The Glossary

Part 5 (S-Z)

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.

* S-type tetrad (Moar, 1993)
	A tetrad in which only one members is fully developed. Example: Leucopogon fasciculatus (Epacridaceae). See also: A-type tetrad, T-type tetrad.

* Saccoid (adj.)(Brugman, 1983)
	A saccus-like expansion of the exine with a complex infrastructure build up by a three dimensional network of sexine elements, extending to and fused with the nexine. See also: camera, protosaccus, pseudosaccus, saccus.

Saccus (pl. sacci, adj. saccate) (Erdtman, 1952)
	A sac formed by an expansion of the exine of a pollen grain and at least partly filled with an alveolate infrastructure. See also: bisaccate, camera, protosaccus, pseudosaccus.

Scabrate (adj.) (sing. scabra, pl. scabrae) (Iversen and Troels-Smith, 1950)
	Describing elements of ornamentation, of any shape, smaller than 1µm in all directions. Examples: Quercus (Fagaceae), Artemisia (Compositae). Comment: Ornamentation elements larger than 1µm are described according to their shape, for example, baculum, clava, gemma, verruca. See also: granulum.

Sclerine (Erdtman, 1952)
	A term encompassing both exine and perine that can be used whether a perine is present or not; sporoderm excluding the intine. See also: sculptine.

Scrobiculus (pl. scrobiculi, adj. scrobiculate) (Potonié, 1934)
	Synonym of punctum.

Sculptine (Erdtman, 1948)
	A term encompassing both exine and perine (sclerine) but excludes the nexine and so provides a neutral term for the sculptured layer when there is doubt whether the pattern belongs to the exine or the perine. See also: sclerine.

* Sculpural density (Balme, 1988)
	The estimated number of sculptural elements in an area of 100 µm2 of the surface of the exine.

Sculpture (Kuprianova, 1948)
	Orthogriphical variant of sculpturing.

Sculpturing (adj. sculptured) (Potonié, 1934)
	The surface relief, or topography, of a pollen grain or spore. Comment: Praglowski (1975) provided a circumscription of this term which would restrict its application to tectate pollen grains. See also: pattern, ornamentation, structure.

Seed- megaspore
	A large, functional megaspore associated with three small, presumably aborted spores present in some fossil lycopsids. Example: Cystosporites.

Semi-erect (adj.) (Reitsma, 1970)
	Synonym of subprolate.

Semitectum (adj. semitectate) (Fægri and Iversen, 1964)
	A partially discontinuous tectum in which the tectal perforations are equal to or wider than the muri and usually larger than 1µm in diameter. Antonym: eutectum. See also: tectum.

Semitransverse (adj.) (Reitsma, 1970)
	Synonym of suboblate.

Sexine (Erdtman, 1952)
	The outer, sculptured layer of the exine, which lies above the nexine. Antonym: nexine. Comment: Although the term ectexine as originally defined (sensu Erdtman, 1943) was more or less synonymous with sexine, it no longer is because ectexine is now invariably used in the redefined sense of Fægri (Fægri (1956). Unlike ectexine, sexine does not include the foot layer. Sexine and nexine are distinguished on purely morphological criteria, whereas ectexine and endexine differ in their staining properties. The two sets of terms are therefore suited for slightly different applications. See also: ectexine.

* Sexine 1, 2, 3, 4, 5 (Reitsma, 1970)
	A system of sexine stratification in which sexine 1 is the innermost and sexine 5 in the outermost layer of the sexine. Comment: Usually the sexine consists of 3 layers (sexine 1 = columellae; sexine 2 = tectum; sexine 3 = sclupture elements).

Shape classes (pl.) (Erdtman, 1943)
	Categories of pollen and spore shape based on the relations between polar axis (P) and equatorial diameter (E). See also: oblate, oblate spheroidal, P/E ratio, peroblate, perprolate, prolate, prolate spheroidal, suboblate, subprolate.

Simplibaculate (adj.) (Erdtman, 1952)
	Synonym of simplicolumellate.

Simplicolumellate (adj.) (Reitsma, 1970)
	With a single row of columellae under each murus. Example: Viburnum opulus (Caprifoliaceae). See also: duplicolumellate, pluricolumellate.

Sinu-aperturate (adj.) (Erdtman, 1952)
	Describing a pollen grain in which the equatorial apertures are situated in the middle of concave sides when seen in polar view. Comment: The term belongs to the system of ambs defined by Erdtman. See also: amb, angulaperturate, planaperturate.

Sole (APLF, 1975)
	Synonym of foot layer.

Special wall (Beer, 1911)
	Pollen mother-cell wall

Spheroidal (Erdtman, 1943)
	Describing the shape of a pollen grain or spore in which the polar axis and the equatorial diameter are approximately equal. Comment: This term belongs to the system of shape classes suggested by Erdtman (1943), and extended in 1952), based on the measurements of the polar axis (P) and equatorial diameter (E). In this system spheroidal is defined as a P/E ratio of 0.88-1.14. See also: oblate spheroidal, prolate spheroidal.

Spina (pl.spinae) (Potonié, 1934)
	Synonym of Spine.

Spine (adj. spiny/spinose) (Erdtman, 1952)
	A general word, applied in palynology to long and tapering pointed elements, exceeding 1µm. Comment: In Erdtman's (1952) definition spines were defined as more than 3µm long (in contrast to spinules which were shorter than this) but since the maximum size of features that may be prefixed with micro- is 1µm the present definition provides consistency with other terms. See also: echinate.

Spinule (adj. spinulose) (Erdtman, 1952)
	Small spines, less than 3µm in length. Comment: The size distinction made by Erdtman between spinules and spines is not consistent with other size criteria used in palynology. See also: spine.

Spiraperturate (adj.) (Erdtman, 1952)
	Describing a pollen grain with one or more spiral apertures. Example: Eriocaulon aquaticum (Eriocaulaceae). Comment: In the pollen classes of Iversen and Troels-Smith (1950), spiraperturate pollen was included in the syncolpate class.

Spore
	A general term for the usually microscopic, unicellular, asexual or sexual reproductive units of cryptogams and fungi (Jackson, 1928). See also: pollen, microspore.

Sporoderm (Bischoff, 1833)
	The entire wall of a pollen grain or spore.

Sporomorph (Erdtman, 1947)
	A general term for spore-like palynomorphs.

Sporopollenin (Zetsche and Vicari, 1931)
	The name given to the acetolysis resistant biopolymers which make up most of the material of the exine.

Square tetrad (Erdtman, 1945b)
	Synonym of tetragonal tetrad.

Stenopalynous (adj.) (Erdtman, 1952)
	Describing plant taxa characterized by only a slight variation in their palynomorphs. Antonym: eurypalynous.

Stephano-
	Synonym of zono-. Comment: Fægri and Iversen (1950) used the prefix (as for example, in stephanocolpate, stephanocolporate, stephanoporate) in their classification of pollen types.

Stratum (APLF, 1975)
	A subdivision of a major layer of the sporoderm.

Striae (pl.) (sing. stria) (Fægri and Iversen, 1950)
	Grooves between elongated sculpturing elements.

Striate (adj.) (Iversen and Troels-Smith, 1950)
	A general descriptive term applied in palynology to elongated, generally parallel elements separated by grooves. Comment: The positive elements of striate ornamentation may also be referred to as muri.

Striato-reticulate (adj.) (Erdtman, 1952)
	Describing a pattern in which parallel or subparallel muri are cross-linked to form a reticulum in the grooves. The connections between the muri lie on a single level or different levels. Example: Gentiana pneumonathe (Gentianaceae).

Structure (adj. structurate, structured) (Potonié, 1934)
	The internal construction of the pollen or spore wall. See also: sculpturing, pattern, ornamentation.

Sub-
	A prefix for under, or less than.

Suberect (adj.) (Reitsma, 1970)
	Synonym of prolate spheroidal.

Subisopolar (adj.) (Walker and Doyle, 1975)
	Describing a pollen grain or spore in which the proximal and distal faces are slightly different. Examples: one face is convex and the other is less convex (Banksia, Proteaceae); on one face viscin threads are connected (Oenothera, Onagraceae).

Sub-layer
	Synonym of stratum.

Suboblate, (adj.) (Erdtman, 1952)
	Describing the shape of a pollen grain or spore in which the ratio between the polar axis and the equatorial diameter is 0.75-0.88. See also: oblate, oblate spheroidal, P/E ratio, peroblate, perprolate, prolate, prolate spheroidal, shape classes, spheroidal, subprolate, subspheroidal.

Subprolate (adj.) (Erdtman, 1952)
	Describing the shape of a pollen grain or spore in which the ratio between the polar axis and the equatorial diameter is 1.14-1.33. See also: oblate, oblate spheroidal, P/E ratio, peroblate, perprolate, prolate, prolate spheroidal, shape classes, spheroidal, suboblate, subspheroidal.

Subspheroidal (adj.) (Erdtman, 1952)
	Describing the shape of a pollen grain or spore in which the ratio between the polar axis and the equatorial diameter is 0.75-1.33. Comment: This shape class includes suboblate, oblate spheroidal, prolate spheroidal and subprolate. See also: oblate, oblate spheroidal, peroblate, P/E ratio, perprolate, prolate, prolate spheroidal, shape classes, spheroidal, suboblate, subprolate.

Subtransverse (adj.) (Reitsma, 1970)
	Synonym of oblate spheroidal.

Subturma (pl. subturmae) (Potonié, 1956)
	A group of form-genera of fossil spores or pollen in the turma-system of Potonié. See also: anteturma, infraturma, turma.

Successiform (Van Campo, 1967)
	Referring to a phyletic series of pollen types with increasing numbers of apertures, ranging from tricolpate to pantocolpate and pantoporate.

Sulculus (pl. sulculi, adj. sulculate) (Erdtman, 1952)
	An elongated latitudinal ectoaperture not situated at a pole. See also: sulcus.

Sulcus (pl. sulci, adj. sulcate) (Erdtman, 1952)
	An elongated latitudinal ectoaperture situated at the distal or proximal pole of a pollen grain. Comment: A sulcus has the same shape as a colpus, but differs in orientation. Sulci are essentially latitudinal apertures whereas colpi are essentially longitudinal apertures. Sulci may be distal (anasulcate), proximal (catasulcate) or extend right around the grain (zonasulcate). See also: ana-, colpus, zona-.

Supra-(prefix)
	A prefix for above. Comment: In palynology this prefix is mostly used for features on top of the tectum, as for example in suprareticulate, suprarugulate, suprastriate.

Supratectal (adj.) (Erdtman, 1969)
	Indicating the position of features, such as spines, on the top of the tectum.

Suture (Potonié and Kremp, 1955)
	Synonym of commissure. See also: laesura.

Sylvestris-type (Rudolph, 1935)
	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. Examples: Pinus sylvestris, Abies (Pinaceae). See also: diploxylonoid, haploxylonoid, Haploxylon-type.

Syn-
	A prefix indicating the fusion or anastomosis of features.

Syncolp(or)ate (adj.) (sensu Erdtman, 1952)
	Describing a pollen grain with two or more simple (or compound) colpi the ends of which anastomose at the pole. Example: Primula farinosa (Primulaceae). Comment: The term was used in a wider sense by Iversen and Troels-Smith (1950) for other forms with fused apertures such as spiraperturate and parasyncolp(or)ate.

Synrugoidate (adj.) (Erdtman, 1935; Jalan and Kapil, 1964)
	Describing a pollen grain with six colpi of which three are long and meeting at one pole and three are short and not meeting at either pole. Example: Schisandra grandiflora (Schisandraceae).

T-shaped tetrad (Walker and Doyle, 1975 )
	A uniplanar tetrad in which two of the members are perpendicular to the other two so that tetrad has the shape of the letter "T". Example: Typha spp. (Typhaceae). See also: tetrad.

* T-type tetrad (Moar, 1993)
	A tetrad in which all four members are fully developed. Example: Pentachondra pumila (Epacridaceae). See also: A-type tetrad, S-type tetrad.

Taenia (pl. taeniae, adj. taeniate) (Leschik, 1956)
	One or more strap-like, more or less parallel strips of exine on the proximal and/or distal sides of the corpus of certain fossil gymnospermous pollen grains. Examples: Striatites, Vittatina. See also: polyplicate.

Tectum (pl. tecta, adj. tectate) (Fægri and Iversen, 1950)
	The layer of sexine, which forms a roof over the columellae, granules or other infratectal elements. See also: eutectum, semitectum, tegillum.

Tectum perforatum (adj. tectate perforate) (Iversen and Troels-Smith, 1950)
	A tectum with perforations smaller than 1µm in diameter. See also: punctum.

Tectum imperforatum (adj. tectate imperforate) (Walker and Doyle, 1975)
	With a continuous tectum, without perforations. Synonym of eutectum.

Tectum solidum (Fægri and Iversen, 1975)
	Synonym of eutectum.

Tegillum (pl. tegilla, adj. tegillate) (Erdtman, 1952)
	Synonym of tectum.

Tenui- (Erdtman, 1952)
	A prefix for thin. Comment: Erdtman often used this prefix in combination with other terms (e.g. tenuimarginate) but thinness is subjective; it is preferable to use alternative descriptions.

Tenuitas (pl. tenuitates) (Potonié, 1934)
	A general term for a thinning, that has been applied to many different situations in palynology. Comment: The term was used by Potonié to describe endoapertures, by Potonié and Kremp (1955) for sulci and has sometimes been used for the distal thin parts in the exine of the Circumpolles group (Corollina).

Tetrad
	A general term for a group of four united pollen grains or spores, either as a dispersal unit or as a developmental stage. Comment: Tetrads may be uniplanar, with all members lying in the same plane (for example, linear, rhomboidal, tetragonal and T-shaped tetrads) or multiplanar, with members in more than one plane (for example, decussate or tetrahedral tetrads). See also: dispersal unit, monad, dyad.

Tetrad mark (Couper and Grebe, 1961)
	The monolete or trilete mark on the proximal face of a spore or, more rarely, a pollen grain. See also: trilete mark, Y-mark.

Tetrad stage
	The period during post-meiotic development when the four microspores or megaspores are united by the presence of a temporary special wall. Comment: The tetrad stage ends at the start of the free spore stage, when the special cell wall is reabsorbed.

Tetragonal tetrad (Grebe, 1971)
	A uniplanar tetrad in which all four members are in contact at the centre of the tetrad so that, in the correct orientation, the adjacent walls form a cross. Example: Uvariastrum hexaloboides (Annonaceae) See also: tetrad.

Tetrahedral tetrad (Grebe, 1971)
	A multiplanar tetrad in which each member is in contact with three others, so that the centers of the grains define a tetrahedron. Example: Erica (Ericaceae). See also: decussate tetrad, tetrad.

Torus (pl. tori) (Thomson and Pflug, 1953)
	An arcuate invagination or protuberance of the exine more or less paralleling the laesura of a spore in the interradial area. See also: kyrtome, labrum.

Transverse (adj.) (Reitsma, 1970)
	Synonym of oblate. Comment: Transverse has also been used to indicate the orientation of latitudinal features. It is also commonly used to indicate a section cut through the equatorial plane of a pollen grain.

Transversal furrow (Iversen and Troels-Smith, 1950)
	Synonym of endocolpus.

Trema (pl. tremata) (Erdtman and Straka, 1961)
	Synonym of aperture. See also: -treme.

-treme (Erdtman and Straka, 1961)
	A suffix synonymous with aperture. Comment: The -treme system of aperture classes suggested by Erdtman and Straka was intended to replace that of Iversen and Troels-Smith (1950). The system included atreme, monotreme, ditreme, tritreme, tetratreme, pentatreme, hexatreme, polytreme, anomotreme, pleotreme and stephanotreme.

Tri-
	A prefix for three.

Trichotomocolpate (adj.) (Erdtman, 1945a)
	Describing a pollen grain with a three-armed colpus. Example: Trapa natans (Trapaceae).

Trichotomosulcate (adj.) (Erdtman, 1952)
	Describing a pollen grain with a three-armed sulcus. Example: Elaeis guineensis (Palmae).

Tricolpate, tricolporate, triporate (adj.) (Iversen and Troels-Smith, 1950)
	Describing pollen grains with three ectocolpi, three compound apertures or three pores. Comment: The terms belong to the system of pollen classes introduced by Iversen and Troels-Smith (1950). See also: monoaperturate, pantoaperturate, zonoaperturate.

Trifolium (pl. trifolia) (Potonié, 1956)
	A three-bladed, proximal feature arising from the proximal pole of a megaspore. Example: Capulitriletes. See also: acrolamella, gula.

Trilete (adj.) (Erdtman, 1943)
	Describing a spore with three laesurae, thus showing a trilete mark. Example: Pteridium (Hypolepidaceae). See also: alete, laesura, monolete.

Trilete mark
	The triradiate mark of a trilete spore.

Triprojectate (adj.) (Mtchedlishvilli, 1961)
	Describing projectate pollen with three projecting arms on which the apertures are situated, as for example in the Triprojectacites group of fossil pollen grains. Example: Aquilapollenites.

Tryphine (Erdtman, 1969)
	A material deposited on the surface of pollen grains by the breakdown of the tapetum and differing from pollenkitt in that it contains membraneous components derived from organelles. See also: pollenkitt.

* Tuberculate (adj.) (Moar, 1993)
	Covered with knobbly projections.

Tubulus (pl. tubuli) (Erdtman, 1952)
	A general term for a small channel, applied in palynology to a channel through the nexine.

Tula (pl. tulae, adj. tulate) (Jansonius and Pocock, 1969)
	A sexinal inflation in gymnospermous pollen at the end of the axis of the distal sulcus or leptoma. Example: Ovalipollis.

Tumescence (pl. tumescences) (Batten and Christopher, 1981)
	A gradual increase in thicknes of the wall layer(s) from a point in the equatorial interradial region to the germinal aperture. Example: Megatriopollis. Comment: This term is used in descriptions of the fossil Normapolles group.

Turma (pl. turmae) (Potonié, 1956)
	An artificial suprageneric grouping of form-genera of fossil spores and pollen. Comment: The following groups are recognized in the system: anteturma, turma, subturma, and infraturma.

* Turriform (Balme, 1988)
	Biform sculptural elements consisting of a capitate basal portion surmounted by a sharply contracted distal spine. Example: Dibolisporites.

Ubisch body (pl. ubisch bodies) (Rowley, 1963)
	A distinctive, orbicular granule of sporopolleninsporopollenin produced by the tapetum, particularly in plants with secretory tapeta. See also: orbicule.

Ulculus (pl. ulculi, adj. ulculate) (Walker and Doyle, 1975)
	A rounded ectoaperture not situated at a pole. Example: Poaceae. See also: ulcus.

Ulcus (pl. ulci, adj. ulcerate) (Erdtman, 1952)
	A rounded ectoaperture situated at the distal or proximal pole of a pollen grain. Examples: Sparganium (Sparganiaceae), Typha (Typhaceae). Comment: An ulcus has the same shape as a pore. Ulci may be distal (ana-ulcerate) or proximal (cata-ulcerate). See also: ana-, cata-, pore, ulculus.

Uniplanar tetrad (Walker and Doyle, 1975)
	A tetrad in which the individual members lie more or less in one plane.

* Urceolate (adj.) (Ferguson et al., 1983)
	Describing a type of ornamentation consisting of urn- shaped elements situated on the footlayer. Example: Pinanga aristata (Palmae).

* Vacuoles (Grebe, 1971)
	Rounded to elongated spaces within an equatorial faeture. Example: Vallatisporites ciliares. Synonym of dissections. Comment: In botanical sence, vacuole is a general term for a with liquid filled cellular component (Jackson, 1928). In latin it means a hollow space, and in this sence this term is used.

Vallum (pl. valla) (Iversen and Troels-Smith, 1950).
	Synonym of murus. Comment: Sometimes used to describe a single, broad, raised feature in certain fossil spores. See also: lira.

Valva (pl. valvae, adj. valvate) (Potonié and Kremp, 1955)
	Radial thickenings in the areas beyond the ends of the laesurae of trilete spores. Example: Triquitrites tribullatus. Comment: The term auriculate (Potonié and Kremp, 1955) refers to an extremely valvate condition.

Velum (pl. vela, adj. velate) (Thomson and Pflug, 1953)
	A feature of a monosaccate pollen grain in which the saccus is convoluted. Example: Tsuga (Pinaceae).

Vermiculate (adj.) (Kosanke, 1950; Harris, 1955)
	A general descriptive term used to describe winding features. Comment: Vermiculate has been used by Kosanke (1950) and Harris (1955) to describe depressions (fossulae). Others use the term for raised structures (muri) in rugulate pollen and spores.

Verruca (pl. verrucae, adj. verrucate) (Iversen and Troels-Smith, 1950)
	A wart-like sexine element, more than 1µm wide, that is broader than it is high and is not constricted at the base. Example: Plantago (Plantaginaceae).

Verrucose (adj.) (Erdtman, 1969)
	Synonym of verrucate.

Vesiculate (adj.) (Iversen and Troels-Smith, 1950)
	Synonym of saccate.

Vestibulum (pl. vestibula) (Potonié, 1934)
	A separation between layers of the exine forming a cavity between the inner and outer pores. For example: Betula (Betulaceae). See also: atrium, fastigium.

Viscin thread (pl. viscin threads)
	A general botanical term (Jackson, 1928), applied in palynology for an acetolysis resistant, sporopollenin thread arising from the exine of a pollen grain, usually from the proximal surface. Example: Oenothera (Onagraceae).

Y-mark (Potonié, 1934)
	Synonym of trilete mark.

Z-layer (Heslop-Harrison, 1979)
	Synonym of exintine.

Zona (pl. zonae, adj. zonate) (Potonié and Kremp, 1955)
	A thin outer structure of a spore that projects at the equator, but does not extend over the distal face or proximal face. Example: Cirratriradites saturni. Comment: The term is especially applied for many representatives of fossil spores. Example: Kraeuselisporites. See also: cingulum, corona.

Zona- (adj. zonate) (Walker and Doyle, 1975)
	Prefix indicating ring-like. Comment: The prefix zona- is used in conjunction with a suffix indicating the type of aperture.

Zona-aperturate (adj.) (Walker and Doyle, 1975)
	Describing a pollen grain with a ring-like aperture. Comment: The range of ring-like apertures includes for example: anazonasulculus (1), a ring-like sulculus in the distal hemisphere; catazonasulculus, a ring-like sulculus in the proximal hemisphere; zonasulculus, a ring-like sulculus around the equator; and zonasulcus (2), a meridional ring-like sulcus perpendicular to the equator.

Zonasulculus (Walker and Doyle, 1975)
	A ring-like sulculus around the equator. Example: Nymphaea violacea (Nymphaeaceae). See also: meridionosulcus.

Zonasulcus (Walker and Doyle, 1975)
	A meridional ring-like sulcus perpendicular to the equator. Example: Laurelia novaezelandiae (Monimiaceae). See also: meridionosulcus.

Zoni- (sensu Erdtman and Vishnu-Mittre, 1956)
	Synonym of zono-.

Zoni- (sensu Walker and Doyle, 1975)
	Prefix indicating a latitudinal orientation.

Zono- (Erdtman and Straka, 1961)
	A prefix indicating features located equatorially. See also: ana-, cata-, stephano-.

Zonoaperturate (adj.) (Erdtman and Straka, 1961)
	Describing a pollen grain with apertures situated only at the equator. Comment: Pollen classes with zonoaperturate pollen include, zonocolpate, zonocolporate, zonoporate. Originally Erdtman and Vishnu-Mittre (1956) introduced the prefix zoni-, but later Erdtman and Straka (1961) changed the ending of the prefix to zono-. See also: stephano-.

Zonorate (adj.) (Erdtman, 1952)
	Describing a pollen grain with a continuous endoaperture (os) around the equator. Synonym of endocingulum.

Zwischenkörper (pl.) (Fritzsche, 1837)
	A lens-shaped body below the aperture of certain unacetolysed pollen grains. Comment: The feature resembles an oncus but is treated as distinct because some pollen grains have both features.

Updated 16 April 1999

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