Historical Biology

An International Journal of Paleobiology

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A news species of conifer wood Brachyoxylon

from the Cretaceous of Eastern China and its
paleoclimate significance

Zikun Jiang, Hao Wu, Ning Tian, Yongdong Wang & Aowei Xie

To cite this article: Zikun Jiang, Hao Wu, Ning Tian, Yongdong Wang & Aowei Xie (2020): A news
species of conifer wood Brachyoxylon from the Cretaceous of Eastern China and its paleoclimate
significance, Historical Biology, DOI: 10.1080/08912963.2020.1755282

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Published online: 22 Apr 2020.

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HISTORICAL BIOLOGY
https://doi.org/10.1080/08912963.2020.1755282

ARTICLE

A news species of conifer wood Brachyoxylon from the Cretaceous of Eastern China and
its paleoclimate significance
Zikun Jianga,b, Hao Wuc, Ning Tiand, Yongdong Wangb,e and Aowei Xief
a
Chinese Academy of Geological Sciences, Beijing, China; bState Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and
Palaeontology, Nanjing, China; cZhejiang Museum of Natural History, Hangzhou, China; dCollege of Palaeontology, Shenyang Normal University,
Shenyang, China; eCenter for Excellence in Life and Paleoenvironment, Chinese Academy of Sciences, Nanjing, China; fInstitute of Geosciences, Division
of Paleontology, University of Bonn, Bonn, Germany

ABSTRACT ARTICLE HISTORY

A new species of conifer wood, Brachyoxylon zhouii sp. nov., is described from the Lower Cretaceous Received 25 February 2020
Guantou Formation of Zhejiang Province, eastern China. The new species is characterised by a mixed Accepted 9 April 2020
type of radial tracheary pitting, araucarioid cross-field pitting, high uniseriate rays, and the occurrence KEYWORDS
of traumatic resin canals. The new finding represents the second species of Brachyoxylon from the Conifer wood; early
Guantou Formation and it contributes to the further understanding of the forest composition of the Cretaceous; eastern China;
Early Cretaceous flora in eastern China.The quantitative analysis of growth ring anatomy indicates that palaeoclimate; Brachyoxylon
the forest composition was evergreen with a Leaf Retention Time (LRT) of 3–15 years. It is deduced
that the Zhejiang Province was dominated by a subtropical to tropical and relatively semiarid climate
during the Early Cretaceous.

Introduction
The Early Cretaceous petrified woods are quite rich in China with
more than 18 localities from seven provinces being reported (Zheng
et al. 2008; Yang et al., 2013). It is noted that most of the Early
Cretaceous wood records are documented from Northeast China
with over 45 species (Zheng et al. 2008), while only a few species
have been recorded in the eastern China region, and all are ascribed
to the genera Brachyoxylon Hollick et Jeffrey (1909) and
Agathoxylon Hartig (1848) (Vozenin-Serra and Pons 1990; Duan
et al. 2002; Tian et al. 2018).
In this study, we report a new coniferous wood species
Brachyoxylon zhouii from the Lower Cretaceous Guantou
Formation in Zhejiang Province, eastern China. The new finding
provides evidence for further understanding of the fossil wood
diversity and the forest composition of the Early Cretaceous flora
in the region. The occurrence of growth rings suggests that
a subtropical to tropical and relatively semiarid climate might be
prevailed during the Early Cretaceous in eastern China, while the
occurrence of false rings in fossil wood points to climate fluctua-
tions or turbulences.
Geological setting
The new material was collected from the northeast of Yongkang
City, central Zhejiang Province, eastern China (28°54′N, 120°3′E)
(Figure 1). The fossil locality belongs to the Yongkang Basin,
a well-known Mesozoic basin with well developed Early
Cretaceous deposits represented by the Yongkang Group.
Traditionally, the Yongkang Group is sub-divided into three
units conformably deposited in ascending order, i.e. the
Guantou Formation, Chaochuan Formation and Fangyan
Formations (Zhejiang BGM, 1989). Recently, some authors (Li
et al. 2018) proposed a new opinion on the Yongkang Group,
making the relationship between these three formations uncer-
tain. Lithologically, the Guantou Formation mainly consists of
variegated sandstones, mudstones, and with a few volcanic beds
(Figure 2), which represent an inland fluvial-lacustrine deposi-
tional environment. Silicified woods and several species of plants,
abundant animal fossils have been discovered from Guantou
Formation, including gastropods, bivalves, ostracods, insects and
fishes (Zhejiang (BGM) 1989). Most of the isotopic dating of
volcanic sequences of the Guantou Formation shows radiometric
ages between 125 Ma to 103 Ma (e.g. Luo and Yu 2004; Li et al.
2014; Ma et al. 2016), thus corresponding to Aptian to Albian of
the Early Cretaceous in age.
Material and methods
The fossil specimen described in this paper was preserved as
a silicified wood fragment with secondary xylem. The technique
used for the anatomical investigation is the classical thin section
method for silicified wood described in Jones and Rowe (1999).
The specimen was cut transversely, radial longitudinally and
tangential longitudinally into several thin sections.
Nomenclatural and taxonomical positions follow those of
Philippe (1993), Bamford and Philippe (2001) and Philippe
and Bamford (2008). Photographs were taken with Scope
Image 9.0 software adapted to a Yongxin BM2000 Microscope.
For growth ring analysis, we use quantitative analysis of growth
ring anatomy approaches, including the Percentage diminution,
Percentage latewood, Ring Markedness Index (RMI), and Skew
of CSDM curves (The Cumulative algebraic Sum of each cell’s
Deviation from the Mean) were calculated using the method by
Falcon-Lang (2000a).

CONTACT Zikun Jiang jiangmaomao_925@sina.com Chinese Academy of Geological Sciences, No. 26 Baiwanzhuang Street, Xicheng District, Beijing 100037,
China; Yongdong Wang ydwang@cags.ac.cn State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Nanjing 210008,
China
© 2020 Informa UK Limited, trading as Taylor & Francis Group

Published online 22 Apr 2020

2 Z. JIANG ET AL.

Figure 1. The fossil location of northeast Yongkang City, central Zhejiang Province, eastern China. (a), Geographical map of China showing the location of Zhejiang
Province; (b), Sketch map of Zhejiang Province showing the fossil locality in the Yongkang Basin(dashed line range region).

All the living trees used in CSDM curves chosen to be Horizon and age
compared with the new species were all from the southern
Guantou Formation, Early Cretaceous
England. For all these trees were derived from trees that had
grown in cultivation rather than in natural stands. Cultivated
species were analysed in order that specimens with a very large
Repository
range of LRTs could be studied within a single climatic setting.
It is unusual for conifers with such a large range of LRTs to be The specimen and slides are housed in the Zhejiang Museum of
found within a single natural community (Falcon-Lang, Natural History, Hangzhou, China.
2000a),
All fossil wood specimens and slides referred to this study are
housed in the Zhejiang Museum of Natural History, Hangzhou, Etymology
China, with a registration number ZMNH M13189.
The specific name is dedicated to Prof. Zhou Zhiyan for his con-
tributions to Chinese Palaeonbotany.
Systematic palaeontology
Class Coniferopsida Gifford et Foster 1989 Description
The studied specimen ZMNH M13189 only preserved secondary
Order Coniferales Engler 1897
wood, without pit and primary xylem. The secondary xylem
presents a tracheidoxylic structure with distinct variably growth
Genus Brachyoxylon Hollick et Jeffrey 1909
rings in width, between 19–50 cells wide. The transitions of
tracheid size from early wood to late wood in are abrupt with
Type species: Brachyoxylon notablile Hollick et Jeffrey 1909
narrow late wood, only 3–4 tangential rows of compressed cells,
much smaller than that in the early wood (Figure 3(a)).
Species: Brachyoxylon zhouii sp. nov. Jiang, Tian, Wu et Wang
Secondary xylem tracheids are polygonal, mostly quadrangular,
slightly round to oval in transverse section, with occasionally
intercellular spaces (Figure 3(b)). Sometimes, some boundaries
Diagnosis are marked by tangential series of traumatic axial canals, with
resin in them (Figure 3(a,b)).
Secondary xylem tracheidoxylic, growth rings distinct, marked by
In radial section, radial wall pitting show mainly biseriate
abrupt transitional tracheids in size from early wood to late wood
(85%), rarely uniseriate (15%), without triseriate; when biseriate,
and occasionally by tangential series of traumatic axial canals;
pittings are alternate (Figure 3(d)) and very rarely opposite
Tracheids pitting in radial walls of mixed type, predominantly
(Figure 3(e)) or sub-opposite (Figure 3(f)); when uniseriate,
alternate when biseriate and contiguous or compressed when uni-
pittings are continuous (Figure 3(c)), partly distant (Figure 4
seriate, partly uniseriate distant, and rarely biseriate opposite;
(a)). The bordered pits on radial walls are ascribed to the mixed
Cross-field pits araucarioid type, with 7–11 closely arranged cupres-
type according to Philippe and Bamford (2008), with biseriate
soid pits per field; horizontal and end walls smooth and without
alternated and uniseriate continuous pits account for 85%, the
pitting; Axial parenchyma absent; Crassulae present; Rays homo-
uniseriate distance ones account for 7%, the biseriate opposite
cellular, mostly uniseriate and rarely biseriate, 3–50 (mostly 15–45)
and sub-opposite ones account for 8%. Horizontal walls and
cells high.
end walls of ray cells are smooth and unpitted (Figure 4(b)).
Cross-fields have numerous bordered pits with the circular or
oval outline, contiguous and placed in 4–5 alternate vertical
Holotype
rows (7–11 cupressoid pits) (Figure 4(c-e)). Pit arrangement is
ZMNH M13189 with 3 thin sides numbered ZJ-0201, ZJ-0202, ZJ-0203 of the araucarioid type sensu IAWA Committee (2004).

Figure 2. Stratigraphic column of the Lower Cretaceous Guantou Formation
in Yongkang County, Zhejiang Province, eastern China (modified after Luo and
Yu 2004). 1. Sandy mudstone; 2. Vitric rhyolitic tuff; 3. Pebbly sandstone; 4.
Mudstone; 5. Argillaceous sandstone; 6. Sandstone; 7. Basalt; 8. Vitric tuff; 9.
Shale; 10. Fossil wood.
In the tangential section, rays are homocellular, mostly uniseri-
ate (Figure 4(f)) and rarely biseriate (Figure 4(f)), 3–50 (mostly
15–45) cells high. Axial parenchyma and tangential pits are absent.
Hypha or septa are occasionally present (Figure 4(f)).
Taxonomic assignment and comparisons
According to Philippe and Bamford (2008), the occurrence of
a mixed type of radial pitting, araucarioid cross-field pitting and
traumatic resin canals indicates that the new species belongs to the
fossil wood genus Brachyoxylon Hollick et Jeffrey.
HISTORICAL BIOLOGY 3
Xylotomical comparisons of our material with species of
Brachyoxylon with traumatic canals are showed in Table 1.
Those Brachyoxylon species which lacking traumatic resin
ducts are excluded in the comparison with the Chinese species
studied here (for detailed information see Tian et al. 2018). It
is noted that Brachyoxylon notabile Hollick et Jeffrey,
B. avramii Iamandei et Iamandei and B. serrae Philippe
et al. posses pits on tracheid tangential walls, which are absent
in the wood studied herein. Brachyoxylon sp. from Tibet,
China is characterised by having triseriate rays (Vozenin-
Serra and Pons 1990). Nevertheless, the new species has
neither tangential tracheary pitting nor multiseriate rays.
B. lagonense (Laudouéneix) Dupéron-Laudouéneix has
a mixed type of biseriate and triseriate tracheary pitting,
which is different from the new species. It seems that
B. woodworthianum Torrey and B. saurinii Boureau et Serra
are more similar to our new species, but they differ from the
new species in radial tracheid pitting, the ray height and pits
number per cross-field. Based on the significant differences
between our specimen and other materials, we therefore pro-
pose a new species Brachyoxylon zhouii sp. nov.
Palaeoclimate implications
Growth rings in fossil woods, influenced by their environ-
ments, received increasing attention from palaeobotanists
and have been used as an important proxy for interpreting
palaeoclimates and palaeoenvironments (e.g. 2011; Chaloner
and Creber 1973; Creber and Chaloner 1984; Francis 1984;
Morgans 1999; Falcon-Lang 2000a; 2000b; 2003; Brea et al.
2008; Shi et al. 2015; 2017). In this study, the quantitative
analysis of growth ring anatomy was used as proxy for reflect-
ing the paleoclimate conditions. The radial diameter of trac-
heid cells referred to six adjacent rings was measured for ring
increment, and these data were used to construct the CSDM
curve (the cumulative algebraic sum of each cell’s deviation
from the mean was then calculated for each growth ring
increment and plotted as a zero-trending curve). We calcu-
lated four parameters: 1. Percentage diminution, 2. Percentage
latewood, 3. Ring Markedness Index (RMI), 4. Skew of CSDM
curves (Figure 5) (Table 2), which are the effective methods to
infer the palaeoclimate and palaeoenvironment (Falcon-Lang
2000a; 2000b).
The CSDM curves of deciduous conifers are dominantly
left-skewed or symmetrical, whilst evergreen conifers have
dominantly right-skewed CSDM curves. The magnitude of
right-skewness in evergreen conifers appears to be positively
related to leaf longevity (Falcon-Lang 2000a). The skew of
CSDM curves of Brachyoxylon zhouii is from 0% to +80%
(mean percentage of skew of CSDM curves +34.83%), which
indicates the new species was evergreen.
The percentage of late wood in conifer woods could be
related to the foliar retention. The late wood development
could be strongly influenced by leaf longevity and intensity of
climate seasonality (Falcon-Lang 2000a; 2000b). In our fossil
specimen, the percentage of late wood is from 10% to 30%
(the mean value is 17.98%), which is similar to the data of
Araucaria araucana. The Ring Markedness Index was calculated
from the percentage of late wood and percentage diminution.
The leaf longevity exerts an important control on growth ring
markedness (Falcon-Lang 2000a). The Ring Markedness Index
of the new species is from 8.45% to 21.03% (mean RMI
14.28%), which resembles the data of Araucaria araucana.
4 Z. JIANG ET AL.

Figure 3. Brachyoxylon zhouii sp. nov. Jiang, Tian, Wu et Wang from the Lower Cretaceous of Zhejiang Province, eastern China. (a), Transverse section showing growth
rings, false rings and traumatic axial ducts, with resin in them, Scale bar = 500 µm; (b), Transverse section showing growth rings (red arrow), Scale bar = 200 µm; (c), Radial
section showing radial wall pitting uniseriate, continuous (red arrow), Scale bar = 100 µm; (d), Radial section showing radial wall pitting biseriate, alternate (red arrow),
Scale bar = 100 µm; (e), Radial section showing radial wall pitting biseriate, opposite (red arrow), Scale bar = 50 µm; (f), Radial section showing radial wall pitting biseriate,
sub-opposite (red arrow), Scale bar = 50 µm.

Compared with the leaf longevity and growth ring markedness
analysed for five conifer trees from southern England, the Leaf
Retention Times (LRTs) of the Brachyoxylon zhouii is
3–15 years (Table 3).
Totally six growth ring types were recognised by Creber and
Chaloner (1984) for extant and fossil trees (Brison et al. 2001).
Our specimen corresponds to the types D to E. The Type D is
represented by rings with a thin band of latewood; earlywood/
latewood boundary is well marked; Type E is featured by rings
be similar to type D, but the transition to latewood is not so
abrupt. The palaeolatitude of Types D and E is widely distrib-
uted in the Early Cretaceous (Brison et al. 2001). The mean
radial diameter of the adjacent six growth ring cells is variable,
which indicates fluctuations of water supply or other environ-
mental disturbances.
During the Early Cretaceous, the age of Brachyoxylon zhouii sp.
nov., several phytoprovinces are recognised, including Siberia-Canada
phytoprovince, European-China phytoprovince, Equatorial phytopro-
vince and Australia phytoprovince (Duan 1986). In China, the
Northeast China region is ascribed to the Siberia-Canada phytopro-
vince, while the Shandong Province, South-East coast and Tibet area
are ascribed to the European-China phytoprovince (Duan et al. 1977;
Duan 1986). During the Early Cretaceous, Zhejiang (the present fossil
locality area), Fujian and Shandong provinces and Tibet region of the
European-China phytoprovince, were dominated by gymnosperm for-
ests (Duan et al. 2002). Among them, the conifer fossils are mainly
represented by Brachyphyllum and Pagiophyllum, while the under-
growth ferns were represented by dominant Ruffordia, Onychiopsis,
and Weichselia with Zamiophyllum and Otozamites were the dominant
representatives of Cycadales and Bennettitales (Duan et al. 2002). The
 HISTORICAL BIOLOGY 5

Figure 4. Brachyoxylon zhouii sp. nov. Jiang, Tian, Wu et Wang from the Lower Cretaceous of Zhejiang Province, eastern China. (a), Radial section showing uniseriate,
continuous and partly distant radial wall pitting (red arrow), Scale bar = 50 µm; (b), Radial section showing horizontal walls and smooth and unpitted end walls of ray cells,
Scale bar = 50 µm. (c, d, e), Radial section showing numerous araucarioid cross-fields pits with circular or oval outline, contiguous and placed in 4–5 alternate vertical rows,
C, scale bar = 50 µm, D, scale bar = 50 µm, E, scale bar = 20 µm. (f), Tangential section showing uniseriate homocellular rays, rarely biseriate rays and hypha or septa (red
arrow), Scale bar=200 µm.

Table 1. Comparative table of species of the genus Brachyoxylon with traumatic resin ducts (modified after Bodnar et al. 2013; Tian et al. 2018).
Species Age Distribution False GR Tg TP Rd TP Axe Pa Cr Rays(height) CFP
Brachyoxylon zhouii sp. nov. Early Cretaceous China P A mixed 1–2 seriate A A mostly 1 seriate (3–50 cells) 7-11 cupressoid
Jiang, Tian, Wu et
Wang
B. notabile Hollick and Late Cretaceous USA A P mixed 1–2 seriate A P 1 seriate (1–8 cells) 5-11 cupressoid
Jeffrey 1909
B. woodworthianum Torrey Early Cretaceous USA A A mixed A A 1-2 seriate (2–20 cells) cupressoid
1923
B. saurinii Boureau and Jurassic (?) Cambodia A A mixed 1–2 seriate A A 1 seriate (1–31 cells) 2-12 cupressoid
Serra 1961
B. lagonense Cretaceous Chad A A mixed 2–3 seriate A A 1 seriate (low) cupressoid
(Laudouéneix 1973)
Dupéron-
Laudouéneix 1991
B. avramii Iamandei and Early Cretaceous Romania A P mixed 1–2 seriate A A 1 seriate (1–21 cells) 1-8 cupressoid
Iamandei 2005
B. serrae Philippe et al. Early Cretaceous Thailand A P mixed 1–2 seriate A A 1 seriate (1–15 cells) 5-16 cupressoid
2011
Brachyoxylon sp. Vozenin- Early Cretaceous China A N mixed 1 seriate A N 1-3 seriate (3–17 cells) 5-12 cupressoid
Serra and Pons 1990
GR Growth rings, Tg TP Tangential tracheid pitting, Rd TP Radial tracheid pitting, Axe Pa Axial parenchyma, Cr Crassulae, CFP Cross-field pits, P Present, A Absent,
N Unknown

spores and pollen grains from the Guantou Formation in Zhejiang spores and pollen, the palaeoclimate of this area is represented by hot
Province (Zhang 1987) reveal a flora constitution of 51.8% of ferns, and subtropical to tropical relatively semiarid conditions, which con-
46.0% of gymnosperms, and 2.2% mosses. Based on the data of the sistent with the previous results deduced from the plant landscape and
6 Z. JIANG ET AL.

Figure 5. CSDM curves of the six adjacent growth rings of Brachyoxylon zhouii sp. nov. Jiang, Tian, Wu et Wang.

Table 2. Results of the four parameters for Brachyoxylon zhouii sp. nov. Jiang, Tian, Wu et Wang.
Ring number Percentage of diminution Percentage latewood Ring Markedness index Percentage of skew
Ring 1 81.3% 12.5% 10% 25%
Ring 2 91% 19.35% 17.61% 16%
Ring 3 70.1% 30% 21.03% 0
Ring 4 81.9% 21.05% 17.24% 58%
Ring 5 84.5% 10% 8.45% 80%
Ring 6 75.5% 15% 11.33% 30%
Mean 80.7% 17.98% 14.28% 34.83%

Table 3. Comparison of the quantification of ring markedness parameters for Brachyoxylon zhouii sp. nov. Jiang, Tian, Wu et Wang with the five extant taxa based on
Falcon-Lang (2000a; 2000b)).
Taxon Percentage latewood (%) Percentage diminution (%) Ring markedness index (%) Range of percentage skews (mean value) (%)
Larix decidua (deciduous) 50.00–54.83 71.55–85.91 35.77–44.36 −40.0 to + 7.7 (- 6.8)
Pinus sylvestris (1–3 years) 41.03–50.00 70.53–77.28 31.56–35.26 −9.1 to +17.9 (+ 5.2)
Picea abies (3–5 years) 25.93–44.19 74.02–84.03 19.90–35.42 0.0 to + 38.2 (+12.0)
Cedrus libani (3–6 years) 30.77–39.58 62.33–72.06 20.22–24.68 +35.7 to + 42.9 (+39.0)
Brachyoxylon zhouii (? years) 10.00–30.00 70.10–91.00 8.45–21.03 0.0 to + 80.0 (+34.83)
Araucaria araucana (3–15 years) 10.00–22.50 28.67–51.79 3.17–10.35 +55.0 to + 80.0 (+66.7)

the palaeosols (Zhang 1987; Duan et al. 2002; Zheng et al. 2008; Li et al. Brachyoxylon, e.g. Agathoxylon (Araucarioxylon) xinchangense,
2009; Zhu et al. 2018; Tian et al. 2018). In this region, only a few fossil Brachyoxylon zhejiangense, B. zhouii and Agathoxylon sp. (Zheng
wood, are reported, including 4 species of Agathoxylon and et al. 2008; Zhu et al. 2018; Tian et al. 2018). The palaeogeographical

distribution and the growth rings analysis of Brachyoxylon – a conifer
wood with long LRTs – also indicated a subtropical to tropical climate.
The method to distinguish between evergreen and deciduous conifer
woods based on growth ring analysis is an attempt in southern China.
This is the first trial used in this area and the third used in China, the
palaeoclimate inferred from the method is consistent with other ana-
lysis e.g. the paleosol and spores and pollen results. This method was
also used in Xinjiang Uygur antonomous Province of China for Lower
Triassic deposits and Maoming Basin, Guangdong Province of China
in Late Pleistocene had been proved to be an effective palaeoecological
tool (Shi et al. 2017; Huang et al. 2019). As a well established palaeocli-
matic analysis tool which has already been frequently used in Europe,
the method should be widespread used in petrified wood in China.
The occurrence of Brachyoxylon from the Lower Cretaceous in
Zhejiang and Tibet indicates that Brachyoxylon was widely distributed
in the European-China area in the Early Cretaceous. The discovery of
the new species not only enriches the understanding of the fossil forest
composition in the Early Cretaceous of Zhejiang Province, but also
better realises the palaeoclimate and palaeoecology through the tree
ring analysis.
Acknowledgments
We would like to thank Prof. Zheng Shaolin, Zhang Wu from Shenyang Center,
China Geological Survey and Prof. Shi Xiao from Jilin University for their
suggestions regarding the manuscript. We also express our gratitude to the
reviewers who improved this manuscript.
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
This work was supported by the National Natural Sciences Foundation of China
(Grant Nos 41772023, 41790454, 41688103, 41402004 and 41302004), the
Strategic Priority Program (B) of CAS (XDB 18000000 and 2600000), the State
Key Program for Basic Research & Development of Ministry of Science &
Technology of China (2016YSC0600406) and the State Key Laboratory of
Palaeobiology and Stratigraphy (Nanjing Institute of Geology and
Palaeontology, CAS) (Grant No. 173113).
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