Paldat Terminology

Illustrated Handbook
on
Pollen Terminology
We encourage you to redistribute the "PalDat - Illustrated Handbook on Pollen Terminology" either in printed or
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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)

Erica herbacea (Ericaceae)

Mimosa pudica (Mimosaceae)
Typha latifolia (Typhaceae)

Tetrad (planar)
tetrad
Dispersal unit of four pollen grains (spores).
Typha latifolia (Typhaceae)

Tetrad (planar)
PalDat - Illustrated Handbook on Pollen Terminology
Page 4 of 62
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)
Hemigraphis primulaefolia (Acanthaceae)
Page 5 of 62
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)
Page 6 of 62
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)
Page 7 of 62
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)
Page 8 of 62

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)
Page 9 of 62
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)
Melittis melissophyllum (Lamiaceae)
Page 10 of 62
Equatorial outline (in polar view, excluding apertural protrusions)

circular
Bidens tripartitus (Asteraceae)

Dry pollen grains
Campanula glomerata (Campanulaceae)
Erica arborea (Ericaceae)

Geranium phaeum (Geraniaceae)
Juncus effusus (Juncaceae)

Lychnis viscaria (Caryophyllaceae)
Melittis melissophyllum (Lamiaceae)
Razisea citrina (Acanthaceae)

Dry pollen grain
Sesleria sadleriana (Poaceae)
Symphytum officinale (Boraginaceae)

Dry pollen grains
Page 11 of 62

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
Page 12 of 62

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)
Page 13 of 62

triangular

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)
Page 14 of 62
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)
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)
Galanthus nivalis (Amaryllidaceae)
Page 15 of 62
Vriesea procera (Bromeliaceae)
cup-shaped
harmomegathy.
Elaeagnus angustifolia (Elaeagnaceae)
Luzula campestris (Juncaceae)

Tetrads (tetrahedral)
interapertural area(s) sunken

harmomegathy.
Bromus erectus (Poaceae)
Cornus sanguinea (Cornaceae)
Page 16 of 62
Corylus avellana (Betulaceae)
Melampyrum arvense (Orobanchaceae)
irregularly infolded
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)
Page 17 of 62
Pinellia ternata (Araceae)
Page 18 of 62
Aperture number
Aperture number
1
Allium flavum (Alliaceae)

sulcate pollen grain
Catopsis floribunda (Bromeliaceae)

Cephalanthera longifolia (Orchidaceae)

ulcerate pollen grain
Monstera deliciosa (Araceae)

Sesleria albicans (Poaceae)

ulcerate pollen grain
Vriesea oleosa (Bromeliaceae)

Colchicum autumnale (Colchicaceae)

porate pollen grain
Crinum augustum (Amaryllidaceae)

colpate pollen grain
Aechmea dealbata (Bromeliaceae)

porate pollen grain
Justicia xylosteoides (Acanthaceae)

colporate pollen grain
Page 19 of 62
Aperture number
3
Achillea millefolium (Asteraceae)

Betula humilis (Betulaceae)

porate pollen grain
Circaea lutetiana (Onagraceae)

porate pollen grain
Hamamelis vernalis (Hamamelidaceae)

Hedera helix (Araliaceae)

Plumbago auriculata (Plumbaginaceae)

Hohenbergia stellata (Bromeliaceae)

porate pollen grains
Impatiens glandulifera (Balsaminaceae)

Scabiosa graminifolia (Dipsacaceae)

porate pollen grain
Alnus incana (Betulaceae)

Page 20 of 62
Aperture number
Nicotiana tabacum (Solanaceae)

Nonea pulla (Boraginaceae)

Spilanthes acmella (Asteraceae)

Legousia speculum-veneris (Campanulaceae)

porate pollen grain
Pulmonaria mollissima (Boraginaceae)


Primula veris (Primulaceae)

Alnus viridis (Betulaceae)

porate pollen grain
Viola arvensis (Violaceae)


porate pollen grain
Page 21 of 62
Aperture number
Salvia coccinea (Lamiaceae)

Sanguisorba officinalis (Rosaceae)

>6
Pollen grain with more than 6 apertures.

porate pollen grains
Agrostemma githago (Caryophyllaceae)

porate pollen grain
Alisma plantago-aquatica (Alismataceae)

porate pollen grain
Cobaea scandens (Polemoniaceae)

porate pollen grain

porate pollen grain
Lychnis flos-cuculi (Caryophyllaceae)

porate pollen grain
Symphytum officinale (Boraginaceae)

Page 22 of 62
Aperture number
inaperturate
Pollen grain without distinct apertures.
Aristolochia clematitis (Aristolochiaceae)
Populus nigra (Salicaceae)
Jatropha podagrica (Euphorbiaceae)
Page 23 of 62
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"
Cucumis sativus (Cucurbitaceae)
Cucumis sativus (Cucurbitaceae)
Myriophyllum spicatum (Haloragidaceae)
Myriophyllum spicatum (Haloragidaceae)
Page 24 of 62
colpate
Pollen grain with elongated apertures (length/width ratio > 2), situated at the
equator or regularly spread over the pollen grain.
Convolvulus tricolor (Convolvulaceae)
Corylopsis platypetala (Hamamelidaceae)
Corylopsis platypetala (Hamamelidaceae)
Lunaria rediviva (Brassicaceae)
Lunaria rediviva (Brassicaceae)
Sherardia arvensis (Rubiaceae)
Stachys annua (Lamiaceae)
Stachys annua (Lamiaceae)
Page 25 of 62
colporate
Pollen grain with compound apertures composed of a colpus (ektoaperture)
combined with an endoaperture of variable size and shape.
Carduus acanthoides (Asteraceae)
Digitalis lutea (Veronicaceae)
Helminthotheca echioides (Asteraceae)
Moltkia petraea (Boraginaceae)
Polygala chamaebuxus (Polygalaceae)
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)
Page 26 of 62
porate
Pollen grain with more or less circular aperture situated at the equator or regularly
spread over the pollen grain.
Cerastium glutinosum (Caryophyllaceae)
Cerastium glutinosum (Caryophyllaceae)
poroidate
Pollen grain with circular or elliptic aperture, with indistinct margin.
Carex distans (Cyperaceae)
Carex distans (Cyperaceae)
Cercidiphyllum japonicum (Cercidiphyllaceae)
Page 27 of 62
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)
Sesleria uliginosa (Poaceae)
Page 28 of 62
Sparganium erectum (Sparganiaceae)
Page 29 of 62
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)

Lolium perenne (Poaceae)
Potentilla erecta (Rosaceae)
Potentilla erecta (Rosaceae)
Page 30 of 62
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

2-porate pollen grain


Page 31 of 62
Galeopsis tetrahit (Lamiaceae)




Viscum album (Viscaceae)

Viscum album (Viscaceae)
Werauhia tarmaensis (Bromeliaceae)

Werauhia tarmaensis (Bromeliaceae)
Page 32 of 62
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)
Digitalis purpurea (Veronicaceae)
Page 33 of 62
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)

Nymphoides peltata (Menyanthaceae)
Pedicularis verticillata (Orobanchaceae)
syncolpate
Pollen grain with anastomosing colpi.
Hydrophyllum canadense (Hydrophyllaceae)
Page 34 of 62
Primula farinosa (Primulaceae)
syncolporate
Pollen grain with anastomosing colpori.
Callistemon coccineus (Myrtaceae)
Cassia pulcherrima (Caesalpiniaceae)
Cuphea purpurea (Lythraceae)
Page 35 of 62
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)
Hibiscus trionum (Malvaceae)
Page 36 of 62
Malva neglecta (Malvaceae)
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)
Page 37 of 62

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)
Stenandrium dulce (Acanthaceae)
Page 38 of 62

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)
Stachys sylvatica (Lamiaceae)
Page 39 of 62

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.
Page 40 of 62
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)
Corylopsis glabrescens (Hamamelidaceae)
Gagea graeca (Liliaceae)
Hyssopus officinalis (Lamiaceae)
Impatiens glandulifera (Balsaminaceae)
Ruellia graecicans (Acanthaceae)
Page 41 of 62

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.
Fumana procumbens (Cistaceae)
Garcia nutans (Euphorbiaceae)
Page 42 of 62
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)
Page 43 of 62
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)
Page 44 of 62
Rubus caesius (Rosaceae)
Rubus caesius (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)
Page 45 of 62
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)
Teucrium pyrenaicum (Lamiaceae)
Page 46 of 62
Teucrium pyrenaicum (Lamiaceae)
Page 47 of 62
Tectum
Tectum
atectate
Pollen grain lacking a tectum.
Cucurbita pepo (Cucurbitaceae)
Rhaphidophora africana (Araceae)
Rhaphidophora africana (Araceae)
eutectate
Pollen grain with a continuous tectum.
Actaea spicata (Ranunculaceae)
Actaea spicata (Ranunculaceae)
Papaver rhoeas (Papaveraceae)
Papaver rhoeas (Papaveraceae)
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)
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
Page 50 of 62
Endexine
spongy
Nelumbo nucifera (Nelumbonaceae)
Ulearum sagittatum var. sagittatum (Araceae)
Zamioculcas zamiifolia (Araceae)
Page 51 of 62
Foot layer
Foot layer
missing
Foot layer missing.
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)
Page 52 of 62
Foot layer
perforate
Foot layer penetrated by thin channels.
Geranium robertianum (Geraniaceae)
Tulipa urumiensis (Liliaceae)
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).
Page 54 of 62
Pollen wall
Pollen wall
alveolate
Infratectum with compartments of irregular size and shape.
Zamioculcas zamiifolia (Araceae)
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).
Epilobium angustifolium (Onagraceae)
Epilobium angustifolium (Onagraceae)
Godetia purpurea (Onagraceae)
Godetia purpurea (Onagraceae)
Page 56 of 62
Wall peculiarities
Kalmia latifolia (Ericaceae)
Kalmia latifolia (Ericaceae)
Page 57 of 62
Cellular condition
Cellular condition
3
Vegetative cytoplasm inclosing two sperm cells.
Galium odoratum (Rubiaceae)
Jasminum nudiflorum (Oleaceae)
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
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
Page 60 of 62
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
Page 61 of 62
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
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
Page 62 of 62

Paldat Terminology

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Illustrated Handbook

Wien, April 2005

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

Acacia myrtifolia (Mimosaceae)

Himantoglossum adriaticum (Orchidaceae)

Platanthera bifolia (Orchidaceae)

Carex remota (Cyperaceae)

Dionaea muscipula (Droseraceae)

Epilobium parviflorum (Onagraceae)

Erica herbacea (Ericaceae)

Mimosa pudica (Mimosaceae)

Typha latifolia (Typhaceae)

Typha latifolia (Typhaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Catalpa ovata (Bignoniaceae)

Dracunculus vulgaris (Araceae)

Ibicella lutea (Martyniaceae)

Iris bucharica (Iridaceae)

Peperomia rubella (Piperaceae)

Phyllanthus spec (Euphorbiaceae)

Cichorium intybus (Asteraceae)

Helminthotheca echioides (Asteraceae)

Scorzonera hispanica (Asteraceae)

Ambrosina bassii (Araceae)

Amorphophallus asterostigmatus (Araceae)

PalDat - Illustrated Handbook on Pollen Terminology

Hemigraphis primulaefolia (Acanthaceae)

Pistia stratiotes (Araceae)

Steudnera assamica (Araceae)

Abies cephalonica (Pinaceae)

Pinus mugo (Pinaceae)

Tsuga canadensis (Pinaceae)

Berberis vulgaris (Berberidaceae)

Crocus speciosus (Iridaceae)

Mimulus guttatus (Veronicaceae)

Thunbergia alata (Acanthaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Shape (single pollen grains)

Aechmea azurea (Bromeliaceae)

Anthurium radicans (Araceae)

Aristolochia arborea (Aristolochiaceae)

Carex humilis (Cyperaceae)

Hordeum murinum (Poaceae)

Laportea divaricata (Urticaceae)

Hohenbergia proctori (Bromeliaceae)

Limnanthes douglasii (Limnanthaceae)

Muscari tenuiflorum (Hyacinthaceae)

Pinus mugo (Pinaceae)

Primula farinosa (Primulaceae)

Salvia involucrata (Lamiaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Shape (single pollen grains)

Tilia cordata (Tiliaceae)

Begonia nelumbifolia (Begoniaceae)

Beloperone guttata (Acanthaceae)

Crossandra flava (Acanthaceae)

Daucus carota (Apiaceae)

Lithospermum officinale (Boraginaceae)

Lotus corniculatus (Fabaceae)

Nandina domestica (Berberidaceae)

Symphytum orientale (Boraginaceae)

PalDat - Illustrated Handbook on Pollen Terminology

Shape (single pollen grains)

Abeliophyllum distichum (Oleaceae)