麻黄属种子表面纹饰的多样性及其系统学意义

麻黄属(Ephedra)的分子系统学研究结果明显不同于基于雌球果成熟苞片质地的传统分类,指出麻黄属的形态特征和分子特征存在冲突。对麻黄属53种植物的种子表面微形态特征进行了研究,结果表明麻黄属的种子表面纹饰包括3种主要类型,即横向片层型(T型)、乳突型(P型)和扁平型(S型)。麻黄属的种子表面纹饰特征既与宏观形态特征不相关,也与分子系统学的研究结果不相关,表明该属微形态特征、宏观形态特征和分子特征之间的更多冲突。在欧洲和东亚发现的与买麻藤类有关的几个大化石带有T型种子表面纹饰,暗示具有T型纹饰的麻黄类植物在早白垩纪就十分多样。     

Ontogeny and Metamorphic Patterns of Female Reproductive Organs of Ephedra sinica Stapf (Ephedraceae)

Ontogeny and metamorphic development of female reproductive organs in Ephedra sinica Stapf were surveyed. At the end of April, female cones began to initialize from the vegetative buds. Pollination occurred in mid-May and seeds matured at the beginning of July. The ontogenetic pattern of female reproductive organs of E.sinica is basically similar to that of E.　distachya L. The foliar nature of the outer envelope of the ovule in Ephedra is corroborated. Reduction of ovule number as a tendency of speciation in the genus is substantiated by the occurrence of tri-ovulate cones coupled with comprehensive characters of the genus. The metamorphic patterns as well as the leaf nature of the outer envelope indicate that female cones of Ephedra are compound while the female reproductive units of the ovulate cone have been reduced from secondary reproductive shoots similar to those of cordaites by phylogenetic transformation, fusion and reduction. Each fertile bract together with its axillary female reproductive unit composed the Bract Scale and Seed Scale Complex of Ephedra .     

草麻黄雌性生殖器官的个体发生和畸形式样研究

用扫描电镜观察并分析了草麻黄（Ephedra sinica Stapf)雌性生殖器官的个体发生及畸形式样。该种雌球花个体发生式样与麻黄属其他具双胚朱球果的类群基本一致。外珠被以一对近轴侧突起物出现以及该突起物与苞片成交互排列的发育式样支持前人关于该构造叶性本质的论述,而内珠被与珠心的发育相关,因此可能是真正的珠被,基于返祖性畸形三胚珠球花的出现及其他形态特征,认为麻黄属胚珠数目减少是该属内物种特化的一个趋势;利用雌球花发育的畸形现象及外珠被的形态学本质提出麻黄属的雌球果为复轴性构造,雌性生殖单位是由类似于科达类的次级生殖枝构造经系统发育变态,融合,简化成现存的生胚珠构造式样,每一可育苞片及其腋内雌性生殖单位共同组成了麻黄属的苞鳞－种鳞复合体。     

Former diversity of Ephedra (Gnetales): evidence from Early Cretaceous seeds from Portugal and North America

BACKGROUND AND AIMS: The extant species of the seed plant group Gnetales (Ephedra, Gnetum and Welwitschia) have been considered a remnant of a much greater, now extinct, diversity due to the pronounced differences in form and ecology among the genera. Until recently, this hypothesis has not been supported by evidence from the fossil record. This paper adds to the expanding information on Gnetales from the Early Cretaceous and describes coalified seeds from Barremian-Albian localities in Portugal and USA. METHODS: The fossils were extracted from sediment samples by sieving in water. Adhering mineral matrix was removed by chemical treatment. Seeds were investigated using light and scanning electron microscopy. Morphology and anatomy of the seeds were documented and compared with those of extant species. KEY RESULTS: The fossils share characters with extant Ephedra, for example papillae on the inner surface of the seed envelope and in situ polyplicate pollen grains that shed the exine during germination. They differ from extant Ephedra seeds in morphological and anatomical details as well as in their smaller size. Two new species of Ephedra are described together with one species assigned to a new genus of Gnetales. Other Ephedra-like seeds, for which pollen and critical morphological details are currently unknown, are also present in the samples. CONCLUSIONS: These Cretaceous seeds document that key reproductive characters and pollen germination processes have remained unchanged within Ephedra for about 120 million years or more. There is sufficient variety in details of morphology to suggest that a diversity of Ephedra and Ephedra-like species were present in the Early Cretaceous flora. Their presence in Portugal and eastern North America indicates that they were widespread on the Laurasian continent. The fossil seeds are similar to seeds of Erdtmanithecales and this supports the previously suggested relationship between Erdtmanithecales and Gnetales.     

Ontogeny of triovulate cones of Ephedra intermedia and origin of the outer envelope ofovules of Ephedraceae

The origin of the two envelopes of ovules of extant species of Ephedraceae has been discussed previously in different ways based only on investigation of bimerous female cones. To generalize the characteristics of the two envelopes, ontogenetic and metamorphic patterns of the two envelopes of ovules of trimerous female cones of Ephedra intermedia Schrenk et C. A. Meyer were studied using scanning electron microscopy. The outer envelope of the ovules of triovulate female cones of Ephedra intermedia is initiated as two adaxial lateral protuberances. Thus, the outer envelope of extant Ephedraceae may have been derived from three foliar components of the second proximal whorl of the secondary reproductive shoot of the ancestral compound female cone. Two adaxial lateral foliar components likely gave rise to the outer envelope by fusion, while the third abaxial foliar component was reduced. These new findings suggest that the Gnetales derived their female cones from trimerous multiaxial female reproductive organs of their progenitors. Some evolutionary aspects of female cones of Ephedra are also discussed.     

吉林延边早白垩世大拉子组植物化石新类型--星学异麻黄

报道产自吉林省延边早白垩世大拉子组植物化石新类型——星学异麻黄(Alloephedra xingxuei gen.et sp.nov.)。大拉子组的时代处于早白垩世的阿普特期-阿尔必期(Aptian-Albian)。化石标本保存了植物的茎枝、雌球花及种子;该种的茎枝分节,节间具细纵槽纹,叶退化,雌球花单个着生于小枝顶部,种子成对且种子顶部宿存珠孔管等特征与现存麻黄科植物最为相近,因此可能属于麻黄科。     

Welwitschiaceae from the Lower Cretaceous of northeastern Brazil

Welwitschiaceae, a family in the Gnetales, is known today from only one extant species, Welwitschia mirabilis. This species is distributed in the Namibian desert, along the western coast of southern Africa, about 10 km inland from the coast. Very little is known about the fossil record of this family. Lower Cretaceous megafossils of various organs, assigned to Welwitschiaceae, are presented here. These fossils include young stems with paired cotyledons attached (Welwitschiella austroamericana n. gen. et sp.), isolated leaves (Welwitschiophyllum brasiliense n. gen. et sp.), and axes bearing male cones (Welwitschiostrobus murili n. gen. et sp.). They were collected in the Crato Formation, which is dated by palynomorphs and ostracods as Late Aptian (114 to 112 million years ago). These sediments are exposed in the Araripe Basin of northeastern Brazil. This study brings together new information of the megafossil record of Welwitschia-like plants and also reports of pollen said to be similar to that of Welwitschia from Lower Cretaceous sediments.     

宁夏贺兰山斑子麻黄荒漠群落特征

斑子麻黄(Ephedra rhytidosperma)群落是贺兰山山麓特有荒漠植物群落。该研究于2019年调查了16个斑子麻黄典型群落, 分析了物种组成的区系特征, 运用TWINSPAN软件对16个斑子麻黄群落进行了分类。结果表明: 1)斑子麻黄群落物种组成以禾本科、菊科、豆科和蒺藜科植物为主, 隶属13科24属27种。2)斑子麻黄荒漠群落生活型以多年生草本(44.44%)和灌木(29.63%)为主; 区系以古地中海成分(44.44%)和东亚成分(33.33%)为主; 水分生态类型以旱生(40.74%)和强旱生(37.04%)为主。3)根据生活型和优势度, 斑子麻黄群落可分为斑子麻黄-半灌木荒漠(Ephedra rhytidosperma - Semi-Shrub Desert)和斑子麻黄-草本荒漠(Ephedra rhytidosperma - Herb Desert) 2个群丛组, 进一步划分为3个群丛: 斑子麻黄-刺旋花-兴安胡枝子 半灌木与草本荒漠(Ephedra rhytidosperma - Convolvulus tragacanthoides - Lespedeza davurica Semi-Shrub and Herb Desert)、斑子麻黄-刺旋花-中亚细柄茅 半灌木与草本荒漠(Ephedra rhytidosperma - Convolvulus tragacanthoides - Ptilagrostis pelliotii Semi-Shrub and Herb Desert)和斑子麻黄-中亚细柄茅 半灌木与草本荒漠(Ephedra rhytidosperma - Ptilagrostis pelliotii Semi-Shrub and Herb Desert)。     

From near extinction to diversification by means of a shift in pollination mechanism in the gymnosperm relict Ephedra (Ephedraceae,Gnetales)

Pollination in gymnosperms is usually accomplished by means of wind, but some groups are insect‐pollinated. We show that wind and insect pollination occur in the morphologically uniform genus Ephedra (Gnetales). Based on field experiments over several years, we demonstrate distinct differences between two Ephedra species that grow in sympatry in Greece in pollen dispersal and clump formation, insect visitations and embryo formation when insects are denied access to cones. Ephedra distachya, nested in the core clade of Ephedra, is anemophilous, which is probably the prevailing state in Ephedra. Ephedra foeminea, sister to the remaining species of the genus, is entomophilous and pollinated by a range of diurnal and nocturnal insects. The generalist entomophilous system of E. foeminea, with distinct but infrequent insect visitations, is in many respects similar to that reported for Gnetum and Welwitschia and appears ancestral in Gnetales. The Ephedra lineage is well documented already from the Early Cretaceous, but the diversity declined dramatically during the Late Cretaceous, possibly to near extinction around the Cretaceous–Palaeogene boundary. The clade imbalance between insect‐ and wind‐pollinated lineages is larger than expected by chance and the shift in pollination mode may explain why Ephedra escaped extinction and began to diversify again.     

The seed cone Eathiestrobus gen. nov.: Fossil evidence for a Jurassic origin of Pinaceae

? Premise of the study: Pinaceae and nonpinoid species are sister groups within the conifer clade as inferred from molecular systematic comparisons of living species and therefore should have comparable geological ages. However, the fossil record for the nonpinoid lineage of extant conifer families is Triassic, nearly 100 million years older than the oldest widely accepted Lower Cretaceous record for Pinaceae. An anatomically preserved fossil conifer seed cone described here extends the stratigraphic range of Pinaceae nearly 30 million years, thus reducing the apparent discrepancy between evidence from the fossil record and inferences from systematic studies of living species. ? Methods: Material was prepared as serial thin sections by the cellulose acetate peel technique, mounted on microscope slides, and viewed and photographed using transmitted light. ? Key results: A large cylindrical cone consisting of bract-scale complexes that diverge from the cone axis in a helical phyllotaxis has bracts and scales that separate from each other in the midregion and are of equal length and of nearly equal width. The cone has two inverted and winged seeds that are attached to the adaxial surface of each cone scale and, thus, represents an early member of the Pinaceae. ? Conclusions: Eathiestrobus mackenziei gen. et sp. nov. extends the fossil record for well-documented members of the family Pinaceae from the Lower Cretaceous to the Kimmeridgian Stage of the Upper Jurassic. This species also clarifies the set of characters that are diagnostic for seed cones of Pinaceae and reveals possible plesiomorphic characters for seed cones of the family.     

Liaoxia Cao et S.Q. Wu (Gnetales): ephedroids from the Early Cretaceous Yixian Formation in Liaoning, northeastern China

Gnetalean compression-impression fossils are described from the Early Cretaceous Yixian Formation, Liaoning Province, north-eastern China, and assigned to six species of Liaoxia Cao et S.Q. Wu. The fossils have opposite-decussate phyllotaxis and cones comprising 2–12 pairs of bracts. Ovulate cones have seeds typically in a distal position. The species differ from each other and from previously described fossils in the absence or presence of leaves, shape of cones and seeds, and shape and position of cone bracts. The species of Liaoxia are probably close relatives of extant species of Ephedra L., but diagnostic reproductive details that could confirm this hypothesis are not preserved. The restricted information in the fossils and the poorly understood morphological diversity of extant Ephedra, prevent assignment of the fossils to any particular subgroup of Ephedra, as well as an explicit exclusion of them from the extant genus.     

Morphogenesis of the reproductive shoots of Welwitschia mirabilis and Ephedra distachya (Gnetales), and its evolutionary implications

For decades, Gnetales appeared to be closely related to angiosperms, the two groups together forming the anthophyte clade. At present, molecular studies negate such a relationship and give strong support for a systematic position of Gnetales within or near conifers. However, previous interpretations of the male sporangiophores of Gnetales as pinnate with terminal synangia conflict with a close relationship between Gnetales and conifers. Therefore, we investigated the morphogenesis of the male reproductive structures of Welwitschia mirabilis and Ephedra distachya by SEM and light microscopy. The occurrence of reduced apices to both halves of the antherophores of W. mirabilis gives strong support for the assumption that the male ‘flowers’ of W. mirabilis represent reduced compound cones. We assume that each half of the antherophore represents a lateral male cone that has lost its subtending bract. Although both halves of the antherophores of Ephedra distachya lack apical meristems, the histological pattern of the developing antherophores supports interpreting them as reduced lateral male cones as well. Therefore, the male sporangiophores of Gnetales represent simple organs with terminal synangia. Although extant conifers do not exhibit terminal synangia, similar sporangiophores are reported for some Cordaitales, the hypothetical sister group of conifers. Moreover, several Paleozoic conifers exhibit male cones with terminal sporangia or synangia. Therefore, we propose that conifers, Cordaitales and Gnetales originated from a common ancestor that displayed simple sporangiophores with a terminal cluster of sporangia.     

A daphniid ephippium (Branchiopoda: Anomopoda) of Cretaceous age

An anomopod ephippium from the Lower Cretaceous of Australia, remarkably similar to that of Simocephalus , shows that daphniids whose ephippia at least were very similar to those of extant species, existed about 120 million years ago. The family clearly originated in pre-Cretaceous times, and the Anomopoda considerably early than this.     

The female reproductive unit of Ephedra (Gnetales): comparative morphology and evolutionary perspectives

Morphological variation in Ephedra (Gnetales) is limited and confusing from an evolutionary perspective, with parallelisms and intraspecific variation. However, recent analyses of molecular data provide a phylogenetic framework for investigations of morphological traits, albeit with few informative characters in the investigated gene regions. We document morphological, anatomical and histological variation patterns in the female reproductive unit and test the hypothesis that some Early Cretaceous fossils, which share synapomorphies with Ephedra, are members of the extant clade. Results indicate that some morphological features are evolutionarily informative although intraspecific variation is evident. Histology and anatomy of cone bracts and seed envelopes show clade‐specific variation patterns. There is little evidence for an inclusion of the Cretaceous fossils in the extant clade. Rather, a hypothesized general pattern of reduction of the vasculature in the ephedran seed envelope, probably from four vascular bundles in the fossils, to ancestrally three in the living clade, and later to two, is consistent with phylogenetic and temporal analyses, which indicate that extant diversity evolved after the Cretaceous–Tertiary boundary. Notwithstanding striking similarities between living and Cretaceous Ephedra, available data indicate that the Mesozoic diversity went almost entirely extinct in the late Cretaceous causing a bottleneck effect in Ephedra, still reflected today by an extraordinarily low level of genetic and structural diversity. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 387–430.     

A new species of Thuja (Cupressaceae) from the Late Cretaceous of Alaska: implications of being evergreen in a polar environment

A branch bearing a number of seed cones of Thuja L. (Cupressaceae) has been recovered from a Late Cretaceous (Turonian) deposit from the North Slope of Alaska. This reproductive material is the oldest known for the genus and is indistinguishable from the seed cones of most of the extant species of Thuja, indicating that the seed cones of this Alaskan fossil Thuja had attained a modern morphological appearance early in the evolutionary history of the genus. From a physiological standpoint, the ability of modern species of Thuja to tolerate cold to freezing conditions and the ability of fossil representatives of the genus to survive periods of extended darkness during the polar winters supports the contention that the polar winters during the Late Mesozoic and early Cenozoic were cold.     

Ecological stasis in Spinicaudata (Crustacea,Branchiopoda)? Early Cretaceous clam shrimp of the Yixian Formation of north‐east China occupied a broader realized ecological niche than extant members of the group

The palaeoecology of Spinicaudata, the dominant group of benthic invertebrates in many pre‐Cenozoic freshwater environments, remains poorly understood. In analogy with extant taxa, it has been oversimplified and often reduced to shallow, temporary environments characterized by few trophic levels, implying ecological stasis from the Devonian to the Recent. We excavated 43 horizons of the Lower Cretaceous Yixian Formation (Anjiagou and Hengdaozi beds) to evaluate whether spinicaudatan ecology can be simplified to such an extent. Sedimentological evidence suggests general perennial conditions during the excavated lake interval. Based on 33 226 specimen counts, we identified three arthropod‐dominated macrobenthic associations and two assemblages. Response curves indicate that the spinicaudatan Eosestheria middendorfii was exceptionally tolerant to environmental gradients, followed, in decreasing order, by mayfly larvae, water boatmen and oligochaetes. Many spinicaudatan‐yielding layers represent death after reproductive senescence rather than mass‐mortality events. Spinicaudatan size varies significantly according to faunal association. A forward stepwise regression model suggests that growth responded to population density and diversity: lower densities and higher diversities triggered fast growth, and vice versa. The proposed strong density effect on carapace size has been corroborated by rearing experiments: as for E. middendorfii, natural log regression provided the best fit for the two extant species Eulimnadia texana and Eocyzicus argillaquus. Hence, E. middendorfii was a tolerant and morphologically variable species adapted to perennial waters and the frequent presence of higher trophic levels. Although there are rare records of extant taxa co‐occurring with fish, the described Early Cretaceous environment is commonly not inhabited by extant Spinicaudata.     

Ancient ontogenies: larval development of the Lower Cretaceous anuran Shomronella jordanica (Amphibia: Pipoidea)

The Lower Cretaceous anuran Shomronella jordanica (Pipoidea) is represented by an assemblage of nearly 300 tadpoles of different ages. The size of the assemblage allows a reconstruction of the larval ontogeny of this species. We describe the ossification sequence and growth rates of S. jordanica and present reconstructions of tadpoles at different stages. The ontogeny of the species seems to be similar to that of extant pipids in many aspects. Larvae are similar in shape and size to those of Xenopus laevis but lack the anterior barbels that are typical for all extant pipids. The ossification sequence is closer to that of pipids than to that of other anurans. We present evidence that suggests that this species was terrestrial as an adult, and we discuss the implications of these data for the evolution of anuran development.     

Early Cretaceous mesofossils from Portugal and eastern North America related to the Bennettitales-Erdtmanithecales-Gnetales group

Four new genera and six new species of fossil seed (Buarcospermum tetragonium, Lignierispermum maroneae, Lobospermum glabrum, L. rugosum, L. stampanonii, Rugonella trigonospermum) are described from five Early Cretaceous mesofossil floras from Portugal and eastern North America. The four genera are distinguished by differences in size, shape, and details of seed anatomy, but all are unusual in having an outer seed envelope with a distinctive anatomical structure that surrounds the nucellus and the integument. The integument is extended apically into a long, narrow micropylar tube. The four new genera are part of a diverse, but previously unrecognized, complex of extinct plants that was widespread in Early Cretaceous vegetation and that coexisted in similar habitats with early angiosperms. The distinctive structure of these seeds, and the strong similarities to other fossil seeds (Ephedra, Ephedripites, Erdtmanispermum, Raunsgaardispermum, and some Bennettitales) already known from the Early Cretaceous, suggests that this newly recognized complex of extinct plants, together with Bennettitales, Erdtmanithecales, and Gnetales (the BEG group), is phylogenetically closely related.     

A Cretaceous dance fly (Diptera: Empididae) from Botswana

Abstract The first fossil dance fly, Empis (s.s.) orapaensis sp.nov., of the subfamily Empidinae (Diptera: Empididae) is described from Africa. It was extracted from middle Cretaceous sediments of Orapa, Botswana.
Empis orapaensis manifests a more plesiomorphous wing venational pattern then the extant species of Empis , and a list of the apomorphic and plesiomorphic characteristics is given. The specimen modifies earlier ideas on the phylogeny of the Empidoidea.
The fossil also provides us with more conclusive evidence on the environment of Cretaceous southern Africa. E.orapanenis probably had a similar life-style to extant members of the group and therefore visited flowers.