Next Article in Journal
Can Biological Traits Serve as Predictors for Fishes’ Introductions, Establishment, and Interactions? The Mediterranean Sea as a Case Study
Next Article in Special Issue
Filmy Ferns (Hymenophyllaceae) and Associated Spike-Mosses (Selaginellaceae) from the Mid-Cretaceous Kachin Amber, Myanmar
Previous Article in Journal
Genetic Characterization and Alternative Preservation Ways of Locally Adapted Sheep Breeds: Cases of Private and Public Sheep Sectors in Tunisia and Italy
Previous Article in Special Issue
Fossil Fruits of Ceratophyllum from the Upper Eocene and Miocene of South China
 
 
Search for Articles:
Title / Keyword
Author / Affiliation / Email
Journal
Article Type
 
 
Advanced Search
 
Section
Special Issue
Volume
Issue
Number
Page
 
Journals
Biology
Volume 11
Issue 11
10.3390/biology11111624
biology-logo
Submit to this Journal Review for this Journal Propose a Special Issue
► Article Menu

Article Menu

share Share announcement Help format_quote Cite question_answer Discuss in SciProfiles
first_page
settings
Order Article Reprints
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:
Background:
Article

Early Cretaceous Keteleerioxylon Wood in the Songliao Basin, Northeast China, and Its Geographic and Environmental Implications

by
Xiao Shi
1,2,*,
Yuewu Sun
3,4,
Fanli Meng
1,
Jianxin Yu
5 and
Zilie Lan
1
1
Department of Geology, College of Earth Sciences, Jilin University, Changchun 130061, China
2
International Center of Future Science, Dinosaur Evolution Research Center, Jilin University, Changchun 130012, China
3
Research Center of Paleontology and Stratigraphy, Jilin University, Changchun 130026, China
4
Bioarchaeology Laboratory, Jilin University, Changchun 130021, China
5
School of Earth Sciences, State Key Laboratory of Biogeology and Environmental Geology, China University of Geosciences, Wuhan 430074, China
*
Author to whom correspondence should be addressed.
Biology 2022, 11(11), 1624; https://doi.org/10.3390/biology11111624
Submission received: 23 September 2022 / Revised: 20 October 2022 / Accepted: 2 November 2022 / Published: 7 November 2022
(This article belongs to the Special Issue Recent Advances in Palaeobotany)
Browse Figures
Review Reports Versions Notes

Abstract

:

Simple Summary

Keteleeria is a small group of Pinaceae, now only distributed in east and southeast Asia, but which was widely distributed in Asia, Europe, and North America in the northern hemisphere during the Late Mesozoic and Cenozoic periods. A new Keteleeria-like wood fossil, Keteleerioxylon changchunense Shi, Sun, Meng et Yu sp. nov., was described in the Early Cretaceous strata about 110 million years ago in Changchun, Jilin Province, Northeast China. The growth rings of wood contain rich palaeoecological and paleoclimatic information. Quantitative analysis of growth rings revealed that the new species is evergreen with leaf longevity of 1–3 years. The growth rings of Keteleerioxylon changchunense indicate that the climate seasonality was pronounced in the Songliao Basin during the Early Cretaceous period. By reviewing Keteleeria and closely related fossil taxa, we depict the probable migration route of Keteleeria. The oldest Keteleeria-like wood was found in the middle Jurassic period in Svalbard, Norway. They were distributed in both the middle and the high latitudes during the Late Jurassic–Cretaceous global warming time, while during the Paleogene and Neogene cooling times, the Keteleeria-like wood became scarce, especially in the Quaternary Glaciation, and until now, they were strictly restricted to the subtropical and tropical regions of east and southeast Asia.

Abstract

The extant Keteleeria is endemic to east and southeast Asia, while Keteleeria-like trees were widely distributed in the northern hemisphere in Earth’s history. In this paper, we reported a novel wood fossil of Keteleerioxylon changchunense Shi, Sun, Meng et Yu sp. nov., collected from the middle part of the Yingcheng Formation, Yingcheng Coal Mine, Changchun City, Jilin Province, northeast China. The quantitative growth-ring analyses of K. changchunense indicate that it was evergreen with a leaf longevity of 1–3 years, which is consistent with the foliar retention of extant Keteleeria. Its high ring-markedness index (RMI) indicates that the climate seasonality was pronounced during the Early Albian period in the Songliao Basin, northeast China. The fossil records of Keteleeria and closely related taxa indicate that this group might have originated from high latitudes in the northern hemisphere, then spread and migrated southward during the Late Jurassic and Cretaceous periods, gradually decreased in the Cenozoic period, and so far only survives in east and southeast Asia.

1. Introduction

Keteleeria was firstly described as a genus in 1866, named by Carrière [1]. The genus Keteleeria Carrière includes ten extant species and two variants, which are endemic to east and southeast Asia and only found in southern China (from Qinling Mountain to Hainan Island), northern Laos, and southern Vietnam [2] (Figure 1). Keteleeria is confined to humid, moderately warm (subtropical) areas with relatively low mountains (200–3000 m above sea level) [3,4,5,6].
Fossil woods with similar anatomy to extant Keteleeria were first described in the Lower Cretaceous of Franz Josef Land with the name Keteleerioxylon [7]. Although Keteleeria and Abies separated in the Paleocene based on molecular clock estimation [8], the oldest representatives of Keteleeria-like (the fossil woods, leaves, and reproductive organs showing similarities to the genus Keteleeria, including Keteleerioxylon, Keteleeria, and Protopiceoxylon or Pinoxylon) wood might have been in the Middle Jurassic [9]; most fossils reLated to the Keteleeria were discovered from the Late Jurassic to Cenozoic strata of Asia, Europe, and North America in the northern hemisphere (e.g., [10,11,12]).
In this study, we describe two new silicified wood specimens belonging to the genus Keteleerioxylon from the early Cretaceous sediments in the Songliao Basin, northeast China, and named Keteleerioxylon changchunense sp. nov. Quantitative growth-ring analyses were made to understand ecological and climatic indications. Meanwhile, the geological and geographical distributions of Keteleeria-like fossil woods are summarized in order to depict the relationship between its migration and paleoclimate changes during the Cretaceous and Cenozoic times.

2. Materials and Methods

2.1. Materials

Two silicified woods (registered nos. JLJY-01 and JLJY-02) were collected from the middle section of the Yingcheng Formation, Yingcheng Coal Mine (44°9′47.53″ N, 125°54′53.92″ E), Changchun City, Jilin Province, northeast China (Figure 2). The location is situated at the southeast margin of the Songliao Basin, the largest Mesozoic–Cenozoic terrestrial oil-bearing basin in northeast Asia. The Lower Cretaceous succession in the Songliao Basin is composed of the Huoshiling, Shahezi, Yingcheng, and Denglouku formations in ascending order [13]. The Yingcheng Formation, dominated by medium-acid volcanic, volcaniclastic rocks, and coal-bearing deposits, can be divided into three parts. The middle section is a set of sedimentary rocks with coal layers yielding spores and pollen, leaves, woods, and insects.
Detailed isotopic dating of volcanic rocks indicates that the age of the middle section is Late Aptian–early Albian, between 115.2 ± 0.4 Ma and 110.0 ± 2.0 Ma [14,15]. Based on palynological data, due to the presence of Tricolpites sp. in the middle part of the Yingcheng Formation where the present silicified wood specimens were collected, the member is considered earliest Albian in age [16,17,18,19].
The specimen JLJY-01 is 45.5 cm long with a maximum diameter of 13.5 cm. The specimen JLJY-02 is 29.4 cm long and 11.0–21.6 cm in diameter. They are gray to black in color with well-preserved secondary xylem. However, no pith or primary xylem is preserved.
All the specimens and slides are housed in the Research Center of Paleontology and Stratigraphy, Jilin University, Changchun, China.

2.2. Methods

To investigate the anatomical characteristics of those silicified woods, microscopic slides of the transverse, radial, and tangential sections were made in the State Key Laboratory of Biogeology and Environmental Geology, China University of Geosciences (Wuhan). The slides were observed using a microscope (Nikon AZ100M) with a digital camera (Nikon DS-Ri2) in the International Center of Future Science, Dinosaur Evolution Research Center, Jilin University. Images were processed by the software Adobe Photoshop CS6 (Version 13. San Jose, CA, USA: Adobe Inc., licensed to Jilin University).
For the quantitative growth ring analysis, the radial diameters of tracheids were measured, and the percentage of diminution, percentage of latewood, ring markedness index (RMI), and percentage of skews of CSDM (cumulative sum of the deviation from mean diameter) curves were calculated following the method of Falcon-Lang [20].

3. Results

ORDER Coniferales.
GENUS Keteleerioxylon I.A. Shilkina, 1960.
Type species: Keteleerioxylon arcticum I.A. Shilkina, 1960.
Keteleerioxylon changchunense Shi, Sun, Meng et Yu sp. nov.
Holotype: Specimen JLJY-02 (Figure 3A); Slides JLJY-02 a1, JLJY-02 a2, JLJY-02 b1, JLJY-02 b2, JLJY-02 c1, JLJY-02 c2.
Repository: Research Center of Paleontology and Stratigraphy, Jilin University, Changchun, China.
Type locality: Yingcheng Coal Mine, Changchun City, Jilin Province, China.
Stratigraphic horizon and age: The middle section of the Yingcheng Formation, earliest Albian (Early Cretaceous).
Etymology: The specific name is derived from Changchun where the specimens were collected.
Diagnosis: Growth rings distinct. Pits on radial walls of tracheids uniseriate to triseriate. Pits circular; uniseriate pits scattered, rarely contiguous slightly; biseriate pits mostly opposite, occasionally alternate; triseriate in a row. Crassulae present. Pits on tangential walls of tracheids absent. Transverse walls of axial parenchyma cells smooth. Rays, (1)6–11(37) cells high, uniseriate, sometimes with one to eight layers of biseriate cells. Horizontal and end walls of ray cells distinctly pitted (Abientineentüpfelung). Ray marginal cells, resembling ray tracheids, with scattered uniseriate pits. Pits, of taxodioid-cupressoid type, 1–3(6) per cross-field, 5–10 μm in diameter. Resin canals, vertical, normal, surrounded by six to eleven thick-walled epithelial cells in a ring. Horizontal resin canals absent.
Description: The two specimens show similar characteristics, but the features of the transverse section are clearer in specimen JLJY-02. Thus, the description here is mainly based on the specimen JLJY-02. Only the secondary xylem was preserved in the two specimens.
The homoxulous pycnoxylic wood consists of tracheids, rays, axial parenchyma cells, and epithelial cells of resin canals. The growth rings are distinct, 0.79–2.85 mm wide, with distinct and relatively straight boundaries (Figure 3B,D). The early-late wood transition is abrupt. In the transverse section, the early wood tracheids are large, thin-walled, and mainly rectangular (sometimes polygonal or circular). The latewood tracheids are thick-walled, and radially compressed, and lumens almost disappeared near the growth-ring margin. The radial tracheid lumens are 0–89 μm in diameter. Intercellular space is absent. Xylem rays mainly consist of uniseriate cells, and 1–10 seriates of tracheids in between.
Pits on the radial walls are borded, circular in shape, (11)12–17(20) μm in diameter. In the late woods, they are uniseriate, partly biseriate, and scattered, rarely slightly contiguous, with an included aperture (circular or elliptical). In the early woods, they are mostly biseriate or triseriate, occasionally uniseriate (Figure 3F,G,H and Figure 4A). When biseriate, the pits are mainly opposite, occasionally they show a tendency toward alternate arrangement (Figure 3G,H); when triseriate, the pits arrange in a row (Figure 4A). The pits are (16)18–22(25) μm in diameter, with circular or elliptical apertures. Crassulae are present (Figure 3F,H and Figure 4A). Axial parenchyma cells are rare, and `their transverse walls are smooth (Figure 4C).
Rays are (1)6–11(37) cells in height, uniseriate, occasionally with one to eight layers of biseriate cells (Figure 4G,H). In the radial section, ray cells are brick-shaped, 19–25 μm in height, and 50–129 μm in length. The horizontal and end walls of the ray parenchyma cells are pitted (Figure 4B,I). In the tangential section, ray cells are 10–28 μm in width, elliptical, circular, or rectangular, but rounded-triangular in marginal ray cells. Marginal ray cells resemble those ray tracheids with scattered uniseriate pits (Figure 4F).
There are 1–3(6) pits per cross-field, of taxodioid-cupressoid type, 5–10 μm in diameter. (Figure 4D,E). Two or three pits arrange in a row, while four to six pits are in a diffuse arrangement.
Vertical resin canals are present only (Figure 3E). Resin canals are circular or oval, radially elongated, 47–206 μm in diameter, and single or in pairs (Figure 3B,C). They are lined with six to eleven thick-walled epithelial cells in a ring and situated in both early and latewood. Horizontal resin canals are absent.
Comparison: The fossil woods described here are characterized by the presence of (only) vertical normal resin canals with thick-walled epithelial cells, axial parenchyma, and the marginal cells of rays resembling ray tracheids. They closely resemble the extant genus Keteleeria [11].
The presence of (only) vertical normal resin canals with thick-walled epithelial cells and the absence of horizontal resin canals are unique characteristics of the extant Keteleeria [7,11,21,22,23,24,25]. Sole vertical resin canals may also occur in the wood of some taxa of the Taxodiaceae and Cupressaceae. However, these canals are traumatic, relatively large, with irregular outlines, and without typical epithelial cells that form a continuous lining around a resin canal [11].
The absence of true ray tracheids is also an important characteristic of Keteleeria [22]. The marginal ray cells of Keteleeria have a slightly greater number of pits on the radial walls and undulate slightly convex external walls. The phenomenon can be also observed in the wood of Abies and Pseudolarix [22]. However, the two taxa do not possess normal resin canals.
The presence of axial parenchyma is also the characteristic of Keteleeria, although it is scarce [21,22,23].
Shilkina proposed the generic name Keteleerioxylon for the fossil woods with the anatomical characteristics of the extant Keteleeria [7]. Pinoxylon (Knowlton) Read or Protopiceoxylon Gothan (the generic name is controversial) shows similar characteristics with Keteleerioxylon in possessing sole vertical resin canals [25,26,27,28,29]. As clearly explained by Philippe and Bamford, Pinoxylon or Protopiceoxylon shows araucarian, or araucarian and abietinean pitting on the radial walls of tracheids, while the pits on the tracheid radial walls of Keteleerioxylon are abietinean, or slightly alternate when biseriate [29]. In the wood of Keteleeria, the tracheid pitting in radial walls is abietinean, but occasionally shows a tendency toward “alternate” [30]. The radial pitting of the studied fossil wood is quite like extant Keteleeria. Thus, the present specimen should be assigned to the genus Keteleerioxylon.
The present fossil woods differ from the extant species of Keteleeria in having uniseriate to triseriate pits on the radial walls of tracheids, in lacking pits on tangential walls of tracheids, and in possessing taxodioid and cupressoid cross-field pits (Table 1).
The fossil wood genus Keteleerioxylon was established from the materials collected from the Early Cretaceous Franz Josef Land, Russia by Shilkina [7]. Until now, four species of Keteleerioxylon have been described. K. arcticum Shikkina, 1960 differs from K. changchunense in having uniseriate to biseriate pits on the radial walls of tracheids, in possessing pits on tangential walls of tracheids, in lacking marginal ray cells of the ray tracheid type, and in having fewer pits per cross-field [7]. K. fokinii Shilkina, 1986 was found in the Early Cretaceous Kirovsk Region of Russia [34]. K. fokinii differs from the present woods in having uniseriate to biseriate pits on the radial walls of tracheids, in the absence of crassula, and in the taxodioid cross-field pits. K. primoryense Blokhina, 2000 from the Oligocene–Miocene of Primorye differs from K. changchunense in having uniseriate to biseriate pits on the radial walls of tracheids, in possessing pits on tangential walls of tracheids, and in the cupressoid cross-field pits [35]. K. kamtshatkiense Blokh. et Afonin, 2006, from the Lower Cretaceous of the Kamchatka Peninsula, Russia, is different from the woods under the description, in having uniseriate to biseriate pits on the radial walls of tracheids, in possessing pits on tangential walls of tracheids, and in the absence of crassula [11]. Jiang et al. reported that the Keteleeria liaoxiense and Keteleeria sp. were discovered from the Middle Jurassic Liaoning Province, but they did not give a description [36].
There are many woods that were identified as Keteleeria. Keteleeria mabetiensis Watari, 1941 was collected from the Lower Miocene of Japan [37,38]. Keteleeria mabetiensis differs from Keteleerioxylon changchunense in possessing pits on tangential walls of tracheids, in knotty transverse walls of axial parenchyma, and taxodioid and piceooid cross-field pits. Keteleeria zhilinii Blokhina et Bondarenko, 2005 was first discovered in the Pliocene of Primorye, Russia [10]. K. zhilinii differs from the woods under study in having uniseriate to biseriate pits on the radial walls of tracheids, in having pits on tangential walls of tracheids, and in the taxodioid cross-field pits. Yang et al. discovered a silicified wood belonging to the genus Keteleeria in the Late Cretaceous of Henan, China [39]. They named these woods after the extant species “Keteleeria fortunei”. “Keteleeria fortunei” is different from Keteleerioxylon changchunense in having uniseriate to biseriate pits on the radial walls of tracheids and in the absence of axial parenchyma. Keteleeria sp. from the early Holocene of Hubei, China, differs from Keteleerioxylon changchunense in having uniseriate to biseriate pits on the radial walls of tracheids and in the taxodioid cross-field pits [40]. Keteleeria sp. from Guangdong, China differs from the woods under study in having uniseriate to biseriate pits on the radial walls of tracheids, in lower rays, and in having fewer pits per cross-field [12].
For the details of each fossil record, please see Table S1.

4. Discussion

4.1. Paleoecological and Paleoclimatic Implications

Quantitative analysis of growth rings can be used to detect whether the conifer species is evergreen or deciduous, and how long the leaf longevity is [20,41]. Through the measuring of radial diameters of five adjacent files of tracheid cells, four parameters were calculated, including (1) percentage of latewood, (2) percentage of cell diminution in a ring increment, (3) RMI, and (4) percentage of skews of CSDM curves (Table 2, Supplementary data).
Deciduous conifers have dominantly left-skewed CSDM or symmetrical curves, whereas evergreen conifers mainly have right-skewed CSDM curves. The CSDM curves of Keteleerioxylon changchunense are from 0 to +20.00% (mean percentage of skew +8.77%), right-skewed, suggesting that this species was evergreen (Figure 5, Table 3).
The percentage of latewood in Keteleerioxylon changchunense is 40.00–50.00%, with a mean of 45.62%; and the percentages of the CSDM curves skews range from 0–20.00% (mean 8.77%). These two parameters are quite close to those of Pinus sylvestris and Picea abies, while the percentage of cell diminution (89.39–97.50%, mean 95.45%) and the RMI (38.92–48.75%, mean 43.51%) are higher than those of Larix decidua (Table 2). Therefore, the leaf retention time of Keteleerioxylon changchunense is considered to be 1–3 years, the most likely approach to the extent of Keteleeria [42].
The CSDM curves are widely used for identifying evergreen or deciduous habits. However, an exception comes from the Late Pleistocene Keteleeria sp. of Guangdong, China, as its CSDM curves showed both left and right skews [12]. However, the radial cells in three growth rings of Keteleeria sp. are less than 15; this might result from false rings triggered by the East Asian monsoon. Therefore, we still think the right skew of CSDM curves is reliable in recognizing evergreen trees like K. changchunense.
The earliest Albian climate is regarded as a greenhouse with high atmospheric carbon dioxide concentration [19]. The global temperature in the Aptian and Albian periods would have been much higher than today [43], and the Arctic area was covered by evergreen broadleaf mixed with deciduous conifer forests in the Late Cretaceous period [44]. The growth ring RMI is considered as an indicator of the climatic seasonality intensity [20]. Keteleerioxylon changchunense shows very marked growth rings (high RMI) (Table 3). This might indicate that the climate seasonality was likely to be strong in the earliest Albian period in the Songliao Basin, Northeast China.

4.2. Phytogeographical Distribution

As mentioned above, Keteleerioxylon and Protopiceoxylon or Pinoxylon show great similarity to the extant Keteleeria. According to Blokhina et al., there is a distinct possibility that they are the remote ancestors of Keteleeria. Although extant Keteleeria is endemic to East and Southeast Asia, the Keteleeria-like fossils (including Keteleeria, Keteleerioxylon, and Protopiceoxylon or Pinoxylon) had been widely distributed in Europe, North America, and Asia in the northern hemisphere since the Middle Jurassic period (Table 4).
The earliest record of a Keteleeria-like fossil was discovered in Middle Jurassic Svalbard, Norway. The Late Jurassic-Early Cretaceous Keteleeria-like trees are widely distributed from western Liaoning, China to the Arctic areas (Figure 6). Their paleolatitudes range from 36° N to 85° N (Figure 6). This indicates that the climate might be warm and suitable for these subtropical thermophilic Keteleeria-like trees in the Late Jurassic-Early Cretaceous. In the Late Cretaceous, Keteleeria-like trees were still persistently distributed in the middle and high latitudes of the Russian far east and northeast China. After the K-Pg mass extinction, records of Keteleeria fossils were rare in the Paleocene [56,63]. In the Eocene, Keteleeria migrated to middle latitudes [6,64]. This migration was contemporary with the Paleocene–Eocene thermal maximum [88,89]. During the Oligocene and Miocene, Keteleeria was widely distributed in the mid-latitudes of North America, Europe, and east Asia in the northern hemisphere. From the Pliocene on, Keteleeria started to retreat southward from Primorye, Russia to the subtropical and tropical monsoon climate regions of South China and Southeast Asia in the middle and low latitudes (~13.8° N–35.5° N) in the Quaternary. Therefore, the palaeogeographical distribution of Keteleeria-like was closely related to the paleoclimate changes (Figure 6).

5. Conclusions

In this study, the species Keteleerioxylon changchunense Shi, Sun, Meng et Yu sp. nov. is described for the first time in the Late Early Cretaceous deposits of Changchun, Northeast China. The quantitative growth ring analysis of the two specimens shows that the species is evergreen, with the foliar long around 1–3 years. This is quite consistent with the habits of extant Keteleeria. The presence of very marked growth rings indicates that it lived in a seasonal climate during the earliest Albian period in the Songliao Basin, northeast China. The palaeogeographical distribution of the Keteleeria-like fossils indicates that they were distributed in both the middle and the high latitudes during the Late Jurassic–Cretaceous global warming time, while they retreated to the middle latitude in the Cenozoic due to global cooling, and they have only survived in east and southeast Asia after the Quaternary glaciations until now.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/biology11111624/s1, Table S1: Comparison of wood anatomical characters of Keteleerioxylon changchunense sp. nov. and closely reLated fossil species. Supplementary data: Cell diameter for CSDM curve.

Author Contributions

Conceptualization, X.S. and Y.S.; field sampling, data processing, and measurement, X.S., Y.S., F.M., J.Y. and Z.L.; writing—original draft preparation, X.S.; writing—review and editing, Y.S. All authors contributed suggestions and revised the article. All authors have read and agreed to the published version of the manuscript.

Funding

This research was supported by the National Natural Sciences Foundation of China (Nos. 92055201, 41972012 and 31700183).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The data presented in this study are available on request from the corresponding author.

Acknowledgments

We are greatly indebted to the editor and the three anonymous reviewers for their critical revision of an earlier version of the manuscript. We thank Ning Tian (Shenyang Normal University) and Zikun Jiang (Chinese Academy of Geological Sciences) for their constructive and helpful suggestions. Xiaogang Wu is thanked for his assistance in making thin sections. We especially thank Wei Sun and Zhen Xu, who helped improve the English.

Conflicts of Interest

The authors declare no conflict of interest.

References

  1. Carriére, E.A. Keteleeria fortunei (Murr.) Carriére comb. nov. Rev. Hortic. 1866, 37, 449–451. [Google Scholar]
  2. Wang, C.-Y.; Ma, S.B.; Lv, J.; Dang, C.-L. Ecological and geographical distribution of Keteleeria and its systematic evolution in China. Guihaia 2012, 32, 612–616. [Google Scholar] [CrossRef]
  3. Farjon, A. A second revision of the genus Keteleeria Carrière (Taxonomic notes on Pinaceae II). Misc. Publ. Univ. Utrecht Herb. 1989, 46, 81–99. [Google Scholar]
  4. Fu, L.-K.; Li, N.; Mill, R.R. Keteleeria Carriére. In Flora of China; Wu, Z.-Y., Raven, P.H., Hong, D.-Y., Eds.; Science Press: Beijing, China, 1999; Volume 4, pp. 42–44. [Google Scholar]
  5. Nguyen, T.H.; Phan, K.L.; Nguyen, D.T.L.; Thomas, P.I.; Farjon, A.; Averyanov, L.; Regalado, J., Jr. Vietnam conifers: Conservation Status Review 2004; Fauna and Flora International Vietnam Programme: Hanoi, Vietnam, 2005; pp. 69–72. [Google Scholar]
  6. Averyanov, L.V.; Nguyen, T.H.; Sinh, K.N.; Pham, T.V.; Lamxay, V.; Bounphanmy, S.; Lorphengsy, S.; Phan, L.K.; Lanorsavanh, S.; Chantthavongsa, K. Gymnosperms of Laos. Nord. J. Bot. 2014, 32, 765–805. [Google Scholar] [CrossRef]
  7. Shilkina, I.A. Wood of a new conifer genus and species from the Lower Cretaceous Deposits of Franz Josef Land. Paleontol. Zhurnal 1960, 1, 113–117. [Google Scholar]
  8. Ran, J.-H.; Shen, T.-T.; Wu, H.; Gong, X.; Wang, X.-Q. Phylogeny and evolutionary history of Pinaceae updated by transcriptomic analysis. Mol. Phylogenetices Evol. 2018, 129, 106–116. [Google Scholar] [CrossRef]
  9. Walton, J. On some fossil woods of Mosozoic and Tertiary age from the Arctic zone. Ann. Bot. 1927, 41, 239–252. [Google Scholar] [CrossRef]
  10. Blokhina, N.I.; Bondarenko, O.V. Fossil Wood of Keteleeria (Pinaceae) from the Pliocene of Southern Primorye (Far East of Russia). Bot. Zhurnal 2005, 90, 937–946. [Google Scholar]
  11. Blokhina, N.I.; Afonin, M.A.; Popov, A.M. Fossil wood of Keteleerioxylon kamtschatkiense sp. nov. (Pinaceae) from the Cretaceous of the Northwestern Kamchatka Peninsula. Paleontol. J. 2006, 40, 678–686. [Google Scholar] [CrossRef]
  12. Huang, L.-L.; Jin, J.-H.; Oskolski, A.A. Mummified fossil of Keteleeria from the Late Pleistocene of Maoming Basin, South China, and its phytogeographical and paleoecological implications. J. Syst. Evol. 2019, 59, 198–215. [Google Scholar] [CrossRef]
  13. Bureau of Geology and Mineral Resource of Jilin Province. Stratigraphy (Lithostratigraphy) of Jilin Province; China University of Geosciences Press: Wuhan, China, 1997. (In Chinese) [Google Scholar]
  14. Ding, R.-X.; Shu, P.; Ji, X.-Y.; Qu, Y.-M.; Cheng, R.-H.; Zhang, B. SHRIMP zircon U-Pb age and geological meaning of reservoir volcanic rocks in Qingshen gas field of the Songliao basin, NE China. J. Jilin Univ. (Earth Sci. Ed.) 2007, 37, 525–530, (In Chinese with English abstract). [Google Scholar] [CrossRef]
  15. Zhang, F.-Q.; Cheng, X.-G.; Chen, H.-L.; Dong, C.-W.; Yu, X.; Xiao, J.; Xu, Y.; Pang, Y.-M.; Shu, P. Zircon chronological and geochemical constraints on the late Mesozoic volcanic events in the southeastern margin of the Songliao Basin, NE China. Acta Petrol. Sin. 2009, 25, 39–54, (In Chinese with English abstract). [Google Scholar] [CrossRef]
  16. Shang, Y.-K.; Wang, S.-Y. Palynomorph assemblages from the Yingcheng Formation, Jiutai, Jilin. Acta Micropalaeontologica Sin. 1991, 8, 91–110, (In Chinese with English abstract). [Google Scholar]
  17. Wang, S.-Y. Palynological assemblages of Yingcheng Formation, Jilin Province. J. Stratigr. 1989, 13, 34–39. (In Chinese) [Google Scholar] [CrossRef]
  18. Jia, J.-T.; Wang, P.-J.; Wan, X.-Q. Chronostratigraphy of the Yingcheng Formation in the Songliao Basin, Cretaceous, NE China. Geol. Rev. 2008, 54, 439–448, (In Chinese with English abstract). [Google Scholar] [CrossRef]
  19. Sun, Y.-W.; Li, X.; Zhao, G.-W.; Liu, H.; Zhang, Y.L. Aptian and Albian atmospheric CO2 changes during oceanic anoxic events: Evidence from fossil Ginkgo cuticles in Jilin Province, Northeast China. Cretac. Res. 2016, 62, 130–141. [Google Scholar] [CrossRef]
  20. Falcon-Lang, H.J. The relationship between leaf longevity and growth ring markedness in modern conifer woods and its implications for palaeoclimatic studies. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2000, 160, 317–328. [Google Scholar] [CrossRef]
  21. Yatsenko-Khmelevskii, A.A. Woods of the Caucasus Mountains; Akademii Nauk Armyanskoi SSSR: Yerevan, Armenia, 1954. (In Russian) [Google Scholar]
  22. Budkevich, E.V. Wood of the Pinaceae; Akad. Nauk.; SSSR: Moscow, Russia, 1961. (In Russian) [Google Scholar]
  23. Chavchavadze, E.S. Wood of the Pinaceae; Akademii Nauk SSSR; SSSR: Leningrad, Russia, 1979. (In Russian) [Google Scholar]
  24. Lin, J.-X.; Liang, E.-Y.; Farjon, A. The occurrence of vertical resin canals in Keteleeria, with reference to its systematic position in Pinaceae. Bot. J. Linn. Soc. 2000, 134, 567–574. [Google Scholar] [CrossRef]
  25. Knowlton, F.H. Fossil wood from Jurassic of the Black Hills. In Twentieth Annual Report of the Director of the United States Geological Survey. Part 2, Gegeral Geology and Paleontology; Walcott, C.D., Ed.; Government Printing Office: Washington, DC, USA, 1900. [Google Scholar]
  26. Read, C.B. Pinoxylon dakotense knowlton from th Cretaceous of the Black Hills. Bot. Gaz. 1932, 93, 173–187. [Google Scholar] [CrossRef]
  27. Gothan, W. Die fossilen Hölzer von König-Karls-Land. K. Sven. Vetensk. Handl. 1907, 42, 1–44. [Google Scholar]
  28. Shengyang Center for Geological Survey, Shenzhen Fairy Lake Botanic Garden. An Introduction to the World’s Petrified Wood; Geological Publishing House: Beijing, China, 2015. [Google Scholar]
  29. Philippe, M.; Bamford, M.K. A key to morphogenera used for Mesozoic conifer-like woods. Rev. Palaeobot. Palynol. 2008, 148, 184–207. [Google Scholar] [CrossRef]
  30. IAWA Committee. List of Microscopic Features for Softwood Identification. IAWA J. 2004, 25, 1–70. [Google Scholar] [CrossRef]
  31. Bailey, I.W. The Cambium and Its Derivative Tissues: VII. Problems in Identifying the Wood of Mesozoic Coniferae. Ann. Bot. 1933, 47, 145–157. [Google Scholar] [CrossRef]
  32. Greguss, P. Xylotomische Bestimmung der Heute Lebenden Gymnospermen; Akadémiai Kiadó-MET Publishing Corp.: Budapest, Hungary, 1955. [Google Scholar]
  33. Chavchavadze, E.S. Age-related changes in morphologicalcharacters of parenchyma in secondary wood of Pines (Pinaceae). In Problems in Comparative Morphology of Seed Plants; Budantsev, L.Y., Ed.; Nauka: Leningrad, Russia, 1975; pp. 165–190. (In Russian) [Google Scholar]
  34. Shilkina, I.A. Fossil woods of northern Eurasia. In Problems in Palaeobotany; Takhtajan, A.L., Ed.; Nauka: Leningrad, Russia, 1986; pp. 150–157. (In Russian) [Google Scholar]
  35. Blokhina, N.I.; Klimova, R.S. Keteleeria from the Tertiary deposits of the Middle Sikhote Alin (Primorye). Paleontol. J. 2000, 34, 105–112. [Google Scholar]
  36. Jiang, Z.-K.; Wang, Y.-D.; Tian, N.; Xie, A.-W.; Zhang, W.; Li, L.-Q.; Huang, M. The Jurassic fossil wood diversity from western Liaoning, NE China. J. Palaeogeogr. 2019, 8, 1–11. [Google Scholar] [CrossRef] [Green Version]
  37. Watari, S. Studies on the fossil woods from the Tertiary of Japan: I. Fossil woods from the river Mabeti, Anatai village, Ninohe district, Iwate Prefecture. J. Jpn. Bot. 1941, 11, 385–416. [Google Scholar]
  38. Watari, S. Some abietinean woods from the Tertiary of Japan. J. Fac. Sci. Univ. Tokyo Sect. III Bot. 1956, 6, 419–437. [Google Scholar]
  39. Yang, J.-J.; Qi, G.-F.; Xu, R.-H. Studies on fossil woods excavated from the Dabie Mountains. Sci. Silvae Sin. 1990, 26, 379–386, (In Chinese with English abstract). [Google Scholar]
  40. Yang, J.-J.; Qi, G.-F.; Xu, R.-H.; Fan, S.-K. Studies on ancient wood of Keteleeria sp. excavated in Wuhan, Hubei Province. Sci. Silvae Sin. 2003, 39, 173–176, (In Chinese with English abstract). [Google Scholar] [CrossRef]
  41. Falcon-Lang, H.J. A method to distinguish between woods produced by evergreen and deciduous coniferopsids on the basis of growth ring anatomy: A new palaeoecological tool. Palaeontology 2000, 43, 785–793. [Google Scholar] [CrossRef]
  42. Zhou, Z.-Z.; Yuan, L.; Wang, Z.-H. Leaf longevity of main evergreen tree species of Guiyang City. Ecol. Sci. 2008, 27, 148–153, (In Chinese with English abstract). [Google Scholar]
  43. Skelton, P.W.; Spicer, R.A.; Kelley, S.P.; Gilmour, I. The Cretaceous World; Cambridge University Press: Cambridge, UK, 2003; pp. 1–360. [Google Scholar]
  44. Spicer, R.A.; Herman, A.B. The Late Cretaceous environment of the Arctic: A quantitative reassessment based on plant fossils. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2010, 295, 423–442. [Google Scholar] [CrossRef] [Green Version]
  45. Seward, A.C. The Jurassic Flora. Part II. Liassic and Oolitic Floras of England; Catalogue of Mesozoic Plants in the British Muscum (Natl. Hist.): London, UK, 1904. [Google Scholar]
  46. Medlyn, D.A.; Tidwell, W.D. A review of the genus Protopiceoxylon with emphasis on North American species. Can. J. Bot. 1979, 57, 1451–1463. [Google Scholar] [CrossRef]
  47. Jiang, H.-E.; Ferguson, D.K.; Li, C.-S.; Cheng, Y.-M. Fossil coniferous wood from the Middle Jurassic of Liaoning Province, China. Rev. Palaeobot. Palynol. 2008, 150, 37–47. [Google Scholar] [CrossRef]
  48. Nishida, M.; Nishida, H. Pinoid woods with resin canals from the Upper Cretaceous of Hokkaido and Saghalien. J. Plant Res. 1995, 108, 161–170. [Google Scholar] [CrossRef]
  49. Mathews, G.B.; Ho, G.A. On the Occurrence of Protopiceoxylon in China. Geobiologia 1945, 2, 27–35. [Google Scholar]
  50. Sze, H.C. Fossil woods of gymnosperm. In Fossil plants of China. Mesozoic Plants from China II; Sze, H.C., Li, X.X., Li, P.J., Eds.; Science Press: Beijing, China, 1963; pp. 316–347. (In Chinese) [Google Scholar]
  51. Wang, Y.D.; Zhang, W.; Saiki, K. Fossil woods from the Upper Jurassic of Qitai, Junggar Basin, Xinjiang, China. Acta Palaeontol. Sin. 2000, 39, 176–185. [Google Scholar]
  52. Shimakura, M. Studies on fossil woods from Japan and adjacent lands, Contribution I. Sci. Rep. Tohoku Imp. University. Ser. 2 (Geol.) 1936, 18, 267–310. [Google Scholar]
  53. Stopes, M.C. Catalogue of the Mesozoic Plants in the British Museum. The Cretaceous Flora, Part II, Lower Greensand (Aptian) Plants of Britain; British Museum of Natural History: London, UK, 1915. [Google Scholar]
  54. Du, N.-Z. Two fossil woods from Heilongjiang Province of China. Acta Bot. Sin. 1982, 24, 383–387, (In Chinese with English abstract). [Google Scholar]
  55. Wang, R.-F.; Wang, Y.F.; Chen, Y.Z. Fossil woods from Late Cretaceous of Heilongjiang Province, Northeast China, and their palaeoenviromental implications. Acta Bot. Sin. 1997, 39, 972–978, (In Chinese with English abstract). [Google Scholar]
  56. Terada, K.; Nishida, H.; Sun, G. Fossil woods from the Upper Cretaceous to Paleocene of Heilongjiang (Amur) River area of China and Russia. Glob. Geol. 2011, 14, 192–208. [Google Scholar] [CrossRef]
  57. Duan, S.-Y. Several fossil woods from Mesozoic of western Liaoning Province, Northeast China. Acta Bot. Sin. 2000, 42, 207–213, (In Chinese with English abstract). [Google Scholar] [CrossRef]
  58. Duan, S.-Y.; Cui, J.-Z.; Wang, X.; Xiong, B.K.; Wang, Y.Q. Fossil woods from the Early Cretaceous of western Liaoning, China. In Wood Anatomy Research, Proceedings of the International Symposium, Tree Anatomy & Wood Formation, Tianjin, China, 24–26 October 1995; Wu, S., Ed.; International Academic Publishers: Beijing, China, 1995; pp. 166–171. [Google Scholar]
  59. Schröter, C. Untersuchung über fossile Hölzer aus der arctischen Zone. Flora Fossilis Arctica 1880, 6, 1–38. [Google Scholar]
  60. Edwards, W.M. On Protopiceoxylon johnseni (Schröeter), a Mesozoic coniferous wood. Ann. Bot. 1925, 39, 1–7. [Google Scholar] [CrossRef]
  61. Ding, Q.-H. Protopiceoxylon mohense sp. nov. from the Jiufengshan Formation in Heilongjiang Province. Chin. Bull. Bot. 2000, 17, 206–209, (In Chinese with English abstract). [Google Scholar]
  62. Miki, S.; Maeda, Y. Fossil cone of Keteleeria cretacea sp. nov. from Upper Cretaceous beds in Awaji, Japan. Chigaku Kenkyu (Geosci. Mag. Tokyo Jpn.) 1966, 17, 248–251. (In Japanese) [Google Scholar]
  63. Dolezych, M.; Reinhardt, L. Protopiceoxylon yukonense sp. nov.–A remarkable extinct conifer from the Mackenzie Delta area, northern Yukon. Yukon Explor. Geol. Yukon Yukon Geol. Surv. 2016, 99–110. [Google Scholar]
  64. The Writing Group of Cenozoic Plants of China. Fossil Plants of China, Vol. 3, Cenozoic Plants from China; Science Press: Beijing, China, 1978. (In Chinese) [Google Scholar]
  65. Terada, K. Fossil wood floras of Early to Middle Miocene in Japan. Ph.D. Thesis, Tohoku University, Sendai, Japan, 1998. (In Japanese). [Google Scholar]
  66. Choi, S.K.; Kim, K.; Jeong, E.K.; Terada, K.; Suzuki, M.; Uematsu, H. Fossil woods from the Miocene in the Yamagata Prefecture, Japan. IAWA J. 2010, 31, 95–117. [Google Scholar] [CrossRef]
  67. Rybalko, V.I.; Ovechkin, V.N.; Klimova, R.S. Cenozoic basaltic rocks of the Amga Group (northeastern Primorye). Sov. Geol. 1980, 12, 59–71. [Google Scholar]
  68. Lakhanpal, R.N. The Rujada flora of west central Oregon. Univ. Calif. Publ. Geol. Sci. 1958, 35, 1–66. [Google Scholar]
  69. Meyer, W.H.; Manchester, S.R. The Oligocene Bridge Creek Flora of the John Day Formation Oregon; University of California Press: Berkeley, CA, USA, 1997. [Google Scholar]
  70. Hofmann, E. Pflanzenreste aus dem Phosphoritvorkommen von Prambachkirchen in Oberdonau. I. Teil. Palaeontographica 1944, 88, 1–24. [Google Scholar]
  71. Klaus, W. Neue fossile Pinaceen-Reste aus dem österreichischen Jungtertiär. Beiträge Zur Paläontologie Von Österreich 1977, 3, 1–120. [Google Scholar]
  72. Kräusel, R. Die tertiäre Flora der Hydrobienkalke von Mainz-Kastel. Paläontologische Z. 1938, 20, 9–103. [Google Scholar] [CrossRef]
  73. Ananova, E.N. Pollen in Neogene Deposits of the Southern Russian Plain; Leningrad State University: Leningrad, Russia, 1974. [Google Scholar]
  74. Ramischvili, I. Ponticheskaya Flora Zapadnoi Gruzii po Dannym Palinologicheskogo Analiza; Metsniereba: Tbilisi, Georgia, 1969. [Google Scholar]
  75. Miki, S. On the change of flora in Eastern Asia since Tertiary Period (I). The clay or lignite beds flora in Japan with special reference to the Pinus trifolia beds of central Hondo. Jpn. J. Bot. 1941, 11, 237–303. [Google Scholar]
  76. Miki, S. Pinaceae of Japan, with special reference to its remains. J. Inst. Polytech. Osaka City Univ. Ser. D 1957, 8, 221–272. [Google Scholar]
  77. Miki, S. Gymnosperms in Japan, with special reference to the remains. J. Inst. Polytech. Osaka City Univ. Ser. D 1958, 9, 125–152. [Google Scholar]
  78. Tanai, T. Neogene floral change in Japan. J. Fac. Sci. Hokkaido Univ. 1961, 11, 119–398. [Google Scholar]
  79. Tanai, T.; Suzuki, N. Miocene floras of southwestern Hokkaido, Japan. In Tertiary Floras of Japan. 2. Miocene Floras; Chane, R.W., Tanai, T., Eds.; The Collaborating Association to Commemorate the 80th Anniversary of the Geological Survey of Japan, Tokyo: Tokyo, Japan, 1963; pp. 1–27. [Google Scholar]
  80. Kamoi, Y.; Kobayashi, I.; Suzuki, K. The middle Miocene Osudo fossil flora in the northern part of Niigata Prefecture. J. Geol. Soc. Jpn. 1978, 84, 15–21. [Google Scholar] [CrossRef]
  81. Ozaki, K. Late Miocene Tatsumitoge flora of Tottori Prefecture, southwest Honshu, Japan (I). Sci. Rep. Yokohama Natl. Univ. Sect. II 1979, 26, 31–56. [Google Scholar]
  82. Wang, Y.-F.; Xiang, Q.-P.; Ferguson, D.K.; Zastawniak, E.; Yang, J.; Li, C.-S. A new species of Keteleeria (Pinaceae) in the Shanwang Miocene flora of China and its phytogeographic connection with North America. Taxon 2006, 55, 165–171. [Google Scholar] [CrossRef]
  83. Kirchheimer, F. Zur Kenntnis der Alttertiärflora von Wiesa bei Kamenz (Sachsen). Planta 1942, 32, 418–446. [Google Scholar] [CrossRef]
  84. Kunzmann, L.; Mai, D.H. Die Koniferen der Mastixioideen-Flora von Wiesa bei Kamenz (Sachsen, Miozän) unter besonderer Berücksichtigung der Nadelblätter. Palaeontogr. B 2005, 272, 67–135. [Google Scholar]
  85. Zalewska, A. The fossil flora of Turow near Botatynia. Prace Mus. Ziemi 1961, 4, 19–49. [Google Scholar]
  86. Hantke, R. Keteleeria hoehnei Kirchh., ein Zapfenrest aus der Unteren Süsswassermolasse des Buechbergs (Kt. Schwyz). Ecol. Geol. Helv. 1973, 66, 739–742. [Google Scholar]
  87. Brown, R.W. Miocene leaves, fruits, and seeds from Idaho, Oregon, and Washington. J. Paleontol. 1935, 9, 572–587. [Google Scholar]
  88. Gingerich, P.D. Environment and evolution through the Paleocene–Eocene thermal maximum. Trends Ecol. Evol. 2006, 21, 246–253. [Google Scholar] [CrossRef]
  89. McInerney, F.A.; Wing, S.L. The Paleocene-Eocene Thermal Maximum: A perturbation of carbon cycle, climate, and biosphere with implications for the future. Annu. Rev. Earth Planet. Sci. 2011, 36, 489–516. [Google Scholar] [CrossRef] [Green Version]
  90. Scotese, C.R.; Song, H.; Mills, B.J.; van der Meer, D.G. Phanerozoic paleotemperatures: The earth’s changing climate during the last 540 million years. Earth-Sci. Rev. 2021, 215, 103503. [Google Scholar] [CrossRef]
Biology 11 01624 g001 550
Figure 1. Distribution of the extant Keteleeria species and variants in the world (Adapted with permission from Ref. [2]).
Figure 1. Distribution of the extant Keteleeria species and variants in the world (Adapted with permission from Ref. [2]).
Biology 11 01624 g001
Biology 11 01624 g002 550
Figure 2. Location of Keteleerioxylon changchunense Shi, Sun, Meng et Yu sp. nov. in the Yingcheng Coal Mine (red star), Changchun City, Jilin Province, Northeast China.
Figure 2. Location of Keteleerioxylon changchunense Shi, Sun, Meng et Yu sp. nov. in the Yingcheng Coal Mine (red star), Changchun City, Jilin Province, Northeast China.
Biology 11 01624 g002
Biology 11 01624 g003 550
Figure 3. Keteleerioxylon changchunense Shi, Sun, Meng et Yu sp. nov. (A). General view of the specimen JLJY-02. (B). Transverse section showing the resin canals (red arrows) and growth ring boundaries. (C). Close-up of the resin canals (red arrow). (D). Close-up of the growth ring boundary. (E). Radial section showing the vertical resin canal. (F). Radial section showing the biseriate pits on the radial walls of tracheids and the crassulae. (G,H). Radial section showing the opposite and alternate biseriate pits and triseriate pits and the crasulae on the radial walls of tracheids. Scale bar: A: the longer scale mark in the card is 1 cm; B = 1 mm; C, D, E = 200 μm; F, G, H = 50 μm.
Figure 3. Keteleerioxylon changchunense Shi, Sun, Meng et Yu sp. nov. (A). General view of the specimen JLJY-02. (B). Transverse section showing the resin canals (red arrows) and growth ring boundaries. (C). Close-up of the resin canals (red arrow). (D). Close-up of the growth ring boundary. (E). Radial section showing the vertical resin canal. (F). Radial section showing the biseriate pits on the radial walls of tracheids and the crassulae. (G,H). Radial section showing the opposite and alternate biseriate pits and triseriate pits and the crasulae on the radial walls of tracheids. Scale bar: A: the longer scale mark in the card is 1 cm; B = 1 mm; C, D, E = 200 μm; F, G, H = 50 μm.
Biology 11 01624 g003
Biology 11 01624 g004 550
Figure 4. Keteleerioxylon changchunense Shi, Sun, Meng et Yu sp. nov. (A). Radial section showing the opposite and alternate biseriate pits and triseriate pits and the crasulae on the radial walls of tracheids. (B). Radial section showing pitted horizonal and end walls of the ray parenchyma cells. (C). Tangential section showing the axial parenchyma cell. (D,E). Radial section showing taxodioid-cupressoid type cross-field pits. (F). Radial section showing the marginal ray cells resembling ray tracheids with scattered uniseriate pits (red arrows). (G). Tangential section showing uniseriate, partly biseriate rays. (H). Close-up of the rays showing uniseriate rays. (I). Tangential section showing the pitted end wall of ray parenchyma cell (red arrow). Scale bar: A, B, D, E, F, I = 50 μm; C, H = 100 μm; G = 200 μm.
Figure 4. Keteleerioxylon changchunense Shi, Sun, Meng et Yu sp. nov. (A). Radial section showing the opposite and alternate biseriate pits and triseriate pits and the crasulae on the radial walls of tracheids. (B). Radial section showing pitted horizonal and end walls of the ray parenchyma cells. (C). Tangential section showing the axial parenchyma cell. (D,E). Radial section showing taxodioid-cupressoid type cross-field pits. (F). Radial section showing the marginal ray cells resembling ray tracheids with scattered uniseriate pits (red arrows). (G). Tangential section showing uniseriate, partly biseriate rays. (H). Close-up of the rays showing uniseriate rays. (I). Tangential section showing the pitted end wall of ray parenchyma cell (red arrow). Scale bar: A, B, D, E, F, I = 50 μm; C, H = 100 μm; G = 200 μm.
Biology 11 01624 g004
Biology 11 01624 g005 550
Figure 5. The CSDM curves of five growth rings and cell diameters of growth ring increment. EW: Earlywood, LW: latewood. The arrows indicate the zeniths of CSDM curves. When the abscissa value of the zenith is greater than half of the cell number, the curve is right-skewed; on the contrary, it is left-skewed.
Figure 5. The CSDM curves of five growth rings and cell diameters of growth ring increment. EW: Earlywood, LW: latewood. The arrows indicate the zeniths of CSDM curves. When the abscissa value of the zenith is greater than half of the cell number, the curve is right-skewed; on the contrary, it is left-skewed.
Biology 11 01624 g005
Biology 11 01624 g006 550
Figure 6. Geographical and geological distribution of Keteleeria-like fossils (red stars) in the world (global average temperature data from [90]).
Figure 6. Geographical and geological distribution of Keteleeria-like fossils (red stars) in the world (global average temperature data from [90]).
Biology 11 01624 g006
Table
Table 1. Comparison of wood anatomical characteristics of Keteleerioxylon changchunense sp. nov. and woods of extant Keteleeria.
Table 1. Comparison of wood anatomical characteristics of Keteleerioxylon changchunense sp. nov. and woods of extant Keteleeria.
Anatomical CharactersKeteleeria davidiana (Bertr.) Beissner [11,22,31,32,33]Keteleeria evelyniana Masters [11,22]Keteleeria fortunei (Andr. Murray) Carr. [11,21,22,32]Keteleerioxylon changchunense sp. nov.
Pits on radial walls of tracheidsUniseriate to tetraseriateUniseriate to biseriateUniseriate to biseriateUniseriate to triseriate
diameter of pits, μm(10)12–15(16)15–2112–15(16)(11)12–17(20)
Pits on tangential walls of tracheidsPresentPresentPresentAbsent
CrassulaePresentPresentNo dataPresent
Uniseriate rays:
height (in cells)
1–30(34)
1–30
1–40
(1)6–11(37)
number of biseriate layers1–8UncommonPresent1–8
marginal ray cells of the ray tracheid typePresentPresentPresentPresent
Transverse walls of axial parenchymaKnotty?KnottySmooth
Number of epithelial cells in vertical resin canals6–7(8–11)66–86–11
Pitting on cross-fields:
number of pits		1–3(6)1–41–4(6)1–3(6)
diameter of pits, μm(4)5–8(12)No data4–65–10
type of pittingTaxodioid, piceoid, and pinoidtaxodioidtaxodioidTaxodioid and cupressoid
Table
Table 2. Results of the quantitative growth ring analysis of Keteleerioxylon changchunense sp. nov.
Table 2. Results of the quantitative growth ring analysis of Keteleerioxylon changchunense sp. nov.
Ring NumberPercentage LatewoodPercentage DiminutionRing Markedness Index (RMI)Percentage Skews
Ring A48.00%97.06%46.42%+4.35%
Ring B50.00%97.50%48.75%0%
Ring C48.00%89.39%42.76%+4.35%
Ring D40.00%97.30%38.92%+20.00%
Ring E42.00%96.00%40.72%+15.15%
Averages45.60%95.45%43.51%+8.77%
Table
Table 3. Comparison of the quantification of ring markedness parameters for Keteleerioxylon changchunense sp. nov. with the five extant taxa (data from [20]).
Table 3. Comparison of the quantification of ring markedness parameters for Keteleerioxylon changchunense sp. nov. with the five extant taxa (data from [20]).
Tree HabitsSpeciesLeaf Retention Time (in years)Percentage of Latewood (%)Percentage Diminution (%)Ring Markedness
Index (RMI) (%)		Range of Percentage Skews (Mean Value) (%)
DeciduousLarix decidua<1 year50.00–54.8371.55–85.9135.77–44.36−40.0 to +7.7 (−6.8)
EvergreenK. changchunense1–3 years40.00–50.0089.39–97.5038.92–48.750.0 to +20.0 (+8.8)
Pinus sylvestris1–3 years41.03–50.0070.53–77.2831.56–35.26−9.1 to +17.9 (+5.2)
Picea abies3–5 years25.93–44.1974.02–84.0319.90–35.420.0 to +38.2 (+12.0)
Cedrus libani3–6 years30.77–39.5862.33–72.0620.22–24.68+35.7 to +42.9 (+39.0)
Araucaria araucana3–15 years10.00–22.5028.67–51.793.17–10.35+55.0 to +80.0 (66.7)
Table
Table 4. Geological distribution of Keteleeria-like fossils (excluding the species without detailed description).
Table 4. Geological distribution of Keteleeria-like fossils (excluding the species without detailed description).
SpeciesLocationAgeType of FossilReferences
Protopiceoxylon wordii WaltonSvalbard, NorwayMiddle JurassicWoodWalton, 1927 [9]
Protopiceoxylon articum SewardFranz Josef Land, RussiaLate JurassicWoodSeward, 1904 [45]
Protopiceoxylon resiniferous Medlyn et TidwellUtah, USALate JurassicWoodMedlyn and Tidwell, 1979 [46]
Protopiceoxylon canadense Medlyn et TidwellBritish ColumbiaLate JurassicWoodMedlyn and Tidwell, 1979 [46]
Pinoxylon (Protopiceoxylon) dacotense knowltonLiaoning Province, China; South Dakota, USA; Sakhalin, RussiaLate Jurassic; Early Cretaceous (?); Late CretaceousWoodJiang et al., 2008 [47]; Knowlton, 1900 [25]; Nishida and Nishida, 1995 [48]
Protopiceoxylon extinctum GothanHebei Province, China; Svalbard, NorwayLate Jurassic; Early Cretaceous (?)WoodGothan, 1907 [27]; Mathew and Ho, 1945 [49]; Sze, 1963 [50]
Protopiceoxylon xinjiangense Wang, Zhang et SaikiXinjiang, ChinaLate JurassicWoodWang et al., 2000 [51]
Protopiceoxylon (Pinoxylon) yabei (Shimakura) SzeJilin, ChinaLate JurassicWoodShimakura, 1936 [52]; Mathews and Ho, 1945 [49]; Sze, 1963 [50]
Keteleerioxylon fokinii ShilkinaKirovsk Region, RussiaValanginian (Early Cretaceous)WoodShilkina, 1986 [34]
Protopiceoxylon edwardsi StopesGreensands, EnglandAptian (Early Cretaceous)WoodStopes, 1915 [53]
Protopiceoxylon amurense DuHeilongjiang Province, ChinaAptian-Albian (Early Cretaceous); Coniacian-Maastrichtian (Late Cretaceous); PaleoceneWoodDu, 1982 [54]; Wang et al., 1997 [55]; Terada et al., [56]
Protopiceoxylon chaoyangensis DuanLiaoning Province, ChinaAptian-Albian (Early Cretaceous)WoodDuan, 2000 [57]
Protopiceoxylon yizhouensis Duan et CuiLiaoning Province, ChinaAptian-Albian (Early Cretaceous)WoodDuan et al., 1995 [58]
Keteleerioxylon arcticum ShilkinaFranz Josef Land, RussiaAptian-Albian (Early Cretaceous)WoodShilkina, 1960 [7]
Keteleerioxylon kamtschatkiense Blokhina et AfoninKamchatka Peninsula, RussiaAptian-Albian (Early Cretaceous); Turonian-Coniacian (Late Cretaceous)WoodBlokhina et al., 2006 [11]
Keteleerioxylon changchunense Shi, Sun, Meng et YuJilin Province, ChinaAlbian (Early Cretaceous)WoodThis paper
Protopiceoxylon johnseni (Schroeter) EdwardsSvalbard, NorwayEarly CretaceousWoodSchröeter, 1880 [59]; Edwards, 1925 [60]
Protopiceoxylon mohense Ding Heilongjiang Province, ChinaEarly CretaceousWoodDing, 2000 [61]
Keteleeria fortunei (Andr. Murray) Carr.Henan Province, ChinaLate CretaceousWoodYang et al., 1990 [39]
Keteleeria cretacea Miki et MaedaAwaji, JapanLate CretaceousConeMiki and Maeda, 1966 [62]
Protopiceoxylon yukonense Dolezych et ReinhardtYukon, CanadaPaleoceneWoodDolezych and Reinhardt, 2015 [63]
Keteleeria sp.British Columbia, CanadaEarly EoceneSeedMathewes et al., 2016 [6]
Keteleeria sp.Liaoning Province, ChinaEoceneConeThe Writing Group of Cenozoic Plants of China, 1978 [64]
Keteleeria mabetiensis WatariIshikawa Prefecture and Akita Prefecture, JapanOligocene and MioceneWoodWatari, 1941, 1956 [37,38]; Terada, 1998 [65]; Choi et al., 2010 [66]
Keteleeria sp.Primorye, RussiaOligocene to Early MioceneLeaves, cones, and seedsRybalko et al., 1980 [67]
Keteleerioxylon primoryense Blokh.Primorye, RussiaOligocene to MioceneWoodBlokhina and Klimova, 2000 [35]
Keteleeria rujadana LakhanpalOregon, USAOligoceneConeLakhanpal, 1958 [68]
Keteleeria ptesimosperma Meyer et ManchesterOregon, USAOligoceneWinged seedsMeyer and Manchester, 1997 [69]
Keteleeria prambachensis (Hofmann) KlausPrambachkirchen, AustriaOligoceneConeHofmann, 1944 [70]; Klaus, 1977 [71]
Keteleeria rhenana KräuselMainz, GermanyEarly MioceneSeedKräusel, 1938 [72]
Keteleeria microreticulata AnanovaTaganrog peninsula, RussiaMiddle MiocenePollenAnanova, 1974 [73]
Keteleeria caucasica RamischviliZugdidi municipalitet, GeorgiaLate MiocenePollenRamischvili, 1969 [74]
Keteleeria davidiana MikiHonshu, JapanMioceneCones and seedsMiki 1941, 1957, 1958 [75,76,77]
Keteleeria ezoana TanaiNiigata and Hokkaido Prefecture, JapanMioceneSeed scales, seeds, and leavesTanai, 1961 [78]; Tanai & Suzuki, 1963 [79]; Kamoi et al., 1978 [80]; Ozaki, 1979 [81]
Keteleeria shanwangensis Wang et al.Shandong Province, ChinaMioceneCone, winged seeds,Wang et al., 2006 [82]
Keteleeria hoehnei KirchheimerSaxony, GermanyMioceneCones, seeds, and needlesKirchheimer, 1942 [83]; Kunzmann and Mai, 2005 [84]
Keteleeria bergeri KirchheimerSaxony, GermanyMioceneCones, seeds, and needlesKirchheimer, 1942 [83]
Keteleeria bergeri KirchheimerTurowo, Poland; Kanton Schwyz, SwitzerlandMioceneConeZalewska, 1961 [85]; Hantke, 1973 [86]
Keteleeria heterophylloides (Berry) BrownIdaho, USAMioceneVegetative shootsBrown, 1935 [87]
Keteleeria zhilinii Blokh. et BondarenkoPrimorye, RussiaPlioceneWoodBlokhina and Bondarenko, 2005 [10]
Keteleeria sp.Maoming, ChinaLate PleistoceneWoodHuang et al., 2019 [12]
Keteleeria sp.Wuhan, ChinaEarly HoloceneWoodYang et al., 2003 [40]
Publisher’s Note: MDPI stays neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Share and Cite

MDPI and ACS Style

Shi, X.; Sun, Y.; Meng, F.; Yu, J.; Lan, Z. Early Cretaceous Keteleerioxylon Wood in the Songliao Basin, Northeast China, and Its Geographic and Environmental Implications. Biology 2022, 11, 1624. https://doi.org/10.3390/biology11111624

AMA Style

Shi X, Sun Y, Meng F, Yu J, Lan Z. Early Cretaceous Keteleerioxylon Wood in the Songliao Basin, Northeast China, and Its Geographic and Environmental Implications. Biology. 2022; 11(11):1624. https://doi.org/10.3390/biology11111624

Chicago/Turabian Style

Shi, Xiao, Yuewu Sun, Fanli Meng, Jianxin Yu, and Zilie Lan. 2022. "Early Cretaceous Keteleerioxylon Wood in the Songliao Basin, Northeast China, and Its Geographic and Environmental Implications" Biology 11, no. 11: 1624. https://doi.org/10.3390/biology11111624

APA Style

Shi, X., Sun, Y., Meng, F., Yu, J., & Lan, Z. (2022). Early Cretaceous Keteleerioxylon Wood in the Songliao Basin, Northeast China, and Its Geographic and Environmental Implications. Biology, 11(11), 1624. https://doi.org/10.3390/biology11111624

Note that from the first issue of 2016, this journal uses article numbers instead of page numbers. See further details here.

Article Metrics

Back to TopTop