Jurassic Daohugou Research Articles

A new fossil sinoalid species from the Middle Jurassic Daohugou beds (Insecta: Hemiptera: Cercopoidea)

Fu, Y., Cai, C. & Huang, D., October 2017. A new fossil sinoalid species from the Middle Jurassic Daohugou beds (Insecta: Hemiptera: Cercopoidea). Alcheringa 42, 94–100. ISSN 0311-5518.A new fossil species, Luanpingia daohugouensis sp. nov., belonging to the family Sinoalidae is described from the Middle to Upper Jurassic Daohugou beds of Inner Mongolia, China, on the basis of two well-preserved complete specimens. The described species of Sinoalidae are reviewed and Jiania gracila is considered a junior synonym of Jiania crebra. The new discovery increases the palaeodiversity of sinoalids from the Daohugou beds. It also indicates stratigraphic correlation between the Daohugou beds, the Haifanggou Formation at Haifeng, Beipiao City, West Liaoning Province, and the Jiulongshan Formation at Zhouyingzi, Luanping County, Hebei Province. All of these units host the ‘early assemblage’ of the Yanliao biota.Yanzhe Fu [yzfu41@163.com], Key Laboratory of Economic Stratigraphy and Palaeogeography, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, PR China; University of Science and Technology of China, Hefei 230026, PR China; Chenyang Cai [cycai@nigpas.ac.cn], Key Laboratory of Economic Stratigraphy and Palaeogeography, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, PR China; Diying Huang* [dyhuang@nigpas.ac.cn], State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, PR China.

Insect herbivory and plant defense on ginkgoalean and bennettitalean leaves of the Middle Jurassic Daohugou Flora from Northeast China and their paleoclimatic implications

Interactions between terrestrial arthropods and plants play a significant role in terrestrial ecosystems. Research on plant–insect interactions through geologic time provides valuable information for studying insect behavior and plant structure, understanding their coevolution, as well as analyzing climate change. In this paper, we choose fossil ginkgoalean and bennettitalean leaves as the plant hosts to study insect herbivory in the Middle Jurassic Daohugou area. Seven damage types of four functional feeding groups have been identified. Of the four functional feeding groups, margin feeding is the most common, indicating an abundance of insects with chewing mouthparts. Ginkgoalean leaves, probably because of their chemical defense, suffered less severe insect damage than bennettitalean leaves. Physical defense has also been observed in various genera of the bennettitalean leaves. Significantly, leaves of Anomozamites had a shaggy indumentum on the abaxial leaf surface and long stiff hairs along the rachis protecting them from insect herbivory. Our results indicate that the climate in the Middle Jurassic of the Daohugou area was relatively warm and humid. This work contributes to the study of plant–insect coevolution in the Daohugou Biota and provides more proxy data for understanding the Middle Jurassic paleoclimate and paleoenvironment in Daohugou area.

A brief introduction to the Middle Jurassic Daohugou Flora from Inner Mongolia, China

The Middle Jurassic Daohugou Biota is named for a fossil assemblage after a village in Ningcheng County, Inner Mongolia, China. Over the last twenty-five years, abundant, exceptionally well-preserved fossils have been reported from various localities in the Daohugou area. While the fauna of the Daohugou Biota has been well-studied, little research has been done on the flora. In this paper, we establish the Daohugou Flora for the first time and document its high diversity of plants which are mainly composed of Filicopsida, Cycadopsida, Ginkgopsida, Czekanowskiales and Coniferopsida. We recognize 49 species belonging to 32 genera, including a new species Sphenobaiera eximia sp. nov. Na, Sun et Wang. An emended diagnosis of Anomozamites villosus Pott, McLoughlin, Wu et Friis is presented. The floristic composition indicates that the Daohugou Flora belongs to the North Floristic Province and it reflects a warm-temperate and humid climate with seasonal temperature and precipitation variations. This report, representing the first systematic treatment of fossil plants from the Daohugou area, is of great significance for understanding the Daohugou Biota. It also provides important information for reconstructing the paleoclimate and paleoenvironment in this area, and lays a solid foundation for studying co-evolution of plants and animals of the Daohugou Biota.

New data on Jurassic Sinoalidae from northeastern China (Insecta, Hemiptera)

AbstractThe Sinoalidae, as one of the three Mesozoic froghopper families, was recently recognized from the latest Middle–earliest Late Jurassic Daohugou Biota of northeastern China. We herein report some new materials from the same horizon and locality, providing some new insights on morphological diversity and evolution of this family.Shufania haninew genus new species indicates that the relative branching position of veins M and CuA of the forewing is highly variable within the Sinoalidae and not appropriate for family-level diagnosis. The venations of three reported sinoalid hind wings are conservative, likely due to its simplified topology and reduced terminal branches. Color patterns of hind wings are likely variable for different sinoalid froghoppers. However, considering that color pattern is easily weakened or even erased by diagenetic processes for imprint fossils, the morphological character is not reliable for distinguishing different sinoalid taxa. Additionally, our new material suggests that the number of lateral spines of the hind tibia can vary intra-individually for sinoalids, just as in some recent froghoppers.

Palaeoboganium gen. nov. from the Middle Jurassic of China (Coleoptera: Cucujoidea: Boganiidae): the first cycad pollinators?

Palaeoboganium jurassicum gen. et sp. nov. is described from the Middle Jurassic Daohugou formation in northern China. Palaeoboganium jurassicum is the first fossil representative of the enigmatic family Boganiidae, currently known from Gondwanan localities in South Africa, Australia and New Caledonia. Based on a phylogenetic analysis of adult external characters, Palaeoboganium is recovered as the sister group of Paracucujus Sen Gupta & Crowson and Metacucujus Endrödy-Younga & Crowson (Paracucujinae), whose adults pollinate cycads in Australia and South Africa. Based on phylogenetic evidence this taxon is suggested to be a potential pollinator of Jurassic cycads.http://zoobank.org/urn:lsid:zoobank.org:pub:7EDB8295-5EF3-40ED-9A84-479AD23611AF

Jurassic lacewings (Insecta: Neuroptera) of Western Siberia

A lacewing assemblage from the Middle Jurassic of the West Siberian Kubekovo locality, the only know Jurassic lacewing fauna of Russia, is discussed. One new genus and two new species of Osmylidae from Kubekovo, Kubekius multiramosus gen. et sp. nov. and Epiosmylus longus sp. nov. are described. Thaumatomerobius mirabilis Ponomarenko, 1985 is transferred to the family Saucrosmylidae, which has only been recorded in the Middle Jurassic of Daohugou.

New Earwigs in Protodiplatyidae (Insecta: Dermaptera) from the Middle Jurassic Jiulongshan Formation of Northeastern China.

Two new genera, each with a new species, Perissoderma triangulum gen. et sp. nov. and Abrderma gracilentum gen. et sp. nov., of the family Protodiplatyidae are described from the latest Middle Jurassic Jiulongshan Formation of Daohugou, Inner Mongolia, China. They are the first records of Protodiplatyidae from the latest Middle Jurassic Daohugou locality. Both new genera are assigned to Protodiplatyidae based mainly on diagnostic characters of antennae, pronotum, tegmina, tarsi and the distinct long, slender, multi-segmented cerci. We summarize the distributions and ages of all described fossil Archidermaptera and infer that the Dermaptera might have originated in Eurasia and started their diversification and migration to a worldwide distribution. The new taxon of Abrderma gracilentum gen. et sp. nov. provides one more example of an early earwig with well-preserved pentamerous tarsi on the hind leg.

Conchostracans of the Middle–Late Jurassic Daohugou and Linglongta beds in NE China

The Yanliao biota is widely distributed in western Liaoning, northern Hebei, and the areas adjacent to eastern Inner Mongolia. The biota is excavated from two typical fossil-bearing beds: the Daohugou fossil-bearing beds in Daohugou, Jiangzhangzi, and Chentaizi, Wuhua Town, Ningcheng County, Inner Mongolia; and the Linglongta fossil-bearing beds in Daxishan, Linglongta Town, Jianchang County, Liaoning Province and Bawanggou, Mutoudeng Town, Qinglong County, Hebei Province. Previously, it was thought that the Daohugou and Linglongta beds are correlatable in age and fossil content and that the conchostracans from both the Daohugou and Linglongta beds could all be attributed to the genus Euestheria. However, clear punctae but no reticulations are observed on all conchostracans collected from the Daohugou beds, which excludes them from Euestheria. In this study, only one species, Triglypta haifanggouensis n. comb., is recognized based on detailed investigations of the ornamentations of hundreds of specimens from the Daohugou beds. In addition to the recently reported Tianzhuestheria jianchangensis, another species, Liaoxiestheria linglongtaensis n. gen. n. sp., is identified in this study based on a large number of specimens from the Linglongta beds in Daxishan. Isotope analysis has shown that the Daohugou and Linglongta beds are approximately 165Ma and 160Ma old, respectively. Therefore, the fossils from the Daohugou beds represent an early assemblage and the fossils from the Linglongta beds represent a late assemblage of the Yanliao biota. This study indicates that the conchostracan assemblage of the Daohugou beds is a typical Triglypta–Qaidamestheria assemblage, which is widely distributed in the Middle Jurassic of North China, and the conchostracan assemblage of the Linglongta beds represents the late, succeeding Triglypta–Qaidamestheria assemblage.

The first fossil salmonfly (Insecta: Plecoptera: Pteronarcyidae), back to the Middle Jurassic.

BackgroundThe fossil record of Plecoptera (stoneflies) is considered relatively complete, with stem-groups of each of the three major lineages, viz. Antarctoperlaria, Euholognatha and Systellognatha (and some of their families) represented in the Mesozoic. However, the family Pteronarcyidae (the salmonflies; including two genera, Pteronarcys and Pteronarcella) has no fossil record to date, and the family has been suggested to have diverged recently.ResultsIn this paper, we report on a set of specimens belonging to a new fossil species of stonefly, discovered from the Middle Jurassic Daohugou locality (China). Our comparative analysis of wing venation and body characters demonstrates that the new species belongs to the Pteronarcyidae, and is more closely related to Pteronarcys than to Pteronarcella. However, it differs from all known species of the former genus. It is therefore assigned to a new genus and named Pteroliriope sinitshenkovae gen. et sp. nov. under the traditional nomenclatural procedure. The cladotypic nomenclatural procedure is also employed, with the resulting combination Pteroliriope nec Pteronarcys sinitshenkovae sp. nov. ConclusionsThe first discovery of a fossil member of the Pteronarcyidae demonstrates that the corresponding lineage is not a very recent offshoot but was already present ca. 165 million years ago. This discovery concurs with the view that divergence of most stonefly families took place very early, probably in the Triassic, or even in the Permian. This contribution demonstrates the need for (re-)investigations of the systematics of fossil stoneflies to refine divergence date estimates for Plecoptera lineages.

New material of Sigmaboilus (Insecta, Orthoptera, Prophalangopsidae) from the Jurassic Daohugou Beds, Inner Mongolia, China

ABSTRACTFive orthopteran specimens from the uppermost Middle–lowermost Upper Jurassic of Daohugou, Inner Mongolia, China are described and attributed to the genus Sigmaboilus Fang, Zhang & Wang, 2007 (Prophalangopsidae); and a new species, S. calophlebius sp. nov., is established herein. The diagnostic characters for Sigmaboilus are revised and a key to species of Sigmaboilus, based on male forewings, is provided. Intraspecific variation in forewings of this genus is also discussed.

Biostratinomy of bivalves from Jurassic and Early Cretaceous lakes of NE China

Bivalves are comparatively rare elements of the benthic faunas of Jurassic and Early Cretaceous lakes of northeastern China. In the past, Arguniella and Sphaerium have been regarded as belonging to the communities that populated the offshore areas of lake Sihetun, represented by the Lower Cretaceous Yixian Formation of western Liaoning. A detailed biostratinomic analysis of these bivalves suggests, however, that they were introduced from marginal lake habitats where they were reworked by storms and occasionally were transported in suspension offshore. Some of these bivalves were dead and settled on the lake floor as single valves or articulated but opened valves (butterfly position). Others were transported alive but died on their arrival at the bottom due to inimical conditions caused by reduced oxygen levels. A similar interpretation is given for articulated valves of Ferganoconcha in thinly bedded siltstones of the Jurassic Daohugou Fossil Beds of Inner Mongolia, which were also carried alive by hyperpycnal flows to their burial site where they died due to lack of oxygen.

New Phoenicopsis leaves (Czekanowskiales) from the Middle Jurassic Daohugou Biota, China and their roles in phytogeographic and paleoclimatic reconstruction

The Middle Jurassic Jiulongshan Formation, distributed in the Daohugou area, Ningcheng County, Inner Mongolia, China, is famous for yielding abundant, well-preserved fossil vertebrates and invertebrates. However, only a few fossil plants have been reported to date. In this paper, one new species of Czekanowskiales, Phoenicopsis (Windwardia) daohugouensis n. sp. is described based upon leaf morphology and epidermal characters. The new species represents the first record of Czekanowskiales from the Daohugou Biota. The discovery of the new species reveals the significance of the Daohugou flora as one of the deciduous forests in the temperate-warm temperate zone of Eurasia during the Middle Jurassic.

A new family of ‘libelluloid’ dragonflies from the Middle Jurassic of Daohugou, northeastern China (Odonata: Anisoptera: Cavilabiata)

Nel, A. & Huang, D.Y., 8.5.2015. A new family of ‘libelluloid’ dragonflies from the Middle Jurassic of Daohugou, northeastern China (Odonata: Anisoptera: Cavilabiata). Alcheringa 39, 525–529. ISSN 0311-5518A new well-preserved Middle Jurassic fossil of Cavilabiata is described and attributed to a new family (Daohugoulibellulidae), genus and species (Daohugoulibellula lini), from the Daohugou beds of China. Together with examples of Juralibellulidae from the same outcrop, they represent the oldest records of the Cavilabiata. The potential closest relative of the new family could be the Late Jurassic Nannogomphidae, suggesting a significant diversity of Cavilabiata during the Middle Jurassic.André Nel [anel@mnhn.fr], Institut de Systématique, Évolution, Biodiversité, ISYEB—UMR 7205—CNRS, MNHN, UPMC, EPHE, Muséum national d’Histoire naturelle, Sorbonne Universités, 57 rue Cuvier, CP 50, Entomologie, F-75005, Paris, France; Diying Huang [dyhuang@nigpas.ac.cn], State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, PR China.

The oldest root-eating beetle from the Middle Jurassic of China (Coleoptera, Monotomidae)

Cai, C.-Y., Ślipiński, A. & Huang, D.-Y., 31.3.2015. The oldest root-eating beetle from the Middle Jurassic of China (Coleoptera, Monotomidae). Alcheringa 39,488–493. ISSN 0311-5518.Jurorhizophagus alienus gen. et sp. nov., a new fossil root-eating beetle, is described and figured based on an exceptionally well-preserved impression fossil from the Middle Jurassic Daohugou beds (ca 165 Ma), Inner Mongolia, northeastern China. It represents the earliest fossil Monotomidae known to date. Jurorhizophagus can not be assigned to either of two subfamilies Monotominae or Rhizophaginae based on the unique combination of many unusual characters, including an 11-segmented antenna with a 3-segmented club, the presence of a distinct frontoclypeal suture and transverse pronotum with a median longitudinal groove. The discovery of a new genus from the Middle Jurassic highlights the antiquity of Monotomidae and provides new information about the phylogenetic relationships between Monotomidae and its allied families.Chen-Yang Cai [caichenyang1988@163.com], Di-Ying Huang [dyhuang@nigpas.ac.cn] (corresponding author), State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, PR China; Adam Ślipiński [adam.slipinski@csiro.au], Australian National Insect Collection, CSIRO National Collections Australia, GPO Box 1700, Canberra ACT 2601, Australia.

Jurassic artematopodid beetles and their implications for the early evolution of Artematopodidae (Coleoptera)

AbstractFossil Artematopodidae are rarely collected and previously confined to middle Eocene Baltic amber. Here we report the first definitive artematopodid, Sinobrevipogon jurassicus gen. et sp.n., from the Middle Jurassic Daohugou beds (c. 165 Ma) in Inner Mongolia, northeastern China. It exhibits a number of defining features of Artematopodidae, including paired carinae on prosternum and an internal apical interlocking tongue on the ventral side of each elytron. However, it differs from any modern Artematopodidae by having the mesocoxal cavitiy closed by the mesepimeron and the anterolateral edge of metanepisternum. The discovery of this new genus represents the earliest fossil record for Artematopodidae, highlighting the antiquity of the family. The systematic positions of Forticatinius Tan & Ren and Tarsomegamerus Zhang are discussed, and the latter is formally transferred to Artematopodidae. Phylogenetic relationships within Artematopodidae were investigated to elucidate the relationships between the two Jurassic genera and Recent genera. Eleven in‐group taxa and two out‐groups were included in a cladistic analysis based on 30 adult characters; the resulting tree recovered the family Artematopodidae in three clades: (i) Electribius authority, (ii) Ctesibius authority + Brevipogon authority + Sinobrevipogon + Tarsomegamerus and (iii) the remaining Recent genera, including Allopogonia authority.

Diagenesis of bivalves from Jurassic and Lower Cretaceous lacustrine deposits of northeastern China

AbstractIn contrast to the numerous excellently preserved arthropods, vertebrates and plants from the Mesozoic lacustrine fossil lagerstätten of northeastern China, which have calcium phosphate or organic skeletons, the preservation of taxa with a calcareous skeleton is fairly poor. Here we investigate, using a scanning electron microscope and energy dispersive X-ray spectrometer, the preservational modes of bivalves from the Jurassic Daohugou Fossil Beds of Inner Mongolia and the Lower Cretaceous Yixian Formation of eastern Liaoning. The Jurassic bivalve Ferganoconcha sibirica is preserved as strongly compressed composite moulds which contain remains of the organic periostracum. In the Yixian Formation, the bivalves Sphaerium anderssoni and Arguniella ventricosa occur as compacted internal, external or composite moulds or are preserved with a silicified shell, and rarely with a shell consisting of iron hydroxides, which had replaced pyrite during late diagenesis/weathering. Silicification produced partly fabric-replacive microcrystalline quartz and partly void-filling megaquartz crystals after the carbonate shell had been dissolved. Films of authigenic aluminosilicate minerals, partly secondarily silicified, cover the exterior and interior shell surfaces. Occasionally, early diagenetic pyrite crystals, now oxidized to iron hydroxides, filled shell cavities forming internal moulds and rarely replaced the bivalve shell. The poor preservation of the bivalves reflects the environment and water chemistry of these lakes, which were heavily influenced by volcanic processes. Frequent ash deposition and decomposition of volcanic glass particles created acidic and alkaline lake and interstitial waters, which led to early diagenetic formation of authigenic aluminosilicate minerals, ferruginous internal moulds, dissolution of shell carbonate and silicification of shells.

A new species of Kalligramma Walther, 1904 (Insecta, Neuroptera, Kalligrammatidae) from the Middle–Upper Jurassic of Daohugou, Inner Mongolia, China

Liu, Q., Khramov, A.V. & Zhang, H.C., 21.4.2015. A new species of Kalligramma Walther, 1904 (Insecta, Neuroptera, Kalligrammatidae) from the Middle–Upper Jurassic of Daohugou, Inner Mongolia, China. Alcheringa 39, XXX–XXX. ISSN 0311-5518.A new species of Kalligramma Walther, 1904 (Insecta, Neuroptera, Kalligrammatidae) is described as Kalligramma delicatum based on a nearly complete forewing from the Middle–Upper Jurassic of Daohugou, Inner Mongolia, China. The new species is characterized by the narrow costal space with sinuous costal veinlets, Rs with eight primary branches, MP with five pectinate branches and 1A forked beyond its mid-length. The forewing of Kalligramma can be distinguished from the hind wing by its ovate rather than subtriangular shape, significantly wider costal space and more numerous crossveins between costal veinlets. The new species is interpreted to represent a forewing based mainly on its shape.Qing Liu [qingliu1029@yeah.net] and Haichun Zhang [hczhang@nigpas.ac.cn], State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, 210008, PR China; Aleksandr V. Khramov [a-hramov@yandex.ru], Borissiak Paleontological Institute of the Russian Academy of Sciences, Moscow, 117997, Russia. Received 10.12.2014; revised 11.2.2015; accepted 17.2.2015.

A new species of Aboilus (Insecta, Orthoptera) from the Jurassic Daohugou beds of China, and discussion of forewing coloration in Aboilus

Wang, H., Li, S., Zhang, Q., Fang, Y., Wang, B. & Zhang, H., 13.02.2015. A new species of Aboilus (Insecta, Orthoptera) from the Jurassic Daohugou beds of China, and discussion of forewing coloration in Aboilus. Alcheringa 39, xxx–xxx. ISSN 0311-5518He Wang* [wanghe0701@163.com], Sha Li* [lisha.paleo@gmail.com], Qi Zhang* [zhqi1105@126.com], Yan Fang [yanfang@nigpas.ac.cn], Bo Wang† [bowang@nigpas.ac.cn] and Haichun Zhang [hczhang@nigpas.ac.cn], State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, PR China.*Also affiliated with University of Chinese Academy of Sciences, Beijing 100049, PR China. †Also affiliated with Steinmann Institute, University of Bonn, Bonn 53115, Germany.A new species of Aboilinae (Orthoptera: Prophalangopsidae), Aboilus perbellus, is described and illustrated based on three well-preserved forewings recovered from the Middle–Upper Jurassic Daohugou beds of Inner Mongolia, China. The new species differs from all congeneric forms in its special forewing coloration and features of its wing venation. To date, three types of forewing coloration have been found among different species of Aboilus at Daohugou, suggesting that these taxa inhabited different ecotopes.

The oldest aphid of the family Oviparosiphidae (Hemiptera: Aphidoidea) from the Middle Jurassic of China

We describe herein a new aphid genus and species Daoaphis magnalata gen. et sp. n., belonging to the extinct family Oviparosiphidae from the Middle Jurassic (ca. 165 Ma) Daohugou beds, Ningcheng County, Inner Mongolia, Northeast China. The new genus is considered to be the earliest certain representative of this family and the body remains of one of the oldest aphids. The evolutionary position of this new genus and distribution of the family are discussed.

The Middle Jurassic palynology of the Daohugou area, Inner Mongolia, China, and its implications for palaeobiology and palaeogeography

Although abundant palaeontological discoveries have been documented in the Daohugou area in Chifeng City, Inner Mongolia, China, no record of the palynoflora has been reported previously. We collected palynological samples from the same level as megafossil plants, and conducted a detailed taxonomic analysis of the pollen and spores in order to augment the megafossil record, to extend documentation of the Middle Jurassic palynoflora, and to examine implications for palaeogeography, palaeoecology and palaeoclimate in north China during the Jurassic. Forty-four genera and 87 species of palynomorphs were identified. Pollen of gymnosperms dominates numerically, followed by spores of pteridophytes. Most taxa, including bisaccate pollen of conifers, monosulcate pollen of Cycadales, Bennettitales or Ginkgoales, along with Cyathidites, Deltoidospora, Granulatisporites, Osmundacidites and Classopollis, are common elements known elsewhere during the Jurassic. Based on comparisons with similar palyno-assemblages in the Jurassic of China, the age of the Daohugou locality is interpreted to be late Middle Jurassic. According to the palaeobotanical and palaeopalynological information, the Daohugou flora is attributed to the warm-temperate to medium temperate zone with seasonal change. By studying the ecological and climatic preferences of the main plant groups, five types of palaeolandscape in the Jurassic of Daohugou are proposed: (1) mesic upland (Pinaceae, Podocarpaceae, Czekanowskiales and Bennettitales), (2) mesic lowland (Pinaceae, Podocarpaceae, Ginkgoales, Czekanowskiales and Cycadales, together with some hygrophilous bryophytes, lycophytes and ferns), (3) dry upland (Czekanowskiales), (4) dry lowland (Czekanowskiales and Cheirolepidiaceae) and (5) waterside (plants growing along streams or near lakes, including bryophytes, lycophytes, ferns and probably some pteridosperms). According to the plant assemblage, the Daohugou flora corresponds to that of the Middle Jurassic Northern Floristic Province of China, which experienced a transition from a humid, warm climate to a hotter climate with seasonal drought.