The Glossary

Part 2 (D-H)

In all drawings, the distal pole is on top, the proximal pole at the bottom, unless otherwise specified.
e = equatorial view, p = polar view; H = high level, L = low level.
The terms printed in bold are preferred.
The colors are according the scheme described in the introduction.
Click on a drawing for a full sized picture.

Decussate tetrad (Walker and Doyle, 1975)
	A multiplanar tetrad of pollen grains or spores arranged in two pairs lying across one another, the pairs (dyads) more or less at right angles to each other. Example: Orophea (Annonaceae).

Demicolpus (pl. demicolpi, adj. demicolpate) (Erdtman, 1952)
	An ectocolpus divided into two parts. Example: Amylotheca (Loranthaceae). See also: bridge.

Di-
	A prefix for two. Comment: Examples: dicolpate, dicolporate, diporate, disulcate, disulculate, diulculate.

Dicolpate, dicolporate, diporate (adj.) (Iversen and Troels-Smith, 1950)
	Describing pollen grains with two ectocolpi, two compound apertures or two pores. Comment: The terms belong to the system of pollen classes introduced by Iversen and Troels-Smith. See also: disulcate, monoaperturate, pantoaperturate, tricolpate, zonoaperturate.

Dicolporate (adj.) (Cranwell, 1953)
	Synonym of diploporate. Comment: The same term refers to a pollen class in the system of Fægri and Iversen (1950).

* Digitate (adj.) (Skvarla and Larson, 1965)
	With bacula or columellae which are branched distally into two or more parts. Example: Polygonum bistorta.

Diorate (adj.) (Erdtman, 1952)
	Synonym of diploporate.

Diploporate (adj.) (Fægri and Iversen, 1964)
	Describing an ectocolpus with two endoapertures. Example: Didymeles (Didymelaceae). See also: endoaperture area.

Diploxylonoid (adj.) (Traverse, 1988)
	Describing bisaccate pollen grains in which the outline of the sacci in polar view is discontinuous with the outline of the corpus so that the grains seem to consist of three distinct, more or less oval parts. See also: haploxylonoid, Haploxylon-type, Sylvestris-type.

Discordant pattern (Fægri and Iversen, 1989)
	A pattern in a tectate pollen grain in which the arrangement of the columellae is different from that of the elements on the tectum. Example: Geranium (Geraniaceae). Antonym: concordant pattern.

Dispersal unit
	The morphological unit in which mature pollen grains or spores are shed, which may range from individuals (monads), to pairs (dyads), groups of four (tetrads), or groups of more than four (polyads). Larger, indeterminate numbers of pollen grains or spores may also be dispersed as pollinia or massulae.

* Dissections (Couper and Grebe, 1961)
	Rounded to elongated cavities in a cingulum or zona. Example: Vallatisporites ciliaris. Synonym of vacuoles.

Distal (adj.)
	A common descriptive term (Jackson, 1928) used in contrast to proximal, applied in palynology to features on the surface that face outward in the tetrad stage (Wodehouse, 1935). Antonym: proximal. See also: ana-. cata-, polarity.

Distal face (Erdtman, 1952)
	That part of a palynomorph that faces outwards the centre of the tetrad, between equator and distal pole. Antonym: proximal face.

Distal pole (Erdtman, 1952)
	The centre of the surface of the distal face. Antonym: proximal pole. See also: distal.

* Disulcate (adj.) (Harley, 1998)
	Describing pollen grains with sulci arranged in pairs. Two types of disulcate pollen are distinguished: equatorial disulcate, with opposing, eaquatorially arranged sulci and distal disulcate with paired sulci lying parallel to the long axis of the pollen grain on the distal face. Examples: Metroxylon salomonense (Palmae) (equatorial disulcate), Chamaerops humilis (Palmae) (distal disulcate). See also: dicolpate, dicolporate, diporate, geminicolpate.

Duplibaculate (adj.) (Erdtman, 1952)
	Synonym of duplicolumellate.

Duplicolumellate (adj.) (Reitsma, 1970)
	With columellae in two rows under each murus. Example: Polygonum persicaria (Polygonaceae). See also: pluricolumellate, simplicolumellate.

Dyad
	General term for two microspores (pollen grains or spores) united as a dispersal unit. Comment: A developmental dyad stage occurs prior to tetrad formation in plants with successive meiosis. See also: dispersal unit.

Echinate (adj.) (sing. echina, pl. echinae) (Wodehouse, 1928)
	Describing pollen and spores with an ornamentation comprising spines longer than 1µm. Comment: (Echinate should only be used in the adjectival form.) (Erdtman (1952) recommended the term spinose, but in his usage spines were defined as longer than 3µm and smaller features as spinules (adj. spinulose). See also: microechinate, spine, spinule.

Echinolophate (adj.) (Wodehouse, 1928)
	Describing a lophate pollen grain with echinate ridges. See also: fenestrate, psilolophate.

Ectexine (sensu Fægri, 1956)
	The outer part of the exine, which stains positively with basic fuchsin in optical microscopy and has higher electron density in conventionally prepared TEM sections. Orthographical variant: ektexine. Comment: Ectexine includes the foot layer (nexine 1), if present. Erdtman introduced the term in 1943, but used sexine in his later publications. See also: sexine, endexine.

Ecto-
	A prefix for outer.

Ectoaperture (Van Campo, 1958)
	An aperture in the outer layer of the sporoderm. Comment: Examples: ectocolpus, ectopore. See also: endoaperture, mesoaperture.

Ectointine (Freytag, 1968)
	Synonym of exintine.

Ektannulus (pl. ektannuli) (Batten and Christopher, 1981)
	An abrupt thickening of the outer wall layer in the region of the exogerminal. Example: Krutzschipollis. Comment: This term is used in descriptions of the fossil Normapolles group for part of an annulus.

Ektexine (Erdtman, 1943)
	Orthographic variant of ectexine. Comment: The original spelling was soon superseded by the latinized spelling "ectexine" in the publications of Erdtman and others.

Ekto-
	Orthographical variant of the prefix ecto-.

Elater
	A spirally twisted, filamentous band attached to certain spores and some fossil pollen grains (Jackson, 1928). Example: Equisetum (Equisetaceae).

Endannulus (pl. endannuli) (Thomson and Pflug, 1953)
	An annulus formed by the endexine of a pollen grain. Example: Atlantopollis. Comment: This term is used in description of the fossil Normapolles group.

Endexine (sensu Fægri, 1956)
	The inner part of the exine which remains relatively unstained with basic fuchsin in optical microscopy and has a lower electron density in conventionally prepared TEM sections. Synonym of nexine 2 (Erdtman, 1960b). Comment: Originally endexine was defined simply as the inner of the two main layers of the exine (Erdtman, 1943) to which Erdtman later applied the term nexine. However, as more recently defined, endexine does not include the foot layer (nexine 1) which is considered part of the ectexine. See also: ectexine.

Endintine (Kress and Stone, 1982)

	The inner, cellulosic zone of the intine which is adjacent to the cytoplasm and in fluorescence microscopy stains positively with PAS calcofluor. Antonym: exintine. Orthographical variant: endointine.

Endo-
	A prefix for inner.

Endoaperture (Van Campo, 1958)
	An aperture in the inner layer of the sporoderm, often the inner aperture of a compound aperture. Comment: Examples are: endocingulum, endocolpus, endopore. See also: ectoaperture, mesoaperture.

Endoaperture area (Verbeek-Reuvers, 1976)
	The region of the nexine of a single compound aperture which encloses a number of endoapertures. Comment: The term is used where the number of endoapertures is more than two and also where the endoapertures are not sharply delimited. Examples: Deplanchea (Bignoniaceae), Phyllanthus spp. (Euphorbiaceae). See also: colpodiporate, diploporate.

Endocingulum (pl. endocingula, adj. endocingulate) (Reitsma, 1966)
	A ring-shaped endoaperture continuous around a pollen grain and lying in the equatorial plane. Example: Polygonum convolvulus (Polygonaceae). Synonym of colpus equatorialis and zonorate.

Endocrack (Oldfield, 1959)
	An irregular groove occurring in the inner surface of the nexine/endexine and readily apparent in acetolysed pollen. Example: Anemone nemorosa (Ranunculaceae).

Endogerminal (adj.) (Batten and Christopher, 1981)
	Describing an aperture in the inner wall layer. Comment: This term, used in descriptions of the fossil Normapolles group, is essentially synonymous with endoaperture.

Endointine (Freytag, 1968)
	Orthographic variant of endintine.

Endoplica (pl. endoplicae) (Thomson and Pflug, 1953)
	A fold or swelling of the inner wall layer. Comment: A term originally used for features in fossil pollen of the genus Triatriopollenites, and later used in the fossil Normapolles group where endoplicae are present as Y-shaped structures centered over the poles.

Endosculpture (Van Campo, 1971)
	Sculpturing occurring on the inner surface of the nexine/endexine.

Endospore
	The innermost layer of a spore wall (Jackson, 1928). Comment: It is probably homologous with the intine of a pollen grain. Also used as a synonym for the corpus in camerate spores. Mycologists use the term for spores formed on the inside of a sporangium. See also: exospore, perispore.

Endosporium (Erdtman, 1943)
	Synonym of intine.

Equator (Wodehouse, 1935)
	The dividing line between the distal and proximal faces of a pollen grain or spore. See also: equatorial diameter, equatorial plane.

Equatorial axis
	Often misappropriately used as a synonym of equatorial diameter.

Equatorial bridge (Moore and Webb, 1978)
	Synonym of bridge.

Equatorial diameter (Erdtman, 1943)
	A line, lying in the equatorial plane, perpendicular to the polar axis and passing through it. Comment: In bilateral palynomorphs the longest line is usually taken to be the equatorial diameter.

Equatorial lacuna (pl. equatorial lacunae) (Wodehouse, 1928)
	A lacuna situated on the equator in the mesocolpial region of a lophate pollen grain. Example: Tragopogon pratensis (Compositae). See also: lacuna.

Equatorial limb
	Synonym of equatorial outline.

Equatorial outline
	General description of the equator when a pollen grain is seen in polar view. Synonym of limb. See also: amb.

Equatorial plane (Fægri and Iversen, 1950)
	The plane perpendicular to the polar axis and lying midway between the poles.

Equatorial ridge (Wodehouse, 1928)
	A ridge lying along the equator in the mesocolpial region, usually applied to lophate pollen grains. Example: Cichorium intybus (Compositae).

Equatorial view (Erdtman, 1943)
	The view of a pollen grain or spore where the equatorial plane is directed towards the observer. Antonym: polar view.

Equiaxe (adj.equiaxal) (Van Campo, 1966)
	Pollen grains with a polar axis equal to the equatorial diameter. See also: breviaxe, longiaxe.

Erect (adj.) (Reitsma, 1970)
	Synonym of prolate.

Etectate (adj.) (Walker, 1976)
	Describing pollen grains interpreted as having lost their tectum during evolutionary development. See also: atectate, intectate, tectum.

Eu-
	A prefix for true. Comment: When applied to patterns of structuring, such as eurugulate, eustriate and eureticulate, the prefix indicates the arrangement of the columellae. This contrasts with patterns on the tectum which are indicated by the prefix supra-. See also: supra-.

Euintine (Kuprianova, 1956)
	Synonym of endintine.

Eurypalynous (adj.) (Erdtman, 1952)
	Describing plant taxa characterized by possession of a great diversity of palynomorphs. Antonym: stenopalynous.

Eutectum (adj.eutectate)
	Describing a pollen grain with a continuous tectum. Synonym of pertectate, tectum imperforatum and tectum solidum. Antonym: semitectum. See also: tectum.

Exine (pl. exines, adj. exinal, exinous) (Fritzsche, 1837)
	The outer layer of the wall of a palynomorph, which is highly resistant to strong acids and bases, and is composed primarily of sporopollenin.

* Exine 1 (Erdtman, 1969)
	Synonym of ectexine.

* Exine 2 (Erdtman, 1969)
	Synonym of endexine.

Exintine (Kress and Stone, 1982)
	An outer (pectic) layer of the intine, lying below the nexine, and which stains positively with alcian blue. Synonym of ectointine. Antonym: endintine.

Exitus (Wodehouse, 1935)
	The site of exit of the pollen tube from the sporoderm. See also: aperture.

Exoexine (Potonié, 1934)
	Synonym of sexine.

Exogerminal (adj.) (Batten and Christopher, 1981)
	Describing an aperture formed in the outer wall layer of the exine. Comment: This term, used in descriptions of pollen of the fossil Normapolles group, is essentially equivalent to ectoaperture.

Exospore
	The outer layer of a spore wall (Jackson, 1928). Comment: A perispore, if present, lies outside the exospore, and is not part of it. Probably homologous with exine in pollen. Mycologists use the term for spores formed on the outside of a sporangium. See also: endospore, perispore.

Exosporium (Erdtman, 1943)
	Synonym of exine.

Fastigium (pl. fastigia, adj. fastigiate) (Reitsma, 1966)
	Cavity in a colporate grain, appearing as a separation of the inner part of the exine from the domed sexine in the region of the endoaperture. Example: Hypericum (hypericaceae). Synonym of caverna. Comment: The term describes a feature morphologically similar to the vestibulum; the latter is restricted to porate pollen grains. See also: vestibulum.

Fenestrate (adj.) (Iversen and Troels-Smith, 1950)
	Describing a class of pollen grains characterized by large, window-like spaces lacking a tectum. Comment: The term is accepted as a category in the classification system of Iversen and Troels-Smith (1950) and was especially established to include lophate pollen grains of the Compositae. Although useful for defining a class of pollen grains the term is not recommended in descriptions. Many such grains can be described as lophate.

Fimbria (pl. fimbriae, adj. fimbriate)
	Long, hair-like appendages (Jackson, 1928). Comment: The term has been used in the descriptions of fossil spores. Example: Radiatisporites radiatus. Synonym of capillus.

Fischer's law/rule (Erdtman, 1952)
	The name given to the widespread arrangement in developmental tetrads of tri-aperturate pollen grains whereby the apertures form in pairs at six points in the tetrad. Example: Ericaceae See also: Garside's law/rule.

Fissura (pl. fissurae) (Potonié, 1934)
	A sharp, straight split that appears during germination in certain inaperturate pollen grains. Examples: Taxodium (Taxodiaceae), Cupressus (Cupressaceae).

Flange
	A general term, used to describe equatorial extensions of spores (Jackson, 1928). Comment: This term is widely used in Palaeozoic spores, but is not precisely defined. See also: capsula, cingulum, corona, patina, zona.

Foot layer (Fægri, 1956)
	The inner layer of the ectexine. Synonym of nexine 1, pedium and sole. See also: ectexine.

Foramen (pl. foramina, adj. forate, which is an abbreviation of foraminate) (Erdtman, 1952)
	Synonym of pore. Comment: This term is sometimes used in its adjectival form forate (e.g. Walker and Doyle, 1975) which is a synonym of pantoporate. A special form of the term is internal foramen, which refers to features in the exine for example of certain Compositae and Acanthaceae pollen grains. See also: internal foramen.

Fossaperturate (adj.) (Erdtman, 1952)
	Refers to an equatorially aperturate, lobate pollen grain with the apertures in the indentations between the lobes. Comment: The term belongs in the system of ambs as described by Erdtman (1952). See also: angulaperturate, lobate, sinu-aperturate.

Fossula (pl. fossulae, adj. fossulate) (Fægri and Iversen, 1950)
	A feature of ornamentation consisting of an elongated, irregular groove in the surface. See also: foveola.

Fossula (Kuprianova, 1948)
	Synonym of areola.

Foveola (pl. foveolae, adj. foveolate) (Erdtman, 1952)
	A feature of ornamentation consisting of more or less rounded depressions or lumina more than 1µm in diameter. The distance between foveolae is greater than their breadth. See also: lumen, punctum.

Frustillum (pl. frustilla, adj. frustillate) (Fægri and Iversen, 1964)
	Synonym of areola.

Furrow
	A common word for an elongate aperture. Comment: Examples: colpus, sulcus, tenuitas.

Furrow membrane (Wodehouse, 1935)
	Synonym of colpus membrane.

Galea (pl. galeae, adj. galeate) (Sullivan, 1964)
	A relatively large element of the outer wall of a spore consisting of a sharply tapering spine and a broad bulbous base.

Garside's law/rule (Garside, 1946; Erdtman, 1952)
	The name given to an unusual arrangement in developmental tetrads of tri-aperturate pollen whereby the apertures form in groups of three at four points in the tetrad. Comment: The name was given by Erdtman to a phenomenon described by Garside, and apparently restricted to Proteaceae. See also: Fischer's law/rule.

Geminicolpate (adj.) (Erdtman, 1952)
	Describing pollen grains with colpi arranged in pairs. See also: pontoperculum.

Gemma (pl. gemmae, adj. gemmate) (Iversen and Troels-Smith, 1950)
	A sexine element which is constricted at its base, higher than 1µm, and that has approximately the same width as its height.

Generative cell
	The cell in a pollen grain which divides to form male gametes (Jackson, 1928).

Geniculum (pl. genicula, adj. geniculate) (Potonié, 1934)
	A bulge in the equatorial exine of the colpus, often associated with a separation of the sexine from the nexine and the rupturing of the latter. Examples: Fraxinus (Olea ceae), Quercus (Faga- ceae). Comment: If the separation forms a cavity the term fastigium should be applied. See also: fastigium.

Germinal aperture (Wodehouse, 1935)
	A hole in the furrow membrane through which the pollen tube emerges. Comment: Examples: germ pore (synonym of pore), germinal furrow (synonym of ectocolpus). Synonym of exitus.

Goniotreme (adj.) (Erdtman and Straka, 1961)
	Synonym of angulaperturate.

Granular exine (Van Campo and Lugardon, 1973)
	A type of exine stratification in which the infratectal layer is composed of more or less rounded, granules rather than of columellae or other structures. See also: columella, alveolate.

Granulate granulatus (adj.) (Erdtman, 1947)
	With granules.

Granule (pl. granules, adj. granular, granulose)
	General word for a small, rounded element.

Granulum (pl.granula, adj.granulate, granulose) (sensu Erdtman, 1952)
	A very small and rounded element of the sexine/ectexine that is less than 1µm in all directions. Comment: A granulum is the diminutive for granum. See also: scabrate.

Granum (pl. grana) (Potonié, 1934)
	The combined elements gemmae and verrucae.

Groove
	A general descriptive word. See also: platea luminosa, striate.

Gula (pl. gulae, adj. gulate) (Potonié and Kremp, 1955)
	A rather ornate projecting, neck-like, extension on the proximal face of a trilete spore. Example: Lagenicula horrida. Comment: This term is mainly used in the description of some fossil megaspores.

Halo (Erdtman, 1952; Fægri and Iversen, 1989)
	A clear zone around a well defined feature such as a spine or an aperture. Examples: Ranunculus acris (Ranunculaceae), Valeriana (Valerianaceae).

Hamulate (adj.) (Krutzsch, 1959)
	Describing a form of rugulate ornamentation consisting of irregularly arranged, winding, or angular rounded muri of varying thickness, which do not form a distinct reticulum, but rather a maze-like pattern. Example: Lycopodiella inundata (Lycopodiaceae). Comment: Mostly used in spore terminology.

Haploxylon-type (Rudolph, 1935)
	Bisaccate pollen in which the outline of the sacci in polar view is more or less continuous with the outline of the corpus, so that the grains appear a more or less smooth ellipsoidal form. Examples: Pinus cembra, Picea (Pinaceae). See also: diploxylonoid, haploxylonoid, Sylvestris-type.

Haploxylonoid (adj.) (Traverse, 1988)
	Describing bisaccate pollen in which the outline of the sacci in polar view is more or less continuous with the outline of the corpus, so that the grains appear a more or less smooth ellipsoidal form. See also: diploxylonoid, Haploxylon-type, Sylvestris-type.

Harmomegathy (adj. harmomegathic) (Wodehouse, 1935)
	The process by which pollen grains and spores change in shape to accommodate variations in the volume of the cytoplasm caused by changing hydration.

H-endoaperture (Punt and Nienhuis, 1976)
	An elaborate endoaperture, consisting of a central part which connects two lateral, longitudinal elongations, forming an "H" shape. Examples: Cornus (Cornacaea), Centaurium (Gentianaceae).

Heterobrochate (adj.) (Erdtman, 1952)
	Describing a reticulum with brochi of different sizes. Examples: Adoxa Moschatelina (Adoxaceae), Forsythia europaea (Oleaceae). See also: brochus, lumen.

Heterocolpate (adj.) (Iversen and Troels-Smith, 1950)
	Describing pollen grains with both simple and compound colpi present. Examples: Lythrum, Peplis (Lythraceae), Myosotis Boraginaceae). See also: pseudocolpus.

Heteropolar (adj.) (Erdtman, 1952)
	Describing pollen or spores in which the distal and proximal faces of the exine are different, either in shape, ornamentation or apertural system. Example: Echium vulgare (Boraginaceae). Antonym: isopolar.

Heterosporous (adj.)
	Describing plants producing both microspores and megaspores (Jackson, 1928). Antonym: homosporous. See also: Isosporous.

Hilum (pl. hila, adj. hilate) (Erdtman, 1952)
	Circular, indistinctly delimited, irregular aperture or thinning in spores. Examples: Aequitriradites verrucosus, Couperisporites tabulatus. Comment: This kind of feature is present in certain bryophytes and fungi and may occur on the proximal or distal face. See also: polumbra

Homobrochate (adj.) (Erdtman, 1952)
	Describing a reticulum with brochi of the same sizes. Example: Armeria maritima (Plumbaginaceae). See also: brochus, lumen.

Homosporous (adj.)
	Describing plants producing only one kind of spores (Jackson, 1928). Synonym of isosporous. Antonym: heterosporous.

Horn (Huynh, 1970)
	An elongated part of an endoaperture, which is curved towards one of the poles. Example: Anagallis arvensis (Primulaceae). See also: H-endoaperture.

Updated 16 April 1999

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