A strobilus (pl.: strobili) is a structure present on many land plant species consisting of sporangia-bearing structures densely aggregated along a stem. Strobili are often called cones, but some botanists restrict the use of the term cone to the woody seed strobili of conifers. Strobili are characterized by a central axis (anatomically a stem) surrounded by spirally arranged or decussate structures that may be modified leaves or modified stems.
Leaves that bear sporangia are called sporophylls , while sporangia-bearing stems are called sporangiophores .
Some members of both of the two modern classes of Lycopodiophyta (Lycopodiopsida and Isoetopsida) produce strobili. In all cases, the lateral organs of the strobilus are microphylls, bearing sporangia. In other lycophytes, ordinary foliage leaves can act as sporophylls, and there are no organized strobili.
The single extant genus of Equisetophyta, Equisetum , produces strobili in which the lateral organs are sporangiophores. Developmental evidence and comparison with fossil members of the group show that the sporangiophores are reduced stems, rather than leaves. Sporangia are terminal.
With the exception of flowering plants, seed plants produce ovules and pollen in different structures. Strobili bearing microsporangia are called microsporangiate strobili or pollen cones, and those bearing ovules are megasporangiate strobili or seed cones (or ovulate cones).
Cycadophyta are typically dioecious (seed strobili and pollen strobili are produced on separate plants). The lateral organs of seed strobili are megasporophylls (modified leaves) that bear two to several marginal ovules. Pollen strobili consist of microsporophylls, each of which may have dozens or hundreds of abaxial microsporangia.
The single living member of the Ginkgophyta, Ginkgo biloba produces pollen strobili, but the ovules are typically borne in pairs at the end of a stem, not in a strobilus. When there are more than a pair of ovules in G. biloba, however, or when fossil taxa bearing large numbers of ovules are examined, it is clear that the paired ovules in the extant species are a highly reduced strobilus.
Pollen strobili of Pinophyta are similar to those of cycads (although much smaller) and Ginkgoes in that they are composed of microsporophylls with microsporangia on the abaxial surface. Seed cones of many conifers are compound strobili. The central stem produces bracts and in the axil of each bract is a cone scale. Morphologically the cone scale is a reduced stem. Ovules are produced on the adaxial surface of the cone scales.
Gnetophyta consists of three genera, Ephedra , Gnetum and Welwitschia . All three are typically dioecious, although some Ephedra species exhibit monoecy. In contrast to the conifers, which have simple pollen strobili and compound seed strobili, gnetophytes have both compound pollen and seed strobili. The seed strobili of Ephedra and Gnetum are reduced, with Ephedra producing only two ovules per strobilus and Gnetum a single ovule.
The flower of flowering plants is sometimes referred to as a bisexual strobilus. Stamens include microsporangia within the anther, and ovules (contained in carpels) contain megasporangia. Magnolia has a particularly strobiloid flower with all parts arranged in a spiral, rather than as clear whorls.
A number of flowering plants have inflorescences that resemble strobili, such as catkins, but are actually more complex in structure than strobili.
It is likely that strobili evolved independently in most if not all these groups. This evolutionary convergence is not unusual, since the form of a strobilus is one of the most compact that can be achieved in arranging lateral organs around a cylindric axis, and the consolidation of reproductive parts in a strobilus may optimize spore dispersal and nutrient partitioning.
The word strobilus is related to the ancient Greek strobilos = whirlwind. The Hebrew word for conifer cone, itstrubal, is an ancient borrowing from the Greek.
According to Liddell & Scott, the Greek: strobilos (στρόβιλος) had many meanings, generally of anything twisted up...hence of the hedgehog,... of an egg-shell,... as a name of various twisted or spinning objects. For example:
1. a kind of seasnail... 2. a top... 3. a whirlpool, a whirlwind which spins upwards... 6. the cone of the fir or pine, fir-apple, pine-cone,… also of the tree itself. [1]
A sporangium ; pl.: sporangia) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other groups form sporangia at some point in their life cycle. Sporangia can produce spores by mitosis, but in land plants and many fungi, sporangia produce genetically distinct haploid spores by meiosis.
Ginkgoales are a gymnosperm order containing only one extant species: Ginkgo biloba, the ginkgo tree. The order has a long fossil record extending back to the Early Permian around 300 million years ago from fossils found worldwide.
Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta, also known as Coniferophyta or Coniferae. The division contains a single extant class, Pinopsida. All extant conifers are perennial woody plants with secondary growth. The great majority are trees, though a few are shrubs. Examples include cedars, Douglas-firs, cypresses, firs, junipers, kauri, larches, pines, hemlocks, redwoods, spruces, and yews. As of 2002, Pinophyta contained seven families, 60 to 65 genera, and more than 600 living species.
Gnetophyta is a division of plants, grouped within the gymnosperms, that consists of some 70 species across the three relict genera: Gnetum, Welwitschia, and Ephedra. The earliest unambiguous records of the group date to the Jurassic, and they achieved their highest diversity during the Early Cretaceous. The primary difference between gnetophytes and other gymnosperms is the presence of vessel elements, a system of small tubes (xylem) that transport water within the plant, similar to those found in flowering plants. Because of this, gnetophytes were once thought to be the closest gymnosperm relatives to flowering plants, but more recent molecular studies have brought this hypothesis into question, with many recent phylogenies finding them to be nested within the conifers.
The gymnosperms are a group of seed-producing plants that include conifers, cycads, Ginkgo, and gnetophytes, forming the clade Gymnospermae. The term gymnosperm comes from the composite word in Greek: γυμνόσπερμος, and literally means 'naked seeds'. The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or on their own as in yew, Torreya, and Ginkgo. The life cycle of a gymnosperm involves alternation of generations, with a dominant diploid sporophyte phase, and a reduced haploid gametophyte phase, which is dependent on the sporophytic phase. The term "gymnosperm" is often used in paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.
A conifer cone or, in formal botanical usage, a strobilus, pl.: strobili, is a seed-bearing organ on gymnosperm plants, especially in conifers and cycads. They are also called, according to the relevant genus, cypress cone, fir cone, pine cone, spruce cone, etc. They are usually woody, and variously conic, cylindrical, ovoid, to globular, and have scales and bracts arranged around a central axis, but can be fleshy and berry-like. The cone of Pinophyta contains the reproductive structures. The woody cone is the female cone, which produces seeds. The male cone, which produces pollen, is usually ephemeral and much less conspicuous even at full maturity. The name "cone" derives from Greek konos, which also gave name to the geometric cone. The individual plates of a cone are known as scales. In conifers where the cone develops over more than one year, the first year's growth of a seed scale on the cone, showing up as a protuberance at the end of the two-year-old scale, is called an umbo, while the second year's growth is called the apophysis.
Afrocarpus is a genus of conifer of the family Podocarpaceae. Two to six species are recognized. They are evergreen trees native to Africa. Afrocarpus was designated a genus in 1989, when several species formerly classified in Podocarpus and Nageia were reclassified.
Equisetidae is one of the four subclasses of Polypodiopsida (ferns), a group of vascular plants with a fossil record going back to the Devonian. They are commonly known as horsetails. They typically grow in wet areas, with whorls of needle-like branches radiating at regular intervals from a single vertical stem.
A sporophyll is a leaf that bears sporangia. Both microphylls and megaphylls can be sporophylls. In heterosporous plants, sporophylls bear either megasporangia and thus are called megasporophylls, or microsporangia and are called microsporophylls. The overlap of the prefixes and roots makes these terms a particularly confusing subset of botanical nomenclature.
Bennettitales is an extinct order of seed plants that first appeared in the Permian period and became extinct in most areas toward the end of the Cretaceous. Bennettitales were amongst the most common seed plants of the Mesozoic, and had morphologies including shrub and cycad-like forms. The foliage of bennettitaleans is superficially nearly indistinguishable from that of cycads, but they are distinguished from cycads by their more complex flower-like reproductive organs, at least some of which were likely pollinated by insects.
Lycopodium clavatum is the most widespread species in the genus Lycopodium in the clubmoss family.
Retrophyllum is a genus of conifers in the family Podocarpaceae. It contains five generally recognized extant species with a disjunct distribution in the Southern Hemisphere, found in Papuasia and also in South America. Retrophyllum are evergreen trees typically occurring in tropical rainforests and cloud forests.
Double fertilization or double fertilisation is a complex fertilization mechanism of angiosperms. This process involves the fusion of a female gametophyte or megagametophyte, also called the embryonic sac, with two male gametes (sperm). It begins when a pollen grain adheres to the stigmatic surface of the carpel, the female reproductive structure of angiosperm flowers. The pollen grain begins to germinate, forming a pollen tube that penetrates and extends down through the style toward the ovary as it follows chemical signals released by the egg. The tip of the pollen tube then enters the ovary by penetrating through the micropyle opening in the ovule, and releases two sperm into the embryonic sac (megagametophyte).
Retrophyllum vitiense is a species of conifer in the family Podocarpaceae. It is a large evergreen rainforest emergent tree native to Fiji, Vanuatu, and the Santa Cruz Islands.
Ephedra aspera is a species of Ephedra known by the common names rough jointfir, boundary ephedra, and pitamoreal. It is native to the southwestern United States from California to Texas and parts of northern Mexico as far south as Zacatecas. It is a resident of varied woodland and scrub plant communities.
Selaginella apoda, commonly known as meadow spikemoss, is a perennial lycophyte native to much of the eastern United States and parts of northeastern Mexico. The life cycle is the shortest of the genus Selaginella, as well as one of the shortest among the lycophytes. Selaginella apoda is found primarily in damp soils in habitats such as swamps, wet fields, open woods and along stream banks. Selaginella apoda presents the potential for case studies involving the plant's adaptability to environmental toxins. It is closely related to Selaginella eclipes and S. ludoviciana, with both of which it has been reported to form hybrids. This group is characterized by relatively flat strobili and large megasporophylls which occur in the same plane as the lateral leaves.
This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary of leaf morphology. For other related terms, see Glossary of phytopathology, Glossary of lichen terms, and List of Latin and Greek words commonly used in systematic names.
A seed plant or spermatophyte, also known as a phanerogam or a phaenogam, is any plant that produces seeds. It is a category of embryophyte that includes most of the familiar land plants, including the flowering plants and the gymnosperms, but not ferns, mosses, or algae.
Dendrolycopodium obscurum, synonym Lycopodium obscurum, commonly called rare clubmoss, ground pine, or princess pine, is a North American species of clubmoss in the family Lycopodiaceae. It is a close relative of other species such as D. dendroideum and D. hickeyi, also treelike. It is native to the eastern United States and southeastern Canada from Georgia to Minnesota to Nova Scotia. It grows in the understory of temperate coniferous and deciduous forests, where it is involved in seral secondary succession, growing in clonal colonies some years after disturbance has occurred. It has also been found in Japan, Taiwan, Korea, Russian Far East, and northeastern China.
Neocalamites is an extinct genus of equisetalean plant. Neocalamites thrived during the Permian and Triassic, and occurs worldwide.