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Pollen Terminology
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Address
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PalDat
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A 1030 Wien
AUSTRIA
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Fax: +43 1 4277 9541
info@paldat.org
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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
Pollen unit
Pollen unit
polyad
Dispersal unit of more than four united pollen grains.
tetrad
Dispersal unit of four pollen grains (spores).
Page 4 of 62
Pollen class
Pollen class
clypeate
Pollen wall, in which the exine is subdivided into shields.
lophate
Pollen wall with lophae, a network-like pattern of ridges (=lophae) formed by the
outer exine surrounding window-like spaces or depressions (=lacunae).
plicate
Pollen grain with plicae, i.e. cirumferential, parallel ridge-like folds.
Page 5 of 62
Pollen class
saccate
Pollen grain with one or more air sacs, i.e. exinous expansions forming an air sac.
spiraperturate
Pollen grain with pollen grain with one or more spiral aperture(s).
Page 6 of 62
oblate
Pollen grain with a polar axis shorter than the equatorial diameter.
Page 7 of 62
prolate
Pollen grain with a polar axis longer than the equatorial diameter.
Page 8 of 62
Page 9 of 62
Shape peculiarities
Shape peculiarities
heteropolar
Pollen grain with different proximal and distal faces.
isopolar
Pollen grain with identical proximal and distal faces.
Page 10 of 62
Page 11 of 62
lobate
Page 12 of 62
quadrangular
Page 13 of 62
Page 14 of 62
boat-shaped
Characteristic shape of sulcate pollen grains caused by an infolding as a
consequence of harmomegathy.
Page 15 of 62
cup-shaped
Characteristic shape of pollen grains caused by an infolding as a consequence of
harmomegathy.
Page 16 of 62
irregularly infolded
Characteristic shape of pollen grains caused by an infolding as a consequence of
harmomegathy.
not infolded
Page 17 of 62
Page 18 of 62
Aperture number
Aperture number
1
Page 19 of 62
Aperture number
3
Page 20 of 62
Aperture number
Page 21 of 62
Aperture number
>6
Pollen grain with more than 6 apertures.
Page 22 of 62
Aperture number
inaperturate
Pollen grain without distinct apertures.
Page 23 of 62
Aperture peculiarities
Aperture peculiarities
annulate
Pollen grain with an annulus or annuli, a ring like thickening of the pollen wall
surrounding a porus or ulcus.
Comment: "anulus" orthographical variant of "annulus"
Page 24 of 62
Aperture peculiarities
colpate
Pollen grain with elongated apertures (length/width ratio > 2), situated at the
equator or regularly spread over the pollen grain.
Page 25 of 62
Aperture peculiarities
colporate
Pollen grain with compound apertures composed of a colpus (ektoaperture)
combined with an endoaperture of variable size and shape.
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.
Page 26 of 62
Aperture peculiarities
porate
Pollen grain with more or less circular aperture situated at the equator or regularly
spread over the pollen grain.
poroidate
Pollen grain with circular or elliptic aperture, with indistinct margin.
Page 27 of 62
Aperture peculiarities
sulcate
Pollen grain with a sulcus, i.e. an elongated aperture situated distally.
ulcerate
Pollen grain with an ulcus, i.e. more or less circular aperture situated distally.
Page 28 of 62
Aperture peculiarities
Page 29 of 62
Aperture peculiarities
Aperture peculiarities
heteroaperturate
Pollen grain with two different types of apertures; only one type presumed to
function a germination site.
operculate
Coherent exine structure covering an aperture.
Page 30 of 62
Aperture peculiarities
Page 31 of 62
Aperture peculiarities
Page 32 of 62
Aperture peculiarities
pantoaperturate
Pollen grain with apertures distributed more or less regularly over the whole
surface.
Page 33 of 62
Aperture peculiarities
stephanoaperturate
Pollen grain with apertures situated at the equator (term usually used for more than
three apertures).
syncolpate
Pollen grain with anastomosing colpi.
Page 34 of 62
Aperture peculiarities
syncolporate
Pollen grain with anastomosing colpori.
Page 35 of 62
clavate
Pollen wall with clavae, i.e. club-shaped element, higher than 1 m.
echinate
Pollen wall with echini, i.e. pointed ornamentation elements longer and/or wider
than 1 m.
Page 36 of 62
fossulate
Pollen wall with fossulae, i.e. irregularly shaped grooves in the surface of a pollen
wall.
Page 37 of 62
frustrate
Special mental condition of palynologists discussing terminology of pollen and
spores.
gemmate
Pollen wall with gemmae, i.e. globular exine elements more than 1 m in diameter.
Page 38 of 62
microechinate
Pollen wall with echini, i.e. pointed ornamentation elements smaller than 1 m.
microreticulate
Pollen wall with network like pattern formed by exine elements (muri), where the
lumina are smaller than 1 m.
Page 39 of 62
psilate
Pollen wall with smooth surface.
Page 40 of 62
reticulate
Pollen wall with network like pattern formed by exine elements (muri), where the
lumina are wider than 1 m.
Page 41 of 62
reticulate-homobrochate
Reticulate pollen wall with lumina of uniform size.
Comment: the term should be used when the feature is remarkably expressed.
reticulo-cristate
Pollen grain with a special type of reticulum; muri with prominent sculpture
elements.
Page 42 of 62
rugulate
Pollen grain with elongated exine elements longer than 1 m; irregularly arranged.
Page 43 of 62
striate
Pollen wall with elongated exine elements separated by grooves predominantly
parallel arranged.
Page 44 of 62
striato-reticulate
Ornamentation intermediate between striate and reticulate.
Page 45 of 62
verrucate
Pollen wall with wart-like elements more than 1m, broader than high.
Page 46 of 62
Page 47 of 62
Tectum
Tectum
atectate
Pollen grain lacking a tectum.
eutectate
Pollen grain with a continuous tectum.
Page 48 of 62
Tectum
semitectate
Pollen grain with a discontinuous tectum, covering less than 50% of pollen grain
surface.
Page 49 of 62
Endexine
Endexine
missing
Endexine missinig.
continuous
Endexine forming a compact layer.
discontinuous
Endexine interrupted.
lamellar
Page 50 of 62
Endexine
spongy
Page 51 of 62
Foot layer
Foot layer
missing
Foot layer missing.
continuous
Foot layer forming a compact layer.
discontinuous
Foot layer interrupted.
Page 52 of 62
Foot layer
perforate
Foot layer penetrated by thin channels.
Page 53 of 62
Infratectum
Infratectum
absent
columellate
Infratectum with rod-like elements.
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.
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.
papilla
Small protuberance typical for Taxodioideae-pollen (Cupressaceae).
viscin threads
Pollen grain with a circumferential aperture (situated more or less equatorially or,
rarely, meridionally).
Page 56 of 62
Wall peculiarities
Page 57 of 62
Cellular condition
Cellular condition
3
Vegetative cytoplasm inclosing two sperm cells.
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
7
7
7
8
9
11
11
12
12
13
13
14
15
15
15
16
16
17
17
19
19
19
20
20
21
21
22
23
24
24
25
26
26
27
27
28
28
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
52
52
52
52
53
Infratectum .................................................................................................................................
absent ...............................................................................................................................
columellate ........................................................................................................................
granular .............................................................................................................................
54
54
54
54
56
56
56
56
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
PalDat - Illustrated Handbook on Pollen Terminology
reticulate-heterobrochate ................................ 42
reticulate-homobrochate ................................. 42
reticulo-cristate ................................................ 42
rugulate ........................................................... 43
saccate .............................................................. 6
semitectate ...................................................... 49
smooth aperture membrane ............................ 33
spheroidal ......................................................... 9
spiraperturate .................................................... 6
spongy ............................................................ 51
stephanoaperturate ......................................... 34
striate .............................................................. 44
striato-reticulate .............................................. 45
sulcate ............................................................. 28
syncolpate ....................................................... 34
syncolporate .................................................... 35
tetrad ................................................................. 4
triangular ......................................................... 14
ulcerate ........................................................... 28
verrucate ......................................................... 46
viscin threads .................................................. 56
Page 62 of 62