INVESTIGATIONS OF NORTH AMERICAN CYCADEOIDS: MONANTHESIA

Delevoryas , Theodore . (Yale U., New Haven, Conn.) Investigations of North American cycadeoids: Monanthesia. Amer. Jour. Bot. 46(9): 657–666. Illus. 1959.—A study of the cycadeoid trunks from the Upper Cretaceous Mesaverde formation of northwestern New Mexico, which are in the collections of the Peabody Museum of Natural History, Yale University, has been completed. These plants had columnar trunks with persistent leaf bases and a cone in the axil of every leaf. The vascular supply of the cone is derived from the fusion of two cortical bundles which arise from two leaf traces, neither of which supplies the subtending leaf. Enough difference appears to exist between these forms and Cycadeoidea to warrant a separate generic designation. Monanthesia is the generic name used, and the single species, M. magnifica, is considered to include all the stems from the New Mexico localities. The pattern of vascularization of cones seems to suggest that they are foliar structures and that Monanthesia is probably a more advanced form than Cycadeoidea.     

Diversity and significance of late cretaceous permineralized plant remains from Hokkaido,Japan

Middle to Late Cretaceous permineralized plants hitherto described from Hokkaido, Japan are summarized. The fossil flora comprises fungi, ferns, gymnosperms and angiosperms. Many modern fern families have been recognized including Anemiaceae, Cyatheaceae, Dennstaedtiaceae, Gleicheniaceae Loxsomaceae, Lygodiaceae and Matoniaceae. Gymnosperms are most abundant in the flora. Some recently-found materials are tentatively introduced with brief comments emphasizing their morphological and taxonomical significance. A bisporangiate flower ofCycadeoidella japonica Ogura shows fine internal anatomy and provides evidence that the cycadeoidalean ovule was a cupulate, unitegmic structure. Vascular tracheids in the synangial wall support the evolution of cycadeoidalean synangia from Paleozoic seed-fern synangia. A new gymnosperm female fructification has a thick envelope comparable to an angiosperm carpel around a large seed. The angiosperms contain various morphologies that require further extensive study.     

INVESTIGATIONS OF NORTH AMERICAN CYCADEOIDS: TRUNKS FROM WYOMING

Delevoryas , Theodore . (University of Illinois, Urbana.) Investigations of North American cycadeoids: trunks from Wyoming . Amer. Jour. Bot. 47(9): 778–786. Illus. 1960.—A study was undertaken of the cycadeoid trunks from Wyoming to more fully understand the morphology and evolutionary significance of these trunks and to determine the validity of the genus Cycadella. The stem named Cycadeoidea (or Cycadella) utopiensis is not the same morphologically as those from Carbon County, Wyoming, and because of its unknown source, it is not a reliable specimen for the study of cycadeoids from the Morrison formation. The Morrison, Carbon County specimens included in Ward's genus Cycadella were shown to have trunk structure, leaf bases, and cones like those in Cycadeoidea. The cone vascular bundle is derived from a fusion of 4 leaf traces. From the fused bundle complex, 4 leaf traces are separated farther out in the cortex, and 1 of these supplies the subtending leaf. In spite of the relatively complex pattern of cone trace formation, there does not seem to be any significant means of distinguishing specimens called Cycadella from the more common Cycadeoidea. Furthermore, it is evident that the genus Cycadeoidea represents a fairly homogeneous group with a wide geographic and geologic range. Pattern of formation of cone traces is interpreted as possible further evidence for regarding cycadeoid cones as portions of foliar systems.     

INVESTIGATIONS OF NORTH AMERICAN CYCADEOIDS: EARLY OVULE ONTOGENY

Ovules on cones of the genus Cycadeoidea from the Cretaceous were found preserved at the pregametophytic stage of nucellar development. At this stage, the ovules are distinct from those of extant gymnosperms in that there is a greater degree of differentiation in both the integuments and the nucellus in Cycadeoidea. Linear clusters of three or four cells, which consistently occur in the middle of the nucellus, are interpreted to be the remains of linear tetrads of megaspores. This example of exceptional preservation allows, for the first time, a comparison of early ontogenetic events in a fossil seed with those of extant seeds.     

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.     

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.     

Some aspects of cycadeoid evolution

First definite remains of the order Cycadeoidales appear at the beginning of the Mesozoic, and extinction occurred at the end of the Cretaceous. Reasoning on the assumption that the characters most frequently present in a group of related plants are primitive ones, an attempt is made to determine what the most primitive cycadeoids looked like and what the probable ancestors were. Most likely the most primitive cycadeoids had slender, elongated, branched stems, with numerous leaves. Pollen was borne in compound sporangia on leaf-like structures. The most likely ancestral form, on the basis of known Paleozoic fossil plants, are the pterido-sperms. Origin of cycadeoid cones is postulated as having occurred in one of at least three ways. One suggestion is that cones are portions of leaf systems, and that 'telescoping' of the stems' resulted in the displacement of the cone in Cycadeoidea to the axil of another leaf. Another suggestion is that cones are fertile axillary branches. A third idea is that eones represent branches arising from leaves.     

Pollen morphology and ultrastructure of the Bennettitales: In situ pollen of Cycadeoidea

The micromorphology and ultrastructure of in situ pollen from Cycadeoidea dacotensis are described from permineralized specimens collected from the Lower Cretaceous of North America. Pollen grains are ovoid and relatively small, averaging 25 μm in length and 12 μm in width. Grains are monosulcate with the exine typically invaginated in apertural regions. Exine ornamentation ranges from punctate to psilate. The exine averages 0.73 μm in thickness and is composed of a light-staining sexine and a dark-staining nexine. The sexine consists of a thin, homogeneous tectum, typically with a well-defined inner boundary, and a thicker granular infratectum. The infratectal granules are relatively uniform in size, however, variation occurs in the arrangement of granules. In some grains, the sexine appears homogeneous because there is little lacunal space between the individual granules. The granular infratectum is in direct contact with the underlying nexine. The nexine is uniform in thickness in both apertural and nonapertural regions, and it lacks lamellae throughout. Pollen morphology and ultrastructure are compared with those of the bennettitalean genus Leguminanthus and the dispersed genus Monosulcites. In addition, the fine structure of Cycadeoidea pollen is compared to that of the gymnosperm groups with which the Bennettitales are regarded to be most closely related, including Gnetales, Pentoxylales, and Eucommiidites-type pollen-producing plants.     

The incertae sedis <Emphasis Type="Italic">Carpathoporella</Emphasis> Dragastan, 1995, from the Lower Cretaceous of Albania: skeletal elements (sclerites,internodes/branches,holdfasts) of colonial octocorals

The incertae sedis Carpathoporella Dragastan, 1995, reported from the Lower Cretaceous of the Western Tethyan domain, is usually interpreted as remains of calcareous algae (Dasycladales or Characeae). New thin-section material from the Aptian of Albania sheds light not only on its biogenic nature but also on the morphological variability of this taxon. In fact, Carpathoporella represents the debris of colonial, bushy, most likely gorgonid octocorals with tuberculated spheroids that may be fused at least near the basal root-like holdfast. Colony branching originates from longitudinally grooved calcareous branches or internodes. Possible relationships to other Upper Cretaceous to Palaeogene genera are discussed and a revised critical inventory of Cretaceous octocorals is presented. Due to the evidenced morphological features, Carpathoporella could either represent an ancestral isidid octocoral of the order Alcyonacea such as Moltkia Steenstrup or, due to the likely primary aragonitic skeletal mineralogy, a representative of Epiphaxum Lonsdale of the order Helioporacea. Due to morphological analogies, the new combination Carpathoporella elliotti (Radoičić) is proposed. In any case, the Lower Cretaceous record from Tethyan peri-reefal shallow-water carbonates is highlighted since numerous skeletal findings of fossil gorgonid Octocorallia were so far only known from Upper Cretaceous and younger strata of outer shelf environments of the boreal realm. The origin of deep-water Upper Cretaceous octocorals from Lower Cretaceous shallow-water taxa such as Carpathoporella is proposed as a possible further example of onshore/offshore evolutionary pattern.     

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.     

Molecular control of early cone development inPinus radiata

Summary A family of genes expressed during early stages of shoot development were isolated fromPinus radiata. A homologue of theLEAFY/FLORICAULA flower meristem-identity genes,NEEDLY (NLY), and three MADS-box genes,PrMADS1, PrMADS2 andPrMADS3 (Pinus radiata MADS-box genes), were expressed at early stages of initiation and differentiation of reproductive (male and female) cone buds, as well as vegetative buds. Expression ofNLY in transgenicArabidopsis thaliana promoted floral fate, demonstrating that it encodes a functional ortholog of theFLORICAUL A/LEAFY genes of angiosperms.Abbreviations DSB dwarf shoot bud - LSTB long-shoot terminal bud - PCB pollen cone bud - SCB seed cone bud - LD long day - SD short day     

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.     

PITYOSTROBUS PUBESCENS,A NEW SPECIES OF PINACEOUS CONES FROM THE LATE CRETACEOUS OF NEW JERSEY

This new species is based on a single semifusinized cone from the Late Cretaceous (Cenomanian) of New Jersey. The cone is flattened but essentially complete. It is 55 mm long and 8 by 13 mm in diam. Scale apices are thin, rounded, and entire, lacking evidence of an umbo or spine. The vascular cylinder of the cone axis is organized as a series of separate strands. The scale is made up of a basal portion that stands out nearly perpendicular to the cone axis and a sharply upturned distal portion. The bract base has a pronounced abaxial keel. Bract and scale traces diverge from the vascular cylinder of the cone axis separately from one another. A poorly developed interseminal ridge is present at the chalazal end of the seeds. An unusual feature of the cone is the presence of a trichome-bearing epidermis on the cone axis, bract-scale complex, and near the scale apex. Resin canals diverge into the bract-scale complex abaxial to the scale trace with branches becoming adaxial to the scale trace outward. A number of features of the new species occur in cones of Abies, Cedrus, Keteleeria, Larix, Picea, Pseudolarix, Pseudotsuga, and Tsuga as well as in the extinct genus Pseudoaraucaria. Features of Pinus are absent. This suggests that Pseudoaraucaria may have served as an ancestral source for modern genera other than Pinus with Pityostrobus pubescens representing an evolutionary intermediate.     

A male cone of Pseudofrenelopsis dalatzensis with in situ pollen grains from the Lower Cretaceous of Northeast China

A Classopollis-containing male cone found in close association with Pseudofrenelopsis dalatzensis from the Lower Cretaceous Dalazi Formation of Yanji Basin, eastern Jilin, China was studied using scanning electron microscopy and described as Classostrobus dalatzensis sp. nov. The small oval cone, borne on the top of a stalk (fertile short shoot), consists of helically arranged microsporophylls, each with a rhomboid and expanded distal head. The short shoot is divided into indistinct nodes and internodes with fine longitudinal striations. Cuticles of the microsporophyll and the internode of the short shoot are generally similar to Pseudofrenelopsis dalatzensis internode cuticles, all of them having papillate outer surfaces. Stomata are similar in structure and arrangement, with papillae of subsidiary cells overhanging the stomatal pit. Epidermal cells are rectangular or isodiametrical, each with a robust papilla on the periclinal wall. Hypodermal cells are present between the stomatal files. The internal structure of the in situ Classopollis pollen grains was studied in detail. Pollen grain is small, with thin sexine, poorly developed nexine and may be immature. This is the first record of in situ pollen grains Classopollis in China of which the internal structure is known in detail.     

INVESTIGATIONS OF NORTH AMERICAN CYCADEOIDS: WELTRICHIA AND WILLIAMSONIA FROM THE JURASSIC OF OAXACA,MEXICO

Specimens of cones of members of the family Williamsoniaceae, order Cycadeoidales, from Jurassic deposits of Oaxaca, Mexico, demonstrate the first bona fide evidence of the pollenbearing Weltrichia from the Mexican Jurassic as well as provide new information on ovulebearing Williamsonia.     

INVESTIGATIONS OF NORTH AMERICAN CYCADEOIDS: POLLINATION MECHANISMS IN CYCADEOIDEA

Wind was at first thought of as the chief pollinating mechanism in Cycadeoidea because of the open angiosperm-flower-like reconstruction of the cone presented by G. R. Wieland in 1906. Selfing seemed a more likely possibility after Delevoryas reinterpreted the cone as a closed structure in 1963. However, the existence of what was considered to be a dioecious group of species made it impossible to consider selfing as ubiquitous throughout the cycadeoids. Furthermore, only ovule-bearing trunks of the species considered to be dioecious were known; no corresponding trunks bearing exclusively microsporangiate cones had ever been found. Recent investigations of a trunk of the type heretofore considered as representative of the dioecious species have revealed that these cones were actually bisporangiate. The discovery that the bisporangiate condition was normal in the cycadeoids eliminates much of the confusion in understanding the system of pollination and allows selfing to be considered the chief means of pollination. Outcrossing probably occurred in very low frequency and could have been the result of a chance wind pollination or of a set of interactions between the cycadeoids and an unidentified animal. Determining the nature of the system of pollination in the cycadeoids makes it possible to speculate on the population structure and evolutionary potential of this important Mesozoic group of plants.     

Multimodal sensory organs in larvae of some insect species

The morphological diversity of antennal sensory organs in larvae of Galeruca sp. (Coleoptera, Chrysomelidae), Limnephilus centralis (Trichoptera, Limnephilidae), and Eristalis tenax (Diptera, Syrphidae) was studied. A sensory cone on the antenna was found in all the species studied. The cuticular part of the antennal cone is pierced with pores. No significant differences in the shape of the antennal cone were revealed. The innervation patterns of the antennal cone varied from single cells in the leaf beetle larvae to groups of cells with a common scolopoid sheath in the fly larvae. The cone combines several types of sensilla that perform mechanoreceptor, olfactory, and gustatory functions. The morphology and ultrastructural organization of the antennal cone in relation to the heterogeneity of the larval habitats are described.     

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.     

Structure and affinities of the petrified plants from the cretaceous of northern Japan and Saghalien XI A cupressoid seed cone from the Upper Cretaceous of Hokkaido

Archicupressus is a new genus of the Cupressaceae based on a permineralized conifer female cone from the Upper Cretaceous of Hokkaido. The type species,Archicupressus nihongii sp. nov., is characterized by peltate bract-scale complexes consisting of a completely fused bract and scale bearing erect seeds. Bract-scale complexes are arranged in whorls of three on the cone axis. Each complex has a bristle-like projection, the bract apex, at the top. Affinities to some genera in the Cupressaceae are discussed. Consecutive number from the previous paper (Ohsawaet al., 1992). Supported by a Grant-in-Aid for Scientific Research from the Ministry of Education, Science and Culture of Japan to Makoto Nishida, No. 02640533.