Morphology and the evolution of cycadeoidales

OneCycadeoidea stem one cycadeoidalean gynoecium and a bisporangiate cone attached to a slender cycadeoidalean trunkCycadeoidella japonica Ogura from the Cretaceous of Japan shows well-preserved internal structure that provides evidence for a better understanding of the morphological architecture of the cycadeoidalean plant. Structural details of the cone were confirmed. The ovule has an intergument enclosing a free nucellus and a thin outer envelope. Both reproductive and vegetative structures support the medullosan affinity of Cycadeoidales. The cone is interpreted as a compressed fertile shoot. Axillary cones characterizing some Cretaceous genera such asCycadeoidea andMonanthesia consist of a lateral shoot subtended by a frond that is the first leaf of the cone shoot itself. The origin of axillary buds in the Cycadeoidales is discussed. Heterochrony may have mediated the morphological changes that resulted in the establishment of the Cycadeoidales.     

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.     

ADVENTIVE EMBRYOGENESIS IN CITRUS AND ITS RELATION TO POLLINATION AND FERTILIZATION

Cytological and histological studies of seeds from three facultative apomictic Citrus cultivars show that adventive embryos develop, as a rule, from the first few cell layers of the nucellus adjacent to the embryo sac in the micropylar half and occasionally from the chalazal end. The adventive embryos initiated in nucellar tissue away from the embryo sac and most of those initiated from the chalazal end of the nucellus do not develop beyond the one-celled stage. When two or more embryos are developing in the same seed, the successful development of a given embryo depends on its location in relation to access to nutrients from the endosperm. The presence of a zygote and triploid endosperm in seeds with adventive embryos, the abortion of seed when endosperm degenerates, and the lack of seed set without pollination indicate that pollination and fertilization are essential for in vivo adventive embryogenesis.     

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.     

SEEDS AND EMBRYOS OF ARAUCARIA MIRABILIS

Seeds and embryos contained within the silicified ovuliferous cone Araucaria mirabilis from the Cerro Cuadrado petrified forest are described. One wingless seed, 0.8–1.3 cm in length and 0.2–0.6 cm in width, is embedded in each ovuliferous scale. Seeds show a three-layered integument and a prominent wavy nucellus which is attached basally to the endotesta. The megagametophyte contains cellular inclusions that may represent starch grains. Ovule vascularization is complex and appears most similar to that of Araucaria bidwillii. Embryos 2 mm long and 0.25 mm wide appear to be in a telo-stage period of development. Shoot apex, cotyledons, root meristem, and calyptroperiblem are present in the embryos in which vascular tissues and secretory elements were beginning to differentiate at the time of fossilization. Embryo ontogeny is considered in light of stages encountered in extant gynmosperm taxa. The absence of the microgametophyte phase in the Cerro Cuadrado collection is discussed.     

The origin of the ovule

Of several theories for the origin of the ovule advanced in this century and based largely on fossil evidence, the telomic concept is widely considered the most plausible. Its principal tenet is the evolution of the integument through fusion of sterile branches or telomes around a terminal megasporangium. The only point of agreement in these theories is that the entire nucellus is a megasporangium that retains a single megaspore and the endosporic female gametophyte. Their differences center on the origin of the integument. A new concept offered here on evidence from ovules of both Paleozoic and extant seed plants significantly alters the telomic theory. It proposes that the nucellus is a sporangiophore of stem origin that bears a terminal megasporangium; that at least some of the fused integumentary telomes were fertile; and that among all features cited to characterize ovules, the unique nature of the retained megaspore alone defines the seed habit. Changes in the seed plant megaspore that extended the period of nutrient absorption over the whole course of female gametophyte development, along with complex physiological changes in the nucellus, were probably achieved along a single phylogenetic line beginning in a Late Devonian population of progymnosperms. For such a combination of events to have occurred more than once is highly unlikely, and, therefore, a monophyletic origin for seed plants is proposed. Several primitive features in ovule structure, some not evidenced since the Lower Carboniferous Period, occur in a mutant form of Arabidopsis thaliana isolated from genetically transformed plants. Their recurrence provides additional support for the proposed concept of ovule origin and also suggests that the genetic mechanisms for expression of primitive features in advanced taxa could be initiated in each case by mutation of a single homeotic gene.     

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.     

濒危植物马蹄参的大孢子发生和雌配子体发育

为探讨马蹄参(Diplopanax stachyanthus Hand.-Mazz.)濒危机制与雌性生殖发育的关系,采用石蜡切片法观察马蹄参大孢子形成和雌配子体的发育过程。结果表明,马蹄参雌蕊单心皮,子房下位,1室,1枚胚珠。胚珠横生于短片状胎座上,具单珠被,厚珠心。单孢原细胞自珠心1~2层表皮细胞处分化;大孢子四分体为直线形。成熟胚囊中,2个极核在受精前融合为次生核,3个反足细胞不发达,较早退化;二核胚囊时期出现二核分裂不均且较小核退化消失的异常发育现象。因此,马蹄参雌配子体发育过程中出现异常现象是造成其结实率低的主要原因。     

ONTOGENY OF THE PALEOZOIC OVULE,CALLOSPERMARION PUSILLUM

The ontogeny of the upper Pennsylvanian age gymnospermous ovule, Callospermarion pusillum, is described from petrifaction specimens collected at the Berryville locality in Illinois. Ovules exhibit a wide range of dimensional and structural features that indicate an extensive developmental sequence. Specimens range from ovules with indistinct zonation of the thin-walled integument to those with thick-walled cells of the sclerotesta. The apex of the fleshy nucellus in some specimens is preserved as a cellular mound, while in others a well-formed cellular pollen chamber is present; still other ovules are characterized by a papery-thin nucellus and pollen chamber wall. The megagametophyte of most specimens is represented by a hollow megaspore membrane that may be restricted to the base of the nucellus, or occupy the entire seed cavity. In a few specimens cellular gametophytes are preserved, and in one ovule archegonia with supposed eggs are also present. Variability in each of the features is compared with ontogenetic changes in comparable structures of living gymnospermous ovules and is correlated with ovule size. A developmental sequence for the fossil ovules is proposed.     

Holocene thecideide brachiopods from the north‐western Pacific Ocean: Systematics,life habits and ontogeny

Abstract

A new species of the lacazelline brachiopod genus Ospreyella is described from shallow cryptic habitats in Palau, western Caroline Islands and Arnd Atoll, Pohnpei, eastern Caroline Islands, north‐western Pacific Ocean. This new taxon, here named Ospreyella palauensis, and representing the third discovery of a species of Ospreyella from the Indo‐Pacific region, is compared with the other two recently described extant species of the genus, O. depressa Lüter from the Great Barrier Reef, Australia, and O. maldiviana Logan from the Maldive Islands, northern Indian Ocean. An ontogenetic sequence is described for the new species and compared with those previously described for other Holocene lacazellines. Additionally, a study of ontogenetic stages in Thecidellina congregata from Saipan in the Northern Mariana Islands shows early‐stage similarities to those of the new species and other lacazellines. These observations on development in extant forms provide important analogies for comparison with ontogenetic stages in ancestral fossil forms and information which may be useful in reconstructing thecideide phylogenetic history.     

Expression of ovule and integument‐associated genes in reduced ovules of Santalales

SUMMARY Santalales comprise mainly parasitic plants including mistletoes and sandalwoods. Bitegmic ovules similar to those found in most other angiosperms are seen in many members of the order, but other members exhibit evolutionary reductions to the unitegmic and ategmic conditions. In some mistletoes, extreme reduction has resulted in the absence of emergent ovules such that embryo sacs appear to remain embedded in placental tissues. Three santalalean representatives (Comandra, Santalum, and Phoradendron), displaying unitegmic, and ategmic ovules, were studied. Observed ovule morphologies were consistent with published reports, including Phoradendron serotinum, which we interpret as having reduced ategmic ovules, consistent with earlier reports on this species. For further understanding of the nature of the ovule reductions we isolated orthologs of the Arabidopsis genes AINTEGUMENTA (ANT) and BELL1 (BEL1), which are associated with ovule development in this species. We observed ovular expression of ANT and BEL1 in patterns largely resembling those seen in the integumented ovules of Arabidopsis. These genes were found to be expressed in the integument of unitegmic ovules and in the surface layers of ategmic ovules, and in some cases, expression of BEL1 was also observed in the surrounding carpel tissue. We hypothesize that ategmic ovules derive from a fusion of the integuments with the nucellus or that the nucellus has taken on some of the characteristics confined to integuments in ancestral species.     

ADVENTIVE EMBRYOGENESIS IN CITRUS I. THE OCCURRENCE OF ADVENTIVE EMBRYOS WITHOUT POLLINATION OR FERTILIZATION

Anatomical studies of unfertilized undeveloped seeds from open- and control-pollinated fruits of ten facultative apomictic Citrus cultivars were carried out with the aid of light and epifluorescence microscopes. With or without pollination, adventive embryos autonomously developed at all positions in the nucellus in all cultivars. The adventive embryos initiated at the chalazal end of the nucellus were more vigorous than those initiated at the micropylar end. Because of the lack of endosperm and poor seed development, however, all adventive embryos within the unfertilized seeds terminated their development at the globular or early cotyledonary stages and were unable to germinate under natural conditions. The capability of unfertilized seeds to develop varied from species to species. Growth of the adventive embryos was dependent on nucellus size, but the growth rate of adventive embryos relative to nucellus size was different in different species. Neither pollination, fertilization nor subsequent zygote and endosperm development further stimulated adventive embryo initiation. Conversely, pollination and subsequent fertilization of other seeds in the same fruit slightly, but significantly, suppressed adventive embryo growth in the unfertilized seeds. These facts concerning adventive embryogenesis in unfertilized seeds indicate that neither pollination nor fertilization is essential for in vivo adventive embryogenesis and that normal endosperm is necessary for perfect development of adventive embryos initiated only in the micropylar half of the nucellus.     

Folivory or fruit/seed predation for Mesopithecus, an earliest colobine from the late Miocene of Eurasia?

Here we compare dental microwear textures from specimens of the fossil genus Mesopithecus (Cercopithecidae, Colobinae) from the late Miocene of Eastern Europe with dental microwear textures from four extant primate species with known dietary differences. Results indicate that the dental microwear textures of Mesopithecus differ from those of extant leaf eaters Alouatta palliata and Trachypithecus cristatus and instead resemble more closely those of the occasional hard-object feeders Cebus apella and Lophocebus albigena. Microwear texture data presented here in combination with results from previous analyses suggest that Mesopithecus was a widespread, opportunistic feeder that often consumed hard seeds. These data are consistent with the hypothesis that early colobines may have preferred hard seeds to leaves.     

罗汉果大孢子发生及雌配子体发育与花部形态、胚珠变化的关系

为弄清罗汉果(Siraitia grosvenorii)大孢子发生、雌配子体发育过程与花部形态特征、胚珠的关系,运用石蜡切片法对罗汉果子房进行了显微观察。结果表明,罗汉果的胚珠倒生,双珠被,厚珠心,大孢子四分体呈线型排列,合点端一个大孢子分化为功能大孢子,成熟胚囊为蓼型。花蕾形态、胚珠变化与大孢子发生、雌配子体的发育时期具有一定相关性,当子房长度为7.0 mm≤L<9.0 mm,珠心呈椭圆形时,约有45.83%的大孢子母细胞处于减数分裂时期。因此,依据罗汉果花部形态可有效确定大孢子发生与雌配子体发育的时期。     

ADVENTIVE EMBRYOGENESIS IN CITRUS (RUTACEAE) II. POSTFERTILIZATION DEVELOPMENT

Cytological and histological studies on postfertilization development of ovules were carried out in six facultatively apomictic Citrus cultivars. At the time of anthesis, adventive embryo initial cells (AEICs) were detected mainly in the cell layers of the nucellus around the chalazal half of the embryo sac. During the approximately 40 days rest period of the AEICs after fertilization, rapid cell division and enlargement in the endosperm and the chalazal half of the nucellus resulted in the split of AEICs into several separated areas forming the micropylar, lateral and chalazal islands surrounding the enlarging embryo sac. Both in diploid seeds with triploid endosperm and triploid seeds with pentaploid endosperm, the AEICs located in the micropylar half successfully developed into adventive embryos. In diploid seeds, almost all AEICs located in the chalazal half did not develop beyond the initial-celled stage, while in the triploid seeds, those located in the chalazal half occasionally developed into cotyledonary embryos. In seeds with aborted endosperm, the AEICs located in the chalazal half often developed into cotyledonary embryos. The chalazal expiants from normal seeds produced a large number of embryos in vitro. Four results can be summarized from these studies on adventive embryogenesis as follows: 1) All AEICs are initiated prior to anthesis. 2) Whether or not the AEICs successfully developed into adventive embryos is dependent upon their position in the seed. 3) The farther the AEICs are located from the micropylar end, the more adventive embryogenesis is suppressed by endosperm. 4) The degree of adventive embryogenesis in the chalazal half is affected by time and extent of malfunction of the endosperm. Under natural conditions, these regulatory systems of adventive embryogenesis contribute to high production of zygotic seedlings in apomictic Citrus species and cultivars.     

A new species of Leptocycas (Zamiaceae) from the Upper Triassic sediments of Liaoning Province,China

A new cycad,Leptocycas yangcaogouensis sp.nov.,was found in sediments from the Late Triassic in western Liaoning,China.The pinnately compound leaves(Pseudoctenis type)are screwed in a crown on the stem top.The leaflets are linear,with parallel veins and decurrent bases on the rachis.The leaf bases are persistent.The cataphylls intermix with the leaves.The female cone is ovoid in shape.The characteristics of the new plant are more similar to those of Leptocycas gracilis,a Triassic cycad from North America,but the new species differs from L.gracilis in the size of its stem(7–8 vs.3–5 cm in diameter,respectively),leaves(length×width 100×16 vs.30×7 cm,respectively)and leaf density along the stem(4–6 vs.1–2 bases/1 cm length,respectively).Both L.gracilis and L.yangcaogouensis,having leaves of the Pseudoctenis type,show a closer relationship to the extant Dioon of Zamiaceae.The present study provides evidence for the origin of the genus Dioon,which may have come from Leptocycas plants of the Triassic.It would be assumed that the extent cycads in Zamiaceae originate from the pteridosperms in the Late Paleozoic and have evolved through the stage of L.gracilis and L.yangcaogouensis in Late Triassic,and reaching the extant Dioon.     

INTERCONNECTED REPRODUCTIVE AND VEGETATIVE REMAINS OF POPULUS (SALICACEAE) FROM THE MIDDLE EOCENE GREEN RIVER FORMATION,NORTHEASTERN UTAH

A new specimen from the Middle Eocene Evacuation Creek Member of the Green River Formation in northeastern Utah shows a twig with several leaves of Populus wilmattae Cockrell and a fruiting raceme attached. This specimen establishes for the first time the type of fruits borne by P. wilmattae and provides additional characters with which to assess its taxonomic and evolutionary status. An associated seed shows attached placental hairs like those of extant species of Populus. The Green River fossil differs from extant Populus species in having basically palmate leaf venation and in bearing its fruiting axis on a young twig. In other aspects, the fossil species is remarkably similar to the extant species Populus mexicana.     

On the structure and evolutionary relationships of the Cerro Cuadrado fossil conifer seedlings*

Seedlings from the Cerro Cuadrado (Jurassic) Petrified Forest in Argentina have been shown to be comparable to extant araucarian seedlings from the sections Eutacta, Bunya and Colombea. Anatomically they are compared to the cone axes and embryos in the fossil cones of Araucaria mirabilis and Pararaucaria patagonica. Two seedlings previously described by Wieland are shown on pith structure to belong to P. patagonica. The remaining turbinate to top-shaped structures are compared with first year extant Bunya and Columbia seedlings and the cone A. mirabilis. Corm-like structures may represent an older seedling stage. From earlier work on the embryonic anatomy and germination stages described here, P. patagonica is deduced to have been epigeal and A. mirabilis hypogeal.     

A new stage in the evolution of the mole rats (Rodentia,Spalacinae) from the Early Miocene of northern Greece

The cheek teeth of Debruijnia kostakii n. sp. from the Early Miocene of Karydia are only slightly smaller than those of the largest extant species of the Spalacinae (Spalax giganteus), but show characteristics that are reminiscent of its Paleogene cricetid ancestor. The peculiar combination of large size and primitive dental morphology of this species bridges the gap between Debruijnia and Pliospalax and thereby supports the hypothesis that these genera form the core of the complex evolutionary history leading to extant Spalax.     

Inflorescence bracts of fossil and extant Tilia in North America,Europe, and Asia: patterns of morphologic divergence and biogeographic history

The enlarged inflorescence bract diagnostic of extant Tilia has an extensive Tertiary fossil record in the Northern Hemisphere. Diversity of bract morphology, and the extent of adnation between peduncle and bract, is reviewed for fossil and extant species of Tilia. An extinct type of bract with an orbicular outline and palmate venation is documented by the fossil species Tilia circularis (Chaney) comb. nov. from the early Oligocene of Oregon and is designated Type A. Living species of the genus have elongate bracts with predominately pinnate venation that are borne in two basic configurations: Type B, with the peduncle fused only to the extreme base of the bract lamina, as in extant Tilia endochrysea Hand.-Mzt. of southern China; and Type C with the peduncle fused medially along the basal one-third of the bract lamina, as in most extant species. Bracts of Type B were widely distributed in the Tertiary of western North America (late Eocene to Miocene) and Europe (early Miocene to Pliocene), while those of Type C are known in the fossil condition only from the middle and late Tertiary of Asia and Pliocene of Europe. The bracts of T. circularis, like those of type B, are borne on relatively long stalks and have the peduncle fused only at the extreme base. The fossil record supports recognition of the following characters as apomorphic in Tilia bract evolution: bracts sessile, peduncle adnate to the upper surface of the bract, and pinnate bract venation.