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Illustrated Handbook

on
Pollen Terminology

We encourage you to redistribute the "PalDat - Illustrated Handbook on Pollen Terminology" either in printed or
electronic version.
In any case it is not allowed to modify it or to distribute it without the copyright notice below.
The "PalDat - Illustrated Handbook on Pollen Terminology" is distributed in the hope that it will be useful for all
persons interested in palynology, but WITHOUT ANY WARRANTY.

Address
Dept. of Palynology and Structural Botany
PalDat
University of Vienna
Rennweg 14
A 1030 Wien
AUSTRIA
Tel: +43 1 4277 54100
Fax: +43 1 4277 9541
info@paldat.org
www.paldat.org

Citation
Halbritter H, Weber M, Zetter R, Frosch-Radivo A, Buchner R, Hesse M. 2009.
PalDat - Illustrated Handbook on Pollen Terminology. Vienna. 62 pp.

Impressum
Copyright holder and publisher
Society for the Promotion of Palynological Research in Austria
PalDat
University of Vienna, Rennweg 14, A 1030 Wien, AUSTRIA
Officer in charge: BUCHNER Ralf
Errors and omissions excepted.

VORWORT
Seit 1994 ist "The Glossary of Pollen and Spore Terminology" von Punt W, Blackmore S, Nilsson S & Le
Thomas A das Standardwerk der palynologischen Terminologie, inkl. den Ergnzungen in der seit 1999 jedoch
nicht mehr berarbeiteten online-Version (http://www.bio.uu.nl/~palaeo/glossary/glos-int.htm).
Schwerpunkt unseres Glossars sind die zahlreichen Abbildungen. Jeder Begriff, der in das Glossar
aufgenommen wurde, ist umfangreich mit elektronenmikroskopischen Bildern illustriert. Dabei wollten wir
besonders die Bandbreite eines Begriffes aufzeigen.
Wir bemhten uns in unserem online-Glossar "PalDat - Illustrated Handbook on Pollen Terminology" um eine
Straffung der Begriffe nach praktischen Gesichtspunkten. Viele Begriffe der Palynologie wurden ja zu einer
Zeit geprgt, als ausschlielich Zeichnungen Grundlage von LM-Beobachtungen zur Verfgung standen.
Daher gibt es aus historischen Grnden immer wieder nomenklatorische Inkonsequenzen, Synonyme,
unterschiedliche Definitionen fr ein und denselben Begriff.
Unser Ziel war es, ein Glossar der wichtigsten Begriffe zu erstellen. Durch bewute Vereinfachung wollten wir
auch erreichen, dass dieses Glossar nicht nur fr Spezialisten verwendbar ist. Dass durch Generalisierung
bzw. Subsummierung unter einem berbegriff manche Begriffe wegfallen, haben wir bewusst in Kauf
genommen. Sollten Begriffe vermisst werden, erschienen sie uns als unwesentlich, oder sie gehren zur
Terminologie der Farnsporen, die in dieser Ausgabe nicht bercksichtigt ist. Gibt es von einem Begriff mehrere
Synonyme, wird nur noch einer der Begriffe verwendet. Wo es notwendig war, haben wir Definitionen
modifiziert. Durch konsequente Anwendung elektronenmikroskopischer Techniken und dem damit
verbundenen erweiterten Wissensstand mussten manche Begriffe auch neu definiert werden.
Wir sehen dieses online-Glossar als eine Mglichkeit, auf einen vernderten Wissensstand flexibel reagieren
zu knnen und freuen uns ber Vorschlge und Anregungen. Schreiben Sie diese an handbook@paldat.org

PREFACE
Since 1994 "The Glossary of Pollen and Spore Terminology" of Punt W, Blackmore S, Nilsson S & Le Thomas
A is the standard publication in palynological terminology. Unfortunately it was not revised following the
additions in 1999 in the online version (http://www.bio.uu.nl/~palaeo/glossary/glos-int.htm).
Emphasis of our glossary are the numerous illustrations. Each term is illustrated with electron microscopic
pictures since we wanted to point out the character range of a term.
We tried to work out in our glossary, the "PalDat - Illustrated Handbook on pollen Terminology" a
rationalization of terms on the basis of practical criteria. Many terms in palynology were coined at a time, when
exclusively LM observations were available. Mainly for historical reasons nomenclatorical inconsequences,
synonyms, different definitions are found for one and the same term.
Our goal was to provide a glossary of the most important terms. By conscious simplification we wanted to write
a glossary which is not only a valuable source for specialists. We accepted consciously that by generalization
and/or subsumption under a general term some terms were omitted. If terms are not considered here then they
appeared us as insignificant, or they belong to the terminology of the fern spores, which are not considered
here. In case that there are several synonyms for a term, only one of the terms is used. Where it was
necessary, we modified definitions. By consistent application of electron microscopic techniques and the
extended knowledge some terms also had to be redefined. We see this online glossary as a possibility of being
able to react to changed knowledge conditions flexibly.
Please send your notes and suggestions to handbook@paldat.org

Wien, April 2005

Halbritter H, Weber M, Zetter R, Frosch-Radivo A, Buchner R, Hesse M

Pollen unit
Pollen unit
polyad
Dispersal unit of more than four united pollen grains.

Acacia myrtifolia (Mimosaceae)

Himantoglossum adriaticum (Orchidaceae)

Platanthera bifolia (Orchidaceae)

Carex remota (Cyperaceae)


Tetrad (pseudomonad)

Dionaea muscipula (Droseraceae)


Tetrad (tetrahedral)

Epilobium parviflorum (Onagraceae)


Tetrad (tetrahedral)

Erica herbacea (Ericaceae)


Tetrad (tetrahedral)

Mimosa pudica (Mimosaceae)

Typha latifolia (Typhaceae)


Tetrad (planar)

tetrad
Dispersal unit of four pollen grains (spores).

Typha latifolia (Typhaceae)


Tetrad (planar)

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Pollen class
Pollen class
clypeate
Pollen wall, in which the exine is subdivided into shields.

Catalpa ovata (Bignoniaceae)

Dracunculus vulgaris (Araceae)

Ibicella lutea (Martyniaceae)

Iris bucharica (Iridaceae)

Peperomia rubella (Piperaceae)

Phyllanthus spec (Euphorbiaceae)

lophate
Pollen wall with lophae, a network-like pattern of ridges (=lophae) formed by the
outer exine surrounding window-like spaces or depressions (=lacunae).

Cichorium intybus (Asteraceae)

Helminthotheca echioides (Asteraceae)

Scorzonera hispanica (Asteraceae)

plicate
Pollen grain with plicae, i.e. cirumferential, parallel ridge-like folds.

Ambrosina bassii (Araceae)

Amorphophallus asterostigmatus (Araceae)

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Hemigraphis primulaefolia (Acanthaceae)

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Pollen class

Pistia stratiotes (Araceae)

Steudnera assamica (Araceae)

saccate
Pollen grain with one or more air sacs, i.e. exinous expansions forming an air sac.

Abies cephalonica (Pinaceae)


Bisaccate, dry pollen grains

Pinus mugo (Pinaceae)


Bisaccate pollen grain

Tsuga canadensis (Pinaceae)


Monosaccate pollen grains

spiraperturate
Pollen grain with pollen grain with one or more spiral aperture(s).

Berberis vulgaris (Berberidaceae)

Crocus speciosus (Iridaceae)

Mimulus guttatus (Veronicaceae)

Thunbergia alata (Acanthaceae)

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Shape (single pollen grains)


Shape (single pollen grains)
irregular

Aechmea azurea (Bromeliaceae)


Dry pollen grains

Anthurium radicans (Araceae)


Dry pollen grains

Aristolochia arborea (Aristolochiaceae)


Dry pollen grain

Carex humilis (Cyperaceae)


Dry tetrads

Hordeum murinum (Poaceae)


Dry pollen grains

Laportea divaricata (Urticaceae)


Dry pollen grains

oblate
Pollen grain with a polar axis shorter than the equatorial diameter.

Hohenbergia proctori (Bromeliaceae)

Limnanthes douglasii (Limnanthaceae)

Muscari tenuiflorum (Hyacinthaceae)

Pinus mugo (Pinaceae)

Primula farinosa (Primulaceae)


Dry pollen grains

Salvia involucrata (Lamiaceae)

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Shape (single pollen grains)

Tilia cordata (Tiliaceae)


Dry pollen grains

prolate
Pollen grain with a polar axis longer than the equatorial diameter.

Begonia nelumbifolia (Begoniaceae)

Beloperone guttata (Acanthaceae)


Dry pollen grains

Crossandra flava (Acanthaceae)

Daucus carota (Apiaceae)

Lithospermum officinale (Boraginaceae)

Lotus corniculatus (Fabaceae)

Nandina domestica (Berberidaceae)


Dry pollen grains

Symphytum orientale (Boraginaceae)

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Shape (single pollen grains)


spheroidal
Pollen grain where the polar axis and the equatorial diameter are equal in length.

Abeliophyllum distichum (Oleaceae)

Daphne alpina (Thymelaeaceae)

Phyteuma spicatum (Campanulaceae)


Dry pollen grains

Ruellia graecicans (Acanthaceae)

Schismatoglottis lancifolia (Araceae)

Stenandrium dulce (Acanthaceae)


Dry pollen grain

Thlaspi arvense (Brassicaceae)

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Shape peculiarities
Shape peculiarities
heteropolar
Pollen grain with different proximal and distal faces.

Aechmea fraseri (Bromeliaceae)

Buglossoides purpurocaerulea (Boraginaceae)

Carex alba (Cyperaceae)


Tetrad (pseudomonad)

Microrrhinum minus (Veronicaceae)

Onosma arenaria (Boraginaceae)

Thesium linophyllon (Santalaceae)

isopolar
Pollen grain with identical proximal and distal faces.

Bellis perennis (Asteraceae)

Knautia drymeia (Dipsacaceae)

Nepeta cataria (Lamiaceae)

Pulmonaria mollissima (Boraginaceae)

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Melittis melissophyllum (Lamiaceae)

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Equatorial outline (in polar view, excluding apertural protrusions)


Equatorial outline (in polar view, excluding apertural protrusions)
circular

Bidens tripartitus (Asteraceae)


Dry pollen grains

Campanula glomerata (Campanulaceae)

Erica arborea (Ericaceae)


Tetrad (tetrahedral)

Geranium phaeum (Geraniaceae)

Juncus effusus (Juncaceae)


Tetrad (tetrahedral)

Lychnis viscaria (Caryophyllaceae)

Melittis melissophyllum (Lamiaceae)

Razisea citrina (Acanthaceae)


Dry pollen grain

Sesleria sadleriana (Poaceae)

Symphytum officinale (Boraginaceae)


Dry pollen grains

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Equatorial outline (in polar view, excluding apertural protrusions)


elliptic

Cabomba palaeformis (Cabombaceae)

Iris histrioides (Iridaceae)

Lamium maculatum (Lamiaceae)


Dry pollen grains

Leucojum vernum (Amaryllidaceae)

Physostegia virginiana (Lamiaceae)


Dry pollen grains

Salvia sclarea (Lamiaceae)

Gunnera chilensis (Gunneraceae)

Oxalis acetosella (Oxalidaceae)


Dry pollen grains

lobate

Artemisia pontica (Asteraceae)

Pinguicula ehlersiae (Lentibulariaceae)


Dry pollen grains

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Equatorial outline (in polar view, excluding apertural protrusions)


polygonal

Alnus glutinosa (Betulaceae)

Anthyllis vulneraria (Fabaceae)

Stellaria holostea (Caryophyllaceae)

Viola arvensis (Violaceae)

Platycodon grandiflorum (Campanulaceae)

quadrangular

Fumaria officinalis (Fumariaceae)

Lithospermum officinale (Boraginaceae)

Pulmonaria officinalis (Boraginaceae)

Viola tricolor (Violaceae)

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Equatorial outline (in polar view, excluding apertural protrusions)


triangular

Beloperone guttata (Acanthaceae)


Dry pollen grains

Callistemon coccineus (Myrtaceae)


Dry pollen grains

Epilobium dodonaei (Onagraceae)

Epilobium montanum (Onagraceae)


Tetrads (tetrahedral)

Prunus padus (Rosaceae)

Stachyurus praecox (Stachyuraceae)

Verbena officinalis (Verbenaceae)

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Dry pollen, infoldings


Dry pollen, infoldings
aperture(s) sunken
Characteristic shape of pollen grains caused by an infolding as a consequence of
harmomegathy.

Antennaria dioica (Asteraceae)

Carex digitata (Cyperaceae)


Dry tetrads (pseudomonads)

Daucus carota (Apiaceae)

Gagea graeca (Liliaceae)

Luzula sylvatica (Juncaceae)


Dry tetrads (tetrahedral)

Myosotis arvensis (Boraginaceae)

Saponaria caespitosa (Caryophyllaceae)

boat-shaped
Characteristic shape of sulcate pollen grains caused by an infolding as a
consequence of harmomegathy.

Asphodeline lutea (Liliaceae)

Convallaria majalis (Ruscaceae)

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Galanthus nivalis (Amaryllidaceae)

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Dry pollen, infoldings

Vriesea procera (Bromeliaceae)

cup-shaped
Characteristic shape of pollen grains caused by an infolding as a consequence of
harmomegathy.

Dracunculus vulgaris (Araceae)

Elaeagnus angustifolia (Elaeagnaceae)

Luzula campestris (Juncaceae)


Tetrads (tetrahedral)

Tsuga canadensis (Pinaceae)

Knautia drymeia (Dipsacaceae)

interapertural area(s) sunken


Characteristic shape of pollen grains caused by an infolding as a consequence of
harmomegathy.

Alnus glutinosa (Betulaceae)

Bromus erectus (Poaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Cornus sanguinea (Cornaceae)

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Dry pollen, infoldings

Corylus avellana (Betulaceae)

Melampyrum arvense (Orobanchaceae)

irregularly infolded
Characteristic shape of pollen grains caused by an infolding as a consequence of
harmomegathy.

Asarum europaeum (Aristolochiaceae)

Betula humilis (Betulaceae)

Dalechampia roezliana (Euphorbiaceae)

Ribes uva-crispa (Grossulariaceae)

Sanguisorba minor (Rosaceae)

Stylochaeton zenkeri (Araceae)

Cirsium canum (Asteraceae)

Onobrychis viciifolia (Fabaceae)

not infolded

Campanula rapunculoides (Campanulaceae)

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Dry pollen, infoldings

Pinellia ternata (Araceae)

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Aperture number
Aperture number
1

Allium flavum (Alliaceae)


sulcate pollen grain

Catopsis floribunda (Bromeliaceae)


sulcate pollen grain

Cephalanthera longifolia (Orchidaceae)


ulcerate pollen grain

Monstera deliciosa (Araceae)


sulcate pollen grain

Sesleria albicans (Poaceae)


ulcerate pollen grain

Vriesea oleosa (Bromeliaceae)


sulcate pollen grain

Colchicum autumnale (Colchicaceae)


porate pollen grain

Crinum augustum (Amaryllidaceae)


colpate pollen grain

Aechmea dealbata (Bromeliaceae)


porate pollen grain

Justicia xylosteoides (Acanthaceae)


colporate pollen grain

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Aperture number
3

Achillea millefolium (Asteraceae)


colporate pollen grain

Betula humilis (Betulaceae)


porate pollen grain

Circaea lutetiana (Onagraceae)


porate pollen grain

Hamamelis vernalis (Hamamelidaceae)


colpate pollen grain

Hedera helix (Araliaceae)


colporate pollen grain

Plumbago auriculata (Plumbaginaceae)


colpate pollen grain

Hohenbergia stellata (Bromeliaceae)


porate pollen grains

Impatiens glandulifera (Balsaminaceae)


colpate pollen grain

Scabiosa graminifolia (Dipsacaceae)


porate pollen grain

Alnus incana (Betulaceae)


colpate pollen grain

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Aperture number

Nicotiana tabacum (Solanaceae)


colporate pollen grain

Nonea pulla (Boraginaceae)


colporate pollen grain

Spilanthes acmella (Asteraceae)


colporate pollen grain

Legousia speculum-veneris (Campanulaceae)


porate pollen grain

Pulmonaria mollissima (Boraginaceae)


colporate pollen grain

Pinguicula ehlersiae (Lentibulariaceae)


colporate pollen grain

Primula veris (Primulaceae)


colpate pollen grain

Alnus viridis (Betulaceae)


porate pollen grain

Viola arvensis (Violaceae)


colporate pollen grain

Fumaria officinalis (Fumariaceae)


porate pollen grain

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Aperture number

Salvia coccinea (Lamiaceae)


colpate pollen grain

Sanguisorba officinalis (Rosaceae)


colporate pollen grain

>6
Pollen grain with more than 6 apertures.

Aechmea azurea (Bromeliaceae)


porate pollen grains

Agrostemma githago (Caryophyllaceae)


porate pollen grain

Alisma plantago-aquatica (Alismataceae)


porate pollen grain

Cobaea scandens (Polemoniaceae)


porate pollen grain

Daphne alpina (Thymelaeaceae)


porate pollen grain

Lychnis flos-cuculi (Caryophyllaceae)


porate pollen grain

Symphytum officinale (Boraginaceae)


colporate pollen grain

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Aperture number
inaperturate
Pollen grain without distinct apertures.

Aristolochia clematitis (Aristolochiaceae)

Dracunculus vulgaris (Araceae)

Pinellia ternata (Araceae)

Populus nigra (Salicaceae)

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Jatropha podagrica (Euphorbiaceae)

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Aperture peculiarities
Aperture peculiarities
annulate
Pollen grain with an annulus or annuli, a ring like thickening of the pollen wall
surrounding a porus or ulcus.
Comment: "anulus" orthographical variant of "annulus"

Bromus erectus (Poaceae)

Bromus erectus (Poaceae)

Circaea lutetiana (Onagraceae)

Circaea lutetiana (Onagraceae)

Cucumis sativus (Cucurbitaceae)

Cucumis sativus (Cucurbitaceae)

Fumaria officinalis (Fumariaceae)

Fumaria officinalis (Fumariaceae)

Myriophyllum spicatum (Haloragidaceae)

Myriophyllum spicatum (Haloragidaceae)

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Aperture peculiarities
colpate
Pollen grain with elongated apertures (length/width ratio > 2), situated at the
equator or regularly spread over the pollen grain.

Convolvulus tricolor (Convolvulaceae)

Convolvulus tricolor (Convolvulaceae)

Corylopsis platypetala (Hamamelidaceae)

Corylopsis platypetala (Hamamelidaceae)

Lunaria rediviva (Brassicaceae)

Lunaria rediviva (Brassicaceae)

Sherardia arvensis (Rubiaceae)

Sherardia arvensis (Rubiaceae)

Stachys annua (Lamiaceae)

Stachys annua (Lamiaceae)

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Aperture peculiarities
colporate
Pollen grain with compound apertures composed of a colpus (ektoaperture)
combined with an endoaperture of variable size and shape.

Beloperone guttata (Acanthaceae)

Carduus acanthoides (Asteraceae)

Daucus carota (Apiaceae)

Digitalis lutea (Veronicaceae)

Helminthotheca echioides (Asteraceae)

Moltkia petraea (Boraginaceae)

Pinguicula ehlersiae (Lentibulariaceae)

Polygala chamaebuxus (Polygalaceae)

Sanguisorba minor (Rosaceae)

leptoma
Thinning of the pollen wall at the distal pole (of a pollen grain) in conifers, presumed
to function as germination area; special case of tenuitas.

Abies cephalonica (Pinaceae)

Tsuga canadensis (Pinaceae)

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Aperture peculiarities
porate
Pollen grain with more or less circular aperture situated at the equator or regularly
spread over the pollen grain.

Aechmea azurea (Bromeliaceae)

Aechmea azurea (Bromeliaceae)

Cerastium glutinosum (Caryophyllaceae)

Cerastium glutinosum (Caryophyllaceae)

Colchicum autumnale (Colchicaceae)

Colchicum autumnale (Colchicaceae)

Epilobium dodonaei (Onagraceae)

Epilobium dodonaei (Onagraceae)

poroidate
Pollen grain with circular or elliptic aperture, with indistinct margin.

Carex distans (Cyperaceae)

Carex distans (Cyperaceae)

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Cercidiphyllum japonicum (Cercidiphyllaceae)

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Aperture peculiarities

Cyperus longus (Cyperaceae)

Guzmania glomerata (Bromeliaceae)

Guzmania glomerata (Bromeliaceae)

sulcate
Pollen grain with a sulcus, i.e. an elongated aperture situated distally.

Allium flavum (Alliaceae)

Catopsis floribunda (Bromeliaceae)

Pycnospatha palmata (Araceae)

Uvularia grandiflora (Liliaceae)


2-sulcate pollen grain

ulcerate
Pollen grain with an ulcus, i.e. more or less circular aperture situated distally.

Cephalanthera longifolia (Orchidaceae)

Luzula campestris (Juncaceae)

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Sesleria uliginosa (Poaceae)

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Aperture peculiarities

Sparganium erectum (Sparganiaceae)

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Aperture peculiarities
Aperture peculiarities
heteroaperturate
Pollen grain with two different types of apertures; only one type presumed to
function a germination site.

Justicia furcata (Acanthaceae)

Lythrum salicaria (Lythraceae)

Myosotis ramosissima (Boraginaceae)

operculate
Coherent exine structure covering an aperture.

Agrostemma githago (Caryophyllaceae)

Dionaea muscipula (Droseraceae)


Tetrad (tetrahedral)

Dionaea muscipula (Droseraceae)

Knautia drymeia (Dipsacaceae)

Lolium perenne (Poaceae)

Lolium perenne (Poaceae)

Lolium perenne (Poaceae)

Potentilla erecta (Rosaceae)

Potentilla erecta (Rosaceae)

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Aperture peculiarities

Veratrum album (Melanthiaceae)

ornamented aperture membrane


Exine elements (smaller than an operculum) covering the aperture surface, often
globular, verrucate or echinate.

Aesculus hippocastanum (Hippocastanaceae)


3-colporate pollen grain

Aesculus hippocastanum (Hippocastanaceae)

Asperula tinctoria (Rubiaceae)

Asperula tinctoria (Rubiaceae)


6-colpate pollen grain

Colchicum autumnale (Colchicaceae)

Colchicum autumnale (Colchicaceae)


2-porate pollen grain

Convolvulus tricolor (Convolvulaceae)


3-colpate pollen grain

Convolvulus tricolor (Convolvulaceae)

Galanthus nivalis (Amaryllidaceae)


1-sulcate pollen grain

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Aperture peculiarities

Galanthus nivalis (Amaryllidaceae)

Galeopsis tetrahit (Lamiaceae)


3-colpate pollen grain

Galeopsis tetrahit (Lamiaceae)


3-colpate pollen grain

Galeopsis tetrahit (Lamiaceae)

Iris histrioides (Iridaceae)


1-sulcate pollen grain

Iris histrioides (Iridaceae)

Onobrychis viciifolia (Fabaceae)

Onobrychis viciifolia (Fabaceae)


3-colpate pollen grain

Viscum album (Viscaceae)


3-colpate pollen grain

Viscum album (Viscaceae)

Werauhia tarmaensis (Bromeliaceae)


1-sulcate pollen grain

Werauhia tarmaensis (Bromeliaceae)

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Aperture peculiarities
pantoaperturate
Pollen grain with apertures distributed more or less regularly over the whole
surface.

Hepatica transsylvanica (Ranunculaceae)

Holosteum umbellatum (Caryophyllaceae)

Lavatera thuringiaca (Malvaceae)

Opuntia paraguayensis (Cactaceae)

Polemonium caeruleum (Polemoniaceae)

Ribes sanguineum (Grossulariaceae)

Sideritis syriaca (Lamiaceae)


dry pollen grains pantoaperturate

smooth aperture membrane


Aperture surface without any exine elements.

Aechmea caesia (Bromeliaceae)

Cyrtosperma beccarianum (Araceae)

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Digitalis purpurea (Veronicaceae)

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Aperture peculiarities

Melampyrum arvense (Orobanchaceae)

Trifolium montanum (Fabaceae)

stephanoaperturate
Pollen grain with apertures situated at the equator (term usually used for more than
three apertures).

Anemone hortensis (Ranunculaceae)

Borago officinalis (Boraginaceae)

Dionaea muscipula (Droseraceae)


Tetrad (tetrahedral)

Nymphoides peltata (Menyanthaceae)

Pedicularis verticillata (Orobanchaceae)

Sherardia arvensis (Rubiaceae)

syncolpate
Pollen grain with anastomosing colpi.

Hydrophyllum canadense (Hydrophyllaceae)

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Aperture peculiarities

Primula farinosa (Primulaceae)

syncolporate
Pollen grain with anastomosing colpori.

Callistemon coccineus (Myrtaceae)

Cassia pulcherrima (Caesalpiniaceae)

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Cuphea purpurea (Lythraceae)

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Ornamentation (LM and EM)


Ornamentation (LM and EM)
baculate
Pollen grain with bacula, rod-like, free standing element, more than 1 m in height
and never pointed.

Erythrochiton brasiliensis (Rutaceae)

Nymphaea alba (Nymphaeaceae)

Viscum laxum (Viscaceae)

clavate
Pollen wall with clavae, i.e. club-shaped element, higher than 1 m.

Geranium phaeum (Geraniaceae)

Ilex aquifolium (Aquifoliaceae)

Linum catharticum (Linaceae)

Plumbago auriculata (Plumbaginaceae)

Ilex aquifolium (Aquifoliaceae)

echinate
Pollen wall with echini, i.e. pointed ornamentation elements longer and/or wider
than 1 m.

Anthemis arvensis (Asteraceae)

Centaurea jacea (Asteraceae)

PalDat - Illustrated Handbook on Pollen Terminology

Hibiscus trionum (Malvaceae)

Page 36 of 62

Ornamentation (LM and EM)

Malva neglecta (Malvaceae)

Pinellia ternata (Araceae)

Ulearum sagittatum var. sagittatum (Araceae)

Zomicarpa riedeliana (Araceae)

Tussilago farfara (Asteraceae)

fossulate
Pollen wall with fossulae, i.e. irregularly shaped grooves in the surface of a pollen
wall.

Erica herbacea (Ericaceae)

Hydrophyllum canadense (Hydrophyllaceae)

Orthilia secunda (Ericaceae)

Rhododendron hirsutum (Ericaceae)

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Ornamentation (LM and EM)


foveolate
Pollen wall with foveolae, i.e. roundish lumina more than 1 m in diameter; distance
between two adjacent lumina larger than their diameter.

Lavandula angustifolia (Lamiaceae)

Streptocalyx poeppigii (Bromeliaceae)

Stylochaeton bogneri (Araceae)

frustrate
Special mental condition of palynologists discussing terminology of pollen and
spores.

Frustrelia mentalis (Frustraceae)

gemmate
Pollen wall with gemmae, i.e. globular exine elements more than 1 m in diameter.

Asarum europaeum (Aristolochiaceae)

Dionaea muscipula (Droseraceae)

PalDat - Illustrated Handbook on Pollen Terminology

Stenandrium dulce (Acanthaceae)

Page 38 of 62

Ornamentation (LM and EM)


granulate
Pollen wall with granula, i.e. structure- or sculpture elements of different size and
shape; smaller than 1 m.
Comment: Not to be confused with "granular", which is a feature of the pollen wall
structure.

Symphytum orientale (Boraginaceae)

Ulmus glabra (Ulmaceae)

microechinate
Pollen wall with echini, i.e. pointed ornamentation elements smaller than 1 m.

Artemisia pontica (Asteraceae)

Campanula patula (Campanulaceae)

Valerianella carinata (Valerianaceae)

microreticulate
Pollen wall with network like pattern formed by exine elements (muri), where the
lumina are smaller than 1 m.

Digitalis lutea (Veronicaceae)

Digitalis purpurea (Veronicaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Stachys sylvatica (Lamiaceae)

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Ornamentation (LM and EM)


perforate
Pollen wall with holes less than 1m in diameter.

Chimonanthus praecox (Calycanthaceae)

Gonatopus angustus (Araceae)

Napoleonaea imperialis (Lecythidaceae)

Dorycnium germanicum (Fabaceae)

Dorycnium germanicum (Fabaceae)

Hedychium gardnerianum (Commelinaceae)

Hippocrepis emerus (Fabaceae)

Hippocrepis emerus (Fabaceae)

Vicia tenuifolia (Fabaceae)

Pulmonaria officinalis (Boraginaceae)

psilate
Pollen wall with smooth surface.

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Ornamentation (LM and EM)

Vicia tenuifolia (Fabaceae)

Vinca minor (Apocynaceae)

reticulate
Pollen wall with network like pattern formed by exine elements (muri), where the
lumina are wider than 1 m.

Ajuga genevensis (Lamiaceae)

Beloperone guttata (Acanthaceae)

Corylopsis glabrescens (Hamamelidaceae)

Gagea graeca (Liliaceae)

Galeopsis tetrahit (Lamiaceae)

Hyssopus officinalis (Lamiaceae)

Impatiens glandulifera (Balsaminaceae)

Ruellia graecicans (Acanthaceae)

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Ornamentation (LM and EM)


reticulate-heterobrochate
Reticulate pollen wall with lumina of different sizes.
Comment: the term should be used when the feature is remarkably expressed.

Anthericum ramosum (Anthericaceae)

Billbergia seidelii (Bromeliaceae)

Forsythia x intermedia (Oleaceae)

Hedera helix (Araliaceae)

Lachenalia aloides (Liliaceae)

Pitcairnia odontopoda (Bromeliaceae)

reticulate-homobrochate
Reticulate pollen wall with lumina of uniform size.
Comment: the term should be used when the feature is remarkably expressed.

Acantholimon glumaceum (Plumbaginaceae)

Hamamelis vernalis (Hamamelidaceae)

reticulo-cristate
Pollen grain with a special type of reticulum; muri with prominent sculpture
elements.

Daphne alpina (Thymelaeaceae)

Fumana procumbens (Cistaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Garcia nutans (Euphorbiaceae)

Page 42 of 62

Ornamentation (LM and EM)

Geranium canariense (Geraniaceae)

Lilium martagon (Liliaceae)

Pachysandra terminalis (Buxaceae)

Phaleria capitata (Thymelaeaceae)

Phaleria capitata (Thymelaeaceae)

Thymelaea passerina (Thymelaeaceae)

rugulate
Pollen grain with elongated exine elements longer than 1 m; irregularly arranged.

Angelica archangelica (Apiaceae)

Angelica archangelica (Apiaceae)

Cabomba palaeformis (Cabombaceae)

Cabomba palaeformis (Cabombaceae)

Nicotiana tabacum (Solanaceae)

Nicotiana tabacum (Solanaceae)

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Ornamentation (LM and EM)

Nymphoides peltata (Menyanthaceae)

Nymphoides peltata (Menyanthaceae)

Ulmus laevis (Ulmaceae)

Ulmus laevis (Ulmaceae)

striate
Pollen wall with elongated exine elements separated by grooves predominantly
parallel arranged.

Acer obtusatum (Sapindaceae)

Acer obtusatum (Sapindaceae)

Helianthemum nummularium (Cistaceae)

Helianthemum nummularium (Cistaceae)

Potentilla inclinata (Rosaceae)

Potentilla inclinata (Rosaceae)

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Ornamentation (LM and EM)

Rubus caesius (Rosaceae)

Rubus caesius (Rosaceae)

Sanguisorba minor (Rosaceae)

Sanguisorba minor (Rosaceae)

Saxifraga vandellii (Saxifragaceae)

Saxifraga vandellii (Saxifragaceae)

striato-reticulate
Ornamentation intermediate between striate and reticulate.

Aesculus glabra (Hippocastanaceae)

Aesculus glabra (Hippocastanaceae)

Aesculus flava (Hippocastanaceae)

Aesculus flava (Hippocastanaceae)

Erodium cicutarium (Geraniaceae)

Erodium cicutarium (Geraniaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 45 of 62

Ornamentation (LM and EM)

Gentiana acaulis (Gentianaceae)

Gentiana acaulis (Gentianaceae)

Gentianella austriaca (Gentianaceae)

Gentianella germanica (Gentianaceae)

Pelargonium carnosum (Geraniaceae)

Solandra longiflora (Solanaceae)

Solandra longiflora (Solanaceae)

verrucate
Pollen wall with wart-like elements more than 1m, broader than high.

Plantago major (Plantaginaceae)

Plantago major (Plantaginaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Teucrium pyrenaicum (Lamiaceae)

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Ornamentation (LM and EM)

Teucrium pyrenaicum (Lamiaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 47 of 62

Tectum
Tectum
atectate
Pollen grain lacking a tectum.

Cucurbita pepo (Cucurbitaceae)

Cucurbita pepo (Cucurbitaceae)

Rhaphidophora africana (Araceae)

Rhaphidophora africana (Araceae)

Cucurbita pepo (Cucurbitaceae)

eutectate
Pollen grain with a continuous tectum.

Actaea spicata (Ranunculaceae)

Actaea spicata (Ranunculaceae)

Papaver rhoeas (Papaveraceae)

Papaver rhoeas (Papaveraceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 48 of 62

Tectum
semitectate
Pollen grain with a discontinuous tectum, covering less than 50% of pollen grain
surface.

Muscari azureum (Hyacinthaceae)

Muscari azureum (Hyacinthaceae)

Ornithogalum kochii (Hyacinthaceae)

Ornithogalum kochii (Hyacinthaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 49 of 62

Endexine
Endexine
missing
Endexine missinig.

Tropaeolum majus (Tropaeolaceae)

continuous
Endexine forming a compact layer.

Galium odoratum (Rubiaceae)

discontinuous
Endexine interrupted.

Glaucium flavum (Papaveraceae)

lamellar

Aristolochia arborea (Aristolochiaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 50 of 62

Endexine
spongy

Dracunculus vulgaris (Araceae)

Nelumbo nucifera (Nelumbonaceae)

Ulearum sagittatum var. sagittatum (Araceae)

Zamioculcas zamiifolia (Araceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 51 of 62

Foot layer
Foot layer
missing
Foot layer missing.

Aristolochia arborea (Aristolochiaceae)

Arophyton buchetii (Araceae)

Pachypodium succulentum (Apocynaceae)

continuous
Foot layer forming a compact layer.

Anaphyllopsis americana (Araceae)

Helwingia japonica (Cornaceae)

discontinuous
Foot layer interrupted.

Anthericum liliago (Anthericaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 52 of 62

Foot layer
perforate
Foot layer penetrated by thin channels.

Geranium robertianum (Geraniaceae)

Tulipa urumiensis (Liliaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 53 of 62

Infratectum
Infratectum
absent

Mahonia aquifolium (Berberidaceae)

columellate
Infratectum with rod-like elements.

Apium nodiflorum (Apiaceae)

Buddleja alternifolia (Buddlejaceae)

Ophiorrhiza longiflora (Rubiaceae)

Podolasia stipitata (Araceae)

granular
Infratectum composed of granula, cluster of granula or elements of different size
and shape (never solid and rod-like).

Alnus glutinosa (Betulaceae)

Ulmus laevis (Ulmaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 54 of 62

Pollen wall
Pollen wall
alveolate
Infratectum with compartments of irregular size and shape.

Zamioculcas zamiifolia (Araceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 55 of 62

Wall peculiarities
Wall peculiarities
internal tectum
A continuous layer inbetween and parallel to foot layer and tectum, separated
from them by columellae.

Nigella arvensis (Ranunculaceae)

papilla
Small protuberance typical for Taxodioideae-pollen (Cupressaceae).

Cryptomeria japonica (Cupressaceae)

Cryptomeria japonica (Cupressaceae)

viscin threads
Pollen grain with a circumferential aperture (situated more or less equatorially or,
rarely, meridionally).

Circaea lutetiana (Onagraceae)

Circaea lutetiana (Onagraceae)

Epilobium angustifolium (Onagraceae)

Epilobium angustifolium (Onagraceae)

Godetia purpurea (Onagraceae)

Godetia purpurea (Onagraceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 56 of 62

Wall peculiarities

Kalmia latifolia (Ericaceae)

Kalmia latifolia (Ericaceae)

Rhododendron hirsutum (Ericaceae)

Rhododendron hirsutum (Ericaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 57 of 62

Cellular condition
Cellular condition
3
Vegetative cytoplasm inclosing two sperm cells.

Galium odoratum (Rubiaceae)

Jasminum nudiflorum (Oleaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Page 58 of 62

Picture Credits

Picture Credits
Aktuna G.
Anthericum liliago, Anthericum ramosum, Convallaria majalis, Leucojum vernum, Muscari azureum, Muscari tenuiflorum, Ornithogalum
kochii

Bombosi P.
Symphytum officinale, Viola tricolor

Halbritter H.
Abeliophyllum distichum, Abies cephalonica, Acacia myrtifolia, Acantholimon glumaceum, Acer obtusatum, Achillea millefolium, Aechmea
azurea, Aechmea caesia, Aechmea dealbata, Aechmea fraseri, Aesculus flava, Aesculus glabra, Aesculus hippocastanum, Agrostemma
githago, Ajuga genevensis, Alisma plantago-aquatica, Allium flavum, Alnus glutinosa, Alnus incana, Alnus viridis, Ambrosina bassii,
Amorphophallus asterostigmatus, Anemone hortensis, Angelica archangelica, Antennaria dioica, Anthemis arvensis, Anthurium radicans,
Anthyllis vulneraria, Aristolochia arborea, Aristolochia clematitis, Artemisia pontica, Asarum europaeum, Asperula tinctoria, Asphodeline
lutea, Begonia nelumbifolia, Bellis perennis, Beloperone guttata, Betula humilis, Bidens tripartitus, Billbergia seidelii, Borago officinalis,
Bromus erectus, Buglossoides purpurocaerulea, Cabomba palaeformis, Callistemon coccineus, Campanula glomerata, Campanula patula,
Campanula rapunculoides, Carduus acanthoides, Carex alba, Carex digitata, Carex distans, Carex humilis, Carex remota, Cassia
pulcherrima, Catalpa ovata, Catopsis floribunda, Centaurea jacea, Cephalanthera longifolia, Cerastium glutinosum, Cercidiphyllum
japonicum, Chimonanthus praecox, Cichorium intybus, Circaea lutetiana, Cirsium canum, Cobaea scandens, Colchicum autumnale,
Convolvulus tricolor, Cornus sanguinea, Corylopsis glabrescens, Corylopsis platypetala, Corylus avellana, Crinum augustum, Crocus
speciosus, Crossandra flava, Cryptomeria japonica, Cucumis sativus, Cuphea purpurea, Cyperus longus, Cyrtosperma beccarianum,
Dalechampia roezliana, Daphne alpina, Daucus carota, Digitalis lutea, Digitalis purpurea, Dionaea muscipula, Dorycnium germanicum,
Dracunculus vulgaris, Elaeagnus angustifolia, Epilobium angustifolium, Epilobium dodonaei, Epilobium montanum, Epilobium parviflorum,
Erica arborea, Erica herbacea, Erodium cicutarium, Erythrochiton brasiliensis, Forsythia x intermedia, Frustrelia mentalis, Fumana
procumbens, Gagea graeca, Galanthus nivalis, Garcia nutans, Gentiana acaulis, Gentianella austriaca, Gentianella germanica, Geranium
canariense, Geranium phaeum, Godetia purpurea, Gonatopus angustus, Gunnera chilensis, Guzmania glomerata, Hamamelis vernalis,
Hedera helix, Hedychium gardnerianum, Helianthemum nummularium, Helminthotheca echioides, Hemigraphis primulaefolia, Hepatica
transsylvanica, Hibiscus trionum, Himantoglossum adriaticum, Hippocrepis emerus, Hohenbergia proctori, Hohenbergia stellata,
Holosteum umbellatum, Hordeum murinum, Hydrophyllum canadense, Ibicella lutea, Ilex aquifolium, Impatiens glandulifera, Iris bucharica,
Iris histrioides, Jatropha podagrica, Juncus effusus, Justicia furcata, Justicia xylosteoides, Kalmia latifolia, Knautia drymeia, Lachenalia
aloides, Lamium maculatum, Laportea divaricata, Lavandula angustifolia, Lavatera thuringiaca, Legousia speculum-veneris, Lilium
martagon, Limnanthes douglasii, Linum catharticum, Lithospermum officinale, Lolium perenne, Lotus corniculatus, Lunaria rediviva, Luzula
campestris, Luzula sylvatica, Lychnis flos-cuculi, Lychnis viscaria, Lythrum salicaria, Malva neglecta, Melampyrum arvense, Melittis
melissophyllum, Microrrhinum minus, Mimosa pudica, Mimulus guttatus, Moltkia petraea, Monstera deliciosa, Myosotis arvensis, Myosotis
ramosissima, Myriophyllum spicatum, Nandina domestica, Napoleonaea imperialis, Nepeta cataria, Nicotiana tabacum, Nonea pulla,
Nymphaea alba, Nymphoides peltata, Onobrychis viciifolia, Onosma arenaria, Opuntia paraguayensis, Orthilia secunda, Oxalis acetosella,
Pachysandra terminalis, Pedicularis verticillata, Pelargonium carnosum, Peperomia rubella, Phaleria capitata, Phyllanthus spec,
Physostegia virginiana, Phyteuma spicatum, Pinellia ternata, Pinguicula ehlersiae, Pinus mugo, Pistia stratiotes, Pitcairnia odontopoda,
Plantago major, Platanthera bifolia, Platycodon grandiflorum, Plumbago auriculata, Polygala chamaebuxus, Populus nigra, Potentilla
erecta, Potentilla inclinata, Primula farinosa, Primula veris, Prunus padus, Pulmonaria mollissima, Pulmonaria officinalis, Pycnospatha
palmata, Razisea citrina, Rhaphidophora africana, Rhododendron hirsutum, Ribes sanguineum, Ribes uva-crispa, Rubus caesius, Ruellia
graecicans, Salvia coccinea, Salvia involucrata, Salvia sclarea, Sanguisorba minor, Sanguisorba officinalis, Saponaria caespitosa,
Saxifraga vandellii, Scabiosa graminifolia, Schismatoglottis lancifolia, Scorzonera hispanica, Sesleria albicans, Sesleria sadleriana,
Sesleria uliginosa, Sherardia arvensis, Sideritis syriaca, Solandra longiflora, Sparganium erectum, Spilanthes acmella, Stachys annua,
Stachys sylvatica, Stachyurus praecox, Stellaria holostea, Stenandrium dulce, Steudnera assamica, Streptocalyx poeppigii, Stylochaeton
bogneri, Stylochaeton zenkeri, Symphytum orientale, Teucrium pyrenaicum, Thesium linophyllon, Thlaspi arvense, Thunbergia alata,
Thymelaea passerina, Tilia cordata, Trifolium montanum, Tsuga canadensis, Tussilago farfara, Typha latifolia, Ulmus glabra, Ulmus laevis,
Uvularia grandiflora, Valerianella carinata, Veratrum album, Verbena officinalis, Vicia tenuifolia, Vinca minor, Viola arvensis, Viscum
album, Viscum laxum, Vriesea oleosa, Vriesea procera, Werauhia tarmaensis, Zomicarpa riedeliana

Hesse M.
Cucurbita pepo

Oberschneider W.
Actaea spicata, Berberis vulgaris, Fumaria officinalis, Glaucium flavum, Mahonia aquifolium, Nigella arvensis, Papaver rhoeas

Svojtka M.
Polemonium caeruleum

Ulrich S.
Galeopsis tetrahit, Hyssopus officinalis

Weber M.
Anaphyllopsis americana, Apium nodiflorum, Arophyton buchetii, Buddleja alternifolia, Galium odoratum, Geranium robertianum, Helwingia
japonica, Jasminum nudiflorum, Nelumbo nucifera, Ophiorrhiza longiflora, Pachypodium succulentum, Podolasia stipitata, Tropaeolum
majus, Tulipa urumiensis, Ulearum sagittatum var. sagittatum, Zamioculcas zamiifolia

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Page 59 of 62

Contents

Contents
Pollen unit .................................................................................................................................... 4
polyad .................................................................................................................................. 4
tetrad ................................................................................................................................... 4
Pollen class ..................................................................................................................................
clypeate ...............................................................................................................................
lophate ................................................................................................................................
plicate ..................................................................................................................................
saccate ................................................................................................................................
spiraperturate ......................................................................................................................

5
5
5
5
6
6

Shape (single pollen grains) ......................................................................................................


irregular ...............................................................................................................................
oblate ..................................................................................................................................
prolate .................................................................................................................................
spheroidal ............................................................................................................................

7
7
7
8
9

Shape peculiarities .................................................................................................................... 10


heteropolar ........................................................................................................................ 10
isopolar .............................................................................................................................. 10
Equatorial outline (in polar view, excluding apertural protrusions) .....................................
circular ...............................................................................................................................
elliptic ................................................................................................................................
lobate ................................................................................................................................
polygonal ...........................................................................................................................
quadrangular .....................................................................................................................
triangular ...........................................................................................................................

11
11
12
12
13
13
14

Dry pollen, infoldings ................................................................................................................


aperture(s) sunken ............................................................................................................
boat-shaped ......................................................................................................................
cup-shaped .......................................................................................................................
interapertural area(s) sunken ............................................................................................
irregularly infolded .............................................................................................................
not infolded ........................................................................................................................

15
15
15
16
16
17
17

Aperture number .......................................................................................................................


1 ........................................................................................................................................
2 ........................................................................................................................................
3 ........................................................................................................................................
4 ........................................................................................................................................
5 ........................................................................................................................................
6 ........................................................................................................................................
>6 ......................................................................................................................................
inaperturate .......................................................................................................................

19
19
19
20
20
21
21
22
23

Aperture peculiarities ...............................................................................................................


annulate ............................................................................................................................
colpate ...............................................................................................................................
colporate ...........................................................................................................................
leptoma .............................................................................................................................
porate ................................................................................................................................
poroidate ...........................................................................................................................
sulcate ...............................................................................................................................
ulcerate .............................................................................................................................

24
24
25
26
26
27
27
28
28

Aperture peculiarities ...............................................................................................................


heteroaperturate ................................................................................................................
operculate .........................................................................................................................
ornamented aperture membrane ......................................................................................
pantoaperturate .................................................................................................................
smooth aperture membrane ..............................................................................................
stephanoaperturate ...........................................................................................................
syncolpate .........................................................................................................................
syncolporate ......................................................................................................................

30
30
30
31
33
33
34
34
35

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Contents
Ornamentation (LM and EM) ....................................................................................................
baculate .............................................................................................................................
clavate ...............................................................................................................................
echinate .............................................................................................................................
fossulate ............................................................................................................................
foveolate ............................................................................................................................
frustrate .............................................................................................................................
gemmate ...........................................................................................................................
granulate ...........................................................................................................................
microechinate ....................................................................................................................
microreticulate ...................................................................................................................
perforate ............................................................................................................................
psilate ................................................................................................................................
reticulate ............................................................................................................................
reticulate-heterobrochate ..................................................................................................
reticulate-homobrochate ...................................................................................................
reticulo-cristate ..................................................................................................................
rugulate .............................................................................................................................
striate ................................................................................................................................
striato-reticulate .................................................................................................................
verrucate ...........................................................................................................................

36
36
36
36
37
38
38
38
39
39
39
40
40
41
42
42
42
43
44
45
46

Tectum ........................................................................................................................................
atectate .............................................................................................................................
eutectate ...........................................................................................................................
semitectate ........................................................................................................................

48
48
48
49

Endexine ....................................................................................................................................
missing ..............................................................................................................................
continuous .........................................................................................................................
discontinuous ....................................................................................................................
lamellar ..............................................................................................................................
spongy ...............................................................................................................................

50
50
50
50
50
51

Foot layer ...................................................................................................................................


missing ..............................................................................................................................
continuous .........................................................................................................................
discontinuous ....................................................................................................................
perforate ............................................................................................................................

52
52
52
52
53

Infratectum .................................................................................................................................
absent ...............................................................................................................................
columellate ........................................................................................................................
granular .............................................................................................................................

54
54
54
54

Pollen wall .................................................................................................................................. 55


alveolate ............................................................................................................................ 55
Wall peculiarities .......................................................................................................................
internal tectum ...................................................................................................................
papilla ................................................................................................................................
viscin threads ....................................................................................................................

56
56
56
56

Cellular condition ...................................................................................................................... 58


3 ........................................................................................................................................ 58

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Register

Register
1 ...................................................................... 19
2 ...................................................................... 19
3 ................................................................ 20, 58
4 ...................................................................... 20
5 ...................................................................... 21
6 ...................................................................... 21
>6 .................................................................... 22
absent ............................................................. 54
alveolate .......................................................... 55
annulate .......................................................... 24
aperture(s) sunken .......................................... 15
atectate ........................................................... 48
baculate .......................................................... 36
boat-shaped .................................................... 15
circular ............................................................ 11
clavate ............................................................. 36
clypeate ............................................................. 5
colpate ............................................................ 25
colporate ......................................................... 26
columellate ...................................................... 54
continuous ................................................. 50, 52
cup-shaped ..................................................... 16
discontinuous ............................................ 50, 52
echinate .......................................................... 36
elliptic .............................................................. 12
eutectate ......................................................... 48
fossulate .......................................................... 37
foveolate ......................................................... 38
frustrate ........................................................... 38
gemmate ......................................................... 38
granular ........................................................... 54
granulate ......................................................... 39
heteroaperturate ............................................. 30
heteropolar ...................................................... 10
inaperturate ..................................................... 23
interapertural area(s) sunken .......................... 16
internal tectum ................................................ 56
irregular ............................................................. 7
irregularly infolded ........................................... 17
isopolar ........................................................... 10
lamellar ........................................................... 50
leptoma ........................................................... 26
lobate .............................................................. 12
lophate .............................................................. 5
microechinate .................................................. 39
microreticulate ................................................. 39
missing ...................................................... 50, 52
not infolded ..................................................... 17
oblate ................................................................ 7
operculate ....................................................... 30
ornamented aperture membrane .................... 31
pantoaperturate ............................................... 33
papilla .............................................................. 56
perforate .................................................... 40, 53
plicate ................................................................ 5
polyad ............................................................... 4
polygonal ......................................................... 13
porate .............................................................. 27
poroidate ......................................................... 27
prolate ............................................................... 8
psilate .............................................................. 40
quadrangular ................................................... 13
reticulate ......................................................... 41
PalDat - Illustrated Handbook on Pollen Terminology

reticulate-heterobrochate ................................ 42
reticulate-homobrochate ................................. 42
reticulo-cristate ................................................ 42
rugulate ........................................................... 43
saccate .............................................................. 6
semitectate ...................................................... 49
smooth aperture membrane ............................ 33
spheroidal ......................................................... 9
spiraperturate .................................................... 6
spongy ............................................................ 51
stephanoaperturate ......................................... 34
striate .............................................................. 44
striato-reticulate .............................................. 45
sulcate ............................................................. 28
syncolpate ....................................................... 34
syncolporate .................................................... 35
tetrad ................................................................. 4
triangular ......................................................... 14
ulcerate ........................................................... 28
verrucate ......................................................... 46
viscin threads .................................................. 56

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