Review of Palaeobotany and Palynology最新文献_第6页

Evaluating Post-K/Pg persistence of Classopollis: Evidence from exceptionally preserved reworked Pollen in Paleogene sub and inter-Volcanic Sediments from Northern Ireland 评价Classopollis的后k /Pg持久性：来自北爱尔兰古近系亚火山和火山间沉积物中异常保存的再加工花粉的证据

IF 1.7 3区地球科学 Q2 PALEONTOLOGY

Review of Palaeobotany and Palynology

Pub Date : 2026-01-01 Epub Date: 2025-08-18 DOI: 10.1016/j.revpalbo.2025.105424

Manuel Vieira , David Jolley

The Cheirolepidiaceae, an extinct family of coniferous plants that thrived during the Mesozoic Era, are well known through their distinctive Classopollis pollen. Although long thought to have declined globally during the Late Cretaceous, isolated occurrences of Classopollis in Paleocene deposits have sparked debate regarding the survival or reworking of this lineage beyond the Cretaceous–Paleogene (K/Pg) boundary. In this study, we document the occurrence of exceptionally well-preserved Classopollis pollen within Paleocene inter-volcanic sedimentary rocks of the Antrim Lava Group in Northern Ireland. Detailed morphological analyses using transmitted light microscopy, scanning electron microscopy (SEM), and transmission electron microscopy (TEM) reveal Classopollis pollen with uniformly thick tectate exines, well-developed baculate infratectal layers, and a dense nano-spinulose supratectal ornamentation. These features can be compared with Barremian-aged English specimens and support a Cretaceous origin. Palynological assemblages also include other reworked Cretaceous taxa and clastic components derived from the underlying Ulster White Limestone Formation, a Santonian–Early Maastrichtian unit exposed within the catchment. The stratigraphic context and preservation state of the Classopollis grains suggest that they were reworked rather than representing in situ Paleocene vegetation. This study supports the interpretation that post-K/Pg occurrences of Classopollis in the Antrim Basin are best explained by sedimentary recycling of Late Cretaceous deposits, adding to the broader discussion on Cheirolepidiaceae persistence and paleoecological relict status in the Paleogene.

Cheirolepidiaceae是一个灭绝的针叶植物家族，在中生代繁荣，因其独特的Classopollis花粉而闻名。尽管人们一直认为，在晚白垩世，Classopollis在全球范围内已经减少，但在古新世沉积物中孤立出现的Classopollis引发了关于该谱系在白垩纪-古近纪（K/Pg）界限之外是否存在或改造的争论。在这项研究中，我们记录了北爱尔兰安特里姆熔岩群古新世火山间沉积岩中保存异常完好的Classopollis花粉的存在。利用透射光镜、扫描电镜和透射电镜对其进行了详细的形态学分析，结果显示，Classopollis花粉具有均匀厚的盾状外壁，发育良好的小管状花序，以及致密的纳米级细刺状的直肠上纹饰。这些特征可以与巴雷米亚时代的英国标本进行比较，并支持白垩纪起源。孢粉组合还包括其他经过改造的白垩纪分类群和碎屑组分，这些碎屑组分来自于下部的阿尔斯特白灰岩组，这是一个暴露在集水区的圣安东尼奥-早期马斯特里赫特单元。Classopollis颗粒的地层背景和保存状态表明它们是被重新加工过的，而不是代表古新世的原位植被。本研究支持了用晚白垩世沉积物的沉积再循环来解释Antrim盆地后k /Pg产状的解释，为进一步探讨Cheirolepidiaceae在古近系的存续和古生态遗存状态提供了理论依据。

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引用次数: 0

Terrestrial end-Guadalupian crisis constrained by mid-latitude Permian palynological data from Jimsar, Junggar Basin, China 准噶尔盆地吉木萨尔中纬度二叠纪孢粉资料约束的瓜达鲁普末期陆相危机

IF 1.7 3区地球科学 Q2 PALEONTOLOGY

Review of Palaeobotany and Palynology

Pub Date : 2026-01-01 Epub Date: 2025-09-05 DOI: 10.1016/j.revpalbo.2025.105445

Xueqin Wang , Yuewu Sun , Xiao Shi

The end-Permian mass extinction and the end-Guadalupian crisis are the most remarkable events in the Permian terrestrial ecosystem. To better understand the end-Guadalupian event in mid-latitude of Pangea in the Northern Hemisphere, palynological investigations are conducted on the terrestrial Permian deposits in the Dalongkou profile of Jimsar, northern Xinjiang, China. Two palynological assemblages are recognized in stratigraphic ascending order, the Cordaitina subrotata–Striatoabieites lipidus–Protowelwitschiapollis exolescus assemblage from the uppermost Lucaogou, Hongyanchi and the lower Quanzijie formations, and the Kraeuselisporites spinulosus–Tuberculatosporites homotubercularis–Potonieisporites turpanensis assemblage from the upper Quanzijie and lowermost Wutonggou formations. Based on the palynological data, the uppermost Lucaogou and Hongyanchi formations are assigned to the Roadian stage, and the lower Quanzijie Formation to the Capitanian stage, and the upper Quanzijie and lowermost Wutonggou formations to the Wuchiapingian stage. Thus, a Wordian-age hiatus is confirmed between the lower Quanzijie Formation and the Hongyanchi Formation on the basis of palynological evidence. The palynofloral succession shows a smooth transition between the Guadalupian and Lopingian palynofloras in the Dalongkou profile of Jimsar, in the Junggar Basin. The end-Guadalupian crisis resulted in a 17% loss of pollen species, indicating that the extinction was on the background level.

二叠纪末的大灭绝和瓜达鲁普期末的危机是二叠纪陆地生态系统中最显著的事件。为了更好地了解北半球盘古大陆中纬度的瓜德鲁普末期事件，对新疆北部吉木萨尔大龙口剖面的陆相二叠系沉积物进行了孢粉学研究。从地层升序上可识别出两个孢粉组合，即最上层芦草沟组、红盐池组和最下层泉子沟组的Cordaitina subrotata-Striatoabieites lipidus-Protowelwitschiapollis exolescus组合，以及最上层泉子沟组和最下层梧桐沟组的Kraeuselisporites spinulosus-Tuberculatosporites homotubercularis-Potonieisporites turpanensis组合。根据孢粉资料，将芦草沟组上部和红盐池组划分为路甸期，将泉子街组下部划分为capitian期，将泉子街组上部和梧桐沟组下部划分为五家坪期。因此，根据孢粉学证据，确定了全子街组下段与红盐池组之间存在一个世界间隙。准噶尔盆地吉木萨尔大龙口剖面的孢粉区系在瓜达鲁普系和洛平系之间有一个平稳的过渡。瓜达鲁普末期的危机导致17%的花粉物种损失，表明灭绝是在背景水平上。

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引用次数: 0

Soil phytolith assemblages reflect palm community composition in western Amazonia 西部亚马逊地区土壤植物岩组合反映了棕榈群落组成

IF 1.7 3区地球科学 Q2 PALEONTOLOGY

Review of Palaeobotany and Palynology

Pub Date : 2026-01-01 Epub Date: 2025-08-28 DOI: 10.1016/j.revpalbo.2025.105443

Crystal N.H. McMichael , Daniel Guerra , Nina H. Witteveen , Britte M. Heijink , Annabel Zwarts , Gabriela Zuquim , Henrik Balslev , Kalle Ruokolainen , Hanna Tuomisto

Humans have domesticated and used palms in tropical forests for millennia. The extent to which past human palm use affects modern tropical systems remains relatively unexplored due to lack of a good fossil record. However, palms produce an array of phytoliths, which are siliceous cell structures that preserve well in the soil even after the organic plant material has decayed. Phytoliths vary in morphology and can be identified to group level and sometimes even to species. We analyzed if palm phytolith assemblages sampled from the soil reflect palm species compositions and abundances in 63 western Amazonian transects. Principal coordinate analysis (PCoA) and Mantel tests showed that palm communities and phytolith assemblages had similar turnover patterns. Negative binomial generalized linear models indicated that the abundances of nine of the 13 common palm genera were significantly related to a combination of soil cation concentrations, grass phytolith percentages, and palm phytolith PCoA axis scores. These results suggest that phytoliths have potential as quantitative indicators of changes in palm abundances in paleoecological and archaeological reconstructions.

几千年来，人类一直在热带森林中驯化和使用棕榈树。由于缺乏良好的化石记录，过去人类使用棕榈树对现代热带系统的影响程度仍然相对未被探索。然而，棕榈树会产生一系列植物岩，这是一种硅质细胞结构，即使在有机植物物质腐烂后，它们也能很好地保存在土壤中。植物岩在形态上各不相同，可以在群水平上识别，有时甚至可以在种水平上识别。我们分析了63个亚马逊西部样带的棕榈植物岩组合是否反映了棕榈物种的组成和丰度。主坐标分析（PCoA）和Mantel试验表明，棕榈群落和植物岩组合具有相似的更替模式。负二项广义线性模型表明，13个常见棕榈属中有9个属的丰度与土壤阳离子浓度、草植体百分比和棕榈植体PCoA轴评分的组合呈显著相关。这些结果表明，植物岩具有作为古生态和考古重建中棕榈丰度变化的定量指标的潜力。

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引用次数: 0

New palynological data from the Devonian of southern Poland (Siewierz area, Podleśna quarry) 波兰南部（siwierz地区，Podleśna采石场）泥盆纪孢粉新资料

IF 1.7 3区地球科学 Q2 PALEONTOLOGY

Review of Palaeobotany and Palynology

Pub Date : 2026-01-01 Epub Date: 2025-10-11 DOI: 10.1016/j.revpalbo.2025.105457

Marcelina Kondas, Paweł Filipiak

The sedimentary succession exposed in the Podleśna quarry near Siewierz (southern Poland) yielded the first confirmed occurrence of Devonian palynomorphs from this locality. Based on the composition of the miospore assemblages—the most abundant palynomorph group—the deposits have been assigned to the EX2 to possibly EX3 subzones of the EX Miospore Zone, corresponding to the rhenanus/varcus to latifossatus/semialternans conodont zones (Givetian). Preliminary palynofacies analysis suggests deposition in a proximal shelf setting.

在siwierz（波兰南部）附近的Podleśna采石场中发现的沉积序列首次证实了该地区的泥盆纪岩形。根据最丰富的微孢子组合组成，矿床被划分为EX2至EX3亚带，对应于小孔/静脉状到纬窝状/半互生牙形带（Givetian）。初步的孢粉相分析表明沉积在近陆架环境中。

引用次数: 0

Effect of steep climate gradient on palynological assemblages on the Arabian part of the Tethys shore in two Carboniferous-Permian ice age time slices 陡峭的气候梯度对两个石炭纪-二叠纪冰期片中特提斯海岸阿拉伯部分孢粉组合的影响

IF 1.7 3区地球科学 Q2 PALEONTOLOGY

Review of Palaeobotany and Palynology

Pub Date : 2026-01-01 Epub Date: 2025-09-15 DOI: 10.1016/j.revpalbo.2025.105448

Michael H. Stephenson , Shuzhong Shen , Junxuan Fan , Linshu Hu , Jin Qi

Palaeoclimate modelling of two time slices, Serpukhovian to Bashkirian (Carboniferous), and Gzhelian to early Sakmarian (latest Carboniferous to Early Permian), indicates marked changes in palynological assemblages from 30°S and 50°S across the Gondwana and Tethys margin. Both time slices indicate a distinct cold climate assemblage dominated by cheilocardioid spores, while a warmer climate with mean annual temperature above 19 °C appears to encourage the growth of plants that produced Euramerican coal belt spores such as Vestispora fenestrata and Reticulatisporites polygonalis. It is possible that the parent plants of cingulicamerate spores such as Lundbladispora spp., though tolerant of cold conditions, were not tolerant of the coldest conditions (for example below 0 °C).

Serpukhovian—Bashkirian（石炭世）和Gzhelian—早Sakmarian（石炭世晚期—早二叠世早期）两个时间片的古气候模拟表明，在30°S和50°S范围内，Gondwana和Tethys边缘的孢粉组合发生了显著变化。这两个时间切片都显示了一个明显的寒冷气候组合，主要是cheilocardioid孢子，而年平均温度高于19°C的温暖气候似乎促进了产生欧美煤带孢子的植物的生长，如Vestispora fenestrata和Reticulatisporites polygonalis。有可能钩孢孢子的亲本植物，如Lundbladispora spp.，虽然能耐受寒冷条件，但不能耐受最冷的条件（例如低于0°C）。

引用次数: 0

Three new exquisitely preserved haplolepideous mosses from the mid-Cretaceous of Kachin, Myanmar 缅甸克钦邦白垩纪中期三种保存完好的单层藓类植物

IF 1.7 3区地球科学 Q2 PALEONTOLOGY

Review of Palaeobotany and Palynology

Pub Date : 2026-01-01 Epub Date: 2025-10-16 DOI: 10.1016/j.revpalbo.2025.105459

Ya Li , Yong-Dong Wang , Peng-Cheng Wu , Lwin Mar Saing , Chun-Xiang Li , Qiang Wang

Dicranidae (haplolepideous mosses) are the second-largest group of mosses with about 4000 species, and thus the radiation of Dicranidae is considered to be a major event in the evolutionary process of mosses. However, the present fossil record of Dicranidae is rather limited for understanding the early diversification of this successful and speciose lineage. Here, we describe three new exquisitely preserved haplolepideous mosses as Calymperites heinrichsianus, Calymperites marginatus and Calymperites striatus based on 32 whole gametophytes and 10 gametophyte fragments embedded in 16 pieces of Kachin amber from the mid-Cretaceous of northern Myanmar. The three fossil species all exhibit a combination of characters, including lanceolate leaves with a simple costa, clasping leaf bases with large rectangular hyaline cells (cancellinae), and subquadrate to short rectangular, smooth to slightly bulging upper lamina cells. These characters are indicative of Calymperaceae, a monophyletic family that is placed in the core Dicranales. Considering the mid-Cretaceous age and morphological differences to extant species, the present three species of Calymperites probably represent stem group members of Calymperaceae. They, along with previously reported three species of Calymperites (Calymperaceae) and Ditrichites aristatus (Ditrichaceae or Dicranaceae), provide palaeobotanical evidence for the early diversification of Dicranidae in the mid-Cretaceous Kachin amber forest of northern Myanmar. Palaeoecological habitat reconstruction indicates that Calymperites from Kachin amber has obviously ecological niche differentiation.

单枝藓科（Dicranidae）是第二大藓类，约有4000种，因此Dicranidae的辐射被认为是苔藓进化过程中的一个重大事件。然而，目前的化石记录对于了解这种成功的物种谱系的早期多样化是相当有限的。本文基于缅甸北部中白垩世克钦琥珀的16块配子体和32个完整配子体和10个配子体碎片，描述了3种新保存完好的单层藓类，分别为heinrichsianus Calymperites、marginatus Calymperites和striatus Calymperites。这三个化石种都表现出以下特征的组合：披针形叶，有一个简单的边缘；紧扣叶基，有大的矩形透明细胞（cancellinae）；近方形到短矩形，光滑到略鼓的上层细胞。这些特征表明萼花科，一个单系的家庭，被放置在核心的Dicranales。考虑到中白垩世时代和与现存物种的形态差异，这3种萼花可能代表萼花科茎群成员。它们与先前报道的三种Calymperites （Calymperaceae）和dirichites aristatus （dirichaceae或Dicranaceae）一起，为缅甸北部克钦琥珀森林中白垩纪中期Dicranidae的早期多样化提供了古植物学证据。古生态生境重建表明，克钦琥珀的钙辉石具有明显的生态位分化。

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引用次数: 0

First records of Pinuxylon and Xenoxylon wood from the Cretaceous in Shandong Province, Northeast China 山东白垩纪初见梭梭木和Xenoxylon木的记录

IF 1.7 3区地球科学 Q2 PALEONTOLOGY

Review of Palaeobotany and Palynology

Pub Date : 2026-01-01 Epub Date: 2025-10-30 DOI: 10.1016/j.revpalbo.2025.105465

Ruiying Hao , Zhanchun Zou , Zikun Jiang , Kemin Xu , Ning Tian , Zhenguo Ning , Aowei Xie , Xiao Shi , Yongdong Wang , Guohua Xu , Fuxia Zhang

Early Cretaceous petrified wood demonstrates remarkably high diversity across both northern and southern phytoprovinces in China. In recent years, an increasing number of fossil wood discoveries have been made in Cretaceous deposits throughout China, while current research on Early Cretaceous fossil wood remains primarily focused on the northeastern region. However, Cretaceous fossil wood occurrences remain exceptionally scarce in Shandong Province compared with other regions. This study reports the first discovery of well-preserved fossil wood from the Lower Cretaceous in the Jiaolai Basin, represented by two taxa, i.e., Pinuxylon shandongense sp. nov. and Xenoxylon latiporosum (Cramer) Gothan. Notably, P. shandongense sp. nov. constitutes the earliest fossil record of the genus Pinuxylon reported. Phenetic analysis of anatomical characters indicates that this new species is most closely related phylogenetically to P. pinastroides (Kraus) Stockmans et willière. Furthermore, anatomical examination of Xenoxylon latiporosum combined with growth-ring analysis of Keteleerioxylon shandongense indicates that the Jiaolai Basin experienced a seasonal climate during the Early Cretaceous, characterized by localized humid zones within a predominantly arid regime. These discoveries markedly expand the fossil records of Xenoxylon and Pinuxylon, providing unprecedented clarity regarding floristic assemblages in the ecotone between northern and southern phytoprovinces. They yield critical insights into both Mesozoic wood evolution and the paleobiogeography of Cretaceous fossil woods in East Asia. Furthermore, our reconstruction of Early Cretaceous paleoenvironments in the Jiaolai Basin, based on growth-ring analysis, establishes fundamental evidence for Cretaceous paleoclimatic reconstructions in this region.

早白垩世的石化木材在中国北部和南部的植物省都显示出非常高的多样性。近年来，在中国各地白垩纪沉积物中发现了越来越多的木化石，但目前对早白垩世木化石的研究仍主要集中在东北地区。然而，与其他地区相比，山东省白垩纪木材化石产地仍然异常稀少。本文报道了在胶莱盆地首次发现保存完好的下白垩统木材化石，以Pinuxylon shandongense sp. 11和Xenoxylon latiporosum (Cramer) Gothan两个分类群为代表。值得注意的是，P. shandonggense sp. nov.是目前报道的最早的Pinuxylon属化石记录。解剖性状的表型分析表明，该新种与P. pinastroides (Kraus) Stockmans et will的系统亲缘关系最为密切。此外，结合对山东油桐生长年轮的分析，对油桐的解剖分析表明，早白垩世胶莱盆地经历了一个季节性气候，以干旱为主的局部湿润带为特征。这些发现显著扩展了Xenoxylon和Pinuxylon的化石记录，为南北植物省之间过渡带的植物区系组合提供了前所未有的清晰度。它们提供了对中生代木材演化和东亚白垩纪化石木材的古生物地理学的重要见解。此外，在生长年轮分析的基础上重建了胶莱盆地早白垩世古环境，为该地区重建白垩世古气候奠定了基础。

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引用次数: 0

A new Coniopteris (Dicksoniaceae) from the Middle Jurassic of the Baojishan Basin, Gansu, China, with discussion on its heteromorphic pinnule characteristics 甘肃宝鸡山盆地中侏罗统一新种针蕨属（针蕨科）及其异型花序特征探讨

IF 1.7 3区地球科学 Q2 PALEONTOLOGY

Review of Palaeobotany and Palynology

Pub Date : 2026-01-01 Epub Date: 2025-11-08 DOI: 10.1016/j.revpalbo.2025.105467

Lan-Xing Wei, Cun-Lin Xin, Zhi -Peng Jiao, Fang-Fang Zhang, Hong Li

Coniopteris (Dicksoniaceae) is a species-rich and diverse Mesozoic fern genus. Based on well-preserved material from the Middle Jurassic Yaojie Formation in the Baojishan Basin, Gansu, China, we combine macromorphology with microstructural characters to describe a new species—Coniopteris baojishanensis Wei et Xin, sp. nov. The frond is lanceolate and at least bipinnate; the ultimate pinnae bear a basal pair of heteromorphic pinnules; the rachis bears a conspicuous longitudinal ridge. Sterile pinnules are elongate-ovate; fertile pinnules are slightly contracted, ovate. A preliminary analysis of the evolutionary distribution of heteromorphic pinnules in Coniopteris indicates first appearance in the Early Jurassic, a peak in the Middle Jurassic, and marked decline by the Cretaceous. We interpret heteromorphic pinnules as an adaptation to moist, low-light understory conditions: by enlarging area or increasing the degree of dissection, they increase light capture and enhance photosynthetic efficiency, thereby improving competitive ability within the community. This study enriches the Chinese fossil record of Coniopteris and provides new clues to the formation of its heteromorphic pinnules.

刺蕨属（刺蕨科）是中生代蕨类植物中种类丰富、种类多样的一个属。根据甘肃宝鸡山盆地中侏罗统姚界组保存完好的材料，结合宏观形态和微观结构特征，描述了一个新种——coniopteris baojishanensis Wei et Xin， sp. 11。末级羽片生一对异型小羽片；轴有明显的纵脊。不育小穗状花序拉长卵形；可育小叶稍收缩，卵形。对锥翼兽异型小尖的演化分布进行初步分析，表明锥翼兽的异型小尖最早出现于早侏罗世，在中侏罗世达到顶峰，在白垩纪显著下降。我们将异型小叶解释为对潮湿、低光照林下条件的适应：它们通过扩大面积或增加解剖程度，增加光捕获和提高光合效率，从而提高群落内的竞争能力。本研究丰富了刺蕨属的中国化石记录，为其异型小尖的形成提供了新的线索。

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引用次数: 0

Modern pollen and stomatal assemblages in Northeastern China: Vegetation–climate relationships and paleoenvironmental implications 中国东北现代花粉和气孔组合：植被-气候关系及其古环境意义

IF 1.7 3区地球科学 Q2 PALEONTOLOGY

Review of Palaeobotany and Palynology

Pub Date : 2026-01-01 Epub Date: 2025-08-25 DOI: 10.1016/j.revpalbo.2025.105441

Jingfan He , Chunhai Li , Shi-Yong Yu , Xiaocen Li , Beibei Li

Decades of research have revealed large uncertainties in paleoclimate reconstructions based on fossil pollen data. To address this issue, modern studies are essential for improving the understanding of the vegetation–climate relationship. In this study, a total of 45 modern surface soil samples were collected along an east–west precipitation gradient and a north–south temperature gradient in Northeast China. These samples were analyzed to investigate modern pollen–vegetation, stomata–parent plant, and pollen–climate relationships. Using Redundancy Analysis (RDA) and random forest modeling, we found that pollen assemblages can effectively distinguish different vegetation types. Furthermore, stomata analysis demonstrated that Pinus and Larix stomata can reliably indicate the local presence of their parent plants. Both RDA and random forest models identified that the mean air temperature of the coldest month was the primary climatic parameter influencing coniferous pollen distribution, while annual precipitation was the dominant factor controlling herb pollen distribution. This study demonstrates that combining pollen assemblages with stomata analysis can remarkably improve the accuracy of vegetation reconstruction in northeastern China, with coniferous and herb pollen serving as reliable respective indicators of coldest month temperature and annual precipitation.

几十年的研究表明，基于化石花粉数据的古气候重建存在很大的不确定性。为了解决这一问题，现代研究对于提高对植被-气候关系的认识至关重要。本研究在东北地区沿东西降水梯度和南北温度梯度采集了45个现代表层土壤样品。对这些样本进行分析，以探讨现代花粉-植被、气孔-亲本植物和花粉-气候的关系。利用冗余分析（RDA）和随机森林模型，我们发现花粉组合可以有效区分不同的植被类型。此外，气孔分析表明，松木和落叶松的气孔可以可靠地指示其亲本植物在当地的存在。RDA模型和随机森林模型均发现，最冷月份的平均气温是影响针叶树花粉分布的主要气候参数，而年降水量是影响草本植物花粉分布的主导因素。本研究表明，将花粉组合与气孔分析相结合可以显著提高东北地区植被重建的准确性，针叶花粉和草本花粉分别作为最冷月温度和年降水量的可靠指标。

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引用次数: 0

Preliminary cuticular studies on plant remains from the Middle Jurassic (Bathonian) of NW Jordan 西北约旦中侏罗统（Bathonian）植物遗存的初步表皮研究

IF 1.7 3区地球科学 Q2 PALEONTOLOGY

Review of Palaeobotany and Palynology

Pub Date : 2026-01-01 Epub Date: 2025-08-22 DOI: 10.1016/j.revpalbo.2025.105437

Hong-Yu Chen , Abdallah Abu Hamad , Yong-Dong Wang , Dieter Uhl

The allochthonous macroflora of the Middle Jurassic (Bathonian) Arda Formation in Northwest Jordan has previously been described based on morphological features of plant remains, without reference to microstructures (i.e., cuticles). This study examines seven types of cuticle from the same flora, all of which belong to gymnosperms. The identified cuticles include bennettitaleans (Otozamites sp. and Bennettitales spp.), and conifers (scale-leaved conifer and probably conifers with longitudinally arranged stomatal rows). The discovery of cuticles provides the first record of conifers in the Middle Jurassic flora of NW Jordan, filling a gap in the previous knowledge. The macroflora, comprising sphenophytes, ferns, bennettitaleans, and conifers (including scale-leaved conifers), shows strong similarities to other Jurassic floras in the Middle East, particularly resembling that of the Middle Jurassic Imran Formation of Makhtesh Ramon in the Negev (Southern Israel). These additions to the known facts about the floral assemblage by microstructural analyses significantly provide more information to support existing reconstructions of the palaeoenvironmental and palaeoclimatic conditions in NW Jordan during the Middle Jurassic.

约旦西北部中侏罗统（Bathonian） Arda组的异域大型植物区系以前是基于植物遗骸的形态学特征而没有参考微观结构（即角质层）来描述的。本研究研究了来自同一植物区系的七种角质层，它们都属于裸子植物。已鉴定的角质层包括针叶树（Otozamites sp.和Bennettitales sp.）和针叶树（鳞叶针叶树和可能具有纵向排列气孔排的针叶树）。角质层的发现提供了约旦西北部中侏罗世针叶树区系的第一个记录，填补了以往认识的空白。大型植物区系包括蕨类植物、蕨类植物、贝尼蒂塔科植物和针叶树（包括鳞片叶针叶树），与中东的其他侏罗纪植物区系非常相似，特别是与内盖夫（以色列南部）Makhtesh Ramon的中侏罗世Imran组的植物区系相似。这些通过显微结构分析对植物组合的已知事实的补充，为支持中侏罗世约旦西北部古环境和古气候条件的现有重建提供了更多的信息。

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