing multiple community-level metrics (e.g., variation in assemblage composition and/or changes in spatial turnover) for the detection of tendencies among the species-specific signals (e.g., altitudinal range shifts).
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On the rocks: Biogeography and floristic identity of rocky ecosystems in eastern South America
Luísa Azevedo, Daniela Cristina Zappi, Daniela Melo Garcia de Oliveira, Leila Meyer, Eimear Nic Lughadha, Rosie Clegg, Leonardo Dias Meireles, Pablo Hendrigo Alves de Melo, R. Toby Pennington, and Danilo M. Neves
J Syst Evol 2024, 62 (2): 305-320.  
doi: 10.1111/jse.13052
The geodiversity of rocky ecosystems includes diverse plant communities with specific names, but their continental-scale floristic identity and the knowledge on the role of macroclimate remain patchy. Here, we assessed the identity of plant communities in eastern Brazil across multiple types of rocky landscapes and evaluated the relative importance of climatic variables in constraining floristic differentiation. We provided lists of diagnostic species and an assessment of the conservation status of the identified floristic groups. We compiled a data set of 151 sites (4498 species) from rocky ecosystems, including campos rupestres, campos de altitude, granitic-gneiss lowland inselbergs, and limestone outcrops. We used unsupervised clustering analysis followed by ANOSIM to assess floristic groups among sites. We performed a random forest variable selection to test whether the identified floristic groups occupy distinct climatic spaces. Six groups (lithobiomes) segregated floristically according to lithology and climate. Alongside campos de altitude and limestone outcrops, inselbergs were divided according to the biome in which they occur (Atlantic Forest or Caatinga), and campos rupestres were largely segregated according to their lithological matrix (ironstone or quartzitic). Plant communities of Caatinga inselbergs were more similar to limestone outcrops, while Atlantic Forest inselbergs communities resembled campos de altitude. The composition of plant communities on outcrops seems to be largely constrained by lithology, but climatic factors are also meaningful for sites with similar lithology. The current network of protected areas does not cover these unique ecosystems and their floristic heterogeneity, with Caatinga inselbergs and limestone outcrops being the least protected.
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Approaching inselberg biodiversity conservation through plant growth and dispersal strategies
Luiz Bondi, Camila M. Patreze, Ricardo P. Louro, and Laura Jane M. Santiago
J Syst Evol 2024, 62 (2): 291-304.  
doi: 10.1111/jse.13041
Climate change is promoting global declines in plant diversity, which are expected to be more critical in islands or island-like ecosystems due to environmental constraints and isolation. The species' vulnerability to climate change (VUL) depends on their ability to cope with changes or mitigate them. Therefore, we investigate the influence of growth and dispersal strategies of species from the Sugarloaf Rock Complex, Brazil, an island-like ecosystem, on their niche breadth (NB), long-dispersal (LD) capacity, and geographical range (GR). Besides, we evaluate the potential use of these strategies as indicators of species' VUL. We found that rock specialists exhibit narrower NB, lower LD capacity, and a more restricted GR when compared to other species. We also found that 63% of rock specialists are found in conservation red-lists and they are more vulnerable to climate change than woody plants. Conversely, self-dispersed plants are expected to be less vulnerable to climate change when compared to species with other dispersal mechanisms. Species vulnerable to climate change are 14 times more likely to be included in conservation red lists, and it might indicate that the species' VUL might also describe the species' vulnerability to other anthropogenic threats. Still, we suggest conservation attention on some species that are expected to be vulnerable to climate change but were not yet included in conservation red lists. We advocate for more efforts to ensure the conservation aspects of different functional groups in which inselbergs might not only offer isolation but also a refuge opportunity.
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The ghost of past climate acting on present-day plant diversity: Lessons from a climate-based delimitation of the tropical alpine ecosystem
Martha Kandziora, Juan M. Gorospe, Luciana Salomon, Diana L. A. Vásquez, Maria Pinilla Vargas, Filip Kolář, Petr Sklenář, and Roswitha Schmickl
J Syst Evol 2024, 62 (2): 275-290.  
doi: 10.1111/jse.13048
Habitat stability is important for maintaining biodiversity by preventing species extinction, but this stability is being challenged by climate change. The tropical alpine ecosystem is currently one of the ecosystems most threatened by global warming, and the flora close to the permanent snow line is at high risk of extinction. The tropical alpine ecosystem, found in South and Central America, Malesia and Papuasia, Africa, and Hawaii, is of relatively young evolutionary age, and it has been exposed to changing climates since its origin, particularly during the Pleistocene. Estimating habitat loss and gain between the Last Glacial Maximum (LGM) and the present allows us to relate current biodiversity to past changes in climate and habitat stability. In order to do so, (i) we developed a unifying climate-based delimitation of tropical alpine regions across continents, and (ii) we used this delimitation to assess the degree of habitat stability, that is, the overlap of suitable areas between the LGM and the present, in different tropical alpine regions. Finally, we discuss the link between habitat stability and tropical alpine plant diversity. Our climate-based delimitation approach can be easily applied to other ecosystems using our developed code, facilitating macro-comparative studies of habitat dynamics through time.
  
Phylogenomics of Puya (Bromeliaceae): Evolution in the Andean slopes and sky island ecosystems
Julián Aguirre-Santoro, Alejandro Zuluaga, Emma Stonesmyth, Julio Betancur, and Rachel S. Jabaily
J Syst Evol 2024, 62 (2): 257-274.  
doi: 10.1111/jse.13062
Phylogenomics enhances our understanding of plant radiations in the biodiverse Andes. Our study focuses on Puya, primarily Andean and a part of the Bromeliaceae family. Using a phylogenomic framework based on the Angiosperms353 probe set for 80 species, we explored Puya′s phenotypic evolution and biogeography. Divergence time analyses and ancestral area estimations suggested that Puya originated in Central Coastal Chile around 9 million years ago (Ma). Subsequently, it dispersed to the dry valleys of the Central Andes and Puna regions between 5–8 Ma, leading to the emergence of major lineages. Key events in the last 2–4 million years include the recolonization of Chilean lowlands and dispersal to the northern Andes via Peru's Jalcas, facilitating passage through the Huancabamba depression. This event gave rise to the high-elevation Northern Andes clade. Using phylogenetic comparative methods, we tested the hypothesis that adaptation to the Andes' island-like high-elevation ecosystems was facilitated by unique leaf and floral traits, life history, and inflorescence morphology. Our findings suggest correlations between inflorescence axis compression, protective bract overlap, and high-elevation living, potentially preventing reproductive structure freezing. Semelparity evolved exclusively at high elevations, although its precise adaptive value remains uncertain. Our framework offers insights into Andean evolution, highlighting that lineages adapted to life in dry ecosystems can easily transition to high-elevation biomes. It also underscores how the island-like nature of high-elevation ecosystems influences phenotypic evolution rates. Moreover, it opens avenues to explore genetic mechanisms underlying adaptation to extreme mountain conditions.
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Plant species richness hotspots and related drivers across spatial scales in small Mediterranean islands
Riccardo Testolin, Fabio Attorre, Vanessa Bruzzaniti, Riccardo Guarino, Borja Jiménez-Alfaro, Michele Lussu, Stefano Martellos, Michele Di Musciano, Salvatore Pasta, Francesco Maria Sabatini, Francesco Santi, Piero Zannini, and Alessandro Chiarucci
J Syst Evol 2024, 62 (2): 242-256.  
doi: 10.1111/jse.13034
Small islands represent a common feature in the Mediterranean and host a significant fraction of its biodiversity. However, the distribution of plant species richness across spatial scales—from local communities (alpha) to whole islands (gamma)—is largely unknown, and so is the influence of environmental, geographical, and topographical factors. By building upon classic biogeographic theory, we used the species–area relationship and about 4500 vegetation plots in 54 Central Mediterranean small islands to identify hotspots of plant species richness and the underlying spatial determinants across scales. To do so, we fitted and averaged eight species–area models on gamma and alpha richness against island area and plot size, respectively. Based on positive deviations from the fitted curves, we identified 12 islands as cross-scale hotspots. These islands encompassed around 70% of species and habitat richness, as well as almost 50% of the rarest species in the data set, while occupying less than 40% of the total island surface. By fitting generalized linear mixed models, we found that gamma richness was mainly explained by island area and was weakly related to mean annual temperature (positively) and annual precipitation (negatively). As for alpha richness, after accounting for the idiosyncratic effect of habitats and islands, plot size and gamma richness remained the only significant predictors, showing a positive relationship. This work contributes to the understanding of the patterns and drivers of plant diversity in Central Mediterranean small islands and outlines a useful methodology for the prioritization of conservation efforts.
  
A test of island plant syndromes using resource-use traits
Andrea C. Westerband, Tiffany M. Knight, and Kasey E. Barton
J Syst Evol 2024, 62 (2): 233-241.  
doi: 10.1111/jse.13032
Despite representing a fraction of the global terrestrial surface area, oceanic islands are disproportionately diverse in species, resulting from high rates of endemicity. Island plants are thought to share a unique phenotype—referred to as an island syndrome—which is thought to be driven by convergent evolution in response to selection by shared abiotic and biotic factors. One aspect of the island plant syndrome that has received relatively little research focus is that island plants are expected to have converged on conservative resource use associated with slow growth rates and weak competitive abilities. Here we tested whether native, woody Hawaiian plant species are phenotypically distinct—with more resource-conservative leaf traits—compared to a globally distributed sample of continental species. Using an archipelago-wide trait data set, we detected that on average, native Hawaiian species had lower leaf nutrient concentrations overall, and lower nutrient concentrations at high leaf mass per area, but no other phenotypic differences compared with continental plants. There was also considerable overlap in the trait spaces of native Hawaiian species and continental species. Our findings indicate that an island plant syndrome for leaf traits is not present in the Hawaiian flora, and that island species can demonstrate extensive variation in their resource-use strategies, on a scale that is comparable with that of continental species worldwide.
  
Early diversification dynamics in a highly successful insular plant taxon are consistent with the general dynamic model of oceanic island biogeography
Jay Edneil C. Olivar, Frank Hauenschild, Hannah J. Atkins, Gemma L.C. Bramley, and Alexandra N. Muellner-Riehl
J Syst Evol 2024, 62 (2): 215-232.  
doi: 10.1111/jse.13049
The general dynamic model (GDM) of oceanic island biogeography views oceanic islands predominantly as sinks rather than sources of dispersing lineages. To test this, we conducted a biogeographic analysis of a highly successful insular plant taxon, Cyrtandra, and inferred the directionality of dispersal and founder events throughout the four biogeographical units of the Indo-Australian Archipelago (IAA), namely Sunda, Wallacea, Philippines, and Sahul. Sunda was recovered as the major source area, followed by Wallacea, a system of oceanic islands. The relatively high number of events originating from Wallacea is attributed to its central location in the IAA and its complex geological history selecting for increased dispersibility. We also tested if diversification dynamics in Cyrtandra follow predictions of adaptive radiation, which is the dominant process as per the GDM. Diversification dynamics of dispersing lineages of Cyrtandra in the Southeast Asian grade showed early bursts followed by a plateau, which is consistent with adaptive radiation. We did not detect signals of diversity-dependent diversification, and this is attributed to Southeast Asian cyrtandras occupying various niche spaces, evident by their wide morphological range in habit and floral characters. The Pacific clade, which arrived at the immaturity phase of the Pacific Islands, showed diversification dynamics predicted by the island immaturity speciation pulse model (IISP), wherein rates increase exponentially, and their morphological range is controlled by the least action effect favoring woodiness and fleshy fruits. Our study provides a first step toward a framework for investigating diversification dynamics as predicted by the GDM in highly successful insular taxa.
  
Slowing taxon cycle can explain biodiversity patterns on islands: Insights into the biogeography of the tropical South Pacific from molecular data
Gunnar Keppel, Francis J. Nge, and Thomas Ibanez
J Syst Evol 2024, 62 (2): 201-214.  
doi: 10.1111/jse.13026
Islands in the tropical Pacific Ocean are renowned for high biodiversity and endemism despite having relatively small landmasses. However, our knowledge of how this biodiversity is formed remains limited. The taxon cycle, where well-dispersed, earlier colonizers become displaced from coastal to inland habitats by new waves of colonizers, producing isolated, range-restricted species, has been proposed to explain current biodiversity patterns. Here, we integrate the outcomes of phylogenetic studies in the region to investigate the sources, age, number of colonizations, and diversification of 16 archipelagos in the tropical and subtropical South Pacific. We then evaluate whether the results support the taxon cycle as a plausible mechanism for these observations. We find that most species in the Pacific arrived less than 5 Mya from geographically close sources, suggesting that colonization by new taxa is a frequent and ongoing process. Therefore, our findings are broadly consistent with the theory of the Taxon Cycle, which posits that ongoing colonization results in the gradual displacement of established lineages. Only the oldest archipelagos, New Caledonia and Fiji, do not conform to this trend, having proportionally less recent colonization events, suggesting that the taxon cycle may slow on older islands. This conclusion is further validated by New Caledonia having lower diversification rate estimates than younger islands. We found that diversification rates across archipelagos are negatively correlated with area and age. Therefore, a taxon cycle that slows with island age appears to be a suitable concept for understanding the dynamic nature and biodiversity patterns of the Pacific Islands.
  
Past, present, and future of mountain and island systems
Alexandra N. Muellner-Riehl, Fabien Anthelme, and Thomas Ibanez
J Syst Evol 2024, 62 (2): 195-200.  
doi: 10.1111/jse.13073
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Issue Information
J Syst Evol 2024, 62 (2): 194-194.  
doi: 10.1111/jse.12972
  
List of Reviewers
J Syst Evol 2024, 62 (1): 193-194.  
doi: 10.1111/jse.13051
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Genetic admixture of Chinese Tajik people inferred from genome-wide array genotyping and mitochondrial genome sequencing
Jing Zhao, Qiao Wu, Xin-Hong Bai, Edward Allen, Meng-Ge Wang, Guang-Lin He, Jian-Xin Guo, Xiao-Min Yang, Jian-Xue Xiong, Zi-Xi Jiang, Xiao-Yan Ji, Hui Wang, Jing-Ze Tan, Shao-Qing Wen, and Chuan-Chao Wang
J Syst Evol 2024, 62 (1): 181-192.  
doi: 10.1111/jse.12957
Chinese Tajiks are an Indo-Iranian-speaking population in Xinjiang, northwest China. Although the complex demographic history has been characterized, the ancestral sources and genetic admixture of Indo-Iranian-speaking groups in this region remain poorly understood. We here provide the genome-wide genotyping data for over 700?000 single-nucleotide polymorphisms (SNPs) and mtDNA multiplex sequencing data in 64 Chinese male Tajik individuals from two dialect groups, Wakhi and Selekur. We applied principal component analysis (PCA), ADMIXTURE, f-statistics, treemix, qpWave/qpAdm, Admixture-induced Linkage Disequilibrium for Evolutionary Relationships (ALDER), and Fst analyses to infer a fine-scale population genetic structure and admixture history. Our results reveal that Chinese Tajiks showed the closest affinity and similar genetic admixture pattern with ancient Xinjiang populations, especially Xinjiang samples in the historical era. Chinese Tajiks also have gene flow from European and Neolithic Iran farmers-related populations. We observed a genetic substructure in the two Tajik dialect groups. The Selekur-speaking group who lived in the county had more gene flow from East Asians than Wakhi-speaking people who inhabited the village. These results document the population movements contributed to the influx of diverse ancestries in the Xinjiang region.
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Exceptionally well-preserved seed cones of a new fossil species of hemlock, Tsuga weichangensis sp. nov. (Pinaceae), from the Lower Miocene of Hebei Province, North China
Ya Li, Carole T. Gee, Zhen-Zhen Tan, Yan-Bin Zhu, Tie-Mei Yi, and Cheng-Sen Li
J Syst Evol 2024, 62 (1): 164-180.  
doi: 10.1111/jse.12952
Tsuga (hemlock) is a small genus of 10 extant species in the Pinaceae, with a disjunct distribution in East Asia and eastern and western North America. Reliable species-level identification of Tsuga fossils depends on the discovery of seed cones with intact bracts, but such cones are rare in the fossil record. Here we describe a new fossil species of hemlock as T. weichangensis sp. nov. based on exquisitely preserved seed cones with nearly complete bracts from the Lower Miocene of Weichang, Hebei Province, North China. This fossil species displays a mosaic of characters between Tsuga and Nothotsuga. The well-developed and slightly exserted bract scales of T. weichangensis are reminiscent of Nothotsuga, but other characters, such as nonleaved peduncles and tongue-shaped bract scales, in addition to monosaccate pollen found at the same fossil locality, suggest an affinity closer to Tsuga. Cladistic analysis based on 15 morphological characters and a molecular backbone constraint supports the assignment of these fossil cones to Tsuga rather than Nothotsuga, and places the fossil species of T. weichangensis in an unresolved polytomy within the genus Tsuga. The occurrence of Tsuga seed cone fossils indicate the paleoclimate in the Miocene of Weichang was warmer and more humid than today's climate, which is consistent with the paleoclimate reconstructed by paleopalynology.
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First recognition of the extinct eudicot genus Palibinia in North America: Leaves and fruits of Palibinia comptonifolia (R.W.Br.) comb. nov. from the Eocene of Utah and Colorado, USA
Steven R. Manchester, Walter S. Judd, and Tatiana Kodrul
J Syst Evol 2024, 62 (1): 149-163.  
doi: 10.1111/jse.13011
Newly investigated leafy twigs bearing axillary fruits from the Eocene Parachute Creek Member of the Green River Formation in eastern Utah, USA, have provided more information on the species previously attributed to the Proteaceae as Banksia comptonifolia R.W.Br. The leaves are simple, estipulate with short petioles, and elongate laminae with prominent angular nonglandular teeth. The laminae have a thick midvein and pinnate craspedodromous secondaries, and are distinctive in the presence of a thick, often coalified, marginal rim. Vegetative and reproductive buds occur in the axils of the leaves. These features indicate that the species belongs to Palibinia Korovin—an extinct Eudicot genus previously known only from the Paleogene of Asia and Europe. Small pedicellate ovoid fruits 1.5–2.2?mm wide are borne in fascicles of three and are seen to be capsules with four apical valves. Despite the specific epithet referring to similarity of the foliage to that of Comptonia (Myricaceae), the fasciculate inflorescence organization with axillary flowers is quite distinct from the catkins characteristic of that family. Assignment to Banksia or other Proteaceae with complex inflorescences and follicular fruits is also problematic. Additionally, MacGinitie′s transfer of the species to Vauquelinia of the Rosaceae is contradicted by the lack of stipule scars on the twig and by differences in leaf venation and floral morphology. We transfer the species to Palibinia comptonifolia (R.W.Br.) comb. nov., but its familial affinity within the Pentapetalae remains uncertain. This new occurrence augments records from the Paleogene of Turkmenistan, Kazakhstan, China, England, and Germany.
  
Functional significance of asymmetrical retention of parental alleles in a hybrid pine species complex
Chang Qu, Hong-Na Kao, Hui Xu, Bao-Sheng Wang, Zhi-Ling Yang, Qi Yang, Gui-Feng Liu, Xiao-Ru Wang, Yan-Jing Liu, and Qing-Yin Zeng
J Syst Evol 2024, 62 (1): 135-148.  
doi: 10.1111/jse.12953
Hybrid genomes usually harbor asymmetrical parental contributions. However, it is challenging to infer the functional significance of asymmetrical retention of parental alleles in hybrid populations of conifer trees. Here we investigated the diversity in the glutathione S-transferase (GST) gene family in a hybrid pine Pinus densata and its parents (Pinus tabuliformis and Pinus yunnanensis). Plant GSTs play major roles in protecting plants against biotic and abiotic stresses. In this study, 19 orthologous groups of GST genes were identified and cloned from these three species. We examined their expression in different tissues, and then purified the corresponding proteins to characterize their enzymatic activities and specificities toward different substrates. We found that among the 19 GST orthologous groups, divergence in gene expression and in enzymatic activities toward different substrates was prevalent. P. densata preferentially retained P. yunnanensis-like GSTs for 17 out of the 19 gene loci. We determined the first GST crystal structure from conifer species at a resolution of 2.19??. Based on this structure, we performed site-directed mutagenesis to replace amino acid residuals in different wild-types of GSTs to understand their functional impacts. Reciprocal replacement of amino acid residuals in native GSTs of P. densata and P. tabuliformis demonstrated significant changes in enzyme functions and identified key sites controlling GSTs activities. This study illustrates an approach to evaluating the functional significance of sequence variations in conifer genomes. Our study also sheds light on plausible mechanisms for controlling the selective retention of parental alleles in the P. densata genome.
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Phylogeography and introgression between Pinus kesiya and Pinus yunnanensis in Southeast Asia
Jie Gao, Kyle W. Tomlinson, Wei Zhao, Baosheng Wang, Ralph Sedricke Lapuz, Jing-Xin Liu, Bonifacio O. Pasion, Bach T. Hai, Souvick Chanthayod, Jin Chen, and Xiao-Ru Wang
J Syst Evol 2024, 62 (1): 120-134.  
doi: 10.1111/jse.12949
Southeast Asia (SEA) has seen strong climatic oscillations and fluctuations in sea levels during the Quaternary. The impact of past climate changes on the evolution and distribution of local flora in SEA is still poorly understood. Here we aim to infer how the Quaternary climate change affects the evolutionary process and range shifts in two pine species. We investigated the population genetic structure and diversity using cytoplasmic DNA markers, and performed ecological niche modeling to reconstruct the species past distribution and to project range shift under future climates. We found substantial gene flow across the continuous distribution of the subtropical Pinus yunnanensis. In contrast, the tropical Pinus kesiya showed a strong population structure in accordance with its disjunct distribution across montane islands in Indochina and the Philippines. A broad hybrid zone of the two species occurs in southern Yunnan. Asymmetric introgression from the two species was detected in this zone with dominant mitochondrial gene flow from P. yunnanensis and chloroplast gene flow from P. kesiya. The observed population structure suggests a typical postglaciation expansion in P. yunnanensis, and a glacial expansion and interglacial contraction in P. kesiya. Ecological niche modeling supports the inferred demographic history and predicts a decrease in range size for P. kesiya under future climates. Our results suggest that tropical pine species in SEA have undergone evolutionary trajectories different from high latitude species related to their Quaternary climate histories. We also illustrate the need for urgent conservation actions in this fragmented landscape.
  
The origin and dispersal of Firmiana danxiaensis among isolated specific landscapes
Su-Fang Chen, Wan-Yi Zhao, Yan-Shuang Huang, Kai-Kai Meng, Kang-You Huang, Rong-Feng Hou, Xiao-Ying Luo, Zai-Xiong Chen, Yuan-Qiu Li, Ren-Chao Zhou, Wen-Bo Liao, and Qiang Fan
J Syst Evol 2024, 62 (1): 102-119.  
doi: 10.1111/jse.12954
China has the most numerous Danxia and Karst landscapes, which serve as special terrestrial islands harboring ample endemic species, though how did these endemic species spread among those isolated sites is still an unresolved issue. To address this question, we explored the phylogeographical structure and demographic history of Firmiana danxiaensis, a tree species endemic to Danxia and Karst landscapes. We collected 295 samples (28 populations) of F. danxiaensis. Plastid genomes were assembled for 25 representative samples. Sanger sequencing of four plastid regions and restriction-site-associated DNA sequencing were performed on the 28 populations. The phylogenetic tree constructed from plastid genomes and restriction site-associated DNA sequencing (RAD-seq) data supported that F. danxiaensis originated from Mount Danxia and Nanxiong Basin, spread to Karst landscapes near Yingde City, and then back to Danxia Mountain and the Nanxiong Basin. In the Nanxiong Basin, the latter arrivals captured the plastid of the former. Population analyses revealed strong population structure among and within Danxia and Karst landscapes, possibly due to low seed and pollen dispersal abilities of the species. The demographic and ecological niche modeling approaches suggested that F. danxiaensis have widely occurred in the southeast of China during the last glacial period, and later retreated to the cliffs of Danxia and Karst landscapes due to temperature rising and competition failure. The declining of the effective population size of the species throughout the postglacial period suggested that global warming, agriculture, and industrial civilizations could have affected the survival of this species, and more measures should be taken to conserve these species.
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Diversity, taxonomy, and history of the tropical fern genus Didymoglossum Desv. (Hymenophyllaceae, Polypodiidae) in Africa
Jean-Yves Dubuisson, Adèle Nivart, Ehoarn Bidault, Vincent Deblauwe, Vincent Droissart, Narcisse G. Kamdem, Germinal Rouhan, Atsushi Ebihara, and Timothée le Péchon
J Syst Evol 2024, 62 (1): 84-101.  
doi: 10.1111/jse.12951
The fern genus Didymoglossum (Hymenophyllaceae) is not so diverse in Africa with seven species at most. However, its local taxonomy is surprisingly still strongly debated, in particular within the Didymoglossum erosum complex interpreted either as a single polymorphic species or as a group of at least three distinct but morphologically very close taxa (D. erosum, Didymoglossum chamaedrys, and Didymoglossum benlii). Investigating these taxonomic issues and more generally the diversity of the genus in Africa and its origin, we conducted a complete anatomo–morphological analysis coupled with a molecular phylogenetic work based on rbcL. Our results support the recognition of all seven species, including Didymoglossum robinsonii that is likely distinct from the Neotropical Didymoglossum reptans to which the African populations were traditionally attributed. We here propose new characters and a novel key to distinguish the seven African species which also include Didymoglossum ballardianum, Didymoglossum lenormandii, and Didymoglossum liberiense. Once the taxonomy is clarified with respect to the distinct evolutionary lineages evidenced, the biogeographic history of the genus in Africa is discussed based on a divergence time estimation and the reconstruction of the ancestral geographic areas. These analyses reveal a Mesozoic (Cretaceous) vicariance event within Didymoglossum which is the second one hypothesized for the family Hymenophyllaceae.
  
Species boundaries and conservation implications of Cinnamomum japonicum, an endangered plant in China
Han-Yang Lin, Yue Yang, Wen-Hao Li, Yu-Xin Luo, Xiao-Hua Bai, Tetsuo Ohi-Toma, Changkyun Kim, Joo-Hwan Kim, and Yun-Peng Zhao
J Syst Evol 2024, 62 (1): 73-83.  
doi: 10.1111/jse.12950
Clear species boundaries are crucial for plans and actions on biodiversity conservation. However, morphological similarities among allied species can result in taxonomic difficulties, thus impeding conservation efforts. In China, Cinnamomum japonicum Siebold is a well-known endangered plant, yet suffers from longstanding taxonomic issues. Here, we explicitly evaluate whether C. japonicum, C. chenii, and C. chekiangense are the same phylogenetic species on the basis of a multi-individual sampling strategy. We identified three sets of low-copy orthologous genes from 19 Lauraceae taxa for phylogenetic inferences. Both the concatenation and coalescent-based phylogenies supported that C. chenii individuals were embedded in the C. japonicum clade, indicating these two taxa are conspecific. Meanwhile, C. chekiangense accessions formed a monophyly which was not sister to C. japonicum. This result, together with the morphological differences that the leaves of C. japonicum are glabrous with a faveolate pattern of venation while those of C. chekiangense have trichomes and inevident lateral veins, led us to consider both as two distinct species. Based on 17 728 neutral single nucleotide polymorphisms (SNPs), the ADMIXTURE analysis suggested that the Chinese C. japonicum populations in Zhoushan Archipelago (=C. chenii) were genetically differentiated from the Japanese and Korean ones. Furthermore, ecological niche modeling predicted that the present distribution area of Chinese C. japonicum is likely to be unsuitable under global warming scenarios. Together with its limited distribution and genetic uniqueness, we recommend that Chinese C. japonicum deserves conservation priorities.
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Resolving relationships despite past hybridization in Aliciella subsection Subnuda (Polemoniaceae)
Theresa C. Saunders, J. Mark Porter, and Leigh A. Johnson
J Syst Evol 2024, 62 (1): 55-72.  
doi: 10.1111/jse.13010
Phylogenetics is crucial in the study of evolutionary processes and events transpiring in the course of species diversification. Phylogenetic studies within kingdom Plantae often reveal hybridization and introgression. Here, we study a subsection rife with historic hybridization and discuss the impacts of such processes on evolutionary trajectories. Aliciella subsection Subnuda comprises seven species of herbaceous plants occurring in Utah, the Navajo Nation, and the Four Corners region of North America. Previous molecular and morphological work left relationships in the subsection unresolved. Here, we use comparative DNA sequencing of nuclear ITS and chloroplast DNA regions and genome-wide RAD-seq data to clarify phylogenetic relationships and examine the role of hybridization in the subsection. We construct haplotype and nucleotype networks from chloroplast and nuclear ITS sequence matrices and compare nuclear and chloroplast phylogenies to identify multiple putative chloroplast capture events. The RAD-seq maximum likelihood phylogeny and multispecies coalescent species tree robustly resolve relationships between six species-level clades. We use STRUCTURE and HyDe on the RAD-seq data to evaluate the influence of hybridization within the subsection. The HyDe results suggest that hybridization has occurred among all species in the subsection at some point in their history. Cytonuclear discordance reveals historic chloroplast capture, and we discuss potential causes of the observed discordance. Our study robustly resolves relationships in Aliciella subsection Subnuda and provides a framework for discussing its speciation despite a history of hybridization and introgression.
  
Phylogenomic insights into the reticulate evolution of Camellia sect. Paracamellia Sealy (Theaceae)
Sheng-Yuan Qin, Kai Chen, Wen-Ju Zhang, Xiao-Guo Xiang, Zheng-Yu Zuo, Cen Guo, Yao Zhao, Lin-Feng Li, Yu-Guo Wang, Zhi-Ping Song, Ji Yang, Xiao-Qiang Yang, Jian Zhang, Wei-Tao Jin, Qiang Wen, Song-Zi Zhao, Jia-Kuan Chen, De-Zhu Li, and Jun Rong
J Syst Evol 2024, 62 (1): 38-54.  
doi: 10.1111/jse.12948
Polyploids are common in Camellia sect. Paracamellia, which contain many important oil crop species. However, their complex evolutionary history is largely unclear. In this study, 22 transcriptomes and 19 plastomes of related species of Camellia were sequenced and assembled, providing the most completed taxa sampling of Camellia sect. Oleifera and C. sect. Paracamellia. Phylogenetic trees were reconstructed with predicted single-copy nuclear genes and plastomes. Phylogenetic trees with nuclear genes demonstrated that C. sect. Oleifera should be merged into C. sect. Paracamellia. Cytonuclear discordance and network analyses suggested hybridizations among polyploid species and relatives. The divergence of major clades in C. sect. Paracamellia was dated to be during the middle to late Miocene from the ancestral Lingnan region, and a rapid diversification during the Quaternary was found, probably through hybridization and polyploidization. The tetraploid Camellia meiocarpa Hu may have originated from hybridization between closely related diploid species. The hexaploid Camellia oleifera C. Abel probably originated from hybridization between closely related diploid and tetraploid (e.g., C. meiocarpa) species. The octoploid Camellia vietnamensis T. C. Huang ex Hu could have originated from hybridization between hexaploid C. oleifera and the closely related diploid species. Hybridization and polyploidization played an important role in generating the rich variation of important fruit traits, especially increased fruit size in polyploid species.
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Time-calibrated phylogeny of Daphne (Thymelaeaceae): Pre-mediterranean temporal origin of the sclerophyllous Daphne gnidium
Cristina Ramos-Capón, Pablo Vargas, Fernando Pomeda-Gutiérrez, and Sara Martín-Hernanz
J Syst Evol 2024, 62 (1): 20-37.  
doi: 10.1111/jse.13018
The sclerophyllous syndrome is characterized by well-defined traits (evergreen coriaceous leaves, inconspicuous flowers, and fleshy fruits). It has been hypothesized that lineages displaying the sclerophyllous syndrome show lower speciation rates than non-sclerophyllous lineages after the establishment of the mediterranean climate. Daphne gnidium displays sclerophyllous traits and some differentiation into three subspecies (gnidium, mauritanica, maritima), but the spatio-temporal origin of this taxonomic group is unknown due to the lack of a time-calibrated phylogeny of the whole genus. Here, we inferred phylogenetic relationships and divergence times of Daphne (32 species, 238 samples) and other genera of Thymelaeaceae (16 genera, 38 species, 34 samples) using the internal transcribed spacer (ITS), which revealed that the current circumscription of Daphne is profoundly polyphyletic because some species are nested within the genera Wikstroemia and Edgeworthia. In contrast, D. gnidium formed a well-supported clade as recognized in taxonomic accounts (subgenus Spachia). We found morphological and phylogenetic support for Daphne mauritanica as a monophyletic lineage sister to D. gnidium. Divergence between D. gnidium and D. mauritanica appears to have predated the establishment of seasonally dry conditions, which supports a pre-mediterranean temporal origin of the lineage. A phylogeographical analysis within D. gnidium based on 66 nrDNA (ITS) and 84 cpDNA (rps16, trnV) sequences agreed with the low differentiation of the species in the Pleistocene despite its large distribution range. Altogether, D. gnidium illustrates one more example of the sclerophyllous syndrome with no speciation after the onset of the mediterranean climate.
  
RAD-seq data provide new insights into biogeography, diversity anomaly, and species delimitation in eastern Asian–North American disjunct clade Benthamidia of Cornus (Cornaceae)
Zhi-Yuan Du, Jin Cheng, and Qiu-Yun (Jenny) Xiang
J Syst Evol 2024, 62 (1): 1-19.  
doi: 10.1111/jse.13037
The big-bracted dogwood clade Benthamidia of Cornus is a typical example of the well-known eastern Asia (EA) and North America (NA) floristic disjunction, with greater species diversity in EA than in NA. The lineage provides an opportunity to explore factors contributing to the plant diversity unevenness between EA and NA and test hypotheses on the origin of disjunct distribution from a phylogenetic perspective. We generated RAD-seq data, conducted phylogenomic and biogeographic analyses for the clade with sampling of all species (9) and subspecies (10) currently recognized in floras. We also assessed species delineation and calculated phylogenetic diversity to evaluate the diversity unevenness between EA and NA. Finally, we examined variation of diversification rates and ecological niches on the phylogeny to explore potential causes underlying the observed diversity pattern. Our results revealed phylogenetic relationships congruent with previous studies and suggested a trans-Beringian ancestral distribution of the clade Benthamidia in the mid-Oligocene, dispersal from Mexico to eastern United States in the mid-Miocene, and early diversification of the EA clade in SW China. Our results also confirmed greater phylogenetic diversity and diversification rate of the EA clade. Species delimitation analysis suggested 17 species in the clade Benthamidia, including all recognized subspecies. By integrating the results of molecular data with morphology, we proposed to retain the subspecies without changing their ranks. Our data suggested increased diversification rate in EA as an intrinsic factor explaining the greater species diversity in the region driven mainly by biogeographic isolation and partially by niche divergence.
  
Issue Information
J Syst Evol 2024, 62 (1): 0-4.  
doi: 10.1111/jse.13050
  
A new red alga preserved with possible reproductive bodies from the 518-million-year-old Qingjiang biota
Rui-Yun Li, Lin-Hao Cui, Dong-Jing Fu, and Xing-Liang Zhang
J Syst Evol 2023, 61 (6): 1091-1101.  
doi: 10.1111/jse.12942
Macroalgae have been a key ecological component of marine ecosystems since the Proterozoic period and are common fossil forms in Cambrian Burgess Shale-type Lagerstätten. However, in most cases, it is difficult to place these early fossil algae into modern groups because little distinctive morphology is preserved. Here, we describe a new form of macroalgae, Qingjiangthallus cystocarpium gen. & sp. nov., from the Qingjiang biota of South China. The new taxon is represented by 546 specimens remarkably preserved with characteristics that allow a phylogenetic placement into crown groups of red algae. Centimeter-sized thalli resemble members of the extant Rhodymeniophycidae (a subclass of the class Florideophyceae), and hence suggest a florideophycean affinity, which indicates that ahnfeltiophycidaen and rhodymeniophycidaen algae may have diverged at least 518Ma, accordant with estimations of molecular studies. The presence of possible cystocarps on Qingjiangthallus thalli suggests that evolutionary innovation of a triphasic life cycle in red algae may have occurred no later than the Early Cambrian. The branching patterns and branch width of Qingjiangthallus are consistent with the coarsely dichotomously branched morphogroup, which was previously present in the Ediacaran, Ordovician, and afterward, but absent in the Cambrian.
  
First fossil evidence of samaras of Ventilago Gaertn. (Rhamnaceae) from India and its implications
Taposhi Hazra, Sampa Kundu, Subir Bera, Tapan Chakraborty, and Mahasin Ali Khan
J Syst Evol 2023, 61 (6): 1079-1090.  
doi: 10.1111/jse.12936
The Ventilago Gaertn. (Rhamnaceae) is widely distributed in pantropical areas of Africa, Asia, and Australia. However, fossil records of this taxon are sparse, which limits understanding of the evolution and biogeographic history of the genus. In the present study, we report and describe two new fossil species of Ventilago, V. siwalika sp. nov. from the Miocene sediments of Himachal Pradesh, western Himalaya, and V. pliocenica sp. nov. from the Pliocene sediments of Jharkhand, eastern India based on single-winged samaras. Ventilago pliocenica is characterized by a prominent midvein, obtuse to sub-round apex with mucronate tip, longitudinal secondary veins extending the full length of the fruit, and reticulate nature of higher-order veins, the presence of equatorial rim, the hypanthium, and short pedicel. On the other hand, V. siwalika is characterized by a prominent midvein, obtuse to sub-round apex with mucronate tip, longitudinal secondary veins extending the full length of the fruit, and reticulate nature of higher-order veins. Our discovery represents the first unambiguous fossil record of single-winged samara of Ventilago from India and provides valuable insights into the evolution of this genus. In this paper, we also review its biogeographic history and add new information to understand its hypothetical migration route. Present and earlier records of Ventilago also suggest that this genus was a common forest element during Neogene (Miocene time) in Asia.
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Herbaceous eudicot Fairlingtonia from the Lower Cretaceous of Jiuquan Basin, Northwest China and its radiation in Laurasia
Bao-Xia Du, Ming-Zhen Zhang, Jing Zhang, Ai-Jing Li, Shao-Hua Lin, Guo-Rong Ma, and Jian-Guo Hui
J Syst Evol 2023, 61 (6): 1065-1078.  
doi: 10.1111/jse.12934
Eudicots exhibit diverse life forms and occupy a wide variety of habitats in the modern terrestrial ecosystems, and the diversification began during the Early Cretaceous; however, few Early Cretaceous fossils are preserved as multiorgan whole plants that can provide sufficient morphological characters for detailed phylogenetic assessment. Here, Fairlingtonia microgyna sp. nov. is reported from the upper Lower Cretaceous of Zhonggou Formation, Hanxia Section, Yumen City, western Gansu Province, Northwest China. The specimen is exceptionally preserved as multiorgan whole plant fossil with fibrous adventitious roots, simple and deeply dissected leaves, solitary and dehiscent capsular fruits attached to the creeping stems. As such, it was interpreted as a herbaceous eudicot. Phylogenetic analyses support a placement within the Papaveraceae, most likely in Papaveroideae, but there are obvious differences in morphological characteristics, which cannot confirm the systematic position within the Papaveraceae. Fossil records of Fairlingtonia from contemporaneous deposits (late Aptian to early Albian) in Northwest China and eastern North America provide direct evidence of the geographical radiation of Fairlingtonia on Laurasia. And the morphological characters of F. microgyna, including creeping leafy branches, fibrous adventitious roots, small and deeply dissected leaves as well as small capsular fruits with tiny seeds probably indicate that it was a colonizer of lake-shore environments under wet and bright conditions and possessed fast-growing and rapid propagation habitats, which allowed it to expand its geographic range with both sexual and asexual reproduction.
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Ancient genome of Empress Ashina reveals the Northeast Asian origin of Göktürk Khanate
Xiao-Min Yang, Hai-Liang Meng, Jian-Lin Zhang, Yao Yu, Edward Allen, Zi-Yang Xia, Kong-Yang Zhu, Pan-Xin Du, Xiao-Ying Ren, Jian-Xue Xiong, Xiao-Yu Lu, Yi Ding, Sheng Han, Wei-Peng Liu, Li Jin, Chuan-Chao Wang, and Shao-Qing Wen
J Syst Evol 2023, 61 (6): 1056-1064.  
doi: 10.1111/jse.12938
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Mitochondrial genome phylogeny reveals the deep-time origin of Gomphomastacinae (Orthoptera: Eumastacidae) and its alpine genera in China
Jun-Jie Ge, Hong-Fei Ying, Sheng-Quan Xu, and Hua-Teng Huang
J Syst Evol 2023, 61 (6): 1047-1055.  
doi: 10.1111/jse.12943
Gomphomastacinae is a grasshopper subfamily in Eumastacidae, with a morphology and distribution distinct from other subfamilies. The alpine genera of Gomphomastacinae that inhabit the Qinghai–Tibet Plateau in China show unique characteristics adapted to high-altitude life. However, their phylogenetic position and biogeographic history remain controversial. Thus, to determine the diversification history of these alpine genera and the origin of the subfamily, we obtained mitochondrial genome sequences from all seven Gomphomastacinae genera distributed in China. The reconstructed phylogeny was well supported and confirmed the phylogenetic position of Gomphomastacinae within Eumastacidae. Time calibration revealed a deep-time origin of the subfamily dating back to the Cretaceous period, and the diversification among alpine genera was also an ancient pre-Miocene event (30–50 Ma). Based on phylogeny and time estimates, the most likely biogeographic scenario is that Gomphomastacinae originated from an ancestral lineage that lived in East Gondwana and dispersed to Central and Western Asia through India. Subsequently, the alpine genera likely diverged along with the uplift of the Qinghai–Tibet Plateau and survived drastic climate change by in situ adaptation to high-altitude dwellings.
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