Paldat Terminology

Illustrated Handbookon

Pollen Terminology

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AddressDept. of Palynology and Structural BotanyPalDatUniversity of ViennaRennweg 14A 1030 WienAUSTRIA

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

ImpressumCopyright holder and publisherSociety for the Promotion of Palynological Research in AustriaPalDatUniversity of Vienna, Rennweg 14, A 1030 Wien, AUSTRIAOfficer in charge: BUCHNER RalfErrors and omissions excepted.

VORWORTSeit 1994 ist "The Glossary of Pollen and Spore Terminology" von Punt W, Blackmore S, Nilsson S & LeThomas A das Standardwerk der palynologischen Terminologie, inkl. den Ergänzungen in der seit 1999 jedochnicht mehr überarbeiteten online-Version (http://www.bio.uu.nl/~palaeo/glossary/glos-int.htm).Schwerpunkt unseres Glossars sind die zahlreichen Abbildungen. Jeder Begriff, der in das Glossaraufgenommen wurde, ist umfangreich mit elektronenmikroskopischen Bildern illustriert. Dabei wollten wirbesonders die Bandbreite eines Begriffes aufzeigen.Wir bemühten uns in unserem online-Glossar "PalDat - Illustrated Handbook on Pollen Terminology" um eineStraffung der Begriffe nach praktischen Gesichtspunkten. Viele Begriffe der Palynologie wurden ja zu einerZeit geprägt, als ausschließlich Zeichnungen Grundlage von LM-Beobachtungen zur Verfügung standen.Daher gibt es aus historischen Gründen immer wieder nomenklatorische Inkonsequenzen, Synonyme,unterschiedliche Definitionen für ein und denselben Begriff.Unser Ziel war es, ein Glossar der wichtigsten Begriffe zu erstellen. Durch bewußte Vereinfachung wollten wirauch erreichen, dass dieses Glossar nicht nur für Spezialisten verwendbar ist. Dass durch Generalisierungbzw. Subsummierung unter einem Überbegriff manche Begriffe wegfallen, haben wir bewusst in Kaufgenommen. Sollten Begriffe vermisst werden, erschienen sie uns als unwesentlich, oder sie gehören zurTerminologie der Farnsporen, die in dieser Ausgabe nicht berücksichtigt ist. Gibt es von einem Begriff mehrereSynonyme, wird nur noch einer der Begriffe verwendet. Wo es notwendig war, haben wir Definitionenmodifiziert. Durch konsequente Anwendung elektronenmikroskopischer Techniken und dem damitverbundenen erweiterten Wissensstand mussten manche Begriffe auch neu definiert werden.Wir sehen dieses online-Glossar als eine Möglichkeit, auf einen veränderten Wissensstand flexibel reagierenzu können und freuen uns über Vorschläge und Anregungen. Schreiben Sie diese an [email protected]

PREFACESince 1994 "The Glossary of Pollen and Spore Terminology" of Punt W, Blackmore S, Nilsson S & Le ThomasA is the standard publication in palynological terminology. Unfortunately it was not revised following theadditions 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 microscopicpictures 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" arationalization of terms on the basis of practical criteria. Many terms in palynology were coined at a time, whenexclusively 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 writea glossary which is not only a valuable source for specialists. We accepted consciously that by generalizationand/or subsumption under a general term some terms were omitted. If terms are not considered here then theyappeared us as insignificant, or they belong to the terminology of the fern spores, which are not consideredhere. In case that there are several synonyms for a term, only one of the terms is used. Where it wasnecessary, we modified definitions. By consistent application of electron microscopic techniques and theextended knowledge some terms also had to be redefined. We see this online glossary as a possibility of beingable to react to changed knowledge conditions flexibly.Please send your notes and suggestions to [email protected]

Wien, April 2005 Halbritter H, Weber M, Zetter R, Frosch-Radivo A, Buchner R, Hesse M

Pollen unit

Pollen unit

polyadDispersal unit of more than four united pollen grains.

Acacia myrtifolia (Mimosaceae) Himantoglossum adriaticum (Orchidaceae) Platanthera bifolia (Orchidaceae)

tetradDispersal unit of four pollen grains (spores).

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)

Typha latifolia (Typhaceae)Tetrad (planar)

PalDat - Illustrated Handbook on Pollen Terminology Page 4 of 62

Pollen class

Pollen class

clypeatePollen 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)

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

Cichorium intybus (Asteraceae) Helminthotheca echioides (Asteraceae) Scorzonera hispanica (Asteraceae)

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

Ambrosina bassii (Araceae) Amorphophallus asterostigmatus (Araceae) Hemigraphis primulaefolia (Acanthaceae)


Pollen class

Pistia stratiotes (Araceae) Steudnera assamica (Araceae)

saccatePollen 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

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

Berberis vulgaris (Berberidaceae) Crocus speciosus (Iridaceae) Mimulus guttatus (Veronicaceae)

Thunbergia alata (Acanthaceae)


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

oblatePollen 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)



Tilia cordata (Tiliaceae)Dry pollen grains

prolatePollen 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)


Shape (single pollen grains)spheroidalPollen 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)


Shape peculiarities

Shape peculiarities

heteropolarPollen 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)

isopolarPollen grain with identical proximal and distal faces.

Bellis perennis (Asteraceae) Knautia drymeia (Dipsacaceae) Melittis melissophyllum (Lamiaceae)

Nepeta cataria (Lamiaceae) Pulmonaria mollissima (Boraginaceae)


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


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)

lobate

Artemisia pontica (Asteraceae) Gunnera chilensis (Gunneraceae) Oxalis acetosella (Oxalidaceae)Dry pollen grains

Pinguicula ehlersiae (Lentibulariaceae)Dry pollen grains


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

Alnus glutinosa (Betulaceae) Anthyllis vulneraria (Fabaceae) Platycodon grandiflorum (Campanulaceae)

Stellaria holostea (Caryophyllaceae) Viola arvensis (Violaceae)

quadrangular

Fumaria officinalis (Fumariaceae) Lithospermum officinale (Boraginaceae) Pulmonaria officinalis (Boraginaceae)

Viola tricolor (Violaceae)


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)


Dry pollen, infoldings


aperture(s) sunkenCharacteristic shape of pollen grains caused by an infolding as a consequence ofharmomegathy.

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-shapedCharacteristic shape of sulcate pollen grains caused by an infolding as aconsequence of harmomegathy.

Asphodeline lutea (Liliaceae) Convallaria majalis (Ruscaceae) Galanthus nivalis (Amaryllidaceae)



Vriesea procera (Bromeliaceae)

cup-shapedCharacteristic shape of pollen grains caused by an infolding as a consequence ofharmomegathy.

Dracunculus vulgaris (Araceae) Elaeagnus angustifolia (Elaeagnaceae) Knautia drymeia (Dipsacaceae)

Luzula campestris (Juncaceae)Tetrads (tetrahedral)

Tsuga canadensis (Pinaceae)

interapertural area(s) sunkenCharacteristic shape of pollen grains caused by an infolding as a consequence ofharmomegathy.

Alnus glutinosa (Betulaceae) Bromus erectus (Poaceae) Cornus sanguinea (Cornaceae)



Corylus avellana (Betulaceae) Melampyrum arvense (Orobanchaceae)

irregularly infoldedCharacteristic shape of pollen grains caused by an infolding as a consequence ofharmomegathy.

Asarum europaeum (Aristolochiaceae) Betula humilis (Betulaceae) Dalechampia roezliana (Euphorbiaceae)

Ribes uva-crispa (Grossulariaceae) Sanguisorba minor (Rosaceae) Stylochaeton zenkeri (Araceae)

not infolded

Campanula rapunculoides (Campanulaceae) Cirsium canum (Asteraceae) Onobrychis viciifolia (Fabaceae)



Pinellia ternata (Araceae)


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

2

Aechmea dealbata (Bromeliaceae)porate pollen grain

Colchicum autumnale (Colchicaceae)porate pollen grain

Crinum augustum (Amaryllidaceae)colpate pollen grain

Justicia xylosteoides (Acanthaceae)colporate pollen grain


Aperture number3

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

Scabiosa graminifolia (Dipsacaceae)porate pollen grain

4

Alnus incana (Betulaceae)colpate pollen grain

Hohenbergia stellata (Bromeliaceae)porate pollen grains

Impatiens glandulifera (Balsaminaceae)colpate pollen grain


Aperture number

Nicotiana tabacum (Solanaceae)colporate pollen grain

Nonea pulla (Boraginaceae)colporate pollen grain

Spilanthes acmella (Asteraceae)colporate pollen grain

5

Alnus viridis (Betulaceae)porate pollen grain

Legousia speculum-veneris (Campanulaceae)porate pollen grain

Pulmonaria mollissima (Boraginaceae)colporate pollen grain

Viola arvensis (Violaceae)colporate pollen grain

6

Fumaria officinalis (Fumariaceae)porate pollen grain

Pinguicula ehlersiae (Lentibulariaceae)colporate pollen grain

Primula veris (Primulaceae)colpate pollen grain


Aperture number

Salvia coccinea (Lamiaceae)colpate pollen grain

Sanguisorba officinalis (Rosaceae)colporate pollen grain

>6Pollen 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


Aperture numberinaperturatePollen grain without distinct apertures.

Aristolochia clematitis (Aristolochiaceae) Dracunculus vulgaris (Araceae) Jatropha podagrica (Euphorbiaceae)

Pinellia ternata (Araceae) Populus nigra (Salicaceae)


Aperture peculiarities


annulatePollen grain with an annulus or annuli, a ring like thickening of the pollen wallsurrounding 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)


Aperture peculiaritiescolpatePollen grain with elongated apertures (length/width ratio > 2), situated at theequator 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)


Aperture peculiaritiescolporatePollen 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)

leptomaThinning of the pollen wall at the distal pole (of a pollen grain) in conifers, presumedto function as germination area; special case of tenuitas.

Abies cephalonica (Pinaceae) Tsuga canadensis (Pinaceae)


Aperture peculiaritiesporatePollen grain with more or less circular aperture situated at the equator or regularlyspread 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)

poroidatePollen grain with circular or elliptic aperture, with indistinct margin.

Carex distans (Cyperaceae) Carex distans (Cyperaceae) Cercidiphyllum japonicum (Cercidiphyllaceae)



Cyperus longus (Cyperaceae) Guzmania glomerata (Bromeliaceae) Guzmania glomerata (Bromeliaceae)

sulcatePollen 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

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

Cephalanthera longifolia (Orchidaceae) Luzula campestris (Juncaceae) Sesleria uliginosa (Poaceae)



Sparganium erectum (Sparganiaceae)




heteroaperturatePollen grain with two different types of apertures; only one type presumed tofunction a germination site.

Justicia furcata (Acanthaceae) Lythrum salicaria (Lythraceae) Myosotis ramosissima (Boraginaceae)

operculateCoherent 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)



Veratrum album (Melanthiaceae)

ornamented aperture membraneExine elements (smaller than an operculum) covering the aperture surface, oftenglobular, 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



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)


Aperture peculiaritiespantoaperturatePollen grain with apertures distributed more or less regularly over the wholesurface.

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 membraneAperture surface without any exine elements.

Aechmea caesia (Bromeliaceae) Cyrtosperma beccarianum (Araceae) Digitalis purpurea (Veronicaceae)



Melampyrum arvense (Orobanchaceae) Trifolium montanum (Fabaceae)

stephanoaperturatePollen grain with apertures situated at the equator (term usually used for more thanthree apertures).

Anemone hortensis (Ranunculaceae) Borago officinalis (Boraginaceae) Dionaea muscipula (Droseraceae)Tetrad (tetrahedral)

Sherardia arvensis (Rubiaceae)

syncolpatePollen grain with anastomosing colpi.

Hydrophyllum canadense (Hydrophyllaceae) Nymphoides peltata (Menyanthaceae) Pedicularis verticillata (Orobanchaceae)



Primula farinosa (Primulaceae)

syncolporatePollen grain with anastomosing colpori.

Callistemon coccineus (Myrtaceae) Cassia pulcherrima (Caesalpiniaceae) Cuphea purpurea (Lythraceae)


Ornamentation (LM and EM)


baculatePollen grain with bacula, rod-like, free standing element, more than 1 µm in heightand never pointed.

Erythrochiton brasiliensis (Rutaceae) Nymphaea alba (Nymphaeaceae) Viscum laxum (Viscaceae)

clavatePollen wall with clavae, i.e. club-shaped element, higher than 1 µm.

Geranium phaeum (Geraniaceae) Ilex aquifolium (Aquifoliaceae) Ilex aquifolium (Aquifoliaceae)

Linum catharticum (Linaceae) Plumbago auriculata (Plumbaginaceae)

echinatePollen wall with echini, i.e. pointed ornamentation elements longer and/or widerthan 1 µm.

Anthemis arvensis (Asteraceae) Centaurea jacea (Asteraceae) Hibiscus trionum (Malvaceae)



Malva neglecta (Malvaceae) Pinellia ternata (Araceae) Tussilago farfara (Asteraceae)

Ulearum sagittatum var. sagittatum (Araceae) Zomicarpa riedeliana (Araceae)

fossulatePollen wall with fossulae, i.e. irregularly shaped grooves in the surface of a pollenwall.

Erica herbacea (Ericaceae) Hydrophyllum canadense (Hydrophyllaceae) Orthilia secunda (Ericaceae)

Rhododendron hirsutum (Ericaceae)


Ornamentation (LM and EM)foveolatePollen wall with foveolae, i.e. roundish lumina more than 1 µm in diameter; distancebetween two adjacent lumina larger than their diameter.

Lavandula angustifolia (Lamiaceae) Streptocalyx poeppigii (Bromeliaceae) Stylochaeton bogneri (Araceae)

frustrateSpecial mental condition of palynologists discussing terminology of pollen andspores.

Frustrelia mentalis (Frustraceae)

gemmatePollen wall with gemmae, i.e. globular exine elements more than 1 µm in diameter.

Asarum europaeum (Aristolochiaceae) Dionaea muscipula (Droseraceae) Stenandrium dulce (Acanthaceae)


Ornamentation (LM and EM)granulatePollen wall with granula, i.e. structure- or sculpture elements of different size andshape; smaller than 1 µm.Comment: Not to be confused with "granular", which is a feature of the pollen wallstructure.

Symphytum orientale (Boraginaceae) Ulmus glabra (Ulmaceae)

microechinatePollen wall with echini, i.e. pointed ornamentation elements smaller than 1 µm.

Artemisia pontica (Asteraceae) Campanula patula (Campanulaceae) Valerianella carinata (Valerianaceae)

microreticulatePollen wall with network like pattern formed by exine elements (muri), where thelumina are smaller than 1 µm.

Digitalis lutea (Veronicaceae) Digitalis purpurea (Veronicaceae) Stachys sylvatica (Lamiaceae)


Ornamentation (LM and EM)perforatePollen wall with holes less than 1µm in diameter.

Chimonanthus praecox (Calycanthaceae) Gonatopus angustus (Araceae) Napoleonaea imperialis (Lecythidaceae)

Pulmonaria officinalis (Boraginaceae)

psilatePollen wall with smooth surface.

Dorycnium germanicum (Fabaceae) Dorycnium germanicum (Fabaceae) Hedychium gardnerianum (Commelinaceae)

Hippocrepis emerus (Fabaceae) Hippocrepis emerus (Fabaceae) Vicia tenuifolia (Fabaceae)



Vicia tenuifolia (Fabaceae) Vinca minor (Apocynaceae)

reticulatePollen wall with network like pattern formed by exine elements (muri), where thelumina are wider than 1 µm.

Ajuga genevensis (Lamiaceae) Beloperone guttata (Acanthaceae) Corylopsis glabrescens (Hamamelidaceae)

Gagea graeca (Liliaceae) Galeopsis tetrahit (Lamiaceae) Hyssopus officinalis (Lamiaceae)

Impatiens glandulifera (Balsaminaceae) Ruellia graecicans (Acanthaceae)


Ornamentation (LM and EM)reticulate-heterobrochateReticulate 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-homobrochateReticulate 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-cristatePollen grain with a special type of reticulum; muri with prominent sculptureelements.

Daphne alpina (Thymelaeaceae) Fumana procumbens (Cistaceae) Garcia nutans (Euphorbiaceae)



Geranium canariense (Geraniaceae) Lilium martagon (Liliaceae) Pachysandra terminalis (Buxaceae)

Phaleria capitata (Thymelaeaceae) Phaleria capitata (Thymelaeaceae) Thymelaea passerina (Thymelaeaceae)

rugulatePollen grain with elongated exine elements longer than 1 µm; irregularly arranged.

Angelica archangelica (Apiaceae) Angelica archangelica (Apiaceae) Cabomba palaeformis (Cabombaceae)

Cabomba palaeformis (Cabombaceae) Nicotiana tabacum (Solanaceae) Nicotiana tabacum (Solanaceae)



Nymphoides peltata (Menyanthaceae) Nymphoides peltata (Menyanthaceae) Ulmus laevis (Ulmaceae)

Ulmus laevis (Ulmaceae)

striatePollen wall with elongated exine elements separated by grooves predominantlyparallel arranged.

Acer obtusatum (Sapindaceae) Acer obtusatum (Sapindaceae) Helianthemum nummularium (Cistaceae)

Helianthemum nummularium (Cistaceae) Potentilla inclinata (Rosaceae) Potentilla inclinata (Rosaceae)



Rubus caesius (Rosaceae) Rubus caesius (Rosaceae) Sanguisorba minor (Rosaceae)

Sanguisorba minor (Rosaceae) Saxifraga vandellii (Saxifragaceae) Saxifraga vandellii (Saxifragaceae)

striato-reticulateOrnamentation intermediate between striate and reticulate.

Aesculus glabra (Hippocastanaceae) Aesculus glabra (Hippocastanaceae) Aesculus flava (Hippocastanaceae)

Aesculus flava (Hippocastanaceae) Erodium cicutarium (Geraniaceae) Erodium cicutarium (Geraniaceae)



Gentiana acaulis (Gentianaceae) Gentiana acaulis (Gentianaceae) Gentianella austriaca (Gentianaceae)

Gentianella germanica (Gentianaceae) Pelargonium carnosum (Geraniaceae) Solandra longiflora (Solanaceae)

Solandra longiflora (Solanaceae)

verrucatePollen wall with wart-like elements more than 1µm, broader than high.

Plantago major (Plantaginaceae) Plantago major (Plantaginaceae) Teucrium pyrenaicum (Lamiaceae)



Teucrium pyrenaicum (Lamiaceae)


Tectum

Tectum

atectatePollen grain lacking a tectum.

Cucurbita pepo (Cucurbitaceae) Cucurbita pepo (Cucurbitaceae) Cucurbita pepo (Cucurbitaceae)

Rhaphidophora africana (Araceae) Rhaphidophora africana (Araceae)

eutectatePollen grain with a continuous tectum.

Actaea spicata (Ranunculaceae) Actaea spicata (Ranunculaceae) Papaver rhoeas (Papaveraceae)

Papaver rhoeas (Papaveraceae)


TectumsemitectatePollen grain with a discontinuous tectum, covering less than 50% of pollen grainsurface.

Muscari azureum (Hyacinthaceae) Muscari azureum (Hyacinthaceae) Ornithogalum kochii (Hyacinthaceae)

Ornithogalum kochii (Hyacinthaceae)


Endexine

Endexine

missingEndexine missinig.

Tropaeolum majus (Tropaeolaceae)

continuousEndexine forming a compact layer.

Galium odoratum (Rubiaceae)

discontinuousEndexine interrupted.

Glaucium flavum (Papaveraceae)

lamellar

Aristolochia arborea (Aristolochiaceae)


Endexinespongy

Dracunculus vulgaris (Araceae) Nelumbo nucifera (Nelumbonaceae) Ulearum sagittatum var. sagittatum (Araceae)

Zamioculcas zamiifolia (Araceae)


Foot layer

Foot layer

missingFoot layer missing.

Aristolochia arborea (Aristolochiaceae) Arophyton buchetii (Araceae) Pachypodium succulentum (Apocynaceae)

continuousFoot layer forming a compact layer.

Anaphyllopsis americana (Araceae) Helwingia japonica (Cornaceae)

discontinuousFoot layer interrupted.

Anthericum liliago (Anthericaceae)


Foot layerperforateFoot layer penetrated by thin channels.

Geranium robertianum (Geraniaceae) Tulipa urumiensis (Liliaceae)


Infratectum

Infratectum

absent

Mahonia aquifolium (Berberidaceae)

columellateInfratectum with rod-like elements.

Apium nodiflorum (Apiaceae) Buddleja alternifolia (Buddlejaceae) Ophiorrhiza longiflora (Rubiaceae)

Podolasia stipitata (Araceae)

granularInfratectum composed of granula, cluster of granula or elements of different sizeand shape (never solid and rod-like).

Alnus glutinosa (Betulaceae) Ulmus laevis (Ulmaceae)


Pollen wall

Pollen wall

alveolateInfratectum with compartments of irregular size and shape.

Zamioculcas zamiifolia (Araceae)


Wall peculiarities

Wall peculiarities

internal tectumA ± continuous layer inbetween and parallel to foot layer and tectum, separatedfrom them by columellae.

Nigella arvensis (Ranunculaceae)

papillaSmall protuberance typical for Taxodioideae-pollen (Cupressaceae).

Cryptomeria japonica (Cupressaceae) Cryptomeria japonica (Cupressaceae)

viscin threadsPollen 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)


Wall peculiarities

Kalmia latifolia (Ericaceae) Kalmia latifolia (Ericaceae) Rhododendron hirsutum (Ericaceae)

Rhododendron hirsutum (Ericaceae)


Cellular condition

Cellular condition

3Vegetative cytoplasm inclosing two sperm cells.

Galium odoratum (Rubiaceae) Jasminum nudiflorum (Oleaceae)


Picture Credits

Picture CreditsAktuna G.Anthericum liliago, Anthericum ramosum, Convallaria majalis, Leucojum vernum, Muscari azureum, Muscari tenuiflorum, Ornithogalumkochii

Bombosi P.Symphytum officinale, Viola tricolor

Halbritter H.Abeliophyllum distichum, Abies cephalonica, Acacia myrtifolia, Acantholimon glumaceum, Acer obtusatum, Achillea millefolium, Aechmeaazurea, Aechmea caesia, Aechmea dealbata, Aechmea fraseri, Aesculus flava, Aesculus glabra, Aesculus hippocastanum, Agrostemmagithago, 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, Asphodelinelutea, 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, Cassiapulcherrima, Catalpa ovata, Catopsis floribunda, Centaurea jacea, Cephalanthera longifolia, Cerastium glutinosum, Cercidiphyllumjaponicum, Chimonanthus praecox, Cichorium intybus, Circaea lutetiana, Cirsium canum, Cobaea scandens, Colchicum autumnale,Convolvulus tricolor, Cornus sanguinea, Corylopsis glabrescens, Corylopsis platypetala, Corylus avellana, Crinum augustum, Crocusspeciosus, 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, Fumanaprocumbens, Gagea graeca, Galanthus nivalis, Garcia nutans, Gentiana acaulis, Gentianella austriaca, Gentianella germanica, Geraniumcanariense, Geranium phaeum, Godetia purpurea, Gonatopus angustus, Gunnera chilensis, Guzmania glomerata, Hamamelis vernalis,Hedera helix, Hedychium gardnerianum, Helianthemum nummularium, Helminthotheca echioides, Hemigraphis primulaefolia, Hepaticatranssylvanica, 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, Lachenaliaaloides, Lamium maculatum, Laportea divaricata, Lavandula angustifolia, Lavatera thuringiaca, Legousia speculum-veneris, Liliummartagon, Limnanthes douglasii, Linum catharticum, Lithospermum officinale, Lolium perenne, Lotus corniculatus, Lunaria rediviva, Luzulacampestris, Luzula sylvatica, Lychnis flos-cuculi, Lychnis viscaria, Lythrum salicaria, Malva neglecta, Melampyrum arvense, Melittismelissophyllum, Microrrhinum minus, Mimosa pudica, Mimulus guttatus, Moltkia petraea, Monstera deliciosa, Myosotis arvensis, Myosotisramosissima, 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, Potentillaerecta, Potentilla inclinata, Primula farinosa, Primula veris, Prunus padus, Pulmonaria mollissima, Pulmonaria officinalis, Pycnospathapalmata, Razisea citrina, Rhaphidophora africana, Rhododendron hirsutum, Ribes sanguineum, Ribes uva-crispa, Rubus caesius, Ruelliagraecicans, 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, Stylochaetonbogneri, 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, Viscumalbum, 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, Helwingiajaponica, Jasminum nudiflorum, Nelumbo nucifera, Ophiorrhiza longiflora, Pachypodium succulentum, Podolasia stipitata, Tropaeolummajus, Tulipa urumiensis, Ulearum sagittatum var. sagittatum, Zamioculcas zamiifolia


Contents

Contents

Pollen unit .................................................................................................................................... 4polyad .................................................................................................................................. 4tetrad ................................................................................................................................... 4

Pollen class .................................................................................................................................. 5clypeate ............................................................................................................................... 5lophate ................................................................................................................................ 5plicate .................................................................................................................................. 5saccate ................................................................................................................................ 6spiraperturate ...................................................................................................................... 6

Shape (single pollen grains) ...................................................................................................... 7irregular ............................................................................................................................... 7oblate .................................................................................................................................. 7prolate ................................................................................................................................. 8spheroidal ............................................................................................................................ 9

Shape peculiarities .................................................................................................................... 10heteropolar ........................................................................................................................ 10isopolar .............................................................................................................................. 10

Equatorial outline (in polar view, excluding apertural protrusions) ..................................... 11circular ............................................................................................................................... 11elliptic ................................................................................................................................ 12lobate ................................................................................................................................ 12polygonal ........................................................................................................................... 13quadrangular ..................................................................................................................... 13triangular ........................................................................................................................... 14

Dry pollen, infoldings ................................................................................................................ 15aperture(s) sunken ............................................................................................................ 15boat-shaped ...................................................................................................................... 15cup-shaped ....................................................................................................................... 16interapertural area(s) sunken ............................................................................................ 16irregularly infolded ............................................................................................................. 17not infolded ........................................................................................................................ 17

Aperture number ....................................................................................................................... 191 ........................................................................................................................................ 192 ........................................................................................................................................ 193 ........................................................................................................................................ 204 ........................................................................................................................................ 205 ........................................................................................................................................ 216 ........................................................................................................................................ 21>6 ...................................................................................................................................... 22inaperturate ....................................................................................................................... 23

Aperture peculiarities ............................................................................................................... 24annulate ............................................................................................................................ 24colpate ............................................................................................................................... 25colporate ........................................................................................................................... 26leptoma ............................................................................................................................. 26porate ................................................................................................................................ 27poroidate ........................................................................................................................... 27sulcate ............................................................................................................................... 28ulcerate ............................................................................................................................. 28

Aperture peculiarities ............................................................................................................... 30heteroaperturate ................................................................................................................ 30operculate ......................................................................................................................... 30ornamented aperture membrane ...................................................................................... 31pantoaperturate ................................................................................................................. 33smooth aperture membrane .............................................................................................. 33stephanoaperturate ........................................................................................................... 34syncolpate ......................................................................................................................... 34syncolporate ...................................................................................................................... 35


Contents

Ornamentation (LM and EM) .................................................................................................... 36baculate ............................................................................................................................. 36clavate ............................................................................................................................... 36echinate ............................................................................................................................. 36fossulate ............................................................................................................................ 37foveolate ............................................................................................................................ 38frustrate ............................................................................................................................. 38gemmate ........................................................................................................................... 38granulate ........................................................................................................................... 39microechinate .................................................................................................................... 39microreticulate ................................................................................................................... 39perforate ............................................................................................................................ 40psilate ................................................................................................................................ 40reticulate ............................................................................................................................ 41reticulate-heterobrochate .................................................................................................. 42reticulate-homobrochate ................................................................................................... 42reticulo-cristate .................................................................................................................. 42rugulate ............................................................................................................................. 43striate ................................................................................................................................ 44striato-reticulate ................................................................................................................. 45verrucate ........................................................................................................................... 46

Tectum ........................................................................................................................................ 48atectate ............................................................................................................................. 48eutectate ........................................................................................................................... 48semitectate ........................................................................................................................ 49

Endexine .................................................................................................................................... 50missing .............................................................................................................................. 50continuous ......................................................................................................................... 50discontinuous .................................................................................................................... 50lamellar .............................................................................................................................. 50spongy ............................................................................................................................... 51

Foot layer ................................................................................................................................... 52missing .............................................................................................................................. 52continuous ......................................................................................................................... 52discontinuous .................................................................................................................... 52perforate ............................................................................................................................ 53

Infratectum ................................................................................................................................. 54absent ............................................................................................................................... 54columellate ........................................................................................................................ 54granular ............................................................................................................................. 54

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

Wall peculiarities ....................................................................................................................... 56internal tectum ................................................................................................................... 56papilla ................................................................................................................................ 56viscin threads .................................................................................................................... 56

Cellular condition ...................................................................................................................... 583 ........................................................................................................................................ 58


Register

Register

1 ...................................................................... 192 ...................................................................... 193 ................................................................ 20, 584 ...................................................................... 205 ...................................................................... 216 ...................................................................... 21>6 .................................................................... 22absent ............................................................. 54alveolate .......................................................... 55annulate .......................................................... 24aperture(s) sunken .......................................... 15atectate ........................................................... 48baculate .......................................................... 36boat-shaped .................................................... 15circular ............................................................ 11clavate ............................................................. 36clypeate ............................................................. 5colpate ............................................................ 25colporate ......................................................... 26columellate ...................................................... 54continuous ................................................. 50, 52cup-shaped ..................................................... 16discontinuous ............................................ 50, 52echinate .......................................................... 36elliptic .............................................................. 12eutectate ......................................................... 48fossulate .......................................................... 37foveolate ......................................................... 38frustrate ........................................................... 38gemmate ......................................................... 38granular ........................................................... 54granulate ......................................................... 39heteroaperturate ............................................. 30heteropolar ...................................................... 10inaperturate ..................................................... 23interapertural area(s) sunken .......................... 16internal tectum ................................................ 56irregular ............................................................. 7irregularly infolded ........................................... 17isopolar ........................................................... 10lamellar ........................................................... 50leptoma ........................................................... 26lobate .............................................................. 12lophate .............................................................. 5microechinate .................................................. 39microreticulate ................................................. 39missing ...................................................... 50, 52not infolded ..................................................... 17oblate ................................................................ 7operculate ....................................................... 30ornamented aperture membrane .................... 31pantoaperturate ............................................... 33papilla .............................................................. 56perforate .................................................... 40, 53plicate ................................................................ 5polyad ............................................................... 4polygonal ......................................................... 13porate .............................................................. 27poroidate ......................................................... 27prolate ............................................................... 8psilate .............................................................. 40quadrangular ................................................... 13reticulate ......................................................... 41

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


Documents

Paldat Terminology