Structure and affinities of the petrified plants from the cretaceous of Northern Japan and Saghalien XIIIYubaristrobus gen. nov., A new Taxodiaceous cone from the upper cretaceous of Hokkaido

Yubaristrobus is a new genus of the Taxodiaceae based on a permineralized seed cone from the Upper Cretaceous of Hokkaido. The type species,Y. nakajimae sp. nov., is characterized by peltate bract-scale complexes consisting of a completely-fused bract and scale. The bract-scale complexes are spirally arranged as in most taxodiaceous genera. Their vascular arrangement is specialized and unique in the Taxodiaceae and suggests a relationship with the Cupressaceae. Consecutive number from the previous paper (Ohsawa, M. Nishida and H. Nishida, 1992b).     

Pinus MUTOI (Pinaceae), a new species of permineralized seed cone from the upper Cretaceous of Hokkaido,Japan

Pinus mutoi is described as a new species on the basis of a permineralized seed cone from the Upper Cretaceous of Hokkaido, Japan. The cone is at least 20 cm long and up to 6 cm in diameter, consisting of a cone axis and numerous cone-scale complexes that are arranged helically around the axis. Two winged seeds are borne on the adaxial surface of each ovuliferous scale. Each complex receives a single trace from the vascular cylinder of the cone axis. In the scale base, all the resin canals occur abaxially to the vascular strand. The spatulate bract of the fossil is unique to the specimen among the cones of both living and fossil Pinus. The central umbo, broad sclerotic cortex of cone axis, and absence of serotinous features of the fossil cone suggest affinity with the subsection Sylvestres of the section Pinus, subgenus Pinus. This is the first record of permineralized preserved Pinus cone from the Cretaceous of Eastern Eurasia.     

Structure and affinities of the petrified plants from the cretaceous of Northern Japan and Saghalien. XII.Obirastrobus gen. nov., petrified pinaceous cones from the Upper Cretaceous of Hokkaido

Two new species of pinaceous cones belonging toObirastrobus gen. nov. are described from the Upper Cretaceous of Hokkaido. Bract-scale complexes are arranged helically and each consists of a small bract and a large scale. The bract and scale are separated. The scale tapers distally and its apex does not become papery. The anatomy of the bracts, scales, and seeds shows a combination of features unique in the Pinaceae and resembling those of the extant genusKeteleeria and the extinct genusPseudoaraucaria. Consecutive number from the previous paper (Ohsawaet al., 1992). Supported by Grant-in-Aid for Scientific Research from the Ministry of Education, Science and Culture Nos. 63540545 and 02640533 to Makoto Nishida.     

A NEW SCIADOPITYACEOUS SEED CONE FROM THE UPPER CRETACEOUS OF HOKKAIDO,JAPAN

A sciadopityaceous seed cone, Sciadopityostrobus kerae, gen. et sp. nov., is described on the basis of a permineralized specimen from the Upper Cretaceous (Cenomanian–Coniacian) of Hokkaido, Japan. The peel method was applied for anatomical observations. The seed cone consists of a cone axis receiving numerous cone scale complexes that are arranged helically. Each complex generally has five inverted ovules that are oriented adaxially. The cone is similar to those of living Sciadopitys verticillata with respect to its peltate cone scale complex, with free apices of both bract and ovuliferous scale, trichomes on the bract, and a trifurcated ovuliferous scale strand. In the fossil, the bract and ovuliferous scale strands fuse with each other in the basal part of the cone scale complex, while S. verticillata bract and ovuliferous scale strands are derived separately from the vascular cylinder and remain separate throughout their length. The present specimen is one of the oldest records of structurally preserved cones that can be assigned to the family Sciadopityaceae.     

Anatomy and relationships of petrified seed cones of the Cupressaceae,Taxodiaceae, and Sciadopityaceae

Morphological features and the affinities of petrified seed cones of the Taxodiaceae, Cupressaceae, and Sciadopityaceae are reevaluated. TheCunninghamia-like plants are the earliest record of the families.Parataiwania explains the divergence ofTaiwania from aCunninghamia-like ancestral form by loss of ovuliferous scale and sclerenchyma.Sequoia-like allies probably diverged from a Cretaceous transitional plant likeYezosequoia and are most diversified in the Late Cretaceous.Yubaristrobus andArchicupressus suggest monophyly ofTaxodium and its allied genera of the Taxodiaceae and the Cupressaceae. The Cupressaceae may be derived from a taxodiaceous plant with orthotropous seeds, acquiring a reniform arrangement of the vascular bundles of the bract-scale complex, decussate or whorled phyllotaxis, and spherical cones consisting of a small number of bract-scale complexes. The Cretaceous fossils of the Sciadopityaceae suggest the ancient origin of the family.     

Ovulate cone,pollination drop,and pollen capture in Sequoiadendron (Taxodiaceae)

In Sequoiadendron ovules are borne inside the ovulate cone, and pollination drops secreted from these ovules collect pollen. We examined: (1) the relation between ovular position and pollen capture; (2) pollen behavior when in contact with a pollination drop; and (3) ultrastructure of ovules during pollination drop secretion. During wet periods a water sheet forms on the surface of the cone due to bract shape and wettability. Pollination drops persist inside the wetted cone, and pollen capture resumes immediately after drying. Pollen landing on a pollination drop is taken inside the drop and carried into the micropyle when the drop contracts. Several notable ultrastructural features appear in the nucellus, integument, chalaza, and bract lamina during pollination-drop secretion. The abaxial surface of the lamina is covered by a membrane that may contribute to the wettable nature of the surface.     

北美香柏雌球果的发育

用扫描电镜(SEM)观察了北美香柏 Thuja occidentalis 雌球果的发育过程。在北京,北美香柏的雌球果是在八月初由营养芽转变而来,雌球果一般有4~6对苞片,中间2~3对可育,每一苞片腋部着生两枚胚珠,在可育苞片腋部最先观察到一扁平的隆起,并在其上分化出两个胚珠原基,接着分化出珠被和珠心,最后形成扁平而两侧对称的胚珠。在北美香柏雌球果发育过程中,约一半的雌球果在2~3对可育苞片中位于下面的1~2对的腋部产生3个胚珠原基,中间一个较小,并在以后的发育中逐渐退化。由此推测北美香柏的雌球果可能是由祖先类群中每一苞片具多于2个胚珠的雌球果演化而来。在光镜下对雌球果维管系统的观察发现,传粉前幼小雌球果的苞片内仅有一束维管束,传粉后随着苞片基部的居间生长,有4—8束维管束在苞片内形成,但是新发育的维管束木质部和韧皮部相对位置与正常叶性器官一致,这与在以往报道的柏科植物成熟雌球果的苞片中均有反向维管束的发育不同。北美香柏雌球果早期发育和维管束分析结果支持傅德志和杨亲二提出的解释裸子植物生殖器官形态演化的“苞鳞-种鳞复合体”理论。关键词北美香柏;雌球果的发育;胚珠分化;SEM     

A NEW TAXODIACEOUS SEED CONE FROM THE OLIGOCENE OF WASHINGTON

Cunninghamiostrobus goedertii is described as a new species based on permineralized seed cones from the Early Oligocene of Washington. The fossils come from the Makah Formation and were found on the northern shore of the Olympic Peninsula. The cone consists of numerous bract-scale complexes arranged helically around a central axis. Each bract-scale complex has a large bract bearing a small flap of tissue adaxially that represents the ovuliferous scale. Up to three seeds were produced on each complex. The vascular trace to the bract-scale complex diverges from the vascular cylinder of the cone axis as a concentric strand. This divides in the outer cortex of the axis to form a large collateral bract tract abaxially and a minute scale trace adaxially. The latter continues outward to supply the ovuliferous scale. The bract trace divides, forming many strands outward which occur in a row with transfusion tissue between them. Many resin canals also occur in the bract. The new cone combines features found in modern cones of Athrolaxis, Cunninghamia, and Taiwania but is most similar to Cunninghamiostrobus yubariensis from the Late Cretaceous of Japan.     

The phylogenetic positions of the conifer genera Amentotaxus,Phyllocladus, and Nageia inferred from 18s rRNA sequences

To determine the evolutionary positions of the conifer genera Amentotaxus, Phyllocladus, and Nageia, we obtained 18S rRNA sequences from 11 new taxa representing the major living orders and families of gymnosperms. With the published Chlamydomonas as an outgroup, phylogenetic analyses of our new data and available sequences indicate that (1) the Gnetales form a monophyletic group, which is an outgroup to the conifers, (2) the conifers are monophyletic, (3) Taxaceae, Cephalotaxaceae, Cupressaceae, and Taxodiaceae form a monophyletic group, (4) Amentotaxus is closer to Torreya than to Cephalotaxus, suggesting that Amentotaxus is better to be classified as a member of Taxaceae, (5) Phyllocladus, Dacrycarpus, Podocarpus, and Nageia form a monophyletic group, and (6) Pinaceae is an outgroup to the other families of conifers. Our finding that Phyllocladus is a sister group of the Podocarpaceae disagrees with the suggestion that the phylloclade of the genus is an ancient structure and that the genus is a terminal taxon within the Podocarpaceae. The genus Nageia is more closely related to Podocarpus than to Dacrycarpus and was derived from within the Podocarpaceae. In conclusion, our data indicate that in conifers, the uniovulate cone occurred independently in Taxacaeae and Cephalotaxaceae, and in Podocarpaceae after the three families separated from Pinaceae, and support the hypothesis that the uniovulate cone is derived from reduction of a multiovulate cone.Correspondence to: S.-M. Chaw     

PINUS BUCHANANII,A NEW SPECIES BASED ON A PETRIFIED CONE FROM THE OLIGOCENE OF WASHINGTON

A silicified seed cone of Oligocene age from the Olympic Peninsula of Washington represents a new species of Pinus. The cone was about 8 cm long and 3 cm at its widest diameter in the living condition. Its scale apices are thickened and each has a dorsal umbo. Internal cone construction confirms the assignment of the new species to Pinus and suggests affinity with the subsections Australes and Ponderosae of the section Pinus, subgenus Pinus. The cone is peculiar in having a stout bract trace that is slightly concave on its adaxial side and in having resin canals that diverge from the axial secretory system toward the bract but constrict markedly and terminate before entering the bract.     

REPRODUCTIVE BIOLOGY OF THE CERRO CUADRADO (JURASSIC) FOSSIL CONIFERS: PARARAUCARIA PATAGONICA

Seventy-two silicified cone specimens of Pararaucaria patagonica Wieland from the Cerro Cuadrado (Jurassic) petrified forest in Patagonia were studied by use of thin sections and reflected light. Traces from the cone axis stele to the bract and ovuliferous scale are separate at their origins. In cross section the abaxially concave ovuliferous scale trace and massive circular bract trace branch half-way out into the cone-scale complex. Two triangular-shaped bundles of fibers follow the scale trace into the laminar part of the ovuliferous scale. Both surfaces of the ovuliferous scale possess hairs which may have assisted in cone closure. Cones typically contain one flattened, winged seed per ovuliferous scale. Multi-layered seed integuments are represented by an inner fleshy layer of large cells filled with a black amorphous material, a sclerotesta consisting of wedges or fan-shaped groups of sclereids oriented at right angles to the seed surface, and a thin, often crushed fleshy layer. Seed wings consist of anastomosing, branched cells resembling glandular hairs and containing many intercellular spaces. Well-preserved polycotyledonous embryos containing up to eight cotyledons are present in several cones. Tissues of the embryo including the shoot apex, calyptroperiblem, hypocotyl, and cotyledons are described. On the basis of anatomical and morphological features, specimens of P. patagonica are compared to members of both extant and extinct families of conifers including the Araucariaceae, Taxodiaceae, Pinaceae, Cupressaceae, and Cheirolepidaceae.     

CONE AND OVULE ONTOGENY IN TAXODIUM AND GLYPTOSTROBUS (TAXODIACEAE – CONIFERALES)

Seed cones in Taxodium distichum and Glyptostrobus pensilis occupy the position of permanent shoots and are initiated in the summer preceding spring pollination. Morphological features are similar in the two genera, reflecting their close taxonomic relationship. Ovule complexes originate as two (rarely more) ovule primordia in the axil of each fertile bract but without any indication of a preceding discrete ovuliferous scale. When the nucellus, integument, and micropyle are well developed, a series of up to ll abaxial lobes forms at the base of each ovule pair. They become fused by basal growth. After pollination the common basal meristem of lobes and bract extends by intercalary growth to form the conspicuous “ovuliferous scale” of the mature cone; the lobes enlarge and exceed the ovules. Despite the topographic similarity in the cones of both genera, there are differences in vasculature such that the vascular traces to the axillary complex originate directly from the axial cylinder in Glyptostrobus but from the bract trace in Taxodium. The complex vasculature of the mature cone develops late and primarily as an expression of intercalary growth.     

Influence of host log age and refuge from natural enemies on colonization and survival of Phoracantha semipunctata

The pine cone weevil, Pissodes validirostris Gyll. (Coleoptera: Curculionidae), attacks seed cones of most Eurasian pine species, except these of Swiss stone pine (Pinus cembra L.). Behavioural responses of adult weevils to cone volatile emissions of Swiss stone pine and to those of a common host, mountain pine (Pinus uncinata Ram.), were compared in an olfactometer. Weevils were significantly attracted by the volatile blend emitted by mountain pine, but Swiss stone pine volatiles elicited an inverse response, with most weevils moving in the opposite direction to the odour source. However, the majority of second instar weevil larvae that were extracted from mountain pine cones and transferred into Swiss stone pine cones were capable of developing to the adult stage. This suggests that Swiss stone pine cones do not contain strong feeding deterrents that could prevent larval development. The possible factors involved in the absence of colonization of Swiss stone pine cones by cone weevils are discussed.     

Use of fluorescent microscopy in defining a small ligule-like ovuliferous scale of a permineralized taxodiaceous cone from the upper cretaceous of Hokkaido,Japan

Fluorescent microscopy was proved to be effective for structural identification of permineralized plant tissues in calcite nodules from the Upper Cretaceous of Hokkaido, Japan. A minute, scale-like projection on the bract of a fossil Taxodiaceous cone is identified as a true ovuliferous scale because it is bordered with a continuous epidermis that exhibits prominent fluorescence. The presence of the ovuliferous scale suggests that the fossil is aTaiwania archetype.     

NEW TAXODIACEOUS SEED CONES FROM THE UPPER CRETACEOUS OF NEW JERSEY

Nephrostrobus cliffwoodensis gen. et sp. nov., Nephrostrobus bifurcatus sp. nov., and Rhombostrobus cliffwoodensis gen. et sp. nov. are described based on anatomical studies of seed cone fragments from the Upper Cretaceous Magothy Formation of New Jersey. These species belong to the Taxodiaceae. As in Sequoia, Sequoiadendron. and Metasequoia, the vascular strands supplying the scale and bract in Nephrostrobus are about equal. These vascular strands are arranged in a reniform pattern resembling that found in Metasequoia, but the orientation differs by 180 degress. Nephrostrobus cliffwoodensis and Nephrostrobus bifurcatus differ from each other in the branching of the complex trace and associated resin canals. Rhombostrobus cliffwoodensis resembles Cunninghamia in the arrangement of vascular strands and associated resin canals in its bract-scale complexes. However, the relative amount of bract and ovuliferous scale making up the complex is more like that in Sequoia. This combination of cone features does not occur in any of the extant genera. Nephrostrobus and Rhombostrobus are not considered to be ancestral to any of the extant taxodiaceous genera, but are extinct members of an ancestral complex from which the extant genera were derived.     

UFO in the Arabidopsis inflorescence apex is required for floral-meristem identity and bract suppression

The UNUSUAL FLORAL ORGANS (UFO) gene of Arabidopsis encodes an F-box protein required for the determination of floral-organ and floral-meristem identity. Mutation of UFO leads to dramatic changes in floral-organ type which are well-characterized whereas inflorescence defects are more subtle and less understood. These defects include an increase in the number of secondary inflorescences, nodes that alternate between forming flowers and secondary inflorescences, and nodes in which a single flower is subtended by a bract. Here, we show how inflorescence defects correlate with the abnormal development of floral primordia and establish a temporal requirement for UFO in this process. At the inflorescence apex of ufo mutants, newly formed primordia are initially bract-like. Expression of the floral-meristem identity genes LFY and AP1 are confined to a relatively small adaxial region of these primordia with expression of the bract-identity marker FIL observed in cells that comprise the balance of the primordia. Proliferation of cells in the adaxial region of these early primordia is delayed by several nodes such that primordia appear “chimeric” at several nodes, having visible floral and bract components. However, by late stage 2 of floral development, growth of the bract generally ceases and is overtaken by development of the floral primordium. This abnormal pattern of floral meristem development is not rescued by expression of UFO from the AP1 promoter, indicating that UFO is required prior to AP1 activation for normal development of floral primordia. We propose that UFO and LFY are jointly required in the inflorescence meristem to both promote floral meristem development and inhibit, in a non-cell autonomous manner, growth of the bract.Shelley R. Hepworth and Jennifer E. Klenz contributed equally to this work.     

遗传改良对杉木针叶和种实性状的影响

为揭示遗传改良对主要造林用材树种叶和种实性状的影响,阐明性状的变异趋势,该研究以杉木第4轮育种的精选树(改良群体)、四省五地的表型优树与古树(未改良群体)为对象,调查了218个无性系的针叶和种实性状指标,采用方差分析和多重对比方法研究遗传改良对杉木及不同类型杉木的表型差异,并通过相关性分析探究遗传改良对杉木针叶和球果部分表型性状的影响,以及利用主成分分析和聚类分析进行了分类。结果表明:(1)未改良群体的针叶长、针叶宽和出籽率分别比改良群体小13.28%、10.81%和33.90%,其他性状表现为未改良群体大于改良群体,差异在10.90%～27.03%之间。未改良群体球果长、球果宽和出籽率的变异系数,分别比改良群体大9.14%、12.73%和15.38%。(2)球果长、球果宽、苞鳞长和苞鳞宽4个性状,在未改良群体中仅有球果长和球果宽(0.931)、苞鳞长和苞鳞宽(0.622)之间呈极显著正相关,经遗传改良后,该4个性状两两之间均呈显著或极显著正相关。(3)四川雅安(SCYA)的球果长和球果宽的性状比改良群体大48.83%和53.26%,安徽黄山(AHHS)的百粒重比改良群体大16.92%。(4)遗传改良导致松张型球果的杉木比例降低,紧包型和反翘型球果的杉木比例增加。综上认为,杉木的遗传改良导致球果大小下降,改变了不同针叶和球果类型的比例,同时会改变针叶性状和种实性状的相关性,将为杉木种质资源评价和未来多目标育种提供依据。     

侧柏雌球果及其胚珠的发育

观察了侧柏（Ｐｌａｔｙｃｌａｄｕｓ ｏｒｉｅｎｔａｌｉｓ（Ｌ．）Ｆｒａｎｃｅ）胚珠的发育过程及后期球果苞片的结构变化。在北京,雌球果原基７月分化。通常一个球果有４对苞片,中部两对可育,靠球果顶端一对各产生一枚胚珠,其下一对各两枚。胚珠的发育顺序是向顶的,下部可育苞片腑部的两枚胚珠源于同一原基。胚珠原基分化成珠心和珠被,在发育过程中,珠被逐渐包围珠心,最后形成２烧瓶状的胚珠。１１月至次年１月,球果处     

侧柏雌球果及其胚珠的发育

Seed cone in Platycladus orientalis (L.) France consists of four or five pairs of decussate bracts. Usually, two pairs of the fertile bracts in the middle of the cone subtend six ovules, which initiate in an acropetal manner. Only one ovule presents on each of the upper fertile bract, while two ovules initiate from a common primordium in the axil of lower bracts. In Beijing, most female cones initiated in July. All parts of the cone formed before dormancy, which occurred during November to the next January. After pollination in March, bract morphology changed dramatically; intercalary growth of the bract base formed a conspicuous protuberance, in which inverted vascular system developed. Furthermore, ovules on different pairs of bracts initiated in an acropetal manner and two ovules in each lower fertile bract initiated from a common primordium, which was different from the basipetal initiation of ovules and independently formed single ovule as reported by Takaso in Calltris.     

Variation in seed packaging of a fleshy‐fruited conifer provides insights into the ecology and evolution of multi‐seeded fruits

• The study of intraspecific seed packaging (i.e. seed size/number strategy) variation across different populations may allow better understanding of the ecological forces that drive seed evolution in plants. Juniperus thurifera (Cupressaceae) provides a good model to study this due to the existence of two subspecies differentiated by phenotypic traits, such as seed size and cone seediness (number of seeds inside a cone), across its range.
• The aim of this study was to analyse seed packaging (seed mass and cone seediness) variation at different scales (subspecies, populations and individuals) and the relationship between cone and seed traits in European and African J. thurifera populations.
• After opening more than 5300 cones and measuring 3600 seeds, we found that seed packaging traits followed different patterns of variation. Large‐scale effects (region and population) significantly contributed to cone seediness variance, while most of the seed mass variance occurred within individuals. Seed packaging differed between the two sides of the Mediterranean Sea, with African cones bearing fewer but larger seeds than the European ones. However, no differences in seed mass were found between populations when taking into account cone seediness. Larger cones contained more pulp and seeds and displayed a larger variation in individual seed mass.
• We validated previous reports on the intraspecific differences in J. thurifera seed packaging, although both subspecies followed the same seed size/number trade‐off. The higher seediness and variation in seed mass found in larger cones reveals that the positive relationship between seed and cone sizes may not be straightforward.We hypothesise that the large variation of seed size found within cones and individuals in J. thurifera, but also in other fleshy‐fruited species, could represent a bet‐hedging strategy for dispersal.