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Article

Understanding the Diversity and Distribution of Lycophytes and Ferns in Northeast China Based on Historical Records

by
Yan Li
1,2,3,4,
Shuai Yu
1,2,4,
Sheng Xu
1,2,4,
Lian Jia
5 and
Xingyuan He
1,2,3,4,6,*
1
CAS Key Laboratory of Forest Ecology and Silviculture, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, China
2
Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, China
3
Arboretum of Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, China
4
Liaoning Shenyang Urban Ecosystem Observation and Research Station, Shenyang 110016, China
5
College of Chemistry and Life Science, Anshan Normal University, Anshan 114007, China
6
University of Chinese Academy of Sciences, Beijing 100049, China
*
Author to whom correspondence should be addressed.
Diversity 2025, 17(3), 204; https://doi.org/10.3390/d17030204
Submission received: 19 January 2025 / Revised: 11 February 2025 / Accepted: 14 February 2025 / Published: 13 March 2025
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Abstract

:
Understanding the species diversity distribution of lycophytes and ferns is crucial for identifying biodiversity hotspots and conservation planning. Northeast China, a biodiversity-sensitive area affected by climate change, lacks comprehensive information on diversity and distribution patterns of these plants. To address this gap, we sorted out all naturally distributed lycophyte and fern species recorded in the region, analyzed their diversity, frequency, and threatened status. Correlation and regression analyses were also conducted with geographic gradients at the county level. Our study identified a total of 143 taxa (species and intraspecific taxa) belonging to 48 genera of 19 families of lycophytes and ferns in Northeast China, with terrestrial (85 spp.) and epilithic (55 spp.) life forms dominating. Species with frequencies below 10.00% comprised 75.52% of the total. Notably, five species were listed as threatened in the Red List of China’s Biodiversity, highlighting the urgency for conservation measures. Overall, species diversity decreased from low to high latitudes, but increased with maximum elevation and elevation range. High diversity areas were concentrated mainly in Da Hinggan Mountains, Xiao Hinggan Mountains, and Changbai Mountains, which correspond to the main mountainous terrain of Northeast China. Changbai Mountains exhibited the highest diversity, establishing itself as a pivotal diversity center for lycophytes and ferns in the region. Exploring the diversity and distribution of lycophytes and ferns is crucial for understanding their interactions with environmental gradients, and thereby supporting significant biodiversity conservation efforts in Northeast China.

1. Introduction

Species diversity is the most basic and critical level of biodiversity, and it is used in many aspects of biodiversity research [1,2,3,4]. A comprehensive understanding of species diversity is a pivotal step in guiding ecological conservation and sustainable utilization strategies, and it serves as a prerequisite for developing effective conservation policies [5,6], carrying profound significance. Among the various plant groups, lycophytes and ferns have ancient origins and include numerous relict and endemic species. They are of great importance in conservation ecology. Most extant lycophytes and ferns have narrow ecological amplitude, poor tolerance to environmental changes, and weak adaptability [7]. Therefore, their sensitivity to environmental changes can indicate alterations in habitat conditions, making them a great group for ecological research [8].
Macro-scale pattern and formation mechanism have always been the core issues in ecology and biogeography [9,10]. The distribution of species diversity is uneven, and the distribution laws vary with different regions and scales. Patterns of species distribution are products of many ecological processes which are mainly influenced by factors such as species development history, geographical configuration, and environmental variables [11,12]. Understanding the distribution pattern of species diversity is of great significance for the identification of biodiversity hotspots and the formulation of conservation strategies.
Some research has been conducted to unravel the distribution patterns and factors of ferns. The global patterns of fern diversity and its influencing factors have been explored and analyzed thoroughly [2,13]. In their research on the distribution patterns of rock ferns in Mediterranean climates, Bystriakova et al. [14] discovered that the distribution of Asplenium ferns was restricted by non-climatic environmental factors on a national scale. Khine et al. [15] assessed species richness patterns of ferns in relation to climatic factors in East and Southeast Asia. The biogeographic pattern of fern richness in the NE Iberian Peninsula at the mesoscale level was analyzed [16], and they found that the species richness is mainly related to terrain and climate. These studies collectively emphasize the complexity of fern diversity patterns across different scales and regions, highlighting their significance as a focal area of research.
Northeast China, positioned as a transitional zone between warm temperate and cold temperate, boasts a unique vegetation ecotone shaped by its distinct climate. This region serves as a sensitive area for biodiversity responses to climatic changes, whether the climate is warming or cooling [17]. Despite its ecological significance, research on lycophyte and fern diversity in Northeast China remains scarce, with limited studies focusing on regionalization of flora [18], flora of lycophytes and ferns [19] and the influence of environmental factors on diversity distribution [20]. However, a comprehensive analysis of the diversity and distribution patterns of lycophytes and ferns in this region, particularly at the macro-scale, remains elusive.
Over the past few decades, botanical surveys in Northeast China have amassed a vast collection of plant specimens, which serve as crucial evidence for identifying plant species and provide invaluable field data support. Thus, utilizing historical records such as specimens and publications to embark on an in-depth study of lycophyte and fern diversity, its distribution patterns, and the exploration of gradient distribution laws, particularly addressing the impact of topographic complexity on species diversity distribution, is not only the optimal strategy but also promises to effectively bridge this knowledge gap. As this study is the first to explore the diversity of lycophytes and ferns in this particular geographic region, it would deepen our understanding of the diversity of this ancient plant group in temperate regions and provide the basis for further conservation strategies as well as for diversity studies in response to climate change and human activities.

2. Materials and Methods

2.1. Study Area

Northeast China, encompassing three provinces (Heilongjiang, Jilin, and Liaoning) and the eastern part of Inner Mongolia (including Hulunbuir City, Tongliao City, Chifeng City, and Hinggan League), was selected as the study area. There are three main mountain ranges (Da Hinggan Mountains, Xiao Hinggan Mountains and Changbai Mountains) running from west to east in this area (Figure 1). The climate exhibits distinct characteristics with cold and dry winters influenced by the Mongolian high pressure and hot, rainy summers affected by oceanic air masses. The annual average temperature ranges from −5 to 11 °C, with annual precipitation varying between 300 and 1000 mm, and relative humidity falling within the bracket of 48.93% to 72.70% [21,22]. Vegetation types include coniferous forests in the cold temperate zone, deciduous broad-leaved forests in the warm temperate zone, and notably, mixed coniferous broad-leaved forests in the temperate zone [23], which serve as ecotones where vegetation types transition.

2.2. Data Selection

Geographical unit selection. For data statistics, the county was selected as the basic geographic unit, based on the administrative division of the Ministry of Civil Affairs of the People’s Republic of China, totaling 216 counties, districts, and banners in Northeast China (hereinafter referred to as counties in order to simplify the names).
Taxa data and life forms. Lycophyte and fern species with natural distribution records in Northeast China were collected from the following monographs and Northeast Biological Herbaria of Institute of Applied Ecology, Chinese Academy of Sciences (IFP). The species names were checked and analyzed by Flora of China [24] and Clavis Plantarum Chinae Boreali-Orientalis (Editio Secunda) [25], and classified according to PPGⅠ classification [26]. With reference to Flora of China, life forms of each species were collected.
Data of threatened species. The threatened and protected levels of each species were collected, with reference to the Red List of China’s Biodiversity (https://www.mee.gov.cn/xxgk2018/xxgk/xxgk01/202305/t20230522_1030745.html (accessed on 20 August 2024)), and the List of National Key Protected Wild Plants in China (https://www.forestry.gov.cn/c/www/gkml/11057.jhtml (accessed on 20 August 2024)).
Distribution information statistics. Distribution data for each species were primarily obtained from 6800 specimens in Northeast Biological Herbaria of Institute of Applied Ecology, Chinese Academy of Sciences (IFP). These specimens were collected over the past 70 years. Additional distribution data were sourced from monographs, such as Flora of China [24], Flora Reipublicae Popularis Sinicae [27], and various local floristic and distribution literature [28,29,30,31,32]. Furthermore, our field survey data from recent years were also included. The presence of each species in a county was recorded as 1, while its absence was recorded as 0.

2.3. Species Diversity Measurements

Species richness was analyzed using Patrick index [33], which has been adopted in many ecological studies [34,35]. Additionally, in order to neutralize the impact of varying county area sizes and to accurately assess the correlation between species number and sample size, the Gleason index [36,37] was employed as a measure of species richness.
Patrick index: DP = S, including Patrick index of family (DPF), Patrick index of genus (DPG), Patrick index of species (DPS).
Gleason index: DG = S/lnA, including Gleason index of family (DGF), Gleason index of genus (DGG), Gleason index of species (DGS).
S is the number of families, genera, and species in each county, and A is the area of each county.

2.4. Frequency Measurement and Frequency Level

The frequency of each species was calculated as follows:
Frequency = the number of counties where a certain plant occurs/the total number of counties × 100%
To calculate the percentage of occurrence within various frequency ranges, the frequencies were categorized into distinct frequency levels: Level A, ranging from 0.01% to 10.00%; Level B, from 10.01% to 20.00%; Level C, from 20.01% to 30.00%; and Level D, from 30.01% to 40.00%.

2.5. Distribution Patterns

Diversity distribution and high value areas. ArcGIS10.2 was utilized to map the diversity distribution of species, identifying areas with high values exceeding the average of Patrick index and Gleason index.
Gradient pattern of species diversity. Pearson correlation and regression analyses were conducted between species diversity (DPS and DGS) and latitude, longitude and elevation (elevation range and maximum elevation of each county). Trends in species diversity along gradients were analyzed using IBM SPSS Statistics 24.

3. Results

3.1. Composition of Lycophytes and Ferns

Our study identified a total of 143 taxa (including species and intraspecific taxa, ‘species’ mentioned hereinafter encompass intraspecific taxa for simplicity) of lycophytes and ferns in Northeast China, which belonged to 48 genera of 19 families. Lycophytes consist of two families, six genera, and 24 species, while ferns comprise 17 families, 42 genera, and 119 species. Among these families (Table 1), Dryopteridaceae emerged as the largest, with its 20 species accounting for 13.99% of the total species. Notably, six families, including Athyriaceae, Lycopodiaceae, Equisetaceae, Selaginellaceae, Woodsiaceae, and Ophioglossaceae, stood out with more than 10 species each, collectively constituting dominant families in Northeast China. These seven families, spanning 19 genera and 92 species, accounted for 39.58% of the total genera and 64.34% of the total species.
In terms of genera (Table 2), Dryopteris took the lead with 15 species, representing 10.49% of the total species. Noteworthy genera with more than nine species each included Equisetum, Selaginella, Woodsia, Athyrium and Asplenium. These 6 dominant genera comprised 67 species, accounting for 12.5% of the total genera and 46.85% of the total species. Additionally, 21 genera, each with only one species, contributed to 43.75% of the total genera and 14.69% of the total species.

3.2. Life Forms of Lycophytes and Ferns

Based on their adaptability to the environment, life forms of lycophytes and ferns were classified into three types: terrestrial, epilithic, and aquatic. The terrestrial type, with species such as Dryopteris crassirhizoma, Athyrium brevifrons, and Deparia coreana, constituted the majority with 85 species, accounting for 59.44% of the total. The epilithic type, including species like Pyrrosia linearifolia, Dryopteris fragrans, and Aleuritopteris argentea, comprised 55 species, accounting for 38.46% of the total. The aquatic type species were Marsilea quadrifolia, Salvinia natans, and Azolla pinnata subsp. asiatica, made up 2.10% of the total with three species.

3.3. Threatened Species

The assessment of lycophytes and ferns native to Northeast China under the Red List of China’s Biodiversity revealed their threatened status. Five species were classified as Threatened, with Huperzia serrata listed as Endangered (EN), and four species—H. miyoshiana, H. selago, Asplenium komarovii, and Cyrtomium falcatum—categorized as Vulnerable (VU), all indicating a dire need for conservation efforts to prevent their extinction. Four species were classified as Near Threatened (NT), highlighting their vulnerability to slipping into a threatened state and the need for continued monitoring. In contrast, 99 species were classified as Least Concern (LC), reflecting their stability within the ecosystems. In addition, 23 species were classified as Data Deficient (DD), and 12 species were not included in the Red List of China’s Biodiversity.
Additionally, six were classified as Class II Protected Species under the List of National Key Protected Wild Plants in China. These include five species of Huperzia: H. asiatica, H. miyoshiana, H. selago, H. appressa, and H. serrata, along with one species of Asplenium komarovii.

3.4. Frequency of Lycophytes and Ferns

The analysis of the frequency of 143 lycophyte and fern species across 216 counties, as depicted in Table S1, revealed a gradual decline in the frequency values from high to low. This transition was characterized by a smooth and continuous pattern, with the highest frequency recorded at 30.56%, indicative of the widespread occurrence of Athyrium brevifrons in 66 counties. Conversely, the lowest frequency stood at 0.46%, signifying a narrow distribution limited to a single county for 15 species, including Deparia pycnosora var. longidens and Athyrium rubripes and others.
Upon further analysis of each frequency level (depicted in Figure 2), it became evident that Level A, with frequencies ranging from 0.01% to 10.00%, comprised 75.53% of the total species. Level B, whose frequencies lied between 10.01% and 20.00%, accounted for 17.48% of the total species. Level C, spanning frequencies from 20.01% to 30.00%, contributed only 6.29%. Notably, there was only one species exceeding the 30.00% frequency threshold. Species belonging to Level C and Level D, which were distributed across 44 or more counties, with a total of 10 species, were considered as relatively common. These included Athyrium brevifrons, Equisetum arvense, Pteridium aquilinum var. latiusculum, Pyrrosia petiolosa, Equisetum hyemale, Selaginella tamariscina, Athyrium sinense, Onoclea sensibilis var. interrupta, Adiantum pedatum, and Aleuritopteris argentea.

3.5. Analysis of Diversity Distribution and High Value Areas

3.5.1. Patrick Index Diversity Distribution and High Value Areas

The distribution of Patrick index diversity exhibited significant variation among the counties in the study (Figure 3). High value areas of DPF (counties with DPF ≥ 5), DPG (DPG ≥ 7), and DPS (DPS ≥ 10), were predominantly concentrated in Da Hinggan Mountains, Xiao Hinggan Mountains, and Changbai Mountains. Counties with higher Patrick index, such as Antu (82 species, 38 genera, and 17 families), Kuandian (67 species, 33 genera, and 16 families), and Shangzhi (58 species, 32 genera, and 16 families), were notably distributed in Changbai Mountains, demonstrating higher Patrick diversity at the species, genus and family levels.

3.5.2. Gleason Index Diversity Distribution and High Value Areas

Similar to Patrick index diversity distribution, Gleason index diversity distribution at the species, genus and family levels followed a comparable pattern. High value areas of DGF (DGF ≥ 0.55), DGG (DGG ≥ 0.72), and DGS (DGS ≥ 1.14) were primarily observed in Da Hinggan Mountains, Xiao Hinggan Mountains, and Changbai Mountains (Figure 3). Counties with higher Gleason index, including Antu (DGF = 1.91, DGG = 4.26, and DGS = 9.09), Anshan (DGF = 2.25, DGG = 3.75, and DGS = 8.24), and Kuandian (DGF = 1.83, DGG = 3.78, and DGS = 7.67), were prominently situated in Changbai Mountains, demonstrating higher Gleason diversity.

3.6. Gradient Distribution Pattern of Species Diversity

Significant relationships emerged from the correlation analysis of species diversity in relation to latitude and elevation. Patrick index exhibited significant correlations with elevation range (r = 0.621, p < 0.001), maximum elevation (r = 0.580, p < 0.001), and latitude (r = −0.157, p = 0.021), but no significant correlation with longitude (r = 0.096, p = 0.159). Gleason index also displayed significant correlations with elevation range (r = 0.600, p < 0.001), maximum elevation (r = 0.556, p < 0.001), and latitude (r = −0.208, p = 0.002), but no significant correlation with longitude (r = 0.092, p = 0.18).
Single-factor regression analyses further confirmed these correlations (Figure 4). Patrick index exhibited significant linear correlations with elevation range (β1 = 0.019 (β1 is slope), R2 = 0.386 (R2 is coefficient of determination), p < 0.001), maximum elevation (β1 = 0.016, R2 = 0.336, p < 0.001), and latitude (β1 = −0.779, R2 = 0.025, p = 0.021), but no significant linear correlation with longitude (β1 = 0.372, R2 = 0.009, p = 0.159). Gleason index demonstrated significant linear correlations with elevation range (β1 = 0.002, R2 = 0.359, p < 0.001), maximum elevation (β1 = 0.002, R2 = 0.309, p < 0.001), and latitude (β1 = −0.12, R2 = 0.043, p = 0.002), but no significant linear correlation with longitude (β1 = 0.041, R2 = 0.008, p = 0.18).
In summary, our correlation and regression analyses revealed a clear species diversity distribution pattern of lycophytes and ferns in Northeast China. Along the latitude gradient, species diversity demonstrated a decreasing trend from low to high latitudes. Conversely, on the elevation gradients, higher maximum elevations and greater elevation ranges corresponded to higher species diversity. No significant correlation was observed between the longitude gradient and species diversity.

4. Discussion

4.1. Comprehensive Analysis of Species Diversity

The diversity of lycophytes and ferns in Northeast China demonstrates their adaptability to the area’s varied topography and climate. Dryopteridaceae and six families (Athyriaceae, Lycopodiaceae, Equisetaceae, Selaginellaceae, Woodsiaceae, and Ophioglossaceae) constitute 64.34% of the total species and are the dominant and common families in Northeast China. The dominance of terrestrial type showcases their versatility in thriving across a spectrum of soil types and microclimates, underscoring their pivotal role in terrestrial ecosystems. Furthermore, the substantial representation of epilithic species, accounting for 38.46% of the total, exemplifies their extraordinary ability to flourish on rocks. These plants are ecologically vital, contributing to soil development, nutrient recycling, and ensuring the stability of rocky habitats. Their continued existence in Northeast China underscores the urgency to preserve natural rock formations and mitigate habitat degradation, thereby safeguarding the future of these unique and indispensable species.
In the 2020 edition of the Red List of China’s Biodiversity, a total of 108 lycophyte and fern species native to Northeast China were assessed. Notably, the categorization of five species as Threatened—one labeled as Endangered and four as Vulnerable—starkly demonstrates the precarious state of these ancient plant lineages and the urgent need for prompt actions to avert their potential extinction. However, 35 species remain with their threat status unassessed, primarily due to insufficient data, which underscores significant deficiencies in our knowledge base and monitoring infrastructure. This underscores the critical importance of data acquisition and analysis in devising effective conservation strategies. Additionally, the inclusion of six species in the List of National Key Protected Wild Plants in China signifies their nationally recognized significance, while also indicating the necessity for even more comprehensive protection measures to ensure their survival.
The frequency research on 143 lycophyte and fern species across 216 counties reveals that 75.52% of these species are within the low-frequency range of 0.01% to 10.00%, highlighting the diversity and prevalence of low-frequency species in Northeast China’s lycophyte and fern flora. This finding echoes Raunkiaer’s Law of Frequency, which postulates the prevalence of species below 20% frequencies. The majority of species falling within the lowest frequency levels (Level A) indicates a widespread distributional narrowing, with many species confined to a single county. This trend underscores the vulnerability of these species to habitat destruction and fragmentation.

4.2. Gradient Distribution Pattern of Species Diversity and Its Diversity Center

In Northeast China, the diversity distribution of lycophyte and fern species exhibits distinct regional variations, a trend that aligns with broader patterns observed in Chinese lycophyte and fern diversity [38,39]. The decreasing trend in species diversity from low to high latitudes is particularly noteworthy and corresponds with the observed relationship between species diversity and latitude. Although the regression equation indicates a significant correlation, the relatively lower R² underscores the intricate influences of elevation and climate factors. It is crucial to note that within the scale of Northeast China, species diversity does not strictly follow a latitudinal distribution due to these influential factors. Conversely, no significant correlation was observed between lycophyte and fern species diversity and longitude. To fully understand this diversity, using both Patrick and Gleason indices is crucial as they offer complementary insights. The Patrick index measures species richness simply and widely, while the Gleason index normalizes for varying area sizes, providing a nuanced assessment and reflecting the species-sample size correlation. Together, they strengthen findings and add significant reference to understanding lycophyte and fern diversity patterns in Northeast China.
The diversity pattern of lycophyte and fern species in Northeast China closely corresponds to the region’s topography. We observed that areas with high species diversity are primarily concentrated in the Da Hinggan Mountains, Xiao Hinggan Mountains, and Changbai Mountains, with Changbai Mountains exhibiting the highest diversity. These three mountain ranges correspond to the distribution areas of forest resources in Northeast China as outlined by Zhang et al. [40]. Our findings further reveal a close relationship between the geographical distribution of lycophytes and ferns in Northeast China and the distribution of forests.
Furthermore, upon analyzing the elevation gradient pattern, we identified a notable alignment between species diversity and terrain factors, specifically elevation range and maximum elevation. Notably, the elevation range displayed a stronger correlation with species diversity. The consistent trend observed in the terrain factors and lycophyte and fern species diversity in Northeast China is as follows: as the elevation gradually ascends, the elevation range expands, contributing to the development of a more intricate and varied terrain, ultimately resulting in a heightened level of species diversity. This trend is consistent with the research findings on Chinese lycophyte and fern species diversity and terrain factors by Yan et al. [39].
Mountainous areas, serving as intricate ecosystems, reflect and encapsulate the natural geography and ecological characteristics of horizontal natural belts. Körner [41] proposed that over short elevation gradients, they host a series of climatically distinct life zones. Furthermore, these mountainous regions possess varied topographies and a wide range of diverse substrates, all of which contribute significantly to greater habitat heterogeneity and high niche diversity. This, in turn, facilitates the evolution and maintenance of diverse species, especially endemic species. Given their rich geodiversity, mountainous areas have emerged as remarkable biodiversity hotspots, estimated to support approximately one-third of terrestrial species diversity [42,43]. Therefore, mountains are generally considered to be biodiversity hotspots.
Moreover, mountainous areas not only safeguarded pristine habitats and diverse vegetation types but also emerge as vital refuges for a myriad of species [44]. This phenomenon is well documented in a study of African ferns, which shows that higher maximum elevations contribute to an increase in diversity, and the distribution of ferns in Africa has been influenced by the presence of refuges [45]. It has been demonstrated in various studies that the identification and protection of biological refugia should be prioritized in environmental protection plans, especially in addressing human-induced climate change [46,47,48]. Myers et al. [5] and Badgley et al. [49] demonstrate that regions with complex terrains encompass many of the world’s current diversity hotspots, providing sanctuary to a high proportion of species prone to extinction. Consequently, these regions possess high conservation value.
Changbai Mountains, with their unique flora and abundant plant species [50], represent the primary mountain ranges hosting coniferous and broad-leaved mixed forests in the northeast temperate zone. Phylogeographic studies highlight the critical role of Changbai Mountains as primary refugia for temperate coniferous and broad-leaved mixed forests during the last glacial maximum [17]. Refugia, acting as habitats for plant survival and proliferation during Quaternary glaciations and interglacial periods, serve as the starting points for population expansion during interglacial or post-glacial periods. Refuge populations exhibit longer evolutionary histories and higher genetic diversity [17,51,52].
The presence of endemic species serves as crucial evidence in identifying ice age refuges [53]. Specifically, in Northeast China, Huperzia asiatica and Parathelypteris changbaishanensis stand out as endemic fern species exclusive to the Changbai Mountains. The high-value area analysis conducted in this study, exploring the diversity distribution of lycophyte and fern species in Northeast China, consistently revealed that Changbai Mountains harbor the highest species diversity. Consequently, designating Changbai Mountains as significant ice age refuges, they emerge as pivotal diversity centers for lycophytes and ferns within the broader context of Northeast China.

5. Conclusions

Our study provides a comprehensive understanding of the lycophyte and fern diversity in Northeast China, offering several key insights. We identified 143 taxa belonging to 48 genera of 19 families in the region. Key findings include the dominance of seven families and six genera, which account for 64.34% and 46.85% of the total species, respectively. The majority of species are terrestrial and epilithic in nature, and with a high proportion (75.52%) having frequencies below 10.00%. Notably, five species are listed as threatened in China’s Biodiversity Red List. Additionally, our study also demonstrates a significant latitude gradient pattern in species diversity, which tends to increase with higher maximum elevation and elevation range. High diversity areas, particularly in Da Hinggan, Xiao Hinggan, and Changbai Mountains, are concentrated in the mountainous regions of Northeast China, characterized by high habitat heterogeneity. Notably, Changbai Mountains emerge as a pivotal diversity center for lycophytes and ferns in the region. Most importantly, our study underscores the importance of considering the species composition and distribution in ecotone areas, highlighting their significance for both understanding and conserving biodiversity.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d17030204/s1. Table S1: The frequency of 143 lycophyte and fern species across counties in Northeast China.

Author Contributions

Conceptualization, Y.L. and X.H.; Methodology, X.H.; Validation, S.Y. and S.X.; Formal analysis, Y.L., S.X. and L.J.; Investigation, Y.L. and S.Y.; Data curation, S.X. and L.J.; Writing, Y.L.; Rewriting and response to reviewers: Y.L. and X.H.; Funding acquisition, X.H. All authors have read and agreed to the published version of the manuscript.

Funding

This work was supported by the National Key R&D Program of China [2022YFF1300500].

Institutional Review Board Statement

Not applicable.

Data Availability Statement

All data are reported in the manuscript. The specimens can be obtained from Northeast Biological Herbaria of Institute of Applied Ecology, Chinese Academy of Sciences.

Acknowledgments

We thank Northeast Biological Herbaria of Institute of Applied Ecology, Chinese Academy of Sciences for the specimens and detailed information.

Conflicts of Interest

The authors report there are no competing interests to declare.

References

  1. Syfert, M.M.; Brummitt, N.A.; Coomes, D.A.; Bystriakova, N.; Smith, M.J. Inferring diversity patterns along an elevation gradient from stacked SDMs: A case study on Mesoamerican ferns. Glob. Ecol. Conserv. 2018, 16, e00433. [Google Scholar] [CrossRef]
  2. Weigand, A.; Abrahamczyk, S.; Aubin, I.; Bita-Nicolae, C.; Bruelheide, H.; Carvajal-Hernández, C.I.; Cicuzza, D.; Erickson Nascimento da Costa, L.; Csiky, J.; Dengler, J.; et al. Global fern and lycophyte richness explained: How regional and local factors shape plot richness. J. Biogeogr. 2020, 47, 59–71. [Google Scholar] [CrossRef]
  3. Suis, M.A.F.; Majuakim, L.; Suleiman, M. Fern species richness patterns and their environmental preferences across elevational gradient on Mount Trus Madi, Sabah, Malaysia. J. Trop. For. Sci. 2021, 33, 58–68. [Google Scholar]
  4. Pan, X.T.; Ye, M.; He, Q.Z.; Zhang, K. Spatial distribution pattern and influencing factors of above-ground biomass and species diversity of grassland in the Altay forest area. Land 2023, 12, 1370. [Google Scholar] [CrossRef]
  5. Myers, N.; Mittermeier, R.A.; Mittermeier, C.G.; da Fonseca, G.A.B.; Kent, J. Biodiversity hotspots for conservation priorities. Nature 2000, 403, 853–858. [Google Scholar] [CrossRef]
  6. Sanginés-Franco, C.; Luna-Vega, I.; Contreras-Medina, R.; Espinosa, D.; Tejero-Díez, J.D.; Gerardo, R. Diversity, endemism, and conservation of ferns (Polypodiales) in the Mexican Mountain Component. J. Mt. Sci. 2015, 12, 891–904. [Google Scholar] [CrossRef]
  7. Lu, S.G. Pteridology; Higher Education Press: Beijing, China, 2007. (In Chinese) [Google Scholar]
  8. Qian, H.; Zhang, J.; Jiang, M.C. Global patterns of fern species diversity: An evaluation of fern data in GBIF. Plant Divers. 2022, 44, 135–140. [Google Scholar] [CrossRef]
  9. Gaston, K.J. Global patterns in biodiversity. Nature 2000, 405, 220–227. [Google Scholar] [CrossRef]
  10. Wang, J.M.; Wang, W.J.; Li, J.W.; Feng, Y.M.; Wu, B.; Lu, Q. Biogeographic patterns and environmental interpretation of plant species richness in desert regions of Northwest China. Biodivers. Sci. 2017, 25, 1192–1201. [Google Scholar] [CrossRef]
  11. Qian, H. Large-scale biogeographic patterns of vascular plant richness in North America: An analysis at the generic level. J. Biogeogr. 1998, 25, 829–836. [Google Scholar] [CrossRef]
  12. Tang, Z.Y.; Fang, J.Y. A review on the elevational patterns of plant species diversity. Biodivers. Sci. 2004, 12, 20–28. [Google Scholar]
  13. Suissa, J.S.; Sundue, M.A.; Testo, W.L. Mountains, climate and niche heterogeneity explain global patterns of fern diversity. J. Biogeogr. 2021, 48, 1296–1308. [Google Scholar] [CrossRef]
  14. Bystriakova, N.; Peregrym, M.; Dragicevic, S. Effect of environment on distributions of rock ferns in the Mediterranean climate: The case of the genus Asplenium in Montenegro. Flora-Morphol. Distrib. Funct. Ecol. Plants 2015, 215, 84–91. [Google Scholar] [CrossRef]
  15. Khine, P.K.; Kluge, J.; Kessler, M.; Miehe, G.; Karger, D.N. Latitude-independent, continent-wide consistency in climate–richness relationships in Asian ferns and Lycophytes. J. Biogeogr. 2019, 46, 981–991. [Google Scholar] [CrossRef]
  16. Pausas, J.G.; Sáez, L. Pteridophyte richness in the NE Iberian Peninsula: Biogeographic patterns. Plant Ecol. 2000, 148, 195–205. [Google Scholar] [CrossRef]
  17. Ye, J.W.; Yuan, Y.G.; Cai, L.; Wang, X.J. Research progress of phylogeographic studies of plant species in temperate coniferous and broadleaf mixed forests in Northeastern China. Biodivers. Sci. 2017, 25, 1339–1349. [Google Scholar] [CrossRef]
  18. Wang, J.Z. The features and regionalization of the Pteridophyte flora in Northeast China. J. Beijing For. Univ. (English Ed.) 1995, 4, 43–52. [Google Scholar]
  19. Li, Y.; Cao, W.; He, H.; Zhang, Y.; Su, D.Y.; Chen, W. Flora of Lycophytes and Ferns in Northeast China. Bull. Bot. Res. 2022, 42, 321–328. [Google Scholar]
  20. Li, Y.; Chang, Y.; He, X.Y.; Xu, S.; Su, D.Y. Effect of environmental factors on the spatial diversity distribution patterns of lycophytes and ferns in Northeast China. Russ. J. Ecol. 2022, 53, 111–122. [Google Scholar]
  21. Zhang, X.W.; Zang, S.Y.; Sun, L. Spatial-temporal variation characteristics of snow cover days in Northeast China in the past 40 years and their relationship with climatic factors. Adv. Earth Sci. 2018, 33, 958–968. [Google Scholar]
  22. Liu, Y.Y.; Li, Y.F.; Xie, J.F.; Zhang, H. Climate change characteristics of free atmospheric humidity and its relationship with temperature and precipitation in Northeast China. Sci. Geogr. Sin. 2016, 36, 628–636. [Google Scholar]
  23. Xu, W.D. The relation between the zonal distribution of types of vegetation and the climate in Northeast China. Acta Phytoecol. Geobot. Sin. 1986, 10, 254–263. [Google Scholar]
  24. Wu, Z.Y.; Raven, P.H.; Hong, D.Y. Flora of China; Lycopodiaceae through Polypodiaceae; Science Press: Beijing, China; Missouri Botanical Garden Press: St. Louis, MO, USA, 2013; Volumes 2–3. [Google Scholar]
  25. Fu, P.Y. Clavis Plantarum Chinae Boreali-Orientalis, 2nd ed.; Science Press: Beijing, China, 1995. [Google Scholar]
  26. PPG, I. A community-derived classification for extant lycophytes and ferns. J. Syst. Evol. 2016, 54, 563–603. [Google Scholar] [CrossRef]
  27. Delectis Florae Reipublicae Popularis Sinicae Agenda Academiae Sinicae Edita. Flora Reipublicae Popularis Sinicae; Science Press: Beijing, China, 2004. [Google Scholar]
  28. Cao, W.; Li, J.Y. Natural Distribution for Plant of Changbai Shan in China; Northeastern University Press: Shenyang, China, 2003. [Google Scholar]
  29. Cao, W.; Li, J.Y.; Fu, P.Y.; Yu, X.H.; Zhu, C.X. Floristics and Distribution of Plants in Da Hinggan Ling China; Northeastern University Press: Shenyang, China, 2004. [Google Scholar]
  30. Cao, W.; Li, J.Y. Floristics and Distribution of Plants in Xiao Hinggan Ling China; Science Press: Beijing, China, 2007. [Google Scholar]
  31. Shrestha, N.; Zhang, X.C. On the presence of North American clubmoss Huperzia lucidula (Lycopodiaceae) in China: An intercontinental disjunction or misidentification. Phytotaxa 2015, 219, 243–252. [Google Scholar] [CrossRef]
  32. Chen, D.K.; Zhou, X.M.; He, H.; Zhang, L.B. Spinulum lioui, a new species referred to Lycopodium neopungens (Lycopodiopsida: Lycopodiaceae) in China. Phytotaxa 2017, 307, 161–163. [Google Scholar] [CrossRef]
  33. Patrick, R. A proposed biological measure of stream conditions, based on a survey of the Conestoga Basin, Lancaster County, Pennsylvania. Proc. Acad. Nat. Sci. Phila. 1949, 101, 277–342. [Google Scholar]
  34. Ma, K.P. Measurement method of community diversity. Chin. Biodivers. 1994, 2, 162–168. [Google Scholar]
  35. Liu, S.L.; Dong, Y.H.; Sun, Y.X.; Li, J.R.; An, Y.; Shi, F.N. Modelling the spatial pattern of biodiversity utilizing the high-resolution tree cover data at large scale: Case study in Yunnan province, Southwest China. Ecol. Eng. 2019, 134, 1–8. [Google Scholar] [CrossRef]
  36. Gleason, H.A. On the relation between species and area. Ecology 1922, 3, 158–162. [Google Scholar]
  37. Gleason, H.A. Species and area. Ecology 1925, 6, 66–74. [Google Scholar]
  38. Zang, D.K. A preliminary study on the ferns flora in China. Acta Bot. Boreali-Occident. Sin. 1998, 18, 459–465. [Google Scholar]
  39. Yan, Y.H.; Zhang, X.C.; Ma, K.P. Pteridophytes in China: Diversity and Distribution; Science Press: Beijing, China, 2013. [Google Scholar]
  40. Zhang, Q.; Ren, R.; Zhao, L. Forest Spatial Distribution and Influencing Factors in Northeast China. J. Northeast For. Univ. 2013, 41, 25–28. [Google Scholar]
  41. Körner, C. Mountain biodiversity, its causes and function. Ambio 2004, 13, 11–17. [Google Scholar] [CrossRef]
  42. Körner, C.; Jetz, W.; Paulsen, J.; Payne, D.; Rudmann-Maurer, K.; Spehn, E.M. A global inventory of mountains for bio-geographical applications. Alp Bot. 2017, 127, 1–15. [Google Scholar] [CrossRef]
  43. Barthlott, W.; Mutke, J.; Rafiqpoor, D.; Kier, G.; Kreft, H. Global centers of vascular plant diversity. Nova Acta Leopold. 2005, 92, 61–83. [Google Scholar]
  44. Wang, X.P.; Wang, Z.H.; Fang, J.Y. Mountain ranges and peaks in China. Biodivers. Sci. 2004, 12, 206–212. [Google Scholar]
  45. Aldasoro, J.J.; Cabezas, F.; Aedo, C. Diversity and distribution of ferns in sub-Saharan African, Madagascar and some islands of the South Atlantic. J. Biogeogr. 2004, 31, 1579–1604. [Google Scholar] [CrossRef]
  46. IPCC. Climate Change 2007: The Physical Science Basis; Summary for policy makers; Contribution of Working Group 1 to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change; Cambridge University Press: Cambridge, UK, 2007. [Google Scholar]
  47. Noss, R.F. Beyond Kyoto: Forest management in a time of rapid climate change. Conserv. Biol. 2001, 15, 578–590. [Google Scholar] [CrossRef]
  48. Hu, Z.J.; Zhang, Y.L.; Liu, L.S.; Yu, H.B. Refugia and their identification methods: A review. Chin. J. Ecol. 2013, 32, 3397–3406. [Google Scholar]
  49. Badgley, C.; Smiley, T.M.; Terry, R.; Davis, E.B.; DeSantis, L.R.G.; Fox, D.L.; Hopkins, S.S.B.; Jezkova, T.; Matocq, M.D.; Matzke, N.; et al. Biodiversity and topographic complexity: Modern and geohistorical perspectives. Trends Ecol. Evol. 2017, 32, 211–226. [Google Scholar] [CrossRef]
  50. Wu, Z.Y. Chinese Vegetation; Science Press: Beijing, China, 1980. [Google Scholar]
  51. Provan, J.; Bennett, K.D. Phylogeographic insights into cryptic glacial refugia. Trends Ecol. Evol. 2008, 23, 564–571. [Google Scholar] [CrossRef]
  52. Keppel, G.; van Niel, K.P.; Wardell-Johnson, G.W.; Yates, C.J.; Byrne, M.; Mucina, L.; Schut, A.G.T.; Hopper, S.D.; Franklin, S.E. Refugia: Identifying and understanding safe havens for biodiversity under climate change. Glob. Ecol. Biogeogr. 2012, 21, 393–404. [Google Scholar] [CrossRef]
  53. Chen, D.M.; Kang, H.Z.; Liu, C.J. An overview on the potential quaternary glacial refugia of plants in China mainland. Bull. Bot. Res. 2011, 31, 623–632. [Google Scholar]
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Figure 1. The study area and mountain ranges in Northeast China.
Figure 1. The study area and mountain ranges in Northeast China.
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Figure 2. The percentage of each frequency level. (Level A, ranging from 0.01% to 10.00%; Level B, from 10.01% to 20.00%; Level C, from 20.01% to 30.00%; and Level D, from 30.01% to 40.00%).
Figure 2. The percentage of each frequency level. (Level A, ranging from 0.01% to 10.00%; Level B, from 10.01% to 20.00%; Level C, from 20.01% to 30.00%; and Level D, from 30.01% to 40.00%).
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Figure 3. Diversity and distribution pattern of ferns in each county of Northeast China. ((A): Patrick index of family (DPF); (B): Patrick index of genus (DPG); (C): Patrick index of species (DPS); (D): Gleason index of family (DGF); (E): Gleason index of genus (DGG); (F): Gleason index of species (DGS)).
Figure 3. Diversity and distribution pattern of ferns in each county of Northeast China. ((A): Patrick index of family (DPF); (B): Patrick index of genus (DPG); (C): Patrick index of species (DPS); (D): Gleason index of family (DGF); (E): Gleason index of genus (DGG); (F): Gleason index of species (DGS)).
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Figure 4. Regression analysis of species diversity with latitude, longitude, and elevation. ((A): Patrick index of species (DPS) and elevation range; (B): Patrick index of species (DPS) and maximum elevation; (C): Patrick index of species (DPS) and latitude; (D): Patrick index of species (DPS) and longitude; (E): Gleason index of species (DGS) and elevation range; (F): Gleason index of species (DGS) and maximum elevation; (G): Gleason index of species (DGS) and latitude; (H): Gleason index of species (DGS) and longitude).
Figure 4. Regression analysis of species diversity with latitude, longitude, and elevation. ((A): Patrick index of species (DPS) and elevation range; (B): Patrick index of species (DPS) and maximum elevation; (C): Patrick index of species (DPS) and latitude; (D): Patrick index of species (DPS) and longitude; (E): Gleason index of species (DGS) and elevation range; (F): Gleason index of species (DGS) and maximum elevation; (G): Gleason index of species (DGS) and latitude; (H): Gleason index of species (DGS) and longitude).
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Table 1. Number of genera and species in families of lycophytes and ferns in Northeast China.
Table 1. Number of genera and species in families of lycophytes and ferns in Northeast China.
FamilyNo. of GenusNo. of SpeciesFamilyNo. of GenusNo. of SpeciesFamilyNo. of GenusNo. of Species
Dryopteridaceae420Aspleniaceae19Osmundaceae33
Athyriaceae316Pteridaceae49Salviniaceae22
Lycopodiaceae *513Polypodiaceae57Davalliaceae11
Equisetaceae111Thelypteridaceae46Hymenophyllaceae11
Selaginellaceae *111Cystopteridaceae25Marsileaceae11
Woodsiaceae111Onocleaceae34
Ophioglossaceae410Dennstaedtiaceae23
Notes: * families belong to lycophytes.
Table 2. Number of species in genera of lycophytes and ferns in Northeast China.
Table 2. Number of species in genera of lycophytes and ferns in Northeast China.
GenusNo. of SpeciesGenusNo. of SpeciesGenusNo. of Species
Dryopteris15Botrypus2Davallia1
Equisetum11Coniogramme2Dendrolycopodium *1
Selaginella *11Cystopteris2Lepisorus1
Woodsia11Dennstaedtia2Marsilea1
Athyrium10Diphasiastrum *2Onoclea1
Asplenium9Diplazium2Oreopteris1
Huperzia *5Matteuccia2Osmunda1
Deparia4Ophioglossum2Osmundastrum1
Adiantum3Parathelypteris2Paragymnopteris1
Aleuritopteris3Spinulum *2Pentarhizidium1
Botrychium3Thelypteris2Phegopteris1
Gymnocarpium3Arachniodes1Pleurosoriopsis1
Lycopodium *3Azolla1Polypodium1
Polystichum3Claytosmunda1Pteridium1
Pyrrosia3Crepidomanes1Salvinia1
Sceptridium3Cyrtomium1Selliguea1
Notes: * genera belong to lycophytes.
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Li, Y.; Yu, S.; Xu, S.; Jia, L.; He, X. Understanding the Diversity and Distribution of Lycophytes and Ferns in Northeast China Based on Historical Records. Diversity 2025, 17, 204. https://doi.org/10.3390/d17030204

AMA Style

Li Y, Yu S, Xu S, Jia L, He X. Understanding the Diversity and Distribution of Lycophytes and Ferns in Northeast China Based on Historical Records. Diversity. 2025; 17(3):204. https://doi.org/10.3390/d17030204

Chicago/Turabian Style

Li, Yan, Shuai Yu, Sheng Xu, Lian Jia, and Xingyuan He. 2025. "Understanding the Diversity and Distribution of Lycophytes and Ferns in Northeast China Based on Historical Records" Diversity 17, no. 3: 204. https://doi.org/10.3390/d17030204

APA Style

Li, Y., Yu, S., Xu, S., Jia, L., & He, X. (2025). Understanding the Diversity and Distribution of Lycophytes and Ferns in Northeast China Based on Historical Records. Diversity, 17(3), 204. https://doi.org/10.3390/d17030204

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