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Pollen and spore keys for Quaternary deposits
in the northern Pindos Mountains, Greece
Pamela I. Chest er & J. Ian Raine
Published online: 05 Nov 2010.
To cite this article: Pamela I. Chest er & J. Ian Raine (2001) Pollen and spore keys f or Quat ernary deposit s in
t he nort hern Pindos Mount ains, Greece, Grana, 40: 6, 299-387, DOI: 10. 1080/ 00173130152987535
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Grana 40: 299±387, 2001
Pollen and spore keys for Quaternary deposits in the
northern Pindos Mountains, Greece
PAMELA I. CHESTER and J. IAN RAINE
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Chester, P.I. & Raine, J.I. 2001. Pollen and spore keys for Quaternary deposits in the northern Pindos
Mountains, Greece. ± Grana 40: 299±387. ISSN 0017-3134. .
Keys for identication of pollen and spores of extant plants in Grevena Province, northwestern Greece
are presented at the lowest commonly achievable taxonomic rank, based on light microscope
observation. As the Province includes a range of vegetation from lowland evergreen oak woods,
through semi-deciduous and deciduous oak and mixed deciduous forests at middle elevations, to higher
altitude beech and pine woods and alpine meadows, the keys cover most common pollen and spores
encountered in Quaternary deposits in upland areas of southeastern Europe. Special attention has been
paid to closely identifying pollen and spores of particular interest to the archaeologist. Based on
compilations of existing data and new observations, separate special keys are included for
Caryophyllaceae, Cyperaceae, Gramineae, Malvaceae, Plantaginaceae, and Rosaceae. A full list of
species included in each pollen type is provided.
Pamela I. Chester, School of Earth Sciences, Victoria University of Wellington, P.O. Box 600, Wellington
(Curr. address: 36 Woodlan d Road, Johnsonville, Wellington) ; J. Ian Raine, Institute of Geological &
Nuclear Sciences, P.O. Box 30368, Lower Hutt; New Zealand. E-mail: pchester@actrix.co.nz
(Manuscript received 15 May 2001; accepted 28 December 2001)
CONTENTS
(Introduction) ............................................................................................................................................................................................. . 300
Material and Methods ................................................................................................................................................................................ . 301
Part 1.
General keys to genera and species
Master key ................................................................................................................................................................................... .
A. Vesiculate .............................................................................................................................................................................. .
B. Compound grains ................................................................................................................................................................. .
B1. Tetrads ......................................................................................................................................................................... .
B2. Polyads ........................................................................................................................................................................ .
C. Bilateral grains ...................................................................................................................................................................... .
D. Subtetrahedral grains (spores) .............................................................................................................................................. .
E. Rotational ellipsoidic (or ovate) grains: main key ............................................................................................................... .
E1.
Polyplicate .................................................................................................................................................................. .
E2.
Inaperturate ............................................................................................................................................................... .
E3.
Trichotomocolpate ..................................................................................................................................................... .
E4.
Monocolpate .............................................................................................................................................................. .
E5.
Monoporate ............................................................................................................................................................... .
E6.
Dicolpate .................................................................................................................................................................... .
E7.
Tricolpate; Main key to subsections .......................................................................................................................... .
E7.1 tectate, psilate .................................................................................................................................................. .
E7.2 tectate, scabrate ............................................................................................................................................... .
E7.3 tectate; baculate, echinate or verrucate ........................................................................................................... .
E7.4 tectate; rugulate or striate ............................................................................................................................... .
E7.5 tectate; supra-reticulate ................................................................................................................................... .
E7.6 per-reticulate ................................................................................................................................................... .
E7.7 intectate; clavate, gemmate and baculate ........................................................................................................ .
E8.
Stephanocolpate ......................................................................................................................................................... .
E9.
Pericolpate .................................................................................................................................................................. .
E10. Dicolporate ................................................................................................................................................................ .
E11. Tricolporate: main key to subsections ....................................................................................................................... .
E11.1 tectate, psilate or scabrate ............................................................................................................................. .
E11.2 tectate, echinate ............................................................................................................................................. .
E11.3 tectate, striate or rugulate ............................................................................................................................. .
E11.4 tectate; foveolate or supra-reticulate ............................................................................................................. .
E11.5 per-reticulate .................................................................................................................................................. .
E11.6 intectate, clavate ............................................................................................................................................ .
Ñ 2001 Taylor & Francis. ISSN 0017-313 4
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302
302
302
304
304
305
306
306
306
306
309
313
313
313
313
314
317
317
319
321
323
323
325
327
327
327
331
332
333
337
342
Grana 40 (2001)
300
P. I. Chester and J. I. Raine
E12
E13
E14
E15
E16
E17
Stephanocolporate ...................................................................................................................................................... .
Pericolporate ............................................................................................................................................................... .
Diporate ...................................................................................................................................................................... .
Triporate ..................................................................................................................................................................... .
Stephanoporate ........................................................................................................................................................... .
Periporate: main key to subsections ........................................................................................................................... .
E17.1 verrucate or echinate ..................................................................................................................................... .
E17.2 rugulate, striate or reticulate ......................................................................................................................... .
E17.3 psilate or scabrate ......................................................................................................................................... .
Syncolpate ................................................................................................................................................................... .
Heterocolpate .............................................................................................................................................................. .
Fenestrate (Compositae: Cichorioideae) ..................................................................................................................... .
342
344
344
345
347
348
348
349
349
351
352
353
Special keys to families
Caryophyllaceae ........................................................................................................................................................................... .
Cyperaceae ................................................................................................................................................................................... .
Gramineae .................................................................................................................................................................................... .
Malvaceae .................................................................................................................................................................................... .
Plantaginaceae .............................................................................................................................................................................. .
Rosaceae ...................................................................................................................................................................................... .
353
357
357
359
360
363
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E18
E19
E20
Part 2.
Part 3.
Pollen types
Lycopsida ..................................................................................................................................................................................... .
Sphenopsida ................................................................................................................................................................................. .
Filicopsida .................................................................................................................................................................................... .
Gymnospermae ............................................................................................................................................................................ .
Angiospermae
Dicotyledones ........................................................................................................................................................................... .
Monocotyledones ..................................................................................................................................................................... .
364
364
364
365
365
379
Index ........................................................................................................................................................................................................... . 382
Acknowledgments ....................................................................................................................................................................................... . 386
References ................................................................................................................................................................................................... . 386
General keys for optical microscope identication of pollen
and spores of extant plants in Grevena Province, northwestern Greece are presented in Part 1. Species extant in the
Grevena region were determined from vegetation surveys
undertaken by the author in the company of Dr Oliver
Rackham, University of Cambridge UK (Rackham: unpubl.
rep.) and several literary sources (Pouqueville 1826, Leake
1835, Tutin et al. 1964±80, Wace & Thompson 1972, Skas
1978, Strid 1986, Polunin 1988, Strid & Tan 1991).
Based on compilations of existing data and new observations, separate special keys are included in Part 2 for
Caryophyllaceae, Cyperaceae, Gramineae, Malvaceae,
Plantaginaceae, and Rosaceae. The keys cover most common
pollen and spores encountered in Quaternary deposits in
upland areas of southeastern Europe.
A full list of plant species included in each pollen and
spore taxon is given in Part 3. Unfortunately the pollen or
spore morphology of many plant species still remains unexamined, but the probable morphology of these species can
be guessed at based on plant taxonomy, allowing them to be
placed in the most likely taxon. These species are listed at
the end of each pollen or spore taxon and can be identied
by the ``?’’ which precedes them. Taxonomic nomenclature
follows the Flora Europaea (Tutin et al. 1964±80), except
where species have been described more recently by Strid
(1986 ) or Strid & Tan (1991). An index to pollen and spore
taxa in the key is provided.
Grana 40 (2001)
Special attention has been paid to identifying closely pollen
and spores of particular interest to the archaeologist, e.g.
cultivated plants, weeds of cultivation, and plants indicating
disturbance of the natural vegetation. Unfortunately, among
the major cereals only Secale cereale (rye) and Zea mays
(maize) can be identied to species level. Triticum (wheat),
Hordeum (barley) and Avena (oats) can be distinguished only
to generic. The genera include several closely related species,
not all of which are or have been cultivated. Other important
cultivated species such as Vicia faba (broad bean) and Pisum
sativum (pea) can be identied specically, but pollen of
Isatis tinctoria (woad ) and Camelina sativa (gold-ofpleasure), and most other crucifers and legumes cannot be
specically determined. The same pertains to many important
weeds and ruderal species. Fenestrate pollen taxa of
Compositae have been given special attention.
Geographic coverage
The landscape of Grevena Province, an inland basin,
resembles Central European landscapes of mountains and
plains. The province ( Text Fig. 1) is bordered by mountains
on three sides; in the west the Pindos massif reaches elevations
of 2250 m asl with many peaks over 2000 m, in the east the
Vourinos reach elevations of 1850 m asl although the peaks
are generally closer to 1500 m, and in the south the lesser
hills of the Chasia Mountains reach 1482 m. The central
plains of the basin undulate between 400 and 700 m asl.
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Pollen and spore keys, Pindos Mts, Greece
Thus Grevena can be divided into ve major topographic
regions: Pindos mountains, Pindos foothills, central plains
or lowlands, ood plains, and the western slopes of the
Vourinos Mountains.
Grevena has a Mediterranean climate characterised by
winter rainfall and a long dry summer season (Trewartha
1990: 8±9). However, inland GrevenaÂ, 140 km from the coast
and at 40ß N latitude, is near the northern limit of this
climatic zone and is inuenced by continental European
climate, so winter temperatures are colder (snow falls at even
the lowest elevations) and summer rainfall is generally greater
than for more southerly or coastal Mediterranean climates
( Walter et al. 1975: 5; Sakellariou 1988: 19 ).
The vegetation is mostly a mixture of continental European
species and more hardy Mediterranean species. The GrevenaÂ
landscape is intensively used and little, if any, unmodied
vegetation remains. Today land use includes agriculture;
pastoralism; removal of trees by cutting; and harvesting of
trees by coppicing, pollarding, and shredding. In spite of
intensive land use, and in contrast to the bareness of much
of Greece (Rackham 1982), the extant vegetation of GrevenaÂ
is dominated by woodland, which covers over half the entire
land area. Nevertheless, the ora of Grevena is very rich in
herbaceous species. Plant communities are varied and complex. With decreasing elevation they range from alpine
vegetation of mountain peaks, mountain pine woods and
beech and r woods of Central European aspect, deciduous
oak woods in the foothills, to lowland Mediterranean-type
woods containing some evergreen species such as Quercus
coccifera and Phillyrea media, and dry steppe reminiscent of
the Hungarian and Russian steppe.
301
in The Northwest European Pollen Flora volumes 1±7 edited by
Punt et al. (1976±1995; in the key these are referred to by ``NW’’
followed by the volume number), Roure (1985), ValdeÂs et al. (1987 ),
and smaller works in the journals Grana, Grana Palynologica, Pollen
et Spores, and Review of Palaeobotany and Palynology. In addition,
reference was made to an extensive collection of pollen and spore
reference material from plants collected in Grevena and from
herbarium specimens. A full list of examined and illustrated specimens is available from the authors. The Part 1 key E20 for fenestrate
pollen, exclusively Compositae Cichorioideae, and special keys in
Part 2 for Caryophyllaceae, Cyperaceae, Gramineae, Malvaceae,
Plantaginaceae, and Rosaceae, are based partly on the detailed
morphological studies referenced at the beginning of each key.
Use of keys and terminology
The keys are intended for use with a light microscope, using mostly
a Ö40 objective and occasionally an oil immersion Ö100 objective
and phase contrast. Pollen and spores are separated at the lowest
commonly achievable taxonomic rank, based on light microscope
observation. The main key of Part 1, and the special keys of Part 2,
are modelled on those of Faegri & Iversen (1989) , and are designed
for specialist pollen analysts. They assume a fair degree of pollenmorphological knowledge.
Except as noted below, pollen-morphological terminology also
follows that of Faegri & Iversen (1989). The terms nexine and sexine
are used to describe the strata of the exine because the sculptured
sexine is easily distinguished from the unsculptured nexine when
viewed under the light microscope, while the ectexine and endexine
layers are diYcult to distinguish under the light microscope (Faegri
& Iversen 1989: 223). The sexine is considered to be made up of,
from outermost layer to the innermost, sculpture element (sexine 3 ),
tectum (sexine 2 ) and columellae (sexine 1). The nexine is made up
of the foot layer (nexine 1) and the endexine (nexine 2) (Moore et al.
1991: 64, Fig. 5.2).
Some important terms describing aperture shape are annulus ,
margo, costa, vestibulum, and fastigium. The denition of these terms
follows Moore et al. (1991:71) :
annulus ± a sudden thickening or thinning of the sexine around
an ectoporus, e.g. Plantago lanceolata;
margo ± a sudden thickening or thinning of the sexine around an
ectocolpus, e.g. Euphorbia;
costa ± thickening or thinning of the nexine around an endoaperture or below the edge of an ectoaperture, e.g. Polygonum
aviculare;
vestibulum ± the cavity formed at the edge of the pore in porate
grains when the sexine and nexine separate, e.g. Betula;
fastigium (geniculus in van Benthem et al. 1984: 87) ± the cavity
formed, when the sexine and nexine separate, around pores in
colporate grains e.g. Hypericum or at the equator when no
pores are present, e.g. Quercus.
Format of the keys
Text-Fig. 1. Location of the study site (Pindos Mts., GrevenaÂ).
MATERIAL AND METHODS
Descriptions of the grains were collected from many sources, the
main literature sources being the major morphological works of
Erdtman (1943, 1952, 1957, 1965, 1969, 1971), Ikuse (1956) , Beug
(1961), Erdtman et al. (1961, 1963), Erdtman & Sorsa (1971),
Faegri & Iversen (1975, 1989) , Nilsson et al. (1977), Belmonte et al.
(1986), Ferrarini et al. (1986), Moore et al. (1991 ), various articles
In principle the keys are binary, but in some places more than two
entries at the same level have been used for simplicity. Aperture
type, number, shape and arrangement of apertures, is the major
entry in the main key. Exine structure and sculpture, as dened by
Faegri & Iversen (1989: 227±9), are considered next. Because these
features vary and are sometimes diYcult to discern, some pollen
taxa occur in more than one place in the key. Cross-references are
provided in these cases. Spores are placed in the aperture classes
originally devised to separate angiosperm pollen grains, for example,
pteridophyte spores with one groove are included under monocolpate
instead of being in a class on their own, that is, monolete. Particular
distinguishing features have been italicized.
Grana 40 (2001)
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P. I. Chester and J. I. Raine
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Measurements given should be considered after the above features
have been considered, and used cautiously. During the course of
researching the morphology of the pollen and spores included in
this key many widely diVering measurements were found in the
literature for the same pollen and spore taxa. Also diVerent chemical
treatments will aVect the size of pollen and spores variously (Reitsma
1969) . No particular attention has been paid to the mounting media
used and the measurements should be considered approximate.
Measurements do however give a rough guide as to whether the
grains are large or small. Occasionally size is used as a distinguishing
feature, as for example in Quercus, where it has been used prudently.
The right-hand column gives the ultimate breakdown to taxa. A
genus name alone refers to species of that genus only; similarly a
species name alone signies that only that species belongs to the
pollen taxon. The addition of the term ``type’’, abbreviated ``t.’’,
implies a pollen morphological type common to several families,
genera or species, as the case may be. The term ``group’’ is used for
subdivisions of a few pollen taxa, following the usage of the Northwest
European Pollen Flora (Punt et al. 1976 ±1995). Taxa illustrated include
those most commonly found taxa, those of archaeological interest,
and representatives of the main morphological classes.
Reference to relevant gure numbers is provided in the body of
the key. Note that some gures are darker because of the use of
safranin stain.
PART 1
General keys
Master key
A2
A1
Grains vesiculate/saccate ....................................................................................................................................... Key A
Grains without sacci.
B2 Compound grains (tetrads, polyads) ............................................................................................................. Key B
B1 Grains single.
C2 Grains bilateral ...................................................................................................................................... Key C
C1 Grains isodiametric or one axis longer/shorter, rarely heteropolar.
D2 Grains subtetrahedral .................................................................................................................. Key D
D1 Grains rotational .......................................................................................................................... Key E
Key A. Vesiculate/ saccate pollen grains
(A2
A1
Three meridional, saccus-like crests ......................................................................................................... Trapa natans)
TRAPACEAE
Distinct, subglobular or pouch-shaped sacci (usually two) with internal, 3-dimensional reticulum.
B2 Proximal surface of the body exine more than 5 mm thick.
C2 Body of the grain 80±100 mm. Sacci more than semiglobular. Fig. 1A .................................................. Abies
PINACEAE
C1 Body of the grain c. 50 mm. Sacci semiglobular. Fig. 1C ..................................................................... Cedrus
PINACEAE
B1 Proximal part of body exine less than 5 mm thick.
D2 Proximal entry angle of saccus very blunt; no obvious constriction between sacci and body of the grain.
E2 Gradual transition between structure of the saccus and that of the wall of the body of the grain. Colpus
membrane smooth. Fig. 1B .................................................................................................... Picea abies
PINACEAE
E1 Abrupt transition between structure of saccus and that of the wall of the body of the grain. Colpus
membrane dotted. Fig. 1D ....................................................................................... Pinus ( Haploxylon)
PINACEAE
D1 Proximal entry angle of saccus sharp; distinct constriction between saccus and body of the grain. Body of
the grain c. 50 mm. Colpus membrane smooth. Fig. 1E±F ............................................... Pinus ( Diploxylon)
PINACEAE
Key B. Compoun d pollen grains
The ``normal’’ position of a tetrahedral tetrad is the one in which the polar axis of one grain is parallel with the optical
axis of the microscope.
Key B1. Tetrads
A2
Tetrads linear, at (quadrangular) or irregular. Grains reticulate or rugulate.
B4 Grains with one pore, c. 3 mm wide, away from join between grains. Tetrad linear or at. Typha spp.
C2 Rugulate. See also Key E5. Fig. 2A ................................................................................ Typha angustifolia t.
TYPHACEAE
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Pollen and spore keys, Pindos Mts, Greece
Fig. 1. Vesiculate (500Ö). A, Abies (e.g. A. pinsapo ); B, Picea abies; C, Cedrus (e.g. C. atlantica); D, Pinus (Haploxylon) (e.g. P. halepensis);
E±F, Pinus (Diploxylon) (e.g. P. leucodermis).
C1
B3
Reticulate, lumina c. 2 mm .............................................................................................. Typha domingensis t.
TYPHACEAE
Tetrad at. Grains with two pores. .............................................................................................. Cytinus hypocistis
RAFFLESIACEAE
Grana 40 (2001)
304
P. I. Chester and J. I. Raine
B2
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A1
Tetrad irregular (sometimes tetrahedral ). Grain with single pore 6 mm wide or two apertures. Reticulate, mesh
3±5 mm wide. Fig. 2B .............................................................................................................................. Epipactis t.
ORCHIDACEAE
B1 Tetrad irregular. Grain with 3±6 pores situated on dividing walls between grains ....................... Periploca graeca
ASCLEPIADACEAE
Tetrads tetrahedral, sometimes irregularly so.
D2 Grains psilate, inaperturate. Exine extremely thin, normally not preserved/recognized in sediments ... Juncaceae
D1 Grains scabrate or verrucate.
E2 Triporate, pores with vestibulum. Grains loosely connected. Cf. Key E15: Triporate. Fig. 2C ......................
Epilobium hirsutum t.
ONAGRACEAE
E1 Tricolpate/tricolporate. Tetrads compact, subglobular. Colpi regular. Pyrolaceae, Ericaceae.
F2 Colpi very short. Costae indistinct ................................................................................. Moneses uniora
PYROLACEAE
F1 Colpi long, usually with margo. Interior walls with few or no perforations. Psilate to irregularly scabrateverrucate.
G2 Costae absent/indistinct. ....................................................................................................... Pyrola
PYROLACEAE
G1 Costae distinct.
H2 Thin exine ............................................................................................... Vaccinium myrtillus
ERICACEAE
H1 Thick exine. Fig. 2D±E ................................................................................... Erica herbacea
ERICACEAE
Key B2. Polyads
Polyads irregular; tetrads usually indistinguishable, densely crowded in massulae.
A2 Monoporate grains. ............................................................................................................................................ Orchis t.
ORCHIDACEAE
A1 Inaperturate grains. .................................................................................................................................... Vincetoxicum
ASCLEPIADACEAE
Key C. Bilateral pollen and spores grains
Some monocolpate and biporate grains are faintly bilateral or pear-shaped (heteropolar). They are found in Key E.
A2 With more than one aperture.
B2 Bi-aperturate: two pores; scabrate ............................................................................................... Cytinus hypocistis
RAFFLESIACEAE
B1 Tetra-aperturate, reticulate ..................................................................................................... Impatiens noli-tangere
BALSAMINACEAE
A1 With one or no aperture.
C2 With columellae. Grain reticulate. Heterobrochate, semitectate, columellae forming muri. Liliaceae,
Amaryllidaceae, etc.: see Key E4.
C1 No columellae. Mostly Pteridophyte spores.
D2 Perine loose, crested or folded, not echinate but in some spores spinulose.
E2 Perine saccate.
F3 Perine folded into numerous vesiculae, surface smooth or minutely vermiculate (SEM ) ...............
Dryopteris lix-mas t.
ASPIDIACEAE
F2 Perine folded into numerous vesiculae, foveolate-reticulate .................... Gymnocarpium dryopteris
ASPIDIACEAE
F1 Perine with widely spaced folds, minutely echinate .......................................... Dryopteris expansa
ASPIDIACEAE
E1 Perine lophate-cristate.
G2 Crests few, 1±4, long, sometimes as long as the spore .............................. Thelypteris phegopteris
THELYPTERIDACEAE
G1 Usually more than 4 crests, which are shorter.
H2 Crests with few or no spinules ......................................................... Athyrium distentifolium
ATHYRIACEAE
H1 Crests densely spinulose.
Grana 40 (2001)
Pollen and spore keys, Pindos Mts, Greece
305
I2
D1 No
J4
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J3
J2
J1
Perine entire or sparsely perforate. Includes Ceterach, Phyllitis. Keys to species are to
be found in Ferrarini et al. (1986). Fig. 2F ................................................ Asplenium t.
ASPLENIACEAE
I1 Perine densely perforate (lacy) ..................................................................... Polystichum
ASPIDIACEAE
loose outer perine or, if present, not folded or crested.
Verrucate, sculpturing elements coarse, more than 3 mm high .............................................. Polypodium
POLYPODIACEAE
Scabrate. (Najas marina might be included here, but its exine is not preserved in acetolysis, or,
presumably, in fossilization).
K2 No perine visible ............................................................................................ Athyrium lix-femina
ATHYRIACEAE
K1 Perine thin, slightly wrinkled ................................................................................ Blechnum spicant
BLECHNACEAE
Psilate. (Like Najas, Ceratophyllum is not preserved in acetolysis).
L2 Wall double with loose outer cover ........................................................................................ Isoetes
ISOETACEAE
L1 Wall one-layered or outer cover not loose, smooth, structureless. Unidentiable fern spores that
have lost their perispore.
Echinate, with echinae and (blunt) papillae.
M2 Mainly papillae .................................................................................................. Cystopteris fragilis
ATHYRIACEAE
M1 Sculpturing elements variable .......................................................................... Thelypteris palustris
THELYPTERIDACEAE
Key D. Subtetrahedral spores
A2
A1
With an outer, loose, folded perine. See also Key E2 .................................................................................... Equisetum
EQUISETACEAE
No loose perine. Cf. Thesium, Key E18.
B3 Grain subglobular.
C3 Spore psilate; aperture obscure, Ôelaters. See also Key E2 ........................................................... Equisetum
EQUISETACEAE
C2 Spore wall undulose to scabrate (perine akes oV ) ..................................................................... Cheilanthes
SINOPTERIDACEAE
C1 Spore baculate-clavate (tending to rugulate distally) ............................................................ Osmunda regalis
OSMUNDACEAE
B2 Spore triangular, semi-lobate. Corners rounded.
D2 Verrucate .................................................................................................................... Anogramma leptophylla
GYMNOGRAMMACEAE
D1 Scabrate-mbriate, usually smooth by loss of ornamentation.
E3 Average diameter 40-50 mm, sides in polar view convex, laesurae margins conspicuously ridged ...........
Adiantum capillus-veneris
ADIANTACEAE
E2 Average diameter 28±36 mm, sides in polar view concave, laesurae margins not ridged. Fig. 2G ...........
Pteridium aquilinum
HYPOLEPIDACEAE
E1 Average diameter <25 mm. Spores similar to Adiantum and Pteridium, but signicantly smaller ...........
Salvinia natans
SALVINIACEAE
B1 Equatorial limb triangular-subcircular. Polar axis shorter than equatorial diameter.
F4 Rugulate/verrucate to reticulate. Equatorial ridge present.
G2 Distally rugulate .................................................................................................................. Pteris cretica
PTERIDACEAE
G1 Distally reticulate with verrucae in lumina ......................................................................... Pteris vittata
PTERIDACEAE
F3 Foveolate/reticulate with rounded muri. No equatorial ridge. Ophioglossum.
H2 Tiny punctae ................................................................................................... Ophioglossum lusitanicum
OPHIOGLOSSACEAE
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F2
F1
H1 Coarsely reticulate .............................................................................................. Ophioglossum vulgatum
OPHIOGLOSSACEAE
Verrucate. Verrucae rounded, sometimes spinulate. Spore 43±58 mm. Sculpturing on proximal face as distinct
as on the distal .............................................................................................................................. Botrychium
OPHIOGLOSSACEAE
Minutely echinate or granulate ...................................................................................................... Selaginella
SELAGINELLACEAE
Key E. Rotational ellipsoidic (or ovate) grains: main key
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A3
A2
A1
No distinct aperture.
B2 With meridional grooves and ridges ................................................................................................. El Polyplicate
B1 No meridional grooves or ridges ..................................................................................................... E2 Inaperturate
With one aperture.
C3 Aperture Y-shaped ................................................................................................................ E3 Trichotomocolpat e
C2 Aperture elongate ........................................................................................................................... E4 Monocolpat e
C1 Aperture circular ............................................................................................................................. E5 Monoporat e
More than one aperture.
D2 No lacunae or, if present, not in a xed geometric pattern.
E2 Apertures not fused.
F3 With colpi, no pores or transverse endexinous colpi (colpate).
G3 Two colpi .................................................................................................................... E6 Dicolpate
G2 Three colpi ................................................................................................................. E7 Tricolpate
G1 More than three colpi.
H2 All colpi meridional ................................................................................ E8 Stephanocolpat e
H1 Some or all colpi not meridional .................................................................... E9 Pericolpate
F2 Both colpi and distinct pores or transverse colpi present (colporate). Usually one pore per colpus,
occasionally missing; in some taxa more than one pore per colpus (not in our material ).
I3 Two colpi ................................................................................................................ E10 Dicolporate
I2 Three colpi ............................................................................................................. E11 Tricolporate
I1 More than three colpi.
J2 All colpi meridional .............................................................................. E12 Stephanocolporat e
J1 Some or all colpi not meridional .................................................................. E13 Pericolporate
F1 Free pores present, no colpi (porate).
K3 Two pores .................................................................................................................. E14 Diporate
K2 Three pores ............................................................................................................... E15 Triporate
K1 More than three pores.
L2 Pores conned to a circular (sub-)equatorial belt ................................... E16 Stephanoporat e
L1 PoresÔuniformly distributed on the surface of the grain .............................. E17 Periporate
El Apertures fused to rings or spirals .......................................................................................... E18 Syncolpate
D1 With lacunae in a xed geometric pattern.
M2 Lacunae elongate, meridional (pseudocolpi) .................................................................... E19 Heterocolpate
M1 Lacunae not elongate .............................................................................................................. E20 Fenestrate
Key E1. Polyplicate
A2
A1
Grooves branched ............................................................................................................................ Ephedra distachya t.
EPHEDRACEAE
Grooves simple. Meridional ridges high, distinct in polar view. Fig. 3A .............................................. Ephedra fragilis
EPHEDRACEAE
Key E2. Inaperturate
Many cryptogram spores, cysts, etc., are also inaperturate. They are, on the whole, distinguished from pollen grains by the
(absence of ) structure of the wall. They are not included in the key.
A5 Striate ................................................................................................................................................... Arisarum vulgare
ARACEAE
A4 Echinate or verrucate.
B3 Echinate. Cf. also Lemnaceae, Nymphaea: Key E5 (Monoporate); and Nuphar, Stratiotes: Key E4 (Monocolpate)
Hydrocharis morsus-ranae
HYDROCHARITACEAE
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Pollen and spore keys, Pindos Mts, Greece
Fig. 2. Compound; Bilateral; Subtetrahedral (1000Ö except C). A, Typha angustifoli a type (e.g. T. latifolia); B, Epipactis type (e.g. E.
helleborine); C, Epilobium hirsutum type (e.g. E. hirsutum), 250Ö; D±E, Erica herbacea; F, Asplenium type (e.g. A. septentrionale); G,
Pteridium aquilinum (e.g. P. aquilinum var. esculentum).
Verrucate. Grain >40 mm. Obscure sulcus .................................................................................. Vallisneria spiralis
HYDROCHARITACEAE
B1 Verrucate. Grain <40 mm. Cf. Plantago, Key E17 (Periporate).
Scabrate or frustillate (Najas is scabrate, bilateral, but its exine is not usually preserved).
C2 Grain heteropolar, with thin areas ( lacunae). Cf. special key. ............................................................... Cyperaceae
B2
A3
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P. I. Chester and J. I. Raine
Isopolar, Ôspherical. No lacunae or perforations.
D5 Scabrate, tiny echinae. Grain 64±99 mm. ............................................................................................. Crocus
IRIDACEAE
D4 Scabrate, tiny echinae. Grain 40±46 mm. Exine 1±1.5 mm. ....................................................... Laurus nobilis
LAURACEAE
D3 Scabrate. Grain 40±52 mm. Exine c. 3 mm. .................................................................................... Aristolochia
ARISTOLOCHIACEAE
D2 Scabrate, micro-verrucate. Grain 18±25 mm. Fig. 3B ....................................................... Orobanche minor t.
OROBANCHACEAE
D1 Scabrate, with minute dark dots. Grain 22±30 mm. Fig. 3C ............................................................... Populus
SALICACEAE
Psilate or gemmate.
E2 Psilate.
F2 Grain more than 50 mm.
G2 Grain c. 60 mm. Obscurely trilete, wall simple or cavate with outer thin layer wrinkled. See also Key
D. Fig. 3D .............................................................................................................................. Equisetum
EQUISETACEAE
G1 Grain c. 90 mm ...................................................................................................... Pseudotsuga menziesii
PINACEAE
F1 Grain less than 50 mm. [Ceratophyllum (grain bilateral, c. 25±30 mm) keys out here, but exine is not usually
preserved. Similarly Juncaceae (spheroidal, very thin-walled)]. Psilate with tiny echinae (often lost). Araceae.
I2 Grain bilateral .................................................................................................................................. Arum
ARACEAE
I1 Grain spheroidal ................................................................................................................. Dracunculus t.
ARACEAE
El Gemmate.
J2 Fossil grains usually two-split. Gemmae scattered, deciduous, rounded, with constricted base, mostly lost in
fossil material. Cf. Pedicularis, Key E16. Fig. 3E±F ......................................................................... Juniperus
CUPRESSACEAE
J1 Gemmae rather crowded, size variable, not deciduous. Fig. 3G ................................................ Taxus baccata
TAXACEAE
Clavate-baculate or reticulate.
K2 Grain bilateral, elliptical. Reticulate ................................................................................... Callitriche obtusangula
CALLITRICHACEAE
K1 Grain rotational.
L3 Rugulose-baculate. Grain 21±32 mm ............................................................................. Limodorum abortivum
ORCHIDACEAE
L2 Isolated clavae/baculae in reticulate pattern .................................................................. Callitriche stagnalis t.
CALLITRICHACEAE
L1 Reticulum distinct.
M2 Columellae distinct.
N3 Diameter of lumina subequal to breadth of muri, some lumina larger (enclosing pores: Key
E17.2), muri simpli- or obscurely duplicolumellate .......................................................... Daphne
THYMELAEACEAE
N2 Diameter of lumina subequal to breadth of muri, muri clearly duplicolumellate. See also Key E5.
Fig. 3H .................................................................................................................... Cephalanthera
ORCHIDACEAE
N1 Diameter of lumina several times the breadth of muri. Potamogetonaceae.
O2 Columellae widely spaced, mostly conned to corners of brochi ... Potamogeton pectinatus
POTAMOGETONACEAE
O1 Columellae denser, also between corners of brochi. Fig. 3J .......... Potamogeton crispus t.
POTAMOGETONACEAE
M1 Columellae indistinct.
P2 Lumina c. 1±2 mm, muri simplicolumellate. Fig. 3I ........................................... Triglochin bulbosa
JUNCAGINACEAE
P1 Lumina c. 4 mm. No apparent columellae ..................................................... Zannichellia palustris
ZANNICHELLIACEAE
C1
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A2
A1
Key E3. Trichotomocolpat e
Triradiate scars are very common in cryptogam spores (cf. Key D). In angiosperm pollen they are found in some taxa, but
in the area none is known, apart from occasional faint traces in some grains of Trapa natans.
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Fig. 3.
(For legend see page 310).
Pollen and spore keys, Pindos Mts, Greece
309
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Key E4. Monocolpate
Compare also Key E5, Monoporate, the pore of which may become colpus-like by collapse of the grain.
A6 Verrucate. Verrucae dimorphic, smaller on the proximal side ................................................................ Gagea pratensis
LILIACEAE
A5 Clavate-baculate. Rounded clavae/baculae up to 2 mm long ......................................................................... Iris pumila
IRIDACEAE
A4 Echinate-baculate.
B2 Grains usually prolate. Echinate; echinae more than 5 mm long ..................................................... Nuphar luteum
NYMPHAEACEAE
B1 Grains subspherical. Baculate-echinate, large projections less than 5 mm long. Aperture sometimes indistinct .......
Stratiotes aloides
HYDROCHARITACEAE
A3 Rugulate to striate. Allium.
C2 Colpus extends to the proximal side of the grain.
D2 Grain more than 50 mm. Fig. 3M ......................................................................................... Allium roseum t.
LILIACEAE
D1 Grain less than 50 mm. Fig. 3L ............................................................................... Allium sphaerocephalon t.
LILIACEAE
C1 Colpus restricted to the distal side of the grain. Grain less than 50 mm. Fig. 3K ................ Allium paniculatum t.
LILIACEAE
A2 Reticulate (brochi equal to or more than 1 mm).
E2 Brochi more than 3 mm diameter.
F2 Heterobrochate. Margo very broad ..................................................................................... Anthericum liliago
LILIACEAE
F1 Homobrochate or brochi not falling into discrete size groups.
G2 Columellae Ôcylindrical. Extremely coarsely reticulate, brochi 4±18 mm ........................ Iris germanica
IRIDACEAE
G1 Muri consisting of distinct transverse elements. No free columellae in lumina. Margo present. Fig. 4A .
Lilium
LILIACEAE
E1 Brochi 1±3 mm diameter.
H3 Colpus extends to the proximal side of the grain. Muscari.
I2 Grain more than 40 mm ................................................................................................ Muscari comosum
LILIACEAE
I1 Grain less than 40 mm ............................................................................................. Muscari neglectum t.
LILIACEAE
H2 Colpus completely covers the distal pole. Colpus membrane covered with irregularly shaped islands of
columellate exine. Grain 44±55 mm ........................................................................... Gynandriris sisyrinchium
IRIDACEAE
H1 Colpus restricted to the distal side of the grain.
J2 Heterobrochate.
K2 Grains long, narrow.
L2 Few large brochi, mostly on the proximal side of the grain. Fig. 3N .......................... Scilla t.
LILIACEAE
L1 Lumina variable, rounded, muri about the same size as lumina ....... Narcissus papyraceus t.
AMARYLLIDACEAE
K1 Grain almost circular in polar view. Perforate-reticulate with muri and lumina almost the same size
over the distal pole, remainder of surface reticulate with lumina twice width of muri. Fig. 4D .....
Asphodelus stulosus t.
LILIACEAE
J1 Homobrochate.
M3 Grain less than 30 mm. Brochi c. 1 mm; reticulum decreases towards colpus Narthecium scardicum
LILIACEAE
M2 Grain 30±50 mm. Margo present. Muri narrow. Lumina up to 3 mm.
Fig. 3. Polyplicate; Inaperturate; Monocolpate (1000Ö). A, Ephedra fragilis; B, Orobanche minor type (e.g. O. minor); C, Populus (e.g. P.
nigra; inset: exine surface); D, Equisetum (e.g. E. arvense); E±F, Juniperus sp. (entire and split grains); G, Taxus baccata; H, Cephalanthera
(e.g. C. damasoniu m); I, Triglochin bulbosa (e.g. Triglochin sp.); J, Potamogeton crispus type (e.g. P. perfoliatus); K, Allium paniculatum type
(e.g. A. schoenoprasu m); L, Allium sphaerocephalon type (e.g. A. vineale); M, Allium roseum type (e.g. A. neapolitanu m); N, Scilla type (e.g.
S. autumnalis).
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Pollen and spore keys, Pindos Mts, Greece
A1
311
N2 Lumina in mesocolpium decrease sharply to a broad colpus margin that is clearly tectateperforate. Grain often circular in outline. Fig. 4B ................................. Butomus umbellatus
BUTOMACEAE
N1 Lumina decrease more gradually to colpus margin, which is not tectate-perforate, i.e. some
lumina are still detectable at the very edge. Grain more often elliptic. Fig. 4C .... Veratrum
LILIACEAE
M1 Grain more than 50 mm.
O2 Pollen often ruptured with ragged colpus edges. No margo. Free columellae in some brochi.
P3 Brochi 2±3 mm ......................................................................... Hermodactylus tuberosus
IRIDACEAE
P2 Brochi 1±2.5 mm ..................................................................................... Iris pseudacorus
IRIDACEAE
P1 Brochi 1±1.5 mm ........................................................................................... Iris spuria t.
IRIDACEAE
O1 Margo present. Columellae distinct. Fig. 4H .......................................................... Fritillaria
LILIACEAE
Microreticulate (brochi less than l mm), psilate or scabrate.
Q2 Colpus extends to the proximal side of the grain. Allium ± see above.
Q1 Colpus restricted to the distal side of the grain.
R2 Grain less than 30 mm.
S2 Columellae imperceptible.
T2 Brochi c. 1 mm .............................................................................................. Narthecium scardicum
LILIACEAE
T1 Psilate (perforate). Fig. 4E ........................................................................................... Galanthus t.
AMARYLLIDACEAE
S1 Columellae distinct. Brochi less than 1 mm ............................................................................. Asparagus
LILIACEAE
R1 Grain more than 30 mm.
U2 Grain shaped like an orange segment.
V4 Microreticulate, brochi c. 1 mm. Grain 50±60 mm ........................................................ Sternbergia
AMARYLLIDACEAE
V3 Perforate to reticulate, brochi <1 mm. Grain 25±50 mm ..................................... Narcissus tazetta
AMARYLLIDACEAE
V2 Perforate. Polar outline Ôcircular ....................................................................... Asphodelus albus
LILIACEAE
V1 Psilate to faintly perforate. Fig. 4G ........................................................................ Polygonatum t.
LILIACEAE
U1 Grain not segment-shaped.
W2 Reticulate, muri broader than width of lumina.
X2 Homobrochate ............................................................................................. Tulipa sylvestris
LILIACEAE
X1 Heterobrochate ............................................................................................ Ornithogalum t.
LILIACEAE
W1 Psilate-scabrate, perforate.
Y3 Grain about 60 mm. Psilate-perforate and micro-echinate. 2 opercula over colpus (giving
appearance of being 3-colpate) ........................................................................... Gladiolus t.
IRIDACEAE
Y2 Grain 47±70 mm. Granulate and perforate ................................................ Urginea maritima
LILIACEAE
Y1 Grain less than 50 mm.
Z2 Colpus edges ragged ............................................................................... Gagea lutea t.
LILIACEAE
Z1 Colpus edges not ragged. Fig. 4F ................................................... Convallaria majalis
LILIACEAE
Fig. 4. Monocolpate, continued (1000Ö). A, Lilium (e.g. L. martagon); B, Butomus umbellatus; C, Veratrum (e.g. V. album); D, Asphodelus
stulosus type (e.g. A. stulosus); E, Galanthus type (e.g. G. nivalis); F, Convallaria majalis; G, Polygonatum type (e.g. P. odoratum );
H, Fritillaria (e.g. F. lusitanica).
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Fig. 4.
312
(For legend see page 311).
P. I. Chester and J. I. Raine
Grana 40 (2001)
Pollen and spore keys, Pindos Mts, Greece
313
Key E5. Monoporate
A3
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A2
A1
Pore large, diameter more than half that of the grain, with operculum.
B2 Grain gemmate ................................................................................................................................ Nymphaea alba
NYMPHAEACEAE
B1 Grain granulate with echinae c. 1 mm projecting above densely crowded granules. See also Key E18. Fig 5A .......
Smilax
LILIACEAE
Pore diameter about half grain diameter. Grain reticulate or rugulate-verrucate.
C2 Grain reticulate. See also Key E2. Fig. 5D ...................................................................................... Cephalanthera
ORCHIDACEAE
C1 Grain with verrucae and short rugulae ................................................................................ Limodorum abortivum
ORCHIDACEAE
Pore smaller, diameter usually much less than half that of the grain.
D2 With distinct annulus. Grain tectate, psilate, scabrate, verrucate. Cf. special key ................................ Gramineae
D1 No annulus. Cf. Juniperus, Key E2.
E4 Grain clavate-baculate .................................................................................................. Callitriche stagnalis t.
CALLITRICHACEAE
E3 Grain psilate, densely perforate with lacunae, one of which forms a rudimentary pore at broad end of usually
pear-shaped grain. Cf. special key ................................................................................................ Cyperaceae
E2 Grain reticulate/rugulate, heterobrochate. Pore not always distinct.
F4 Microreticulate ( lumina <1 mm). Fig. 5B .............................................................. Sparganium erectum
SPARGANIACEAE
F3 Reticulate, lumina 1±2 mm ............................................................................ Sparganium angustifolium
SPARGANIACEAE
F2 Reticulate, lumina c. 2 mm ..................................................................................... Typha domingensis t.
TYPHACEAE
F1 Rugulate. Often in tetrads ( Key B1 ). Fig. 5C ........................................................ Typha angustifolia t.
TYPHACEAE
E1 Grain echinate, echinae c. 1.5 mm with broad conical base. Pore may be indistinct. Cf. algal cysts and other
microfossils of unknown origin ..................................................................................................... Lemnaceae
Key E6. Dicolpate
A2
A1
Columellae distinct. Per-reticulate. Margo present ............................................................................... Tamus communis
DIOSCOREACEAE
Columellae indistinct. No margo.
B2 Grain sometimes syncolpate. Grain densely microechinate .................................................................... Hypecoum
PAPAVERACEAE
(B1 Grain usually two-split (syncolpate), colpus edges ragged. Structure and sculpturing very faint or nil. Cf. Key
E18. Fig. 5E±F ...................................................................................................................................... Pedicularis
SCROPHULARIACEAE
Key E7. Tricolpate; Main key to subsections
A2
Tectate.
B5 Psilate (may be perforate tectum) ............................................................................................................
B4 Scabrate ....................................................................................................................................................
B3 Baculate, echinate, or verrucate ...............................................................................................................
B2 Rugulate-striate ........................................................................................................................................
B1 Supra-reticulate .........................................................................................................................................
A1 Semitectate or intectate.
C2 Per-reticulate .............................................................................................................................................
C1 Intectate (clavate, gemmate and baculate) ...............................................................................................
The crested grains of Trapa are not included in this key; see Key A.
Key
Key
Key
Key
Key
E7.1
E7.2
E7.3
E7.4
E7.5
Key E7.6
Key E7.7
Key E7.1 Tricolpate, psilate ( may be perforate tectum)
A3
A2
Colpi very short with nexinous ridges crossing underneath, never gaping ..................................... Monotropa hypopitys
PYROLACEAE
Colpus without equatorial constriction.
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B4
A1
Exine equal to or thicker than 3 mm. Columellae shorter and exine thinner next to colpus.
C2 Exine 4±6 mm thick. Columellae branched. Grain 45±67 mm. Fig. 5I .......................................... Convolvulus
CONVOLVULACEAE
C1 Exine c. 3 mm thick. Columellae simple.
D2 Grain >30 mm ............................................................................................................. Putoria calabrica
RUBIACEAE
D1 Grain # 30 mm. Cuscuta.
E2 In centre of mesocolpium sexine about as thick as nexine. Fig. 5J±K ......... Cuscuta epithymum t.
CONVOLVULACEAE
E1 In centre of mesocolpium sexine thicker than nexine ..................................... Cuscuta europaea t.
CONVOLVULACEAE
B3 Exine thinner in middle of intercolpium. Colpus narrow, straight, edges sharp. Grain c. 20 mm ........ Melampyrum
SCROPHULARIACEAE
B2 Exine 2±3 mm, even thickness. Edge of colpus clear, not ragged. Grain 21±30 mm ............... Phytolacca americana
PHYTOLACCACEAE
B1 Exine equal to or thinner than 2 mm. Edge of colpus ragged.
F2 Exine thinner next to colpus, micro-echinate. Columellae distinct and/or irregularly distributed.
G2 Colpi long.
H2 Grain globular-prolate. Tectum without perforations. Columellae present in colpus, grouped. See
also Key E7.2. Fig. 5G±H ............................................................................. Veronica triphyllos t.
SCROPHULARIACEAE
H1 Grain globular-oblate. No polar thickening of exine. Tectum with perforations. Columellae in
colpus, if present, scattered.
I2 Distinctly micro-echinate. Number of perforations and columellae approximately equal.
Columellae in a dense, regular pattern ........................................................................ Spergula
CARYOPHYLLACEAE
I1 Not echinate. Perforations fewer than columellae. Columellae irregularly distributed ...............
Spergularia t.
CARYOPHYLLACEAE
G1 Colpi short (poroid ) .................................................................................................. Corrigiola litoralis
CARYOPHYLLACEAE
F1 Exine same thickness throughout , c. 1 mm. Surface smooth.
J2 Grains oblate to spheroidal. Polar area small, index below 0.3.
K2 Grain c. 30 mm. Structure imperceptible ......................................................... Myricaria germanica
TAMARICACEAE
K1 Grain 40±50 mm. Structure distinct. Sometimes 4-colpate. Fig. 5N±O ............................ Euphrasia
SCROPHULARIACEAE
J1 Grains subglobular/prolate, 30±40 mm. Surface uniform.
L3 Supratectal granules grouped into circles. Fig. 5P±Q .................................................... Odontites t.
SCROPHULARIACEAE
L2 Supratectal granules grouped only as rugulae ....................................................... Bellardia trixago
SCROPHULARIACEAE
L1 Minutely supercially strio-reticulate. Incl. Orobanche purpurea ................................ Rhinanthus t.
SCROPHULARIACEAE
Colpus with equatorial constriction. Cf. Key E11.1.
M2 Colpus with prominent, undulating edges. Special key ........................................................................... Rosaceae
M1 Colpus edges at, straight.
N2 Microreticulate. Columellae distinct. See also Keys E7.6 & E11.5. Fig. 5L±M ............... Saxifraga stellaris
SAXIFRAGACEAE
N1 Exine not reticulate.
O2 Columellae imperceptible. Exine thin ................................................................................ Viola hirta t.
VIOLACEAE
O1 Columellae distinct.
P2 Columellae coarse ............................................................................................ Saxifraga bulbifera
SAXIFRAGACEAE
P1 Columellae irregularly dispersed, minute. Perforations scattered, irregular Myricaria germanica
TAMARICACEAE
Key E7.2 Tricolpate, scabrate
A2
Colpus with equatorial constriction.
B2 Costae colpi distinct. Equatorial limb triangular. Cornus.
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A1
Grain more than 40 mm, scabrate. Tectum perforate. See also Key E11.1. Fig. 6A ................... C. sanguinea
CORNACEAE
C1 Grain less than 35mm, with scattered microverrucae. Tectum imperforate ......................................... C. mas
CORNACEAE
B1 No distinct costae colpi. Columellae crowded, minute ........................................................... Myricaria germanica
TAMARICACEAE
Colpus without equatorial constriction.
D3 Nexine very thin. Sexine thicker than nexine, distinctly columellate, tectate. Usually syncolpate ........ Soldanella
PRIMULACEAE
D2 Nexine distinct. Exine thick, at any rate in the polar area.
E3 Exine thick in mesocolpia (3.5±5 mm); structure of sexine complex, with 2 layers of columellae. See also Key
E7.3. Fig. 6B±C .................................................................................................................................... Nigella
RANUNCULACEAE
E2 Exine more than 3 mm thick, crescentic. Nexine heavy. Columellae distinct, their mutual distance 1±2 times
diameter. Distinct, regularly and widely spaced microechinae. Colpus membrane with columella-like granules.
F2 Grain >40 mm. Some echinae larger than 1 mm. Exine thicker at poles (6 mm) .................. Centranthus
VALERIANACEAE
F1 Grain <40 mm. Only microechinae. Valerianella.
G2 Sexine as thick as nexine in mesocolpium; columellae more scattered and somewhat larger at poles
than elsewhere ............................................................................................... Valerianella rimosa t.
VALERIANACEAE
G1 Sexine thicker than nexine in mesocolpia; columellae not diVerentiated at poles. Fig. 6J±K .........
Valerianella locusta t.
VALERIANACEAE
E1 Exine thinner except in polar area.
H2 Exine sometimes thickest adjacent (but not actually at) poles; endocracks absent. Granulate colpus
operculum. Colpus edges straight. Fig. 6L ................................................................................ Teucrium
LABIATAE
H1 Exine thickest at poles; endocracks present. Colpus with granulate membrane, but not operculate.
I3 Colpus narrow, infolded, margins very prominent with adhering granules, traversed by prominent
endocracks; colpus membrane not usually visible. Columellae distinct ................................ Adonis
RANUNCULACEAE
I2 Colpus broad; margins ragged, not prominent, traversed by endocracks (but not prominently);
colpus membrane covered with numerous distinct granules. Columellae distinct ........ Consolida t.
RANUNCULACEAE
I1 Colpus broad, slightly sunken; margin ragged and indistinct, traversed by faint endocracks; colpus
membrane granular. Columellae indistinct. Fig. 6H±I ...................................................... Caltha t.
RANUNCULACEAE
D1 Not so.
J3 Without spinules.
K2 Exine perforate. Grain 27±36 mm. See also Key E7.1. Fig. 5G±H ........................ Veronica triphyllos t.
SCROPHULARIACEAE
K1 Not perforate. Grain 17-23 mm ............................................................................... Orobanche ramosa t.
OROBANCHACEAE
J2 Low ridges at base of spinules form a reticuloid pattern. Exine with multiple perforations. Spinules (scabrae)
regularly distributed. Detailed key to species within this type (P. dubium, P. hybridum, P. rhoeas, P.
somniferum) in NW2. See also Key E8. Fig. 6D±E .............................................................. Papaver rhoeas t.
PAPAVERACEAE
J1 No ridges between bases of spinules.
L2 Structure and sculpturing very ne. Grain oblate .................................................. Myricaria germanica
TAMARICACEAE
L1 Structure and sculpturing distinct. Spinules/verrucae irregularly distributed, size variable.
M3 Tectum Ô smooth with very scattered verrucae ............................................................... Glinus t.
MOLLUGINACEAE
Fig. 5. Monoporate; Dicolpate; Tricolpate psilate (1000Ö except I, N, O). A, Smilax (e.g. S. aspera); B, Sparganiu m erectum; C, Typha
angustifolia type (e.g. T. latifolia); D, Cephalanthera (e.g. C. damasoniu m); E±F, Pedicularis (e.g. P. verticillata), split and entire grains;
G±H, Veronica triphyllos type (e.g. V. serpyllifolius); I, Convolvulus (e.g. C. arvensis), 500Ö; J±K, Cuscuta epithymum type (e.g. C.
epithymum); L±M, Saxifraga stellaris; N±O, Euphrasia (e.g. E. salisburgensi s), 500Ö; P±Q, Odontites type (e.g. Parentucellia viscosa).
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Fig. 5.
(For legend
see page 315).
Pollen and spore keys, Pindos Mts, Greece
M2
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M1
317
Tectum without perforations. Columellae distinct, uniform, rather crowded. Colpus
narrow. Quercus.
N3 Indistinctly scabrate/rugulate. Polar diameter 22±28 mm. See also Key E7.4. Fig. 7D±F .......
Quercus coccifera t.
FAGACEAE
N2 Scabrate. Polar diameter 30±37 mm. Fig. 6M±N ................................................. Q. trojana t.
FAGACEAE
N1 Distinctly scabrate/verrucate. Fig. 7A±C ......................................................... Q. frainetto t.
FAGACEAE
Tectum with perforations, (sub-)verrucate-echinate. Colpus open, membrane granulate. Grains
often pericolpate, cf. Key E9. Columellae in open pattern, forming a bright area (``halo’’) around
each spinule. Perforations follow the same pattern. See key to species and groups in NW6. Incl.
Ranunculus p.p., Pulsatilla, Anemone, Ceratocephalus, & Clematis. Fig. 6F±G Ranunculus acris t.
RANUNCULACEAE
Key E7.3 Tricolpate; baculate, echinate or verrucate
A3
A2
A1
Baculate, baculae c. 2.5 mm long, in groups of 2 and 3. Exine 2±4 mm, tectum complete, infratectum columellate. Grain
42±60 mm ............................................................................................................................................ Plumbago europaea
PLUMBAGINACEAE
Echinate. Grain large, 40 mm or more.
B2 Echinae prominent, widely separated.
C2 Echinae 1±1.5 mm long, supported on shield-shaped verrucae. Valeriana.
D2 Echinae considerably shorter than wall thickness, height greater than width. Fig. 7G±I ........................
Valeriana oYcinalis t.
VALERIANACEAE
D1 Echinae about as wide as tall .................................................................................... Valeriana tuberosa
VALERIANACEAE
(C1 Echinae coarse, not on shield-shaped verrucae. Polar area large. Equatorial limb subtriangular. Lonicera.
Tricolporate: see Key E11.2.)
B1 Echinae smaller, of diVerent size classes. Exine 5±10 mm. Columellae branched. Colpus short, broad, cf. Key E15
(Triporate). Dipsacaceae p.p. NW2.
E3 Halo surrounding a very short colpus. See also Key E15. Fig. 7N±O .......................................... Dipsacus t.
DIPSACACEAE
E2 Structural and sculpturing elements regularly distributed. Exine thicker at poles. Fig. 7J±K ..........................
Scabiosa columbaria t.
DIPSACACEAE
E1 Not so .................................................................................................................................... Succisa pratensis
DIPSACACEAE
Grains smaller.
F3 Echinae conical, low. Tectum thick. Colpi short, indistinct. Compositae, see also Key E11.2. Fig. 7L±M .............
Xanthium
COMPOSITAE
F2 Grains with prominent mixed echinae and microechinae, irregularly and widely spaced ............... Actaea spicata
RANUNCULACEAE
F1 Grains microechinate, echinae evenly distributed. See also Key E7.2 ......................................................... Nigella
RANUNCULACEAE
Key E7.4 Tricolpate, rugulate or striate
A2
A1
Exine very thick, 5±7 mm. Striae coarse, multilevelled. Colpi wide. Fig. 8A±B .................................................. Erodium
GERANIACEAE
Exine thickness and striation normal and ne.
B2 Exine with or without perforations. Edges of colpi undulating, often prominent. Rosaceae, see special key.
B1 Exine perforate. Edges of colpi straight or ragged.
Fig. 6. Tricolpate scabrate (1000Ö). A, Cornus sanguine a; B±C, Nigella (e.g. N. sativa); D±E, Papaver rhoeas type (e.g. P. rhoeas); F±G,
Ranunculus acris type (e.g. R. repens); H±I, Caltha type (e.g. C. palustris); J±K, Valerianella locusta type (e.g. V. carinata); L, Teucrium (e.g.
T. scordium); M±N, Quercus trojana type (e.g. Q. cerris).
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C3
C2
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C1
319
Very nely rugulate-striate. Fig. 8C±D ............................................................. Veronica anagallis-aquatic a t.
SCROPHULARIACEAE
Rugulate±scabrate. Vallae short.
D2 Vallae in reticuloid pattern, may be perfectly reticulate. Colpus margins ragged. See also Keys E7.6 &
E8. Fig. 8E±F ................................................................................................................................ Oxalis
OXALIDACEAE
D1 Vallae very faint. Colpus margins straight. See also Key E7.2. Fig. 7D±F ............ Quercus coccifera t.
FAGACEAE
Distinctly striate.
E3 Sculpturing ne, striae predominantly meridional. Grain 30 mm or more. Fig. 8L±M ...............
Acer
ACERACEAE
E2 Direction of striae variable.
F2 Grain smaller than 25 mm. Sculpturing ne ........................................................ Trollius europaeus
RANUNCULACEAE
F1 Grain more than 25 mm. Sculpturing coarse. Refer NW2. See also Key E11.3. Fig. 8J±K ..............
Saxifraga oppositifolia t.
SAXIFRAGACEAE
E1 Striation diVuse. Vallae in very low relief. S. granulata type of NW2.
G2 Vallae with granulae ................................................................................. Saxifraga tridactylites t.
SAXIFRAGACEAE
G1 No granulae on vallae ............................................................................ Saxifraga sempervivum t.
SAXIFRAGACEAE
Key E7.5 Tricolpate, supra-reticulate
A2 Colpus without equatorial constriction or poroid area. No costae.
B2 Meridional limb compressed oval, grain subcylindric, usually prolate-perprolate. Colpus narrow, not boatshaped.
C3 Muri broader than lumina; lumina of even size, c. 1 mm .................................................................. Acanthus
ACANTHACEAE
C2 Muri narrow; all brochi large, c. 1.5 mm ........................................................................................ Onobrychis
LEGUMINOSAE
C1 Muri narrow, brochi small, c. 0.5 mm, or a median eld of small brochi in each intercolpium ........................
Hedysarum coronarium
LEGUMINOSAE
B1 Meridional limb circular, oval or rhomboidal, not compressed. Colpi boat-shaped, often ruptured. Labiatae p.p.
D2 LuminaÔuniform in size all over the grain (may be so small as to be microreticulate).
E3 ÔMicroreticulate. Grain >25 mm. Fig 8I ................................................................. Stachys sylvatica t.
LABIATAE
E2 Reticulate.
F2 Thick muri. Lumina somewhat variable in size, singly perforate (punctae c. 1 mm). Grain
<25 mm. Fig. 8G±H ....................................................................................................... Marrubium
LABIATAE
F1 Thin muri (<0.5 mm). Grain >25 mm.
G2 Baculae in lumina. Fig. 8O ............................................................................ Stachys arvensis
LABIATAE
G1 Numerous punctae perforating each lumina. Fig. 8N ....................................... Stachys recta
LABIATAE
E1 Doubly reticulate ........................................................................................................................ Phlomis
LABIATAE
D1 Lumina largest in the centre of each mesocolpium decreasing in size gradually towards colpi. Fig. 8P .........
Scutellaria t.
LABIATAE
A1 Equatorial part of colpus constricted or ruptured. Costae interrupted at equator.
H2 Poroid area large, usually ruptured. Branches of costae colpi short. ......................................................... Genista t.
LEGUMINOSAE
Fig. 7. Tricolpate scabrate (continued); Tricolpate echinate (1000Ö except G±I, J±K, N±O). A±C, Quercus frainetto type (e.g. Q. robur);
D±F, Quercus coccifera type (e.g. Q. coccifera); G±I, Valeriana oYcinalis type (e.g. V. oYcinalis), 500Ö; J±K, Scabiosa columbaria type
(e.g. S. atropurpure a), 500Ö; L±M, Xanthium (e.g. X. strumarium); N±O, Dipsacus type (e.g. D. fullonum), 500Ö.
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Pollen and spore keys, Pindos Mts, Greece
321
H1 Poroid area small. Branches of costae colpi long. See Key E11.5 ........................................................... Hypericum
GUTTIFERAE
Key E7.6 Tricolpate, per-reticulate
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(A3
Tri-sulcate, one sulcus oval and occupying the distal pole of the grain, the other two semi-circular and located at the
margins of the proximal face ............................................................................................................. Tulipa praecox t.)
LILIACEAE
A2 Colpi without equatorial constriction.
B2 Grain large, more than 50 mm, thick-walled.
C3 Finely and uniformly reticulate, lumina c. 1 mm.
D2 Colpus membrane granulate ............................................................................... Leontice leontopetalum
BERBERIDACEAE
D1 Not so. Muri broader than lumina .......................................................................................... Acanthus
ACANTHACEAE
C2 Coarsely reticulate, brochi 4±14 mm. Columellae not branched, the heads standing above the muri spinulose .
Armeria maritima t.
PLUMBAGINACEAE
C1 Columellae branched, fused to a reticulum some distance below their heads, forming coarse clavae on top. See
key to groups in NW6. Fig. 9A±B ................................................................................................... Geranium
GERANIACEAE
B1 Grain smaller.
E2 Brochi small, less than 1 mm (microreticulate).
F2 Heterobrochate, brochi larger in mesocolpia.
G2 Colpi long, margins diVuse, membrane rugate Ôgranules. Ôcircular in polar view, prolate.
Infratectum clearly columellate ............................................................................................ Reseda
RESEDACEAE
G1 Colpi short. Angulaperturate, triangular in polar view; suboblate .................... Comandra elegans
SANTALACEAE
F1 Homobrochate.
H2 Grain larger than 25 mm.
I2 Exine c. 4 mm thick ...................................................................................... Cuscuta monogyna
CONVOLVULACEAE
I1 Exine 2 mm or less. Colpus margin indistinct. Colpus membrane granulate. Columellae
indistinct .................................................................................................... Chelidonium majus
PAPAVERACEAE
H1 Grain smaller than 25 mm.
J2 Exine thicker than 1 mm, thinner at colpus margin and in apocolpia. Polar exine not
diVerentiated. Columellae distinct in surface view.
K3 Exine 2±4 mm thick. Colpus margin distinct. Colpus membrane not granulate. .. Arabis
CRUCIFERAE
K2 Exine 2 mm or less. Colpus margin distinct. Colpus membrane not granulate. Fig. 9C±E
Hornungia t.
CRUCIFERAE
K1 Exine 1±3 mm. Colpus membrane granulate. Muri broad ................................. Tamarix
TAMARICACEAE
J1 Exine 1 mm or less, tectate-perforate at poles. Columellae indistinct in surface view.
L2 Exine c. 1 mm thick. Polar area tectate-perforate, not thickened or reticulate. See also
Keys E7.1 & E11.5 ............................................................................. Saxifraga stellaris
SAXIFRAGACEAE
L1 Exine less than 1 mm thick. Polar area exine thicker, tectate with few perforations.
Obscurely tricolporate, with 2±6 endocolpi ...................... Chrysosplenium alternifolium
SAXIFRAGACEAE
E1 At least some brochi more than 1 mm.
M2 Colpus long.
N2 Heterobrochate, brochi reduced in size towards colpi.
O2 Margo distinct, semitectate. Brochi c. 2 mm. GrainÔspheroidal. Fig. 9O±P ........ Helleborus
RANUNCULACEAE
O1 Margo thin, tectate.
P2 Endoaperture apparent. Grain prolate.
Q2 Brochi 1.5±2 mm. Grain 16±20 mm. See also Key E11.5. Fig. 9I±J . Salix triandra
SALICACEAE
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Fig. 8. Tricolpate rugulate/striate and supra-reticulate (1000Ö except A±B, J±K ). A±B, Erodium (e.g. E. cicutarium), 500Ö; C±D, Veronica
anagallis-aquatica type (e.g. V. anagallis-aquatic a); E±F, Oxalis (e.g. O. acetosella); G±H, Marrubium (e.g. M. vulgare); I, Stachys sylvatica
type; J±K, Saxifraga oppositifoli a type (e.g. S. paniculat a); L±M, Acer (e.g. A. pseudoplatanu s), 500Ö; N, Stachys recta; O, Stachys arvensis;
P, Scutellaria type (e.g. Ajuga genevensis).
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Q1
A1
Brochi 2±3 mm. Grain 16±27 mm. See also Key E11.5. Fig. 9M±N Salix fragilis t.
SALICACEAE
P1 Endoaperture very indistinct. GrainÔspheroidal. See also Keys E7.4 & E8 ..... Oxalis
OXALIDACEAE
N1 Brochi not reduced in size towards colpi. No distinct margo. Columellae very distant, only their
heads joined into a reticulum.
R2 Exine rather thick. Columellae clear in top view, length decreases next to colpi. Reticulum
with narrow muri. Colpus without granules or with a delicate structure. Fig 9F±H .............
Sinapis t.
CRUCIFERAE
R1 Muri broad. Colpus with scattered granules ........................................................... Glaucium
PAPAVERACEAE
M1 Colpus short. Polar area rather large, index c. 0.4.
S2 Colpus broad, with scattered granules. Grain c. 22 mm. Fig. 9K±L .................. Platanus orientalis
PLATANACEAE
S1 Colpus narrow, covered with granules, broader towards equator, geniculate. Fraxinus.
T2 Equatorial outline angular, sides sometimes nearly straight. Nexine about as thick as sexine.
Reticulum ne, brochiÔ1 mm. Grain 18.5 ±23.5 mm ............................... Fraxinus angustifolia
OLEACEAE
T1 Equatorial outline circular. Nexine distinctly thinner than sexine. Reticulum slightly
coarser, brochi >1 mm. Grain 20±29.5 mm .............................................. Fraxinus excelsior t.
OLEACEAE
Colpi with equatorial constriction. See Key E11.5 (Tricolporate).
Key E7.7 Tricolpate, intectate ( clavate, gemmate and baculate)
A2
A1
Size of clavae variable.
B2 Intercolpium and polar area with scattered, large, free clavae in addition to many small ones, more or less fused.
Colpus not distinct.
C2 Grain c. 60 mm. Equatorial outline circular. Fig. 10C±D ......................................................... Viscum album
LORANTHACEAE
C1 Grain c. 20 mm. Equatorial outlineÔtriangular .......................................................... Arceuthobium oxycedri
LORANTHACEAE
B1 All clavae free. Small clavae scattered beneath the large ones. Near the colpus there are only small clavae, forming
a margo. See also Key E11.6. Fig. 10A±B ....................................................................................... Ilex aquifolium
AQUIFOLIACEAE
Clavae dimorphic or uniform. Linaceae.
D2 Clavae dense, almost contiguous, ne, 1±1.5 mm, dimorphic, the smaller ones dominant. Fig. 10E±F ....................
Linum usitatissimum t.
LINACEAE
D1 Clavae /gemmae distinctly separate.
E2 Grain smaller than 25 mm. Colpus margins regular, not appearing ruptured. Gemma of Ôuniform diameter
Radiola linoides
LINACEAE
E1 Grain large, >35 mm. Colpus margins very irregular, ruptured. If dimorphic, both size classes subequally
represented.
F3 Grain c. 80 mm. Clavae uniform. [Rogers 1984 suggests pericolpate] ..................... Linum hologynum t.
LINACEAE
F2 Grain smaller, c. 50-65 mm. Endocolpus lolongate, broad, distinct ......................... Linum austriacum t.
LINACEAE
F1 Grain 35±70 mm. Endoaperture not, or hardly visible, except at ends. Includes L. catharticum, L.
tenuifolium, L. trigynum, L. strictum, L. hirsutum ± See detailed key to groups and species in NW3.
Fig. 10G±H ...............................................................................................................................................
Linum catharticum t.
LINACEAE
Key E8. Stephanocolpat e
A2
Psilate-scabrate.
B3 Grain large, 60±80 mm. 9±11 broad colpi. Finely psilate-scabrate .............................................. Sesamum indicum
PEDALIACEAE
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P. I. Chester and J. I. Raine
Fig. 9. Tricolpate per-reticulate (1000Ö except A±B, C±E ). A±B, Geranium (e.g. G. robertianum), 500Ö; C±E, Hornungia type (e.g. Alyssum
murale), 500Ö; F±H, Sinapis type (e.g. Cardamine hirsuta); I±J, Salix triandra; K±L, Platanus orientalis; M±N, Salix fragilis type (e.g. S.
fragilis); O±P, Helleborus (e.g. H. hyemalis).
B2
B1
Grain medium size.
C2 25±28 mm. 11±12 colpi. Fig. 10K ......................................................................................... Sherardia arvensis
RUBIACEAE
C1 20±35 mm. 4 or 6 colpi. Detailed key to species within this type (P. dubium, P. hybridum, P. rhoeas, P.
somniferum) in NW2. See also Key E7.2 ............................................................................. Papaver rhoeas t.
PAPAVERACEAE
Grain small.
D2 Grain 15±29 mm. 6±10 narrow colpi. Ornament of circular perforations and scabrae. Fig. 10Q±R ...............
Galium t.
RUBIACEAE
D1 Grain 15±21 mm. C. 6 colpi. Ornament of small circular perforations regularly distributed, together with
paired rugae giving a unique band and point pattern ....................................................................... Valantia
RUBIACEAE
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A1
325
Reticulate.
E2 Polar area large, index higher than 0.6.
F3 Supra-reticulate; no columellae discernible in muri. Colpi 4. Grain oblate in equatorial view, subrectangular
in polar view .................................................................................................................. Impatiens noli-tangere
BALSAMINACEAE
F2 Reticulate-perforate. Colpi 4. Grain obtuse-rectangular or quadrangular in equatorial and polar views ........
Sideritis
LABIATAE
F1 Per-reticulate.
G3 5±10 colpi .................................................................................................................................... Primula
PRIMULACEAE
G2 4±6 colpi. Usually tricolpate, see Keys E7.4 & E7.6 .................................................................... Oxalis
OXALIDACEAE
G1 4 colpi.
H2 Columellae indistinct ...................................................................................... Fraxinus angustifolia
OLEACEAE
H1 Muri simpli- or duplicolumellate ..................................................................... Fraxinus excelsior t.
OLEACEAE
E1 Polar area medium-small, index lower than 0.5.
I2 Supra-reticulate.
J2 6 or more colpi. Free columellae in lumina. Fig. 10L .............................................................. Prunella t.
LABIATAE
J1 4 colpi. No free columellae ( lumina punctate) .......................................................... Stachys angustifolia
LABIATAE
I1 Per-reticulate. Columellae restricted to muri.
K3 Muri scabrate. Microreticulate ................................................................................................ Crucianella
RUBIACEAE
K2 Muri beaded.
L2 Reticulum double (cf. Valdes et al. 1987). Fig. 10M±N ...................................... Salvia oYcinalis t.
LABIATAE
L1 Reticulum single, Ôrugulate. Fig. 10O±P .............................................................. Salvia verticillata
LABIATAE
K1 Muri not beaded. Fig. 10I±J ..................................................................................................... Mentha t.
LABIATAE
Key E9. Pericolpate
A3
A2
A1
Reticulate.
B2 Lumina deep, with free, coarse clavae. 30 short colpi in dodecahedral arrangement .............. Polygonu m amphibium
POLYGONACEAE
B1 No free elements in lumina. 4±6 colpi. Cf. Tricolpate, Keys E7.5 and E7.6.
Echinate. Echinae small, irregularly distributed. Tectum perforate. Central area of intercolpium with coarse columellae
in reticuloid pattern. 12 colpi.
C2 Exine thickest in mesocolpium, thinning towards colpi. See also Key E18. Fig. 10S±T ................ Montia fontana
PORTULACACEAE
C1 ExineÔeven thickness ................................................................................................................ Portulaca oleracea
PORTULACACEAE
Psilate, microechinate, scabrate, verrucate.
D3 Irregularly distributed spinules, verrucae or scabrae of varying size. Tectum with perforations (phase, SEM ):
some Papaveraceae, Ranunculaceae more usually inaperturate or tricolpate ( Keys E2, E7 ) may key out here.
E2 Psilate. 6- or 12-colpate, usually syncolpate ................................................................... Corydalis ochroleuca
PAPAVERACEAE
E1 Verrucate. 6-colpate, not syncolpate ................................................................................... Corydalis solida t.
PAPAVERACEAE
Fig. 10. Tricolpate intectate; Stephanocolpate; Pericolpate (1000Ö except E±F, G±H, I±J, M, N±O). A±B, cf. Ilex aquifolium (e.g. I.
latifolia); C±D, Viscum album; E±F, Linum usitatissimum type (e.g. L. bienne), 500Ö; G±H, Linum catharticum type (e.g. L. catharticum),
500Ö; I±J, Mentha type (e.g. M. aquatica), 500Ö; K, Sherardia arvensis; L, Prunella type (e.g. P. vulgaris), 500Ö; M±N, Salvia oYcinalis
type (e.g. M: S. verbenacea, N: Salvia glutinosa), 500Ö; O±P, Salvia verticillata, 500Ö; Q±R, Galium type (e.g. G. verum); S±T, Montia fontana.
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327
D2 Psilate-scabrate with uniform minute spinules and perforations ............................................................... Spergula
CARYOPHYLLACEAE
D1 Psilate, spinules absent ....................................................................................................................... Spergularia t.
CARYOPHYLLACEAE
Key E10. Dicolporate
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Prolate, equatorially constricted (bone-shaped ). Colpi 2/3 length of grain; ora rectangular, lalongate. Exine tectate, psilate.
Includes Tordylium maximum ........................................................................................................................... Echinophora t.
UMBELLIFERAE
Key E11. Tricolporate: main key to subsections
The transition colpate±colpate with equatorial constriction±colporate is not easy to dene in categorical terms. Consultation
of Key E7 is recommended if Key E11 does not lead to a result.
A3 Tectate.
B4 Psilate or scabrate .................................................................................................................................... Key E11.1
B3 Echinate ................................................................................................................................................... Key E11.2
B2 Striate or rugulate ................................................................................................................................... Key E11.3
B1 Foveolate or supra-reticulate ................................................................................................................... Key E11.4
A2 Semitectate. Per-reticulate ............................................................................................................................... Key E11.5
A1 Intectate. Clavate ............................................................................................................................................. Key E11.6
Key E11.1 Tricolporate. Psilate-scabrate
A2
A1
With costae aequatoriales and transverse colpi.
B4 Equatorial limb intersemiangular. Exine more than 4 mm thick, crescentic. Columellae coarse, branched. Centaurea
sect. Cyanus.
C2 Meridional limb compressed oval. Costae aequatoriales sharply projecting in meridional optical section.
Fig. 11A±C .................................................................................................................................... C. cyanus t.
COMPOSITAE
C1 Meridional limb oval. Costae aequatoriales less prominent. Fig. 11D .................................... C. napulifera t.
COMPOSITAE
B3 Equatorial limb subcircular. Exine thinner, not crescentic. Columellae ner. Costae aequatoriales less prominent.
Grain 22±39 mm, colpi long. See also Key E13. Fig. 11H±J ............................................... Polygonu m aviculare t.
POLYGONACEAE
B2 Equatorial limb intermediate between B4 and B3. Exine slightly crescentic ........................................ Bilderdykia
POLYGONACEAE
B1 Equatorial limb triangular. Exine 1.25±1.5 mm, not crescentic, short columellae. Costae aequatoriales stronger
near ectoapertures, colpi short and narrow with parallel sides, often forking slightly at ends. Grain 45±56 mm. See
also Key E12 ............................................................................................................................. Anchusa arvensis t.
BORAGINACEAE
No costae aequatoriales.
D3 Heteropolar. Exine thin, psilate or punctate, columellae not discernible.
E2 Pear-shaped.
F2 Grain 10±16 mm. Exine with scattered punctae <1 mm bordering the apertures ................... Alkanna t.
BORAGINACEAE
F1 Grain 17±24 mm. Exine comprehensively but faintly micro-suprareticulate. See also Key E11.4.
Fig. 11N±O .................................................................................................................................. Echium
BORAGINACEAE
E1 GrainÔspheroidal, one apocolpium very small, the other larger. Exine psilate.
G2 Grain >15 mm. Colpi constricted at equator; endoapertures indistinct. Fig 11P±Q . Gratiola oYcinalis
SCROPHULARIACEAE
G1 Grain 10±15 mm. Costae colpi present; endoapertures large and distinct, 8-shaped. Fig. 11R±T ...........
Cyclamen
PRIMULACEAE
D2 Meridional limb compressed oval.
H2 Grain distinctly prolate.
I2 Grain larger than 20 mm. Exine >1 mm. For detailed key see NW4; additional taxa described by
Cerceau-Larrival (1963 ± Tordylium oYcinale; 1967 ± Cachrys); van Zeist & Bottema (1977 ± Malabaila) .
Figs. 11E±G & K±M ........................................................................................................... Umbelliferae
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I1 Grain <20 mm. Exine <1 mm ...................................................................................... Androsace villosa
PRIMULACEAE
H1 Not so distinctly prolate.
J2 Exine more than 3 mm thick. Grain more than 50 mm, dimorphic, one type with denser structure. Tectum
with scattered perforations. Columellae branched, coarser at poles.
K2 Columellae evenly distributed, gaps between columellae less than their diameters .........................
Polygonum viviparum
POLYGONACEAE
K1 Columellae irregularly distributed, more widely spaced ................................ Polygonum bistorta t.
POLYGONACEAE
J1 Exine thin.
L3 Grain more than 30 mm. Pore edges distinctly protruding. Scabrate, polar area suprareticulate ......
Anthyllis
LEGUMINOSAE
L2 Grain 20±30 mm.
M2 Pore edges distinctly protruding. Fig. 11U±W ...................................................... Dorycnium
LEGUMINOSAE
M1 Pores not protruding ........................................................................... Ornithopus compressus
LEGUMINOSAE
L1 Grain less than 20 mm, psilate. Fig. 11D¾±E¾ ....................................................................... Lotus t.
LEGUMINOSAE
D1 Meridional limb oval or circular.
N2 Intercolpium attened or concave in polar view.
O2 Exine thicker and columellae coarser at poles. Colpus narrow, slit-shaped. Grain subglobular ............
Hydrocotyle vulgaris
UMBELLIFERAE
O1 Thickness of exine uniform. Colpus not slit-shaped.
P2 With transverse colpus.
Q2 Scabrate. Grain c. 35±45 mm, angulaperturate. Tectum perforate.
R2 Longitudinally oriented rugulae. .............................................................. Rhus coriaria
ANACARDIACEAE
R1 Micro-foveolate/fossulate. Equatorial sexinal bridge over colpus ...... Styrax oYcinalis
STYRACACEAE
Q1 Psilate. Tectum not perforate.
S2 Grain more than 20 mm.
T2 Irregular outline. H-shaped endoaperture. Fig. 11B¾±C¾ ............................ Verbena
VERBENACEAE
T1 Smooth outline. Endoaperture not H-shaped, fastigiate. Fig. 11A¾ .........................
Solanum nigrum t.
SOLANACEAE
S1 Grain smaller than 20 mm. Endoaperture fastigiate. Fig. 11F¾±G¾ .. Solanum dulcamara
SOLANACEAE
P1 No transverse colpus.
U2 Equatorial limb semilobate. Psilate ............................................................................ Frangula
RHAMNACEAE
U1 Equatorial limb semiangular. Scabrate.
V3 Grain 58±64 mm, subprolate, circular equatorial outline. Colpi long. Exine c. 3 mm,
tectum perforate .............................................................................. Lagenaria siceraria
CUCURBITACEAE
V2 Grain more than 40 mm, angulaperturate. See also Key E7.2 ............ Cornus sanguinea
CORNACEAE
V1 Grain c. 25 mm, angulaperturate .................................................................. Cornus mas
CORNACEAE
N1 Intercolpium convex.
W2 Pore edges distinctly protruding.
X2 Endopore equatorially elongated.
Y2 Grain 23 mm or more.
Z2 Psilate to sparsely punctate ......................................................................... Osyris alba
SANTALACEAE
Z1 Densely punctate/microreticulate (see E11.5 ) .................................. Ricinus communis
EUPHORBIACEAE
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Y1
W1
Grain less than 25 mm.
A¾3 Microechinate, vermiculate ........................................................................ Filipendula
ROSACEAE
A¾2 Psilate, tectum not perforate. Fig. 11X±Z ........................................... Castanea sativa
FAGACEAE
A¾1 Scabrate or indistinctly suprareticulate.
B¾2 Operculate. Fig. 25A±C ......................................................... Sanguisorba minor
ROSACEAE
B¾1 No operculum. Incl. Medicago sativa p.p. See also Key E11.4. Fig. 14M±N ......
Trifolium montanum t.
LEGUMINOSAE
X1 Endopore circular. Colpus slit-shaped.
C¾2 With vestibulum. Colpus very short, about twice the diameter of the pore ..........................
Ludwigia palustris
ONAGRACEAE
C¾1 No vestibulum.
D¾2 Microechinate, columellae in vermiculoid pattern .................................... Filipendula
ROSACEAE
D¾1 Psilate. Colpi very short. Grain spheroidal to oblate ............. Elaeagnus angustifolia
ELAEAGNACEAE
Edges of pore not protruding, endopore meridionally elongated, isodiametric or indistinct.
E¾2 Grain perforate. Colpus slit-shaped, endopore small, completely covered.
F¾2 With costae endopori. See key to species and groups in NW5. Incl. Oxyria digyna. See also
Keys E11.5 & E13. Fig. 11H¾±I¾ ............................................................................ Rumex t.
POLYGONACEAE
F¾1 No costae endopori.
G¾2 Grain 33±45 mm .............................................................................. Spartium junceum
LEGUMINOSAE
G¾1 Grain 16±19 mm. Colpus interrupted at equator ............................ Samolus valerandi
PRIMULACEAE
E¾1 Not so.
H¾2 Distinctly prolate. Grain <30 mm.
I¾2 Scabrate.
J¾2 Scabrate/suprareticulate. Equatorial limb circular with deeply intruding colpi.
Colpi long, wide, with densely granulate membrane. Exine thins towards colpi.
Globularia
GLOBULARIACEAE
J¾1 Scabrate/granulate. Intercolpia attened or slightly convex. Colpi long and
narrow, with psilate membrane ................................................... Diospyros lotus
EBENACEAE
I¾1 Psilate. Crassulaceae.
K¾2 Grain >27 mm. Colpus membrane slightly granulate ........... Jovibarba heuVelii
CRASSULACEAE
K¾1 Grain <27 mm .................................................................................... Umbilicus
CRASSULACEAE
H¾1 Subprolate to spheroidal.
L¾2 Colpi long, broad. Psilate to scabrate. H-endoaperture with equatorial bridge of
sexine over colpus. Colpus membrane with coarse granules; margins not intruding,
irregular .............................................................................................. Robinia pseudoacacia
LEGUMINOSAE
L¾1 Colpi short, slit-like.
M¾2 Grain 44±50 mm. Scabrate/rugulate. Endoaperture circular, with thickened sexine
(marginate). Fig. 11J¾±L¾ .................................................................................. Fagus
FAGACEAE
Fig. 11. Tricolporate psilate/scabrate (1000Ö except D, J¾±L¾). A±C, Centaurea cyanus type (e.g. C. cyanus); D, Centaurea napulifera type
(e.g. C. triumfetti), 500Ö; E±G, Umbelliferae (e.g. Sison amomum); H±J, Polygonum aviculare type (e.g. P. aviculare); K±M, Umbelliferae
(e.g. Bupleurum lancifolium); N±O, Echium (e.g. E. vulgare); P±Q, Gratiola oYcinalis; R±T, Cyclamen (e.g. C. hederifolium); U±W,
Dorycnium (e.g. D. graecum); X±Z, Castanea sativa; A¾, Solanum nigrum type (e.g. S. nigrum); B¾±C¾, Verbena (e.g. V. oYcinalis); D¾±E¾,
Lotus type (e.g. L. aegaeus); F¾±G¾, Solanum dulcamara; H¾±I¾, Rumex type (e.g. R. acetosella); J¾±L¾, Fagus (e.g. Fagus sylvatica), 500Ö.
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Pollen and spore keys, Pindos Mts, Greece
M¾1
331
Grain 20±28 mm. Scabrate/granulate. Endoaperture circular, with thickened
nexine (costate) .................................................................................... Emex spinosa
POLYGONACEAE
Key E11.2 Tricolporate. Echinate
This Key mainly comprises Compositae Asteroideae. Their exine is in typical cases multi-layered and consists of, from the
inside outwards: (1) foot layer = nexine; sexine composed of (2) inner columellae layer, usually with scattered, coarse, often
branched columellae; (3) paratectum formed by partial fusion of heads of the inner columellae; (4) outer columellae layer,
columellae thinner and more numerous than in the inner layer; (5) (outer) tectum, which is very thin and densely perforated.
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A2
A1
Columellae straight, not branched.
B2 Diameter of echinae and columellae equal. Echinae few, scattered. Lonicera.
C2 Sexine about as thick as nexine, columellae in surface view rather indistinct. Grain usually smaller than
60 mm ....................................................................................................................................... L. xylosteum t.
CAPRIFOLIACEAE
C1 Sexine much thicker than nexine, columellae in surface view distinct. Grain usually larger than 60 mm.
D2 Echinae up to 2.5 mm. Ends of ectocolpi usually obtuse ..................................................... L. alpigena
CAPRIFOLIACEAE
D1 Echinae up to 1.5 mm. Ends of ectocolpi usually acute. Fig. 12A±B ......................... L. periclymenum t.
CAPRIFOLIACEAE
B1 Echinae much coarser than columellae, conical, broad at base. Short colpi, prominent endopores ........ Abutilon t.
MALVACEAE
Columellae branched, echinae conical from base. Asteroideae.
E2 Columellae indistinct in optical section. Fig. 12E±F .............................................................................. Solidago t.
COMPOSITAE
E1 Columellae prominent in optical section.
F2 Echinae at least partly joined by low ridges.
G2 Echinae joined by smooth low ridges which form a coarse reticulum .......................... Picnomon acarna
COMPOSITAE
G1 Echinae large, partly conuent at bases. Inner columella layer variable, thicker under echinae ...............
Centaurea amplifolia
COMPOSITAE
F1 Echinae discrete, not joined by ridges.
H2 Inner columella layer equal to or thicker than the outer.
I3 Echinae vestigial.
J2 Grains large, prolate with longitudinal thickenings of intercolpia (intersemiangulate).
Fig. 12L±M ................................................................................................................. Echinops
COMPOSITAE
J1 Grains small, spherical to oblate. Fig. 12N±O ......................................................... Artemisia
COMPOSITAE
I2 Echinae blunt, length equal to or smaller than basal diameter. Prolate. Fig. 12C±D ...... Carlina t.
COMPOSITAE
I1 Echinae long, sharp. Intercolpium in optical section distinctly crescentic. Fig. 12G±H .. Achillea t.
COMPOSITAE
H1 Inner columellae layer thinner than outer.
K2 Echinae vestigial .............................................................................................. Centaurea graeca t.
COMPOSITAE
K1 Echinae distinct.
L2 Grains c. 30 mm or smaller. Spherical. Colpi short. Echinae short (1±2 mm). See also
Key E7.3 .................................................................................................................. Xanthium
COMPOSITAE
L1 Grains larger. Colpi long.
M3 Prolate. Spines short. Fig. 12I ................................................... Centaurea solstitialis t.
COMPOSITAE
Fig. 12. Tricolporate echinate (1000Ö except A±B, L±M ). A±B, Lonicera periclymenum type (e.g. L. periclymenum), 500Ö; C±D, Carlina
type (e.g. C. acaulis); E±F, Solidago type (e.g. Senecio vulgaris); G±H, Achillea type (e.g. A. millefolium); I, Centaurea solstitialis type (e.g.
C. solstitialis); J±K, Cirsium type (e.g. C. creticum); L±M, Echinops (e.g. E. microcephalus), 500Ö; N±O, Artemisia (e.g. A. vulgaris).
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Pollen and spore keys, Pindos Mts, Greece
M2
M1
333
Spheroidal to subprolate. Numerous warts between spines. Size 40±62 mm ....... Crupina
COMPOSITAE
Oblate. Spines long. Incl. Carduus. Fig. 12J±K ............................................. Cirsium t.
COMPOSITAE
Key E11.3 Tricolporate. Striate-rugulate
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A2
A1
Operculate. See Special key ............................................................................................................................... Rosaceae
No operculum.
B2 Endoaperture a transversal colpus. Costae transversales present. Anacardiaceae.
C2 Colpus long, transverse colpus well dened.
D2 Nexine and sexine of equal thickness. Fig. 13C±D ........................................................... Schinus molle
ANACARDIACEAE
D1 Nexine markedly thinner than sexine ................................................................................ Rhus coriaria
ANACARDIACEAE
C1 Colpus shorter, transverse colpus not so well dened, sometimes poroid. Nexine thinner than sexine ...........
Cotinus coggygria
ANACARDIACEAE
B1 Endoaperture not distinctly transversal. No costae transversales.
E3 Finely verrucate. Colpus very short, about twice the diameter of the pore ........................ Ludwigia palustris
ONAGRACEAE
E2 Rugulate.
F2 Colpus short. Distinctly rugulate. Fig. 13A±B .......................................................... Datura stramonium
SOLANACEAE
F1 Colpus long. Finely rugulate-perforate. Fig. 13G±H .................................................. Vitex agnus-castus
VERBENACEAE
E1 Striate-rugulate. Colpus much longer than the diameter of the pore.
G2 Pore indistinct.
H2 Edge of colpus undulating. See Special key ..................................................................... Rosaceae
H1 Edge of colpus straight.
I2 Striae very distinct. Columellae ne.
J2 Costae colpi distinct. Strio-reticulate .................................................. Ailanthus altissima
SIMAROUBACEAE
J1 Costae colpi absent or very thin. See also Key E7.4 ................ Saxifraga oppositifolia t.
SAXIFRAGACEAE
I1 Striae indistinct .................................................................................. Saxifraga tridactylites t.
SAXIFRAGACEAE
G1 Pore distinct.
K2 Grain 25 mm or larger.
L2 Pore covered. Striae ne. Fig. 13E±F ............................................................... Helianthemum
CISTACEAE
L1 Pore not or only partly covered.
M2 All or almost all striae meridional.
N2 Grain c. 40 mm. Endoaperture colpoid.
O2 Fastigium large and distinct. Striate in mesocolpium, reticulate or microreticulate at poles ........................................................................... Atropa t.
SOLANACEAE
O1 No fastigium. Faint striae throughout, or Ôreticulate in mesocolpium and
tectate at poles .............................................................................. Nicotiana
SOLANACEAE
N1 Grain less than 40 mm. Endoaperture poroid.
P2 Poles rounded-pointed. Endoaperture margins smooth, or with a few coarse
granules. P/E>1. Fig. 13I±K .............................................. Gentiana lutea t.
GENTIANACEAE
P1 Poles rounded/attened. Endoaperture margins coarsely granular. P/E usually
<1. See also Key E11.3:R1. Fig. 13N±O ................................... Centaurium
GENTIANACEAE
M1 Some striae not meridional.
Q2 Intercolpium median with predominantly meridional striae.
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Striae run together at pores and poles to form a reticulum ..........................
Tuberaria guttata
CISTACEAE
R1 Striae mostly meridional in mesocolpia, sometimes abruptly changing
direction, sometimes transverse next to colpi. See also Key E11.3:P1.
Fig. 13N±O ................................................................................. Centaurium
GENTIANACEAE
Q1 Intercolpium median rugulate.
S3 Grain 47±56 mm. Intercolpium edges meridionally striated. Vestibulae present
Haplophyllum
RUTACEAE
S2 Grain <40 mm. Intercolpium edges meridionally striated. No vestibulae. See
also Key E12. Fig. 13P±Q ........................................... Blackstonia perfoliata
GENTIANACEAE
S1 Grain <c. 30 mm. Intercolpium edges with groups of non-meridional striae.
T2 Grain 19±26 mm. Faint costae colpi; H-endoaperture .............................
Cicendia liformis
GENTIANACEAE
T1 Grain 22±32 mm. Distinct costae colpi; endoaperture lalongate, not
H-shaped ..................................................................... Ruta chalepensis
RUTACEAE
K1 Grain smaller than 25 mm.
U2 Colpus with coarse echinae ................................................................ Aesculus hippocastanum
HIPPOCASTANACEAE
U1 Colpus membrane without sculptural elements.
V2 Sculpturing of intercolpium indistinct.
W2 Grain prolate, rectangular in equatorial view. Exine c. 1.5 mm.
X2 Rugulate pattern over all of grain. Each elliptic or 8-shaped endoaperture
commonly with ``horns’’ into mesocolpia ................ Coronilla scorpioides t.
LEGUMINOSAE
X1 Rugulate pattern may disintegrate into small reticulations. Each elliptic or
8-shaped endoaperture without ``horns’’ ................................... Hippocrepis
LEGUMINOSAE
W1 Grain prolate, oval in equatorial view.
Y2 Exine 3±5 mm, thins towards colpi. Intercolpium edges with groups of
non-meridional striae ....................................................... Cicendia liformis
GENTIANACEAE
Y1 Exine thinner, even thickness. Striae very indistinct (phase contrast) ..........
Crassula
CRASSULACEAE
V1 Sculpturing of intercolpium uniform, distinct.
Z2 Columellae distinctly visible .............................................................. Sempervivum
CRASSULACEAE
Z1 Columellae indistinct-imperceptible.
A¾2 Endoapertures H-shaped. Fig. 13L±M ............................................... Sedum
CRASSULACEAE
A¾1 Endoapertures circular or oval ....................................... Androsace maxima
PRIMULACEAE
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R2
Key E11.4 Tricolporate. Suprareticulate-foveolate
A2
A1
Heteropolar. Pores near the broad end. See also Key E11.1 ................................................................................. Echium
BORAGINACEAE
Isopolar.
B3 Grain peroblate. Foveolate. Colpi very short. Costae colpi heavy. Exine more than 3 mm thick. Tilia.
C2 4±6 brochi/10 mm .................................................................................................................. Tilia platyphyllos
TILIACEAE
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335
Fig. 13. Tricolporate striate/rugulate (1000Ö except A±B, E±F ). A±B, Datura stramonium, 500Ö; C±D, Schinus molle; E±F, Helianthemum
(e.g. H. nummularium), 500Ö; G±H, Vitex agnus-castus; I±K, Gentiana lutea type (e.g. G. lutea); L±M, Sedum (e.g. S. acre); N±O,
Centaurium (e.g. C. erythraea); P±Q, Blackstonia perfoliata.
C1
B2
B1
8±11 brochi/10 mm. Fig. 14I ..................................................................................................... Tilia cordata t.
TILIACEAE
Grain oblate, triangular. Exine 4 mm. Colpi short, endopores large.
D2 Finely rugulate/foveolate. Lumina with granules ............................................................................ Phaseolus
LEGUMINOSAE
D1 Regular reticulum, brochi 4±7 mm. Incl. Vigna .............................................................................. Psoralea t.
LEGUMINOSAE
Grain not peroblate. Colpus not very short.
E2 Grain prolate. Exine more than 2 mm thick. Columellae very coarse.
F2 Grain oval. Columellae uniform.
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G2
E1
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Costae colpi thick; margo of thin sexine along colpi. Grain subprolate, pores often protruding.
Exine c. 3 mm, with densely packed columellae; foveolate to reticulate. Fig. 14C±D .... Euphorbia
EUPHORBIACEAE
G1 Costae colpi absent, colpi long; endoaperture slightly lalongate. Grain subprolate to spheroidal,
60 mm; circular in polar view. Exine c. 3 mm, with well-spaced columellae; microrugulate to
reticulate ............................................................................................................ Lagenaria siceraria
CUCURBITACEAE
F1 Grain compressed oval. Columellae coarser and more scattered at poles.
H2 Channels between columellae at apocolpium Ô regular in width, narrow, no wider than columella
diameter; branched columellae extending no more than 1/2 way down grain from poles ...............
Polygonum viviparum
POLYGONACEAE
H1 Channels between columellae at apocolpium irregular in width, often broader than columella
diameter; branched columellae often extending from poles almost to equator ................................
Polygonum bistorta t.
POLYGONACEAE
Grains variable. Exine less than 2 mm thick. Columellae ne or indistinct.
I2 Equatorial limb subangular. Grain subspherical-oblate. Polar area very small. Costae transversales and
pores distinct. Fig. 14A±B ....................................................................................................... Rhamnus t.
RHAMNACEAE
I1 Equatorial limb circular or semiangular. Grain subspherical or prolate. Polar area medium to large.
Mainly Leguminosae.
J3 Reticulum indistinct.
K2 Grain prolate. Pore distinct. Costae colpi present.
L2 Endoaperture lalongate, 8-shaped, covered by a thin, structureless exine. Costae colpi
well developed. Fig. 14E±F ........................................................................ Vicia cracca t.
LEGUMINOSAE
L1 EndoapertureÔcircular, not covered ...................................................... Punica granatum
PUNICACEAE
K1 Grain subspherical. Pore covered, more or less distinct.
M3 Grain more than 35 mm. Pore indistinct. Exine thin. Colpus with costae and equatorial
bridge. Fig. 14H ...................................................................................... Medicago sativa
LEGUMINOSAE
M2 Grain more than 30 mm. Exine thin ................................................................. Astragalus
LEGUMINOSAE
M1 Grain <35 mm. Exine thick.
N2 Grain 30±35 mm, compressed oval in equatorial view. Endoaperture lolongate. See
also Key E11.1. Fig. 14M±N ............................................... Trifolium montanum t.
LEGUMINOSAE
N1 Grain <30 mm, rhomboidal in equatorial view. Colpus with equatorial bridge.
Endoaperture a colpus ............................................................. Solanum melongena
SOLANACEAE
J2 Reticulum distinct, some lumina blocked.
O2 Polar area large ............................................................................................... Coronilla emerus
LEGUMINOSAE
O1 Polar area medium .................................................................................................... Astragalus
LEGUMINOSAE
J1 Reticulum distinct. No lumina blocked.
P2 More than 13±15 equatorial brochi per intercolpium.
Q2 Lumina with free columellae ............................................................................ Astragalus
LEGUMINOSAE
Q1 No free columellae in lumina.
R2 Grain more than 27 mm. Fig. 14K .............................................................. Ononis t.
LEGUMINOSAE
R1 Grain smaller than 27 mm ............................................................... Galega oYcinalis
LEGUMINOSAE
P1 Less than 13±15 equatorial brochi per intercolpium.
S2 Meridional limb compressed oval. Colpus with heavy costae. Pore distinct, covered by a
thin, almost structureless sexine.
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T2
Grain subspheroidal. Lumina with perforations. Breadth of muri less than diameter
of lumina ........................................................................................... Lathyrus sativus
LEGUMINOSAE
T1 Grain prolate.
U2 Grain more than 50 mm.
V2 With distinct perforations in lumina ................................... Pisum sativum
LEGUMINOSAE
V1 No perforations in lumina ........................................................ Vicia faba
LEGUMINOSAE
U1 Grain smaller than 50 mm.
W2 Lumina with distinct perforations. Fig. 14L ............ Lathyrus pratensis t.
LEGUMINOSAE
W1 No perforations in lumina.
X2 Ectocolpi usually × length of polar axis. Fig. 14O ... Vicia sativa t.
LEGUMINOSAE
X1 Ectocolpi rather short (c. 25 mm) ............................................. Lens
LEGUMINOSAE
S1 Meridional limb circular or more or less rhomboidal . Costae colpi variable. Pore covered
by sexine with distinct structure. Trifolium.
Y2 Grain more than 40 mm. Fig. 14J ............................................................. T. repens t.
LEGUMINOSAE
Y1 Grain smaller than 40 mm. Fig. 14G ....................................................... T. dubium t.
LEGUMINOSAE
Key E11.5 Tricolporate. Per-reticulate ± frustillate
A3
A2
A1
Reticulum indistinct or frustillate-perforate.
B3 Incomplete reticulum, composed of irregular muri .............................................................. Cistus monspeliensis t.
CISTACEAE
B2 Columellae united only by their topmost parts. Colpus with equatorial bridge. Mercurialis.
C2 Exine more than 2 mm thick. Columellae grouped/irregularly distributed. Colpus membrane not with a row
of granules ........................................................................................................................ Mercurialis perennis
EUPHORBIACEAE
C1 Exine less than 1.5 mm thick. Columellae evenly distributed. Colpus operculum formed by a (single) row of
granules ............................................................................................................................ Mercurialis annua t.
EUPHORBIACEAE
B1 Irregularly reticulate-frustillate-perforate, lumina larger in the polar area. Colpus narrow, slightly wider at both
ends. Pore small, covered. Distinctive hexagonal outline in polar view. Fig. 15G±I .......................... Vitis vinifera
VITACEAE
Reticulate-striate. Columellae coarse, isolated at bottom, forming reticulum at middle level and striations at top; duplior simplicolumellate muri. Fig. 15E±F ..................................................................................................... Gentiana verna
GENTIANACEAE
Reticulate.
D2 Brochi more than 1 mm.
E2 Sexine thick, usually more than 2 mm. Columellae coarse, distinct.
F2 Muri distinctly duplicolumellate. Reticulum irregular; brochi smaller towards colpus. Muri beaded,
breadth equal to or less than diameter of lumina. With transverse colpus. Fig. 15O±Q ... Hedera helix
ARALIACEAE
F1 Muri usually simplicolumellate.
G3 Lumina with distinct free columellae, those in muri crowded. Transverse colpus short.
H2 Reticulum very coarse except in the polar area. Fig 15J±L .......................... Sambucus ebulus
CAPRIFOLIACEAE
H1 Reticulum ner and more irregular. Reticulum in polar area not diVerent, but ner in narrow
margo to ectocolpus. Sometimes 2- or 4-colporate. Fig. 15C±D .......................
Viburnum
CAPRIFOLIACEAE
Fig. 14. Tricolporate foveolate/supra-reticulate (1000Ö). A±B, Rhamnus type (e.g. R. alpinus); C±D, Euphorbia (e.g. E. amygdaloides); E±F,
Vicia cracca type (e.g. V. hirsuta); G, Trifolium dubium type (e.g. T. dubium); H, Medicago sativa; I, Tilia cordata type (e.g. T. cordata); J,
Trifolium repens type (e.g. T. repens); K, Ononis type (e.g. O. spinosa); L, Lathyrus pratensis type (e.g. L. sylvestris); M±N, Trifolium
montanum type (e.g. T. montanum); O, Vicia sativa type (e.g. V. sativa).
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339
G2 No free columellae in lumina. Endoaperture a transverse colpus. Fig. 15A±B . Ligustrum vulgare
OLEACEAE
G1 No free columellae in lumina. Those in muri clearly separated. Endoaperture poroid.
I2 Grain more than 40 mm.
J2 Endoaperture circular, indistinct. Fig. 15M±N ................................................. Jasminum
OLEACEAE
J1 Endoaperture circular, distinct.
K2 Muri spinose. Fig. 16A .................................................................. Cistus salvifolius
CISTACEAE
K1 Muri not spinose. Exine thicker at apocolpium; sexine>nexine. Fig. 15E±F ..........
Gentiana verna
GENTIANACEAE
I1 Grain less than 40 mm, spheroidal to oblate.
L2 H-endoaperture. Exine of even thickness; sexineÔas thick as nexine Gentianella ciliata
GENTIANACEAE
L1 Lolongate endoaperture, 6 mm. Exine thins towards colpi .......... Euonymu s europaeus t.
CELASTRACEAE
Exine thin, usually less than 2 mm.
M2 Columellae distinct in surface view, coarse.
N2 Lumina 1.5±2 mm. See also Key E7.6 ..................................................................... Salix triandra
SALICACEAE
N1 Lumina 2±3 mm. See also Key E7.6 ....................................................................... Salix fragilis t.
SALICACEAE
M1 Columellae indistinct in surface view, ne.
O2 Equatorial limb subangular.
P2 Grain >30 mm, prolate, polar area pointed. Lumina c. 1.5 mm, meridionally elongated in
apocolpia ...................................................................................................... Dictamnus albus
RUTACEAE
P1 Grain 18±22 mm, prolate. Lumina irregular in size, c. 1 mm. Endoaperture indistinct.
Fig. 16F±H ..................................................................................................... Sambucus nigra
CAPRIFOLIACEAE
O1 Equatorial limb circular.
Q2 Greatest diameter of grain >20 mm. Transverse colpus/pore distinct.
R2 Costae aequatoriales; ``horns’’ on endoaperture ........................... Anagallis arvensis t.
PRIMULACEAE
R1 Not so.
S2 Distinctly duplicolumellate. Fig. 16B±E ................................................ Lysimachia
PRIMULACEAE
S1 Simplicolumellate or columellae indistinct.
T2 Muri broad. With transverse colpus.
U2 Columellae indiscernible. Margo heavy .............................. Lysimachia
PRIMULACEAE
U1 Columellae distinct. Margo thin ............................. Anagallis tenella t.
PRIMULACEAE
T1 Muri narrow. Endoaperture poroid or indistinct.
V3 Grain large, P= 45±73 mm, prolate, equatorial outline subangular.
Brochi c. 1.5±2 mm, decreasing in size towards colpi. Endoaperture a
pore ........................................................................................... Bryonia
CUCURBITACEAE
V2 P<40 mm, prolate, equatorial outline variable, commonly interlobate.
Brochi c. 1 mm. Colpus equator constricted, endoaperture indistinct. See
also F¾2, below. Fig. 16I±K ........................................... Scrophularia t.
SCROPHULARIACEAE
V1 P= 19±25 mm, equatorial and polar outlines circular. Lumina 1.5±3.5 mm,
Fig. 15. Tricolporate per-reticulate (1000Ö except M±N ). A±B, Ligustrum vulgare; C±D, Viburnum (e.g. V. tinus); E±F, Gentiana verna;
G±I, Vitis vinifera; J±L, Sambucus ebulus; M±N, Jasminum (e.g. J. fruticans), 500Ö; O±Q, Hedera helix.
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Fig. 15.
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smaller towards poles and colpi. Endoaperture a pore. Fig. 16L±M .....
Parnassia palustris
PARNASSIACEAE
Q1 Grain smaller, P<20 mm.
W2 Endopore indistinct. Lumina not decreasing towards colpi. See also Key E12.
Fig. 16T±U ..................................................................................................... Phillyrea
OLEACEAE
W1 Endoaperture lalongate colpus, usually tapering, sometimes with small horns; margins
sharp; costae Ôdistinct. Lumina decreasing in size towards colpi and apocolpia ........
Anagallis tenella t.
PRIMULACEAE
D1 Micro-reticulate; brochi less than 1 mm.
X2 Costae aequatoriales; ``horns’’ on endoaperture. Grain 20±25 mm ................................. Anagallis arvensis t.
PRIMULACEAE
X1 No costae aequatoriales.
Y2 Colpus slit-shaped
Z2 Endoaperture a transverse colpus.
A¾4 Grain large: P = 40 mm, E = 25 mm. Endoaperture distinct, with abruptly pointed ends ........
Andrachne telephioides
EUPHORBIACEAE
A¾3 P= 14±17 mm, E = 13±16 mm. Endoaperture distinct, with tapering ends
...........................
Asterolinon linum-stellatum
PRIMULACEAE
A¾2 P= 23±28 mm, E = 22±28 mm. Endoaperture distinct, with blunt ends ...... Ricinus communis
EUPHORBIACEAE
A¾1 P= 17±22 mm, E = 15±20 mm. Endoaperture indistinct ................................. Ruta montana t.
RUTACEAE
Z1 Endoaperture poroid.
B¾2 P= 18±22 mm, E = 18±22 mm. Endopore indistinct, without costae ................ Fraxinus ornus
OLEACEAE
B¾1 Various sizes. Endopore covered, with distinct costae. Incl. Oxyria digyna. See key to species
and groups in NW5, also Keys E11.1 & E13 ......................................................... Rumex t.
POLYGONACEAE
Y1 Colpus not slit-shaped.
C¾2 Exine 2 mm or thicker. Transverse endocolpus prominent, oblong. Ectocolpus sunken. P =
13±28 mm ........................................................................................................... Peganum harmala
ZYGOPHYLLACEAE
C¾1 Exine thinner than 2 mm.
D¾3 EndoapertureÔcircular.
E¾3 Muri supra-microechinate ............................................................................... Fumana
CISTACEAE
E¾2 Muri not microechinate. Prolate. Colpus equatorially constricted. Very similar ± refer
to reference slides.
F¾4 P= 15±23 mm, E = 10±21 mm ................................................. Elatine alsinastrum
ELATINACEAE
F¾3 P= 26 mm, E = 18 mm. Fig. 16Q±S ......................................... Cercis siliquastrum
LEGUMINOSAE
F¾2 P<40 mm. See also V2, above ..................................................... Scrophularia t.
SCROPHULARIACEAE
F¾1 P= 15±23 mm, E = 11±15 mm. Fig. 16N±P ................... Hypericum perforatum t.
GUTTIFERAE
E¾1 Muri not microechinate. Colpus fusiform, not constricted, with smooth membrane.
P= 20±25 mm, subprolate, equatorial outline circular. Fig. 16V±X ................ Kickxia
SCROPHULARIACEAE
D¾2 Endoaperture lalongate.
G¾2 Endoaperture 11±16 mm broad, with short costae near colpus. P = 28±41 mm.
Fig. 16Y±A¾ ..................................................................................... Hyoscyamus niger
SOLANACEAE
G¾1 Endoaperture 7±10 mm broad, without costae. P= 21±24 mm . Hypericum hircinum t.
GUTTIFERAE
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D¾1
Endoaperture area with 2±6 small, Ôdistinct, lalongate narrow endocolpi. Saxifragaceae
p.p.
H¾2 P>16.5 mm, E>14 mm. See also Keys E7.1 & E7.6 ....................... Saxifraga stellaris
SAXIFRAGACEAE
H¾1 P<16.5 mm, E<14 mm ................................................... Chrysosplenium alternifolium
SAXIFRAGACEAE
Key E11.6 Tricolporate. Intectate-clavate
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Ilex aquifolium is the only taxon properly keying out here, although Linum spp. may be considered to have a lolongate
endoaperture paralleling the smaller ectoaperture ± see Key E7.7 ( Tricolpate, clavate).
Key E12. Stephanocolporat e
All grains are psilate-scabrate except where otherwise indicated. The transverse colpus in Boraginaceae is blunt-ended, porelike. The number of colpi varies individually; numbers indicated represent averages.
A2
A1
More than 4 colpi. Cf. also Key E19.
B2 Grain larger than 20 mm.
C2 Endoapertures colpoid.
D2 ( Endo-)colpus aequatorialis present. No costae aequatoriales.
E2 More than 10 colpi. Equatorial ridge.
F2 Polar area with several isodiametric lacunae. Polygal a.
G2 7 lacunae in apocolpium (2±5 mm) ............................................ Polygala monspeliaca t.
POLYGALACEAE
G1 15 lacunae in apocolpium (4±7 mm) ................................................. Polygal a vulgaris t.
POLYGALACEAE
F1 Polar area coarsely vermiculate, no round lacunae. Fig. 17A±C ........................... Utricularia
LENTIBULARIACEAE
E1 9±10 colpi. No equatorial ridge. Fig. 17K±M ....................................................... Borago oYcinalis
BORAGINACEAE
D1 Endocolpi distinctly separate.
H2 7±11 meridional colpi. Grain compressed-oval, P = 23±32 mm .................................... Symphytum
BORAGINACEAE
H1 Mostly 7 meridional colpi. Grain bone-shaped, P= 14±25 mm ...................................... Cerinthe t.
BORAGINACEAE
C1 Endoapertures poroid or obscure.
I2 6±7 colpi.
J2 Microreticulate. Grain spherical ........................................................................................ Pinguicula
LENTIBULARIACEAE
J1 Psilate to striate. Grain prolate. Tricolporate with broad colpal operculi, thus appearing 6-colporate ..
Sanguisorba oYcinalis
ROSACEAE
I1 5 colpi (more commonly 4).
K2 Grain more than 40 mm, subprolate, angulaperturate. Tectate, supercially scabrate except in polar
area which has an intectate eld of coarse columellae. See also Q2, below .......... Viola arvensis t.
VIOLACEAE
K1 Grain 30±40 mm, spheroidal, circular in polar view. Reticulate ............................................. Citrus
RUTACEAE
B1 Grain smaller than 20 mm.
L2 Grain bone-shaped. Meridional colpi very short, hardly discernible. Protruding poroid area with free granules
~operculum.
M2 Colpi 4±6. P= 14±17 mm .................................................................................... Buglossoides arvensis t.
BORAGINACEAE
M1 Colpi 7. P= 14±25 mm ............................................................................................................ Cerinthe t.
BORAGINACEAE
L1 Grain compressed oval, P = 18±21 mm. Meridional colpi distinct, 7±8 ............................ Neatostema apulum
BORAGINACEAE
Four colpi.
N2 Grain larger than 40 mm.
O2 Transverse colpus distinct.
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Fig. 16. Tricolporate per-reticulate, continued (1000Ö except A). A, Cistus salvifolius, 500Ö; B±E, Lysimachia (e.g. L. nummularia); F±H,
Sambucus nigra; I±K, Scrophularia type (e.g. S. nodosa); L±M, Parnassia palustris; N±P, Hypericum perforatum type (e.g. H. perforatum);
Q±S, Cercis siliquastrum; T±U, Phillyrea (e.g. P. latifolia); V±X, Kickxia (e.g. K. spuria); Y±A¾, Hyoscyamus niger.
P2
O1
Grain prolate, oval in equatorial view. Psilate. See also Key E11.1. Fig. 17D±E ... Anchusa arvensis t.
BORAGINACEAE
P1 Grain oblate. Echinate. See key to species in Key E11.2 .......................................................... Lonicera
CAPRIFOLIACEAE
No transverse colpus. Viola.
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Q2
Grain depressed oval (barrel-shaped) to polyhedric. Polar area with an intectate eld of coarse
columellae. See also K2, above. Fig. 17H±J .................................................................. Viola arvensis t.
VIOLACEAE
Q1 Grain oval. Exine structure uniform ............................................................................ Viola riviniana t.
VIOLACEAE
N1 Grain smaller than 40 mm.
R3 Grain elliptic or circular in equatorial view.
S2 H-endoaperture. Reticulum striate near colpus. See also Key E11.3 ................... Blackstonia perfoliata
GENTIANACEAE
S1 Endoaperture indistinct in surface view. Finely reticulate.
T2 Colpi slightly sunken in polar view. P= 15±25 mm. ( Usually tricolporate ± see Key E11.5 ) ...........
Phillyrea
OLEACEAE
T1 Colpi not sunken. P = 23±29 mm ........................................................................... Ceratonia siliqua
LEGUMINOSAE
R2 Grain compressed oval. Equatorial exine reticulate. Transverse colpus present.
U2 Exine thinner in polar area. Equatorial reticulum coarse ............................................................. Nonea
BORAGINACEAE
U1 Exine thickness uniform. Grain more than 20 mm.
V2 Meridional colpi with margo of coarse columellae ............................................. Pulmonaria mollis
BORAGINACEAE
V1 Meridional colpi not with coarse columellae.
W2 Colpi short. Equator constricted. Psilate to punctate. Fig. 17F±G ..... Anchusa oYcinalis t.
BORAGINACEAE
W1 Colpi long. Equator not constricted. Reticulate ......................................................... Citrus
RUTACEAE
R1 Grain constricted oval. No transverse colpi. Pores (but not constriction) nearer one pole.
X2 Poles normally of equal width. Grain >13 mm ........................................ Buglossoides purpurocaerulea
BORAGINACEAE
X1 One pole usually considerably wider than the other. Grain <13 mm .............................. Lithospermum
BORAGINACEAE
Key E13. Pericolporate
Tri- or stephanocolporate grains sometimes appear with more apertures than normal, and are then usually imperfectly
isometric. The most important cases are those set out below.
A2
A1
Echinate.
B2 Apertures arranged in spirals .................................................................................................................. Gossypium
MALVACEAE
B1 Apertures not arranged in spirals. Cf. Key E11.2 .............................................................. Compositae Asteroideae
Psilate or reticulate.
C2 Psilate. Costae aequatoriales present. See also Key E11.1 ................................................. Polygonu m aviculare t.
POLYGONACEAE
C1 Psilate with perforations and/or reticulate. Colpus slit-shaped. Pore covered. No costae aequatoriales. Incl. Oxyria
digyna. See key to species and groups in NW5, also Keys E11.1 & E11.5 .............................................. Rumex t.
POLYGONACEAE
Key E14. Diporate
A2
A1
Grains elongate with pores near ends.
B2 Scabrate (usually in planar tetrads) .................................................................................................... Cytinus ruber
RAFFLESIACEAE
B1 Reticulate.
C2 Reticulate, muri much narrower than lumina. Pore 10 mm, edges ragged. P = 30±32 mm, E = c. 60±85 mm. See
also Key E15. Fig. 17N ................................................................................................................... Colchicum
LILIACEAE
C1 Perforate to microreticulate, muriÔ as wide as lumina. Pore 5 mm, not with ragged edges. P = 17±24 mm,
E = c. 30±45 mm. ............................................................................................................. Merendera sobolifera
LILIACEAE
Grains subspheroidal, Ô radially symmetrical.
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Pollen grains of Ficus t here, but given its highly specialized pollination mechanism it is unlikely that pollen would ever
be found in sediments except in exceptional circumstances. The pollen grain is psilate and very small (P= 10 mm, E = 16 mm).
D2 Pori distinctly protruding in optical section, annuli broad and distinct ...................................... Humulus lupulus
CANNABACEAE
D1 Pori not or only slightly protruding, annuli narrow or absent.
E2 Pori slightly or not protruding, circular in outline, small (diameter 1±2.5 mm). Grains small (E = 13±20 mm).
See also Key E15 (triporate) ....................................................................................................... Urtica dioica
URTICACEAE
E1 Pori not protruding, usually irregularly elliptic, rather large (diameter 2.5±5 mm). Grains larger (E =
18±37 mm). See also Key E15. Fig. 17O ....................................................................................... Morus alba
MORACEAE
Key E15. Triporate
A2
A1
With large, cylindrical vestibulum, more than 10 mm deep. Onagraceae.
B2 Grain 50 mm or larger. See also Key E16. Fig. 18A ....................................................... Epilobium angustifolium t.
ONAGRACEAE
B1 Grain smaller than 50 mm. Fig. 18B .............................................................................................. Circaea lutetiana
ONAGRACEAE
Vestibulum smaller, conical, or absent.
C3 Reticulate. Grains large, >50 mm.
D2 Pores annulate, not ragged ....................................................................................................... Cucumis melo
CUCURBITACEAE
D1 Pore margins ragged (usually diporate, Key E14) ......................................................................... Colchicum
LILIACEAE
C2 Distinctly echinate.
E2 Grain more than 50 mm. Exine thick, more than 4 mm. Echinae dimorphic, of two size classes. Dipsacaceae.
F2 Pore with thickened annulus surrounded by halo of thinner exine. Echinae short, broad-based, scattered.
G2 ExineÔeven thickness.
H2 Microechinae between echinae not distinct. Halo broad. Grain spheroidal. See also Key
E7.3 ....................................................................................................................... Dipsacus t.
DIPSACACEAE
H1 Microechinae distinct. Halo narrow. Grain oblate ..................................... Scabiosa sicula t.
DIPSACACEAE
G1 Exine much thicker at poles than at equator, often partially detached, forming sacci ......................
Pterocephalus t.
DIPSACACEAE
F1 Annulus indistinct, no halo. Echinae slender, crowded; microechinae distinct .......................... Knautia
DIPSACACEAE
E1 Grain smaller. Echinae uniform, scattered. Campanulaceae.
I2 Echinae > or = 1 mm. Grain >25 mm.
J2 Echinae c. 1 mm. Pore annuli 3.5±4 mm. (Sometimes stephanoporate, see Key E16.) Fig. 18C ..........
Campanul a rotundifolia
CAMPANULACEAE
J1 Echinae coarser, longer. Pore annuli thicker. Usually 4-porate ........................................ Phyteuma
CAMPANULACEAE
I1 Echinae <1 mm.
K2 Tectum rugulate-perforate. Grain >25 mm ...................................................... Campanul a erinus t.
CAMPANULACEAE
K1 Tectum psilate-perforate. Grain <25 mm ............................................................................. Jasione
CAMPANULACEAE
C1 Psilate-scabrate.
L2 Equatorial limb almost circular, without protruding pore edges.
M2 Scabrate; micro-sculpturing faintly rugulate with minute echinae.
N2 Columellae not visible, even at high magnication; exine 1±2 mm. Morus.
O2 E = 19±25 mm. Outline in polar view circular. ( Usually diporate, elliptical, see Key E14).
Fig. 18D ............................................................................................................... Morus alba
MORACEAE
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346
Fig. 17. Stephanocolporate; Diporate (1000Ö except H±J, N ). A±C, Utricularia (e.g. U. vulgaris); D±E, Anchusa arvensis type (e.g. A.
azurea); F±G, Anchusa oYcinalis type (e.g. A. oYcinalis); H±J, Viola arvensis type (e.g. V. tricolor), 500Ö; K±M, Borago oYcinalis; N,
Colchicum (e.g. C. autumnale), 500Ö; O, Morus alba.
E = 26±37 mm. Outline in polar view oftenÔangular. Pores not always equidistant, 3±4 ......
Morus nigra
MORACEAE
N1 Columellae very distinct, scattered; exine c. 2 mm. Grain more than 20 mm ........................ Celtis
ULMACEAE
Psilate, columellae delicate.
O1
M1
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P2
L1
Exine very thin, but thickened around pores, with minute projections; pore margins a little
protruding.
Q2 Spinules scattered. 2±5 pores, usually 3±4. Fig. 18E ......................................... Urtica dioica
URTICACEAE
Q1 Spinules closer together, more regularly distributed. (Urtica urens has 3±4 pores, Pilea
microphylla 2±3). Fig. 18F .............................................................................. Urtica urens t.
URTICACEAE
P1 Exine slightly thicker, but not thickened around pores; pore margins not protruding. Projections
slightly coarser.
R2 Collumellae indistinct. 3±4 pores, small, circular, margins regular and distinct. Porus membrane
not granulate. Fig. 18I±J .......................................................................................... Parietaria
URTICACEAE
R1 Columellae more distinct, shorter in mesocolpium and distinctly longer at poles. 3 pores,
slightly elongate, margins irregular and indistinct. Porus membrane granulate .........................
Corrigiola litoralis
CARYOPHYLLACEAE
Equatorial limb semiangular, or with protruding pore edges.
S3 Psilate. Grain square in equatorial view ........................................................................ Vicia lathyroides
LEGUMINOSAE
S2 Scabrate, Ô verrucose or granular.
T2 Grain very large, prolate, polar axis c. 205 mm. Pores, trumpet-shaped with thickened edges,
protrude from circular equatorial outline. Subtectal sexine ``spongy’’, not columellate ...................
Morina persica
DIPSACACEAE
T1 Grains 20±40 mm, spheroidal to oblate.
U2 Pores not protruding, annuli narrow. Outline in polar view often Ô angular (pores not
always equidistant) ............................................................................................... Morus nigra
MORACEAE
U1 Pores protruding, annuli broad. Incl. Cannabis sativa and Humulus lupulus, which may be
distinguished on subtle characters of pore morphology, see NW4. Also stephanoporate ( Key
E16). Fig. 18H ....................................................................................................... Humulus t.
CANNABACEAE
S1 Micro-sculpturing and structure more or less rugulate, with scattered minute spinules on top of low
vallae (phase!). Betulaceae.
V2 Equatorial limb circular with protruding pores. Nexine and tectum diverge in the pores, forming
a vestibulum.
W2 Vestibulum distinct. Tectum sharply thickened at the pore. Fig. 18L ............... Betula pendula
BETULACEAE
W1 Vestibulum not distinct. Tectum not appreciably thickened at the pore. Incl. some grains of
Carpinus, which is usually 4-porate. Fig. 18K ......................................................... Ostrya t.
CORYLACEAE
V1 Equatorial limb semiangular, pore not protruding. Nexine and tectum do not diverge in the pore.
Depth of pore not much greater than thickness of the regular exine. Fig. 18G .................... Corylus
CORYLACEAE
Key E16. Stephanoporate
A4
A3
A2
Echinate. Echinae distinct, regularly distributed. Campanulaceae. See Key E15: E1. Species of Phyteuma, Asyneuma,
and Legousia are usually 4-porate, but occasional grains of some Campanula spp. are also stephanoporate.
B2 Echinae <1 mm. Incl. Legousia hybrida ............................................................................................... Asyneuma t.
CAMPANULACEAE
B1 Echinae >1 mm ......................................................................................................................................... Phyteuma
CAMPANULACEAE
Finely micro-suprareticulate and microechinate; exine 2.5 mm with distinct columellae. 4±8 pores without annuli. P =
23±26 mm ..................................................................................................................................... Theligonum cynocrambe
THELIGONACEAE
Coarsely and softly rugulate or suprareticulate. Peroblate. 4±7 pores. Ulmus.
C3 Rugulate sculpture on both poles. Angular outline. 5±7 pores, mostly 6 .............................................. U. procera
ULMACEAE
C2 Rugulate sculpture reduced on proximal pole. Angular outline. 4±6 pores, mostly 4±5. Fig. 18N±O U. glabra t.
ULMACEAE
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A1
Rugulate sculpture reduced on proximal pole. Rounded outline, well-developed annuli lacking. 4±6 pores,
mostly 5 .................................................................................................................................................. U. minor t.
ULMACEAE
Psilate, scabrate or very delicately rugulate.
D2 Grain globular. Equatorial limb circular. Pores not distinctly protruding.
E2 Rugulate, spinulose. 3±6 pores .............................................................................................................. Celtis
ULMACEAE
E1 Psilate. Columellae delicate.
F3 Pores with large cylindrical vestibulum (>10 mm deep). Grain more than 50 mm. ( Usually triporate, see
Key E15) ........................................................................................................ Epilobium angustifolium t.
ONAGRACEAE
F2 Pores annulate, protruding; no vestibulum. Mostly 3-porate, see Key E15 ........................... Humulus t.
CANNABACEAE
F1 Pores annulate, but not protruding or vestibulate.
G2 Exine very thin, structure very delicate. See Key E15: P2 for distinction of species which are
occasionally stephanoporate ................................................................................................. Urtica
URTICACEAE
G1 Exine thicker, structure coarser.
H2 Pores 3 or 4. Fig. 18I±J .......................................................................................... Parietaria
URTICACEAE
H1 Pores 6±12 ................................................................................................. Urtica pilulifera t.
URTICACEAE
D1 Grain oblate. Equatorial limb more or less angular and/or pore edges protruding.
I2 Neighbouring pores (annuli) connected with thickened bands (arci). Pore with thick annulus. 4±6 pores.
Fig. 18M ................................................................................................................................... Alnus glutinosa
BETULACEAE
I1 No arci.
J2 4±6 ``pores’’ meridionally elongated with endexinous annuli. Myriophyllum.
K2 Annulus distinct, protruding. Pores circular or elliptic ................................................. M. spicatum
HALORAGACEAE
K1 Annulus externally at, not protruding. Pores colpoid .......................................... M. verticillatum
HALORAGACEAE
J1 Exine thickness uniform. Pores circular, without endexinous annuli.
L2 Equatorial limb angular (square) ............................................................................ Vicia lathyroides
LEGUMINOSAE
L1 Equatorial limb more or less circular.
M2 Grain more than 40 mm. Microsculpturing faintly rugulate with minute spinules on vallae.
Incl. some grains of Ostrya .................................................................................... Carpinus t.
CORYLACEAE
M1 Grain less than 40 mm. Microechinate. Pores with narrow annuli, not always equidistant ......
Morus nigra
MORACEAE
Key E17. Periporate: Main key to subsections
Verrucate or echinate ............................................................................................................................................... Key E17.1
Rugulate, striate or reticulate .................................................................................................................................. Key E17.2
Psilate or scabrate .................................................................................................................................................... Key E17.3
Key E17.1 Periporate. Verrucate or echinate
A2
A1
Verrucate. Special key .............................................................................................................................. Plantaginaceae
Echinate.
B2 Grains large, >60 mm.
C2 c. 75 pores. Grain c. 80±130 mm. Sexine between echinae composed of crowded baculae. Echinae long
(8±11 mm), with rounded tips, situated at the corners of polygona l ridges formed by longer baculae, in the
centre of each of which are situated the rather indistinct pores. Smaller baculae surround the pores. Fig. 19A.
Ipomoea
CONVOLVULACEAE
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B1
Echinae not linked by ridges.
D2 Pores 4±12, large (diameter 15±26 mm), not spirally arranged, with operculi. Grain 90±190 mm. Echinae
regularly spaced over whole grain, including over the opercula, which bear a single central echina.
Echinae not based on cushions. Numerous microprojections (baculae or microechinae) between echinae.
For detailed key to species see Ayala-Nieto et al. 1988 .......................................................... Cucurbita
CUCURBITACEAE
D1 Pores 9 to >100, in some species arranged in a spiral, without echinate opercula. Grain >60 mm.
Echinae usually sharp, in some species elevated on cushions of thicker exine. See special key ...............
Malvaceae
Grains <60 mm.
E2 Pores with annuli. Special key ......................................................................................... Caryophyllaceae p.p.
E1 Pores without annuli.
F2 24±28 indistinct pores usually covered by non-operculate membranes with echinae. Grain 44±65 mm.
Echinae conical, c. 3 mm long, irregularly spaced, with broad irregular bases which sometimes coalesce in
groups of 2±3 ....................................................................................................... Ranunculus arvensis t.
RANUNCULACEAE
F1 Pores 10±15 (diameter 4±6 mm). Grain 26±36 mm. Echinae sharp, about as tall as broad, irregularly
distributed but without broad bases and not coalescent. Fig. 19D .......................... Sagittaria sagittifolia
ALISMATACEAE
Key E17.2 Periporate. Rugulate, striate or reticulate
A2
A1
Vermiculate-rugulate. Columellae ne. Fewer than 50 pores ............................................................ Buxus sempervirens
BUXACEAE
Reticulate.
B2 Pores with annuli. Special key ................................................................................................. Caryophyllaceae p.p.
B1 Annuli absent or indistinct.
C3 Reticulum regular, brochi 4±8 mm, muri simplicolumellate. Pore in each lumen: 60±75 pores. Fig. 19B ...........
Tribulus terrestris
ZYGOPHYLLACEAE
C2 Reticulum regular, brochi >4 mm, muri duplicolumellat e. Not every lumina with a pore: 12±27 pores. Lumina
with pores are smaller than those without. Fig. 19C ................................................. Polygonu m persicaria t.
POLYGONACEAE
C1 Reticulum imperfect, brochi smaller and not polygonal. Fewer than 50 pores.
D2 Pore indistinct, included in a lumen. Lumen with a pore slightly larger than those without. Columellae
fused at top into wedge-shaped elements in surface view, which are arranged to form a reticulum
(``croton’’ pattern) ....................................................................................................................... Daphne
THYMELAEACEAE
D1 Pore larger than brochi. Microreticulate. Columellae not fused into wedge-shaped elements ................
Buxus sempervirens
BUXACEAE
Key E17.3 Periporate. Psilate or scabrate
A2
A1
Grain irregular.
B2 Usually pear-shaped. Pores indistinct. Special key. ................................................................................ Cyperaceae
B1 Irregular angular or cube-shaped. Grain <20 mm. No echinae. 4±7 pores .......................................... Herniaria t.
CARYOPHYLLACEAE
Grain regularly ellipsoidic.
C2 Microechinate, dotted.
D2 Columellae ne and evenly distributed, or not visible.
E2 Pores aggregated towards one pole. Fig. 19J ....................................................................... Juglans regia
JUGLANDACEAE
E1 Pores Ô evenly distributed.
F2 Columellae clear, exine surface undulating. Pores 5±6 ......................................................... Celtis
ULMACEAE
Fig. 18. Triporate; Stephanoporate; Syncolpate; Heterocolpate (1000Ö except B). A, Epilobium angustifoliu m type (e.g. E. angustifolium);
B, Circaea lutetiana, 500Ö; C, Campanula rotundifoli a; D, Morus alba; E, Urtica dioica; F, Urtica urens type (e.g. U. urens); G, Corylus
(e.g. C. avellana); H, Humulus type (e.g. Cannabi s sativa); I±J, Parietaria (e.g. P. diVusa); K, Ostrya type (e.g. O. carpinifolia); L, Betula
pendula; M, Alnus glutinosa; N±O, Ulmus glabra type (e.g. U. glabra); P±Q, Myrtus communis; R±T, Lythrum salicaria type (e.g. L. salicaria);
U±V, Myosotis type (e.g. M. arvensis).
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D1
C1
Not
M2
M1
Columellae invisible, exine surface smooth. Pores 6±12, usually 7±9 ................ Urtica pilulifera t.
URTICACEAE
Columellae coarse and/or irregularly distributed. Pores evenly distributed.
G2 Annulus distinct. Special key ................................................................................. Caryophyllaceae p.p.
G1 Annulus absent or indistinct.
H2 Surface undulating. Special key ............................................................................... Plantaginaceae
H1 Surface even.
I2 Grain more than 25 mm. Indistinctly intrareticulate.
J2 Grain sub-polyhedric. Exine between pores distinctly crescentic. Pores round.
K2 Pores with annuli. See also T1, below. Fig. 19E .......................................... Alisma
ALISMATACEAE
K1 No annuli ................................................................................. Damasonium alisma
ALISMATACEAE
J1 Grain (sub-)globular. Exine not distinctly crescentic. Pores irregular, indistinct .............
Papaver argemone t.
PAPAVERACEAE
I1 Grain 10±25 mm. No intrareticulum. Thalictrum.
L3 Grain >17 mm. Columellae distinct. Porus margins clearly dened, regular . T. lucidum
RANUNCULACEAE
L2 Grain >17 mm. Columellae distinct. Porus margins poorly dened, irregular . T. minus
RANUNCULACEAE
L1 Grain <17 mm. Columellae usually very indistinct. Fig. 19F±G ......... T. aquilegifolium
RANUNCULACEAE
microechinate, dotted.
Areas without tectum, surrounding and connecting pores ..................................................................... Ribes
GROSSULARIACEAE
Tectum not reduced.
N2 Grain large (70±80 mm). Exine thick. Fig. 19I ...................................................................... Calystegia
CONVOLVULACEAE
N1 Grain smaller.
O2 Annulus present.
P2 Annulus distinct, but often narrow and without structure.
Q2 Tectum without perforations. Number of pores usually higher than 50. Incl.
Amaranthaceae and Sagina (Caryophyllaceae). Fig. 21G ................ Chenopodiacea e t.
Q1 Tectum perforate (indistinct in Sagina). Pores fewer than 50.
R2 6±12 large pores. Tectum very thick. Annulus prominent. Columellae very ne and
short. Fumaria.
S2 Usually 12 pores .............................................................. Fumaria parviora
PAPAVERACEAE
S1 Usually 6 pores ............................................................. Fumaria densiora t.
PAPAVERACEAE
R1 Not so. Special key ............................................................... Caryophyllaceae p.p.
P1 Annulus diVuse. Columellae widely dispersed in interporia, dense on annuli. Grain usually
polyhedric. Intrareticulate. Alismataceae.
T2 Grain less than 25 mm ................................................................ Baldellia ranunculoide s
ALISMATACEAE
T1 Grain more than 25 mm. See also K2, above ...................................................... Alisma
ALISMATACEAE
O1 No annulus.
U2 Grain 23±42 mm. 4±11 pores. Exine punctate/microreticulate. Fig. 19H ................. Pistacia
ANACARDIACEAE
U1 Grain 10±25 mm. 4±12 pores.
V2 Grain rounded. Distinct minute echinae. See Key E17.3: H1 .................... Thalictrum
RANUNCULACEAE
V1 Grain sub-polyhedric. Indistinctly dotted. Incl. Polycnemum (Chenopodiaceae) ..........
Paronychia t.
CARYOPHYLLACEAE
Key E18. Syncolpate
Colpi sometimes fuse at one pole also in grains not considered here to be syncolpate.
A2
Heteropolar.
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A1
Tricolporate with large pores, the three colpi joining at one pole only.
C2 Coarsely reticulate ....................................................................................................... Thesium divaricatum t.
SANTALACEAE
C1 Psilate to nely verrucate ....................................................................................................... Thesium alpinum
SANTALACEAE
B1 Single encircling colpus, closer to one pole. Microechinate. See also Key E5. ........................................... Smilax
LILIACEAE
Isopolar.
D3 Colpi meridional.
E2 Two colpi fused to a ring. Grain usually split, edges of colpus ragged. Cf. Juniperus, Key E2.
F2 Grain 17±21 mm. Scabrate (densely microechinate) ................................................................ Hypecoum
PAPAVERACEAE
F1 Grain 26±32 mm. Psilate perforate-perfossulate. See also Key E6. ........................................ Pedicularis
SCROPHULARIACEAE
E1 More than two colpi.
G2 Colpi bifurcate, fused with neighbouring colpi, delimiting a large polar eld. Grain larger than 30 mm.
Coarsely striated ...................................................................................................... Nymphoide s peltata
MENYANTHACEAE
G1 Colpi 3, not bifurcate.
H4 Grain striate. Tricolporate. Prolate, c. 45 mm ..................................................... Arachis hypogaea
LEGUMINOSAE
H3 Grain scabrate. No endoapertures. Grain subprolate, 18±21 mm .................................. Soldanella
PRIMULACEAE
H2 Grain microverrucate. Grain oblate .............................................................. Loranthus europaeus
LORANTHACEAE
H1 Grain psilate. Oblate, triangular in polar view. Short transverse colpus present. Fig. 18P±Q .......
Myrtus communis
MYRTACEAE
D2 Colpi spirally oriented (various arrangements). Grain spheroidal.
I3 Grain microechinate, 60±100 mm .......................................................................................................... Crocus
LILIACEAE
I2 Grain with scattered clavae on top of irregularly granulate exine, 110±130 mm ........................ Iris planifolia
IRIDACEAE
I1 Grain irregularly perforate, 30±50 mm. Exine usually splits ............................................... Berberis vulgaris t.
BERBERIDACEAE
D1 Peri-syncolpate. Exine divided into angular plates, usually falling apart.
J3 Exine microechinate, 1.5±3 mm thick, columellae coarse. Colpi 12 or more. Grain spheroidal, 20±35 mm. See
also Key E9 .............................................................................................................................. Montia fontana
PORTULACACEAE
J2 Exine psilate, pitted, 1.5 mm thick. Colpi 6 to 16. Grain spheroidal, 30±40 mm. ............. Mahonia aquifolium
BERBERIDACEAE
J1 Exine psilate, pitted, 2 mm thick. Colpi 6 or 12. Grain spheroidal or a convex tetrahedron, 40±50 mm ...........
Corydalis ochroleuca
PAPAVERACEAE
Key E19. Heterocolpate
Grains with three colpi with pores which alternate with three similar without pores.
A2
Colpi approximately equally long, or aporate colpi longer than porate colpi. Grains attened at poles, often dumbbellshaped (constricted at equator), or compressed oval in equatorial view. Polar area index large, above 0.5. Boraginaceae.
Also see detailed key in NW2.
B2 Endoapertures join together at equator in a narrow colpus aequatorialis. ``Aporate’’ colpi longer than those in
which the endoapertures are more strongly developed ................................................................... Cynoglossum t.
BORAGINACEAE
B1 Endoapertures discrete, not joined at equator.
C2 Grain >30 mm, not constricted at equator ................................................................................. Heliotropium
BORAGINACEAE
C1 Grain <20 mm, constricted at equator.
D2 Isopolar, endoapertures at equator. Fig. 18U±V ................................................................... Myosotis t.
BORAGINACEAE
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A1
Heteropolar, two endoapertures closer to one pole and one closer to the other ....................... Lappula
BORAGINACEAE
Aporate colpi distinctly shorter than the porate ones. Polar area smaller. Grain circular to oval in equatorial view, not
attened at poles, or constricted at equator.
E2 Exine scabrate-psilate ...................................................................................................................... Vinca herbacea
APOCYNACEAE
E1 Non-porate colpi separated from porate colpi by bands of thick, tectate striate exine. Lythrum.
F2 Grain asymmetrical, with apertures often arranged irregularly. Non-porate colpi wide areas of intectate,
granular exine ......................................................................................................................... Lythrum portula
LYTHRACEAE
F1 Grain symmetrical, with apertures clearly zonally arranged. Non-porate colpi do not occupy wide areas, are
narrower and shorter than porate colpi. Fig. 18R±T ....................................................... Lythrum salicaria t.
LYTHRACEAE
Key E20. Fenestrate
Grains echinate, lophate (ridged). COMPOSITAE Cichorioideae ( Tomb 1975; Blackmore 1981, 1982, 1984 (NW4), 1986;
Blackmore & Jarvis 1986; DõÂez 1987; Moore et al. 1991:91 ±93).
A2
A1
Ectoapertures pores, surrounded by continuous lophae; paraporal lacunae 2 per mesocolpium; total of 15 lacunae ....
Scolymus
Ectoapertures colpi, broadly rounded to angular and often divided into lacunae by constrictions or short paired ridges.
B2 Colpi each divided into 2 abporal lacunae by a central constriction or pair of ridges.
C2 One equatorial lacuna per mesocolpium. Paraporal lacunae absent, interporal lacunae present, total of 15
lacunae ........................................................................................................................................... Tragopogon
C1 Two equatorial lacunae per mesocolpium. Abporal lacunae extending to an expanded spiny polar area, or
to a polar lacuna; total of 18±20 lacunae.
D2 Polar lacunae absent, extensive echinate polar areas present; total of 18 lacunae ..................................
Scorzonera laciniata t.
D1 Polar lacunae 1 per pole, large, hexagonal; total of 20 lacunae ........................ Scorzonera hispanica t.
B1 Colpi each divided into 1 poral and 2 abporal lacunae by 2 pairs of short continuous ridges. Paraporal lacunae
2 per mesocolpium.
E2 Polar lacunae absent, polar area ranging from a triradiate ridge to an extensive echinate region; total of 15
lacunae (20 in occasional 4-aperturate grains).
F2 Polar area not more than a triradiate ridge usually with one row of echinae, abporal and paraporal
lacunae extending almost to the poles. Fig. 20A±C ................................................................ Lactuca t.
F1 Polar area more extensive.
G2 Grains small, usually E<25 mm. Fig. 20D±G ............................................................. Hieracium t.
G1 Grains larger, E>25 mm.
H2 Polar area small with 1±(2)±4 isolated central echinae. Fig. 20H±I ................... Cichorium t.
H1 Polar area extensive, 4±(5.5)±15 isolated central echinae. Fig. 20J±K ............. Taraxacum t.
E1 Polar lacunae 3 in each of the extensive polar areas; total of 21 lacunae (28 in occasional 4-aperturate
grains). Fig. 20L±M ......................................................................................................................... Sonchus t.
PART 2
Special keys to families
CARYOPHYLLACEA E (periporate)
[Cf. Chanda 1962, Melzheimer 1975, McNeill & Crompton 1978, Candau 1987, Faegri & Iversen 1989:283, Moore et al.
1991:110-112 , Punt & Hoen 1995 (NW7)]
The section Sperguleae has colpate grains, cf. Key E7.1.
A2
Grain c. 15 mm.
Fig. 19. Periporate (1000Ö except A). A, Ipomoea (e.g. I. purpurea), 500Ö; B, Tribulus terrestris; C, Polygonu m persicaria type (e.g. P.
persicaria); D, Sagittaria sagittifolia; E, Alisma (e.g. A. plantago-aquatic a); F±G, Thalictrum aquilegifolium; H, Pistacia (e.g. P. lentiscus);
I, Calystegia (e.g. C. sepium); J, Juglans regia.
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A1
355
B2 3 pores (or colpi). Ellipsoidal ..................................................................................................... Corrigiola litoralis
B1 4±7 pores. Irregular angular or cube-shaped ........................................................................................ Herniaria t.
Grain usually larger than 15 mm.
C2 Grains spheroidal-rounded.
D2 Grain >50 mm (c. 63 ). >30 pores (up to 48), Ô8 mm, annulus. Fig. 21A ................
Agrostemma githago
C1
D1 Grains <50 mm.
E3 Reticulate, microreticulate, semitectate or infrareticulate; lumina wider than width of columellae or
muri.
F2 Pores >20.
G3 Reticulate to microreticulate, muri simplicolumellate. Fig. 21C ........................ Silene alba t.
G2 Microreticulate to punctate, columellae irregularly arranged ....................... Lychnis viscaria
G1 Infrareticulate, columellae regularly arranged in reticulate pattern.
H2 Scabrate. See also J2, below ................................................... Dianthus deltoides t. p.p.
H1 Psilate.
I2 Operculum of fused strings, columellae ne, pores 27±40 ......... Lychnis os-cuculi
I1 Operculum a disc with granules, columellae coarse, pores usually <30. See also
K2, below ..................................................................................... Silene vulgaris t.
F1 Pores 10±20.
J2 Distinctly scabrate (microechinate). Fig. 21D .................................. Dianthus deltoides t. p.p.
J1 Psilate.
K2 Pores >18. Fig. 21E ...................................................................... Silene vulgaris t. p.p.
K1 Pores 10±14 ............................................................................... Minuartia setacea t. p.p.
E2 Punctate (puncta <width of collumellae); puncta distinct or indistinct, but never wider than width of
columellae.
L2 Pores <15, columellae ne and crowded ................................................. Minuartia setacea t. p.p.
L1 Pores 15 or more.
M2 Columellae in mesoporial area spaced.
N2 A few columellae attached to the annulus. Fig. 21K ................ Holosteum umbellatum
N1 Many small columellae attached to annulus ............................. Moehringia trinervia t.
M1 Columellae distinctly crowded.
O2 Distance between two pores shorter than pore diam., annulus included. Fig. 21J ........
Arenaria serpyllifolia
O1 Distance between two pores larger than pore diam., annulus included. Fig. 21B ..........
Lychnis coronaria
E1 Tectum without perforations. Number of pores usually higher than 50. Incl. Amaranthaceae and Sagina
(Caryophyllaceae). Fig. 21G ....................................................................................... Chenopodiacea e t.
Grains spheroidal-polyhedral. Pores (+ annuli) in depressions, surrounded by a zone without big columellae; the
rest of the grain with a pentagonal pattern of ridges.
Q2 Ridges scabrate or with indistinct sculpturing.
R2 Pores 6±12 (2.5±4 mm). Grain 8±25 m ................................................................................ Paronychia t.
R1 Pores usually 12 or more. Grain >25 mm.
S2 Outside annulus a distinct margo present.
T2 Grain 20±42 mm. Pores 14±23 (c. 4 mm). Margo small, <3.5 mm broad, with small columellae.
Polygonal to circular. Fig. 21I ............................................................... Scleranthus annuus t.
T1 Grain c. 26±42 mm. Pores 12±16 (c. 5 mm). Margo large, distinct, >3.5 mm broad, without
small columellae. Polygonal (6-angular) .................................................. Scleranthus perennis
S1 No distinct margo outside annulus present.
U2 Annulus broad (>1 mm); crowded, coarse columellae which decrease in size towards the
pores. In LM it is clearly visible, that the columellae tend to stand in irregular circles, thus
forming a ne infrareticulum .................................................................. Vaccaria pyramidata
U1 Annulus narrow; coarse columellae in the middle of the mesoporial area distinctly sparsely
spaced, smaller and more densely arranged towards the pores; pollen often very large.
Fig. 21F ............................................................................................... Cerastium fontanum t.
Q1 Ridges echinate. Grain c. 36±49 mm. Pores 12 (4±6 mm). Margo distinctly sunken with a few columellae.
Columellae in mesoporium coarse and irregularly crowded. Distinctly polygonal. Fig. 21H. Stellaria holostea
Fig. 20. Fenestrate (1000Ö). A±C, Lactuca type (e.g. L. virosa); D±G, Hieracium type (e.g. H. pilosella); H±I, Cichorium type (e.g. C.
intybus); J±K, Taraxacum type (e.g. Lapsana communis); L±M, Sonchus type (e.g. S. oleraceus).
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357
CYPERACEAE
(FernaÂndez 1987b, Faegri & Iversen 1989: 284)
Grains with perforate/granulate tectum with lacunae, i.e. areas where the tectum is broken up into small frustillae, usually
forming an areolar sculpturing. The form and arrangement of lacunae is of great diagnostic value. One or two of the lateral
lacunae may be smaller than the others; the indices given below do not apply to these.
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A3
A2
Lateral lacunae oblong, length twice as long as the breadth or more. Grain ovoid or pear-shaped.
B2 Length of lateral lacunae more than half that of the grain.
C2 Grain 36±43 mm .................................................................................................................. Schoenus nigricans
C1 Grain 27±37 mm. Fig. 22B .................................................................................................... Cyperus longus t.
B1 Length of lateral lacunae less than half of that of the grain. Grain more than 40 mm. Fig. 22A ............... Scirpus
Lateral lacunae more or less circular or slightly elongated. Grain ovoid or pear-shaped.
D2 2±3 small lacunae, length less than 20% that of the grain. Grain more than 40 mm, the narrow end usually
projecting like a beak or a nger ................................................................................................. Cladium mariscus
D1 4±6 lateral lacunae, their length normally more than 20% of that of the grain. Grain smaller than 40 mm,
exceptionally (Carex hirta) 40±50 mm. The narrow end rounded: Carex type.
E2 Lacunae poroid. Incl. Scirpus p.p. ...................................................................................... Carex hallerana t.
A1
E1 Lacunae colpoid. Fig. 22C ....................................................................................................... Carex acca t.
Lacunae irregular, rather large. Grain almost isodiametric, less than 20 mm. Structure rather coarse, columellae distinct
Cyperus michelianus
GRAMINEAE
(Andersen 1979, Andersen & Bertelsen 1972, Beug 1961, Peltre et al. 1987, Faegri & Iversen 1989: 284±286)
Owing to the great importance of grasses, both in natural plant communities and in cultivated areas, and also the importance
of certain taxa as cultivation indicators, it would be highly desirable to be able to identify grasses to species. However, the
paucity of distinguishing characters limits separation. The Gramineae as a family are easily recognized, but also very
monotonous in pollen morphology. The main identifying characters used in this key are: diameter of grain, of pore and of
annulus (the latter two being the most reliable). The measurements are taken from grains embedded in glycerol jelly. Where
data from Andersen’s key (grains embedded in silicone oil ) has been used, a conversion factor of 1.2 has been applied.
However, for the same species, conversion factors may vary between 1.1 and 1.3 for grain size and between 1.1 and 1.5 for
pore size.
To make the distinction between the two cereal types, Triticum and Avena, the less easily seen columellae pattern is used.
Two diVerent patterns can be detected. In single-grain structure columellae are individually free, and more or less crowded
and aggregated. In the double-grain structure some or all columellae are united into small frustillae. These patterns are also
used to make the less important distinction between the two wild grass types of grasslands, Dactylis and Festuca. The
patterns can be more easily seen using phase contrast.
A2
A1
Grain >40 mm. Incl. mostly cereals and very few wild grasses. Cerealia type.
B2 Grain >60 mm. Fig. 23A .......................................................................................................................... Zea mays
B1 Grain between 40 and 60 mm.
C2 P/ E ratio higher than 1.25. Exine distinctly scabrate. Pore lateral ........................................... Secale cereale
C1 P/ E ratio less than 1.25. Pore (sub-)apical.
D2 Annulus diameter larger than 12 mm. Exine verrucate.
E2 Exine with single-grain structure. Fig. 23B .................................................................... Triticum t.
E1 Exine with double-grain structure. Fig. 23C ......................................................................... Avena
D1 Annulus diameter between 10 and 12 mm. Exine scabrate. Incl. some wild grasses.
F2 Outer limit of annulus sharp. Incl. Elymus, Triticum monococcum. Fig. 23D .............. Hordeum t.
F1 Outer limit of annulus not sharp ................................................................................ Agropyron t.
Grain smaller than 40 mm. Annulus diameter less than 10 mm. Incl. most wild grasses.
G2 Grain smaller than 26 mm. Fig. 23E±F ................................................................................... Phragmites australis
Fig. 21. Caryophyllaceae (1000Ö). A, Agrostemma githago; B, Lychnis coronaria (separate inset at high focus shows punctae); C, Silene
alba; D, Dianthus deltoides type (e.g. D. deltoides); E, Silene vulgaris type (e.g. S. vulgaris subsp. maritima); F, Cerastium fontanum type
(e.g. C. fontanum subsp. triviale; separate inset at high focus shows punctae); G, Chenopodiaceae type (e.g. Chenopodium album); H,
Stellaria holostea (succession of focal levels); I, Scleranthus annuus type (e.g. S. annuus ); J, Arenaria serpyllifolia (succession of focal levels);
K, Holosteum umbellatum (separate inset at highest focus).
Grana 40 (2001)
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Fig. 21.
358
(For legend see page 357).
P. I. Chester and J. I. Raine
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Pollen and spore keys, Pindos Mts, Greece
359
Fig. 22. Cyperaceae, A±C (1000Ö): A, Scirpus (e.g. S. holoschoenu s); B, Cyperus longus type (e.g. Cyperus longus); C, Carex acca type
(e.g. Cyperus rotundu s); Malvaceae, D±H (all 250Ö): D, Hibiscus (e.g. H. trionum); E±F, Malva pusilla; G±H, Malva sylvestris type (e.g.
M. sylvestris).
G1 Grain between 26 and 40 mm.
H2 Double-grain structure. Fig. 23G±H .............................................................................................. Dactylis t.
H1 Single-grain structure.
I2 Exine thick. Fig. 23K±L ........................................................................................................... Festuca t.
I1 Exine thin. Some aggregation opposite the pore. Fig. 23I±J .................................................. Glyceria t.
MALVACEAE
[Ganguly & Chanda 1974, FernaÂndez & Candau 1981, Christensen 1986, FernaÂndez 1987a, Culhane & Blackmore
1988 (NW5)]
A3
A2
A1
Tricolporate ( Key E11.2 ) ................................................................................................................................ Abutilon t.
Pericolporate (apertures spirally arranged; Key E13) ..................................................................................... Gossypium
Periporate (apertures spirally or evenly arranged ).
B2 Supratectal processes all conical echinae.
C2 Bases of echinae subglobose; porus margins often indistinct ............................................. Althaea oYcinalis
C1 Bases of echinae not subglobose; porus margins distinct.
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P. I. Chester and J. I. Raine
Fig. 23. Gramineae (N.B. 600Ö). A, Zea mays; B, Triticum type (e.g. T. aestivum); C, Avena (e.g. A. sterilis); D, Hordeum type (e.g. Elymus
repens); E±F, Phragmites australis; G±H, Dactylis type (e.g. Melica ciliata); I±J, Glyceria type (e.g. Bromus hordeaceus); K±L, Festuca type
(e.g. F. ovina).
<100 pori per grain. Refer to reference material!
E3 46±93 pori (3±9 mm). Echinae c. 18 mm. Fig. 22D ............................................................. Hibiscus
E2 60±80 pori (c. 3 mm). Echinae 4±14 mm ................................................................ Malva moschata
E1 78±81 pori (c. 10 mm). Echinae 22 mm ........................................................ Abelmoschus esculentus
D1 >100 pori per grain; distance between pori c. 11 mm.
F2 Echinae tapering, forming more than half of the thickness of the exine and frequently curved
towards the apex; grains larger than 110 mm .......................................................... Althaea hirsuta
F1 Echinae generally straight sided, forming approximately half of the thickness of the exine; grains
60±125 mm. Fig. 22G±H ..................................................................................... Malva sylvestris t.
Supratectal processes other than conical echinae present including clavae, verrucae or bluntly rounded echinae.
G2 Number of pori >200; echinae long, conical, acute, interspersed with smaller rounded clavae; porus diameter
1±2 mm; sexine-1 and sexine-2 <1 mm ................................................................................................ Alcea t.
G1 Number of pori <150; processes either conical or blunt echinae, or rounded verrucae, in various proportions;
porus diameter 2±3 mm; sexine-1 and sexine-2 >1 mm. Fig. 22E±F .......................................... Malva pusilla
D2
B1
PLANTAGINACEAE
(Cf. Clarke & Jones 1980 (NW2), Saad 1986, Ubera et al. 1988, Faegri & Iversen 1989: 286)
A3
Pore with distinct annulus.
B2 Pore operculate (operculum may be lost, but then no granules on pore membrane). Markedly protruding annulus.
Grana 40 (2001)
Pollen and spore keys, Pindos Mts, Greece
8±12 pores, 1±3 mm. Distinctly scabrate, verrucate, undulate. Grain 18±33 mm. Fig. 24A±B ...................................
Plantago lanceolata t.
B1 Pore rarely operculate, but with isolated granules.
C2 Markedly protruding annulus. Verrucate, scabrate, undulate. 5±8 pores, 1±3 mm. Grain 17±31 mm. Fig. 24C±D
P. coronopus
C1 Exine scabrate, undulate. 5±9 pores, 2±4 mm. Grain 20±29 mm ..................................................... P. alpina t.
Pore with incipient annulus.
D2 Pore operculate.
E2 Exine psilate, slightly undulate. Grain 22±33 mm. 6±12 pores, 2±4 mm. Fig. 24E±F ..................... P. albicans
E1 Exine microechinate, not undulate. Grain 19±24 mm. 8±9 pores, 2±3 mm .................................... P. argentea
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A2
361
Fig. 24. Plantaginaceae (1000Ö, darker images are of stained specimens). A±B, Plantago lanceolata type (e.g. P. lanceolata); C±D, P.
coronopus ; E±F, P. albicans; G±H, P. amplexicaulis; I±J, P. arenaria; K±L, P. major; M±N, P. media; O±Q, P. afra.
Fig. 25. Rosaceae (1000Ö). A±C, Sanguisorb a minor; D±E, Rosa (e.g. R. arvensis); F±I, Potentilla (e.g. F±G, P. reptans; H-I, P. argentea);
J±L, Rubus type (e.g. R. caesium); M±P, Alchemilla type (e.g. A. xanthochlora); Q, Agrimonia type (e.g. A. eupatoria); R±T, Crataegus (e.g.
C. monogyna ).
Grana 40 (2001)
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Fig. 25.
362
(For legend see page 361).
P. I. Chester and J. I. Raine
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Pollen and spore keys, Pindos Mts, Greece
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A1
363
D1 Pore not operculate, membrane granulate. Exine undulate-verrucate. Grain 22±28 mm. 4±6 pores, 3±4 mm.
Fig. 24G±H .................................................................................................................................... P. amplexicaulis
Pore without distinct annulus, with granules, no operculum.
F2 Grain >25 mm.
G2 8±9 pores, 3±4 mm. Verrucate, scabrate. Grain 27±36 mm ................................................................ P. atrata
G1 4±6 pores.
H2 Verrucae distinct. Pores not sharply dened, 2±4 mm. Grain 21±28 mm. Fig. 24I±J ............. P. arenaria
H1 Verrucae indistinct. Pores 2±4 mm. Grain 22±32 mm ............................................................. P. bellardii
F1 Grain <25 mm. Pores not sharply delimited. Microechinate, or verrucate/undulate.
I3 Distinctly micro-echinate. Pores 4±7, 1±2 mm. Grain 14±25 mm. Fig. 24O ±Q ...................................... P. afra
I2 Verrucae very coarse, scabrae distinct. Pores 7±11, 2±5 mm. Grain 18±27 mm. Fig. 24M±N ............. P. media
I1 Verrucae small, scabrae very weak. Pores 5±9, 1±3 mm. Grain 16±22 mm. Fig. 24K±L ...................... P. major
ROSACEAE
(Cf. Teppner 1966, Faegri & Iversen 1989: 287±288, Moore et al. 1991: 156±158)
A2
A1
With tectate operculum.
B2 Psilate±scabrate±echinate, oblate-globular. Sanguisorba.
C2 Operculum narrow, colpus very short. Fig. 25A±C ........................................................... Sanguisorba minor
C1 Operculum as broad as the intercolpium, colpus long ................................................ Sanguisorba oYcinalis
B1 Striate.
D2 Vallae coarse. Perforations not visible. Grain size c.20 mm.
E2 Vallae broad, diVuse.
F2 Polar area attened ............................................................................................... Fragaria vesca t.
F1 Polar area not attened. See also G1, below .................................................................. Potentilla
E1 Vallae narrow, distinct.
G2 Colpus ends rounded ........................................................................................... Fragaria viridis t.
G1 Colpus ends acute. Fig. 25F±I ........................................................................................ Potentilla
D1 Vallae ne. Tectum perforate.
H2 Vallae transversal, extremely ne. Operculum extremely narrow. Fig. 25Q ....................... Agrimonia t.
H1 Vallae submeridional to subtransversal. Operculum short. See also O2, below. Fig. 25D±E ........ Rosa
No tectate operculum, but colpus membrane occasionally with loose columellae.
I3 Microechinate-echinate. No perforations ................................................................................................ Filipendula
I2 Rugulate-microstriate/psilate.
J3 Rugulate-microstriate. No costae colpi nor granules on colpus membrane. Size c. 50 mm Mespilus germanica
J2 Rugulate to psilate, costae colpi and granules usually present ................................................... Cotoneaster t.
J1 Psilate. Polar projection usually subtriangular. Fig. 25M±P ......................................................... Alchemilla t.
I1 Striate.
K2 Vallae coarse. Colpus with distinct bridge .............................................................................................. Geum
K1 Vallae ne.
L2 Vallae paired.
M3 Striation weak. Size 29±42 mm. Fig. 25R±T ..................................................................... Crataegus
M2 Striation denite. Size 30±36 mm ........................................................................................... Pyrus
M1 Size 42±58 mm ....................................................................................................... Cydonia oblonga
L1 Vallae simple.
N2 Vallae indistinct. Perforations large. Fig. 25J±L ................................................................ Rubus t.
N1 Vallae distinct.
O2 Vallae high, frequently short. See also H1, above ........................................................... Rosa
O1 Vallae low and narrow.
P2 Vallae straight, moderately branched.
Q2 Perforations distinct. Vallae without dense structure. Fig. 26A±C ........... Sorbus t.
Q1 Perforations may be unobservable. Vallae with dense structure. Fig. 26D±E ..........
Prunus avium t.
P1 Vallae curved, branched, anastomosing.
R2 Vallae short, strongly curved. Fig. 26F±H .................................... Prunus spinosa t.
R1 Vallae moderately curved. With costae colpi, size c. 27 mm. Fig. 26I±L ....... Malus
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P. I. Chester and J. I. Raine
Fig. 26. Rosaceae, continued (1000Ö). A±C, Sorbus type (e.g. S. aucuparia); D±E, Prunus avium type (e.g. P. avium); F±H, Prunus spinosa
type (e.g. P. cerasifera); I±L, Malus (e.g. M. sylvestris).
PART 3
Composition of pollen and spore taxa
A full list of plant species included in each family is provided
below. The taxa for which we have pollen or spore data are
in boldface; followed by constituent plant species in the
Greek ora. The pollen or spore morphology of many plant
species remains unexamined, but the probable pollen morphology of these species are assumed based on plant taxonomy, allowing them to be placed in the most likely genus.
These species are listed at the end of each taxon and are
identied by a `?’. Taxonomic nomenclature, and family
arrangement follows Flora Europaea ( Tutin et al. 1964±80 ),
except where species have been described more recently by
Grana 40 (2001)
Strid (1986) or Strid & Tan (1991). For pollen and spores,
a genus without species indicates that the pollen can only be
identied to genus; a species name signies that only that
species has pollen of a particular type. The term `type’
(abbreviated `t.’ in the keys of Parts 1 and 2) implies a
pollen type which includes species from several families,
genera or species, as the case may be. The pollen types
take the names of associated plant taxa. The term `group’
is used for subdivisions of a few plant taxa, following the
usage of Northwest European Pollen Flora (Punt et al.
1976±1995).
Pollen and spore keys, Pindos Mts, Greece
PTERIDOPHYTA
LYCOPSIDA
SELAGINELLACEAE
Selaginella
S. denticulata, S. helvetica
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ISOETACEAE
Isoetes
I. lacustris
?Isoetes heldreichii
SPHENOPSIDA
EQUISETACEAE
Equisetum
E. arvense, E. uviatile, E. palustre, E. ramosissimum,
E. sylvaticum, E. telmateia
FILICOPSIDA
OPHIOGLOSSACEAE
Ophioglossum lusitanicum
Ophioglossum vulgatum
Botrychium
B. lunaria, B. simplex
ATHYRIACEAE
Athyrium lix-femina
Athyrium distentifolium
Cystopteris fragilis
ASPIDIACEAE
Polystichum
P. aculeatum, P. lonchitis, P. setiferum
?Polystichum x illyricum
Dryopteris lix-mas type
Dryopteris lix-mas, D. villarii, Gymnocarpium
robertianum
?Cystopteris dickieana, Dryopteris x tavelii
Dryopteris expansa
Gymnocarpium dryopteris
BLECHNACEAE
Blechnum spicant
POLYPODIACEAE
Polypodium
P. australe, P. vulgare
SALVINIACEAE
Salvinia natans
OSMUNDACEAE
Osmunda regalis
SINOPTERIDACEAE
Cheilanthes
C. persica
?Cheilanthes catanensis, C. fragrans, C. marantae
ADIANTACEAE
Adiantum capillus-veneris
PTERIDACEAE
Pteris cretica
Pteris vittata
GYMNOGRAMMACEAE
Anogramma leptophylla
HYPOLEPIDACEAE
Pteridium aquilinum
THELYPTERIDACEAE
Thelypteris palustris
Thelypteris phegopteris
ASPLENIACEAE
Asplenium type
Asplenium adiantum-nigrum , A. cuneifolium, A. ssum,
A. fontanum, A. lepidum, A. obovatum, A. onopteris,
A. petrarchae, A. ruta-muraria, A. septentrionale,
A. trichomanes, A. viride,Ceterach oYcinarum, Phyllitis
sagittata, P. scolopendrium
?Asplenium bornmuelleri, A. javorkeanum,
A. macedonicum
365
SPERMATOPHYTA
GYMNOSPERMAE
PINACEAE
Abies
A. alba, A. borisii-regis, A. cephalonica
Pseudotsuga menziesii
Picea abies
Cedrus
C. atlantica, C. deodara
Pinus subgenus Diploxylon
Pinus heldreichii, P. leucodermis, P. nigra, P. sylvestris
Pinus subgenus Haploxylon
Pinus peuce
CUPRESSACEAE
Juniperus
J. communis, J. excelsa, J. foetidissima, J. oxycedrus,
J. sabina
TAXACEAE
Taxus baccata
EPHEDRACEAE
Ephedra fragilis
Ephedra distachya type
Ephedra distachya, E. major
ANGIOSPERMAE
Dicotyledones
SALICACEAE
Salix triandra
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Salix fragilis type
Salix alba, S. aurita, S. caprea, S. fragilis, S. pedicellata,
S. pentandra, S. purpurea, S. viminalis
?S. aegyptiaca, S. amplexicaulis, S. cinerea, S. elaeagnos,
S. retusa
Populus
P. alba, P. canescens, P. nigra, P. tremula
Urtica urens type
Urtica urens, Pilea microphylla
Urtica pilulifera type
Urtica dubia, U. pilulifera
Parietaria
P. diVusa, P. lusitanica, P. oYcinalis
?Parietaria cretica
JUGLANDACEAE
Juglans regia
SANTALACEAE
Comandr a elegans
Osyris alba
Thesium divaricatum type
Thesium divaricatum, T. humile
?Thesium arvense, T. bavarum, T. bergeri,
T. brachyphyllum , T. linophyllon , T. parnassi
Thesium alpinum
BETULACEAE
Alnus glutinosa
Betula pendula
CORYLACEAE
Carpinus type
Carpinus betulus (mostly), C. orientalis (p.p.), Ostrya
carpinifolia p.p.
Ostrya type
Carpinus betulus p.p., C. orientalis (mostly), Ostrya
carpinifolia (mostly)
Corylus
C. avellana, C. colurna, C. maxima
FAGACEAE
Fagus
F. moesiaca, F. sylvatica
Castanea sativa
Quercus coccifera type
Quercus coccifera, Q. ilex
Quercus trojana type
Quercus cerris, Q. macrolepis, Q. trojana
?Quercus brachyphylla
Quercus frainetto type
Quercus frainetto, Q. pubescens, Q. virgiliana
?Quercus dalechampii, Q. pedunculiora, Q. polycarpa
ULMACEAE
Ulmus minor type
Ulmus minor
?Ulmus canescens
Ulmus procera
Ulmus glabra type
Ulmus glabra, U. laevis
Celtis
C. australis
?Celtis caucasia
MORACEAE
Morus nigra
Morus alba
Ficus carica
CANNABACEAE
Cannabi s sativa
Humulus lupulus
URTICACEAE
Urtica dioica
Grana 40 (2001)
LORANTHACEAE
Viscum album
Arceuthobium oxycedri
Loranthus europaeus
ARISTOLOCHIACEAE
Aristolochia
A. clematitis, A. longa
?Aristolochia macedonica, A. pallida, A. rotunda
RAFFLESIACEAE
Cytinus hypocistis
Cytinus ruber
POLYGONACEAE
Polygonu m bistorta type
Polygonu m bistorta
?Polygonum alpinum
Polygonu m viviparum
Polygonu m amphibium
Polygonu m aviculare type
Polygonu m arenarium, P. arenastrum, P. aviculare,
P. equisetiforme, P. patulum
?Polygonum longipes
Polygonu m persicaria type
Polygonu m hydropiper, P. lapathifolium, P. mite,
P. persicaria
?Polygonum salicifolium
Bilderdykia
B. convolvulus, B. dumetorum
Rumex type
Oxyria digyna
Rumex acetosella group
Rumex acetosella, R. tenuifolius
Rumex acetosa group
Rumex acetosa, R. thyrsiorus
?Rumex arifolius, R. gussonii
Rumex crispus p.p.
Rumex conglomeratus group
Rumex conglomeratus , R. pulcher, R. scutatus
?Rumex nepalensis, R. tuberosus
Rumex obtusifolius group
Rumex alpinus, R. crispus p.p., R. obtusifolius,
R. patientia p.p.
Pollen and spore keys, Pindos Mts, Greece
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?Rumex balcanicus, R. cristatus, R. hydrolapathum ,
R. kerneri
Rumex palustris group
Rumex crispus p.p., R. palustris, R. patientia p.p.
?Rumex dentatus
Rumex bucephalophoru s group
Rumex bucephalophorus , R. crispus p.p.
?Atraphaxis billardieri
Emex spinosa
CHENOPODIACEAE
Chenopodiacea e type
Chenopodiaceae: Atriplex halimus, A. hastata,
A. hortensis, A. littoralis, A. oblongifolia, A. patula,
A. rosea, A. tatarica, Beta vulgaris, Camphorosma
monspeliaca, Chenopodium album, C. ambrosioides,
C. bonus-henricus, C. cifolium, C. foliosum, C. glaucum,
C. hybridum, C. murale, C. opulifolium, C. polyspermum,
C. rubrum, C. urbicum, C. vulvaria, Corispermum nitidum,
Noaea mucronata, Petrosimonia brachiata, P. triandra,
Salsola kali, S. soda
Amaranthaceae: Amaranthus albus, A. deexus,
A. hybridus, A. lividus, A. retroexus
Caryophyllaceae: Sagina apetala, S. procumbens,
S. saginoides, S. subulata
PHYTOLACCACEAE
Phytolacca americana
MOLLUGINACEAE
Glinus type
Glinus lotoides, Mollugo cerviana
PORTULACACEAE
Montia fontana
Portulaca oleracea
CARYOPHYLLACEAE
Agrostemma githago
Arenaria serpyllifolia
?Arenaria biora, A. conferta, A. cretica, A. licaulis,
A. leptoclados, A. serpentini
Cerastium fontanum type
Cerastium alpinum, C. arvense, C. brachypetalum,
C. cerastoides, C. diVusum, C. fontanum, C. glomeratum,
C. pumilum, C. semidecandrum, Moenchia erecta,
Myosoton aquaticum, Stellaria alsine, S. graminea,
S. media, S. neglecta, S. nemorum
?Cerastium banaticum, C. decalvans, C. dinaricum,
C. dubium, C. gracile, C. moesiacum, C. rectum,
C. smolikanum, C. vourinense, Moenchia graeca,
M. mantica
Corrigiola litoralis
Dianthus deltoides type
Dianthus deltoides, D. sylvestris, Petrorhagia prolifera,
P. saxifraga
?Dianthus bi orus, D. brachyzonus, D. capitatus,
D. cruentus, D. degenii, D. diVusus, D. drenowskianus ,
D. formanekii, D. giganteus, D. gracilis, D. haematocalyx,
D. leptopetalus, D. minutiorus, D. myrtinervius,
D. pallens, D. petraeus, D. pinifolius, D. pontederae,
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D. stenopetalus, D. tristis, D. viscidus, Petrorhagia cretica,
P. fasciculata, P. glumacea, P. illyrica
Herniaria type
Herniaria cinerea, H. glabra, H. hirsuta, H. incana,
H. nigrimontium, H. parnassica
Holosteum umbellatum
Lychnis coronaria
Lychnis os-cuculi
Lychnis viscaria
Minuartia setacea type
Bufonia paniculata, Minuartia setacea
?Bufonia tenuifolia, Minuartia globulosa, M. glomerata
Moehringia trinervia type
Cucubalus baccifer, Minuartia verna, Moehringia trinervia
?Minuartia juniperina, Moehringia pentandra
??Bolanthus graecus, B. thessalus, Drypis spinosa,
Minuartia baldaccii, M. eurytanica, M. hirsuta,
M. garckeana, M. graminifolia, M. recurva, M. stellata,
Saponaria bellidifolia, S. calabrica, S. glutinosa, S. sicula
Paronychia type
Paronychia argentea, P. capitata, P. echinulata
Chenopodiaceae : Polycnemum arvense, P. heuVelii,
P. majus
?Paronychia albanica, P. cephalotes, P. kapela,
P. macedonica, P. polygonifolia , P. taurica
Scleranthus perennis
Scleranthus annuus type
Scleranthus annuus
?Scleranthus uncinatus
Silene alba type
Silene alba, S. conica
Silene vulgaris type
Silene armeria, S. dichotoma, S. italica, S. nutans,
S. otites, S. vulgaris
?Silene apetala, S. asterias, S. balcanica, S. bupleuroides,
S. caesia, S. cephallenia, S. chromodonta, S. ciliata,
S. colorata, S. compacta, S. congesta, S. cretica,
S. damboldtiana , S. densiora, S. fabarioides,
S. frivaldszkyana, S. gallica, S. genistifolia, S. gigantea,
S. haussknechtii, S. lerchenfeldiana, S. linifolia, S. lydia,
S. multicaulis, S. oligantha, S. paradoxa, S. parnassica,
S. pindicola, S. pusilla, S. radicosa, S. roemeri,
S. saxifraga, S. schwarzenbergeri, S. skorpilii, S. supina,
S. thessalonica, S. trinervia, S. ventricosa, S. viridiora,
S. waldsteinii
Spergula
Spergula arvensis, S. pentandra
Spergularia type
Polycarpon tetraphyllum, Spergularia diandra, S. marina,
S. media, S. rubra
Stellaria holostea
Vaccaria pyramidata
NYMPHAEACEAE
Nymphae a alba
Nuphar lutea
CERATOPHYLLACEAE
Ceratophyllum
C. demersum, C. submersum
Grana 40 (2001)
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RANUNCULACEAE
Helleborus
H. odorus
?Helleborus cyclophyllus
Nigella
N. arvensis, N. damascena, N. sativa
Trollius europaeus
Actaea spicata
Consolida type
Consolida orientalis, C. regalis, Delphinium ssum,
D. peregrinum
?Consolida brevicornis, Delphinium balcanicum,
D. hellenicum
Adonis
Adonis aestivalis, A. annua, A. ammea, A. microcarpa
Caltha type
Aquilegia vulgaris, Caltha palustris, Myosurus minimus
?Aquilegia amaliae, A. ottonis
Ranunculus acris type
Ranunculus acris group
Ranunculus acris, R. bulbosus, R. lanuginosus,
R. muricatus, R. nemorosus, R. polyanthemos , R. repens,
R. sardous
?Ranunculus constantinopolitanus , R. serbicus, R. velutinus
Ranunculus lingua
Ranunculus aquatilis group
Clematis ammula, Ranunculus aquatilis, R. peltatus,
R. pseudomontanus , R. rionii, R. sphaerospermus,
R. trichophyllus
?Ranunculus lateriorus
Pulsatilla halleri
Anemone nemorosa group
Anemone apennina, A. nemorosa
?Anemone blanda
Clematis vitalba group
Clematis vitalba, C. viticella
Ranunculus auricomus group
Ceratocephalus falcatus, Ranunculus auricomus,
R. cassubicus, R. fallax, R. monophyllus
Ranunculus caria
Ranunculus ophioglossifolius
Ranunculus sceleratus group
Ranunculus platanifolius, R. sceleratus
?Ranunculus brevifolius, R. brutius, R. demissus,
R. marginatus, R. neapolitanus, R. sartorianus
Ranunculus arvensis type
Ranunculus arvensis
?Ranunculus chius, R. gracilis, R. miliarakesii,
R. millefoliatus, R. millii, R. psilostachys, R. rumelicus,
R. spruneranus, R. subhomophyllu s
Thalictrum aquilegifolium
Thalictrum minus
Thalictrum lucidum
BERBERIDACEAE
Berberis vulgaris type
Berberis vulgaris
?Berberis cretica
Mahonia aquifolium
Leontice leontopetalum
Grana 40 (2001)
LAURACEAE
Laurus nobilis
PAPAVERACEAE
Chelidonium majus
Papaver dubium
Papaver hybridum
Papaver rhoeas type
Papaver rhoeas
?Papaver apulum, P. lecoqii
Papaver somniferum
Glaucium
G. corniculatum, G. leiocarpum
Papaver argemone type
Papaver argemone, Roemeria hybrida
Fumaria densiora type
Fumaria densiora, F. kralikii, F. oYcinalis, F. rostellata
?Fumaria petteri, F. schleicheri
Fumaria parviora
Corydalis solida type
Corydalis bulbosa, C. solida
Corydalis ochroleuca
Hypecoum
H. imberbe, H. pendulum
CAPPARIDACEAE
?Cleome ornithopodioide s
CRUCIFERAE
Hornungia type
Alyssum alyssoides, A. bertolonii, A. caliacrae,
A. chalcidicum, A. chlorocarpum, A. corymbosoides,
A. corymbosum, A. cuneifolium, A. desertorum,
A. diVusum, A. doereri, A. foliosum, A. heldreichii,
A. minus, A. minutum, A. montanum, A. murale,
A. obtusifolium, A. petraeum, A. pichleri, A. pulvinare,
A. repens, A. saxatile, A. sibiricum, A. smolikanum,
A. strigosum, A. tortuosum, A. umbellatum, Arabidopsis
thaliana, Berteroa incana, Capsella bursa-pastoris,
Cardaria draba, Descurainia sophia, Hornungia petraea,
Lepidium campestre, Thlaspi alpestre
?Aethionema saxatile, Berteroa mutabilis, B. obliqua,
B. orbiculata, Capsella grandiora, C. rubella, Lepidium
graminifolium, L. hirtum, L. perfoliatum, L. sativum,
Thlaspi bulbosum, T. epirotum, T. goesingense,
T. microphyllum, T. ochroleucum, T. perfoliatum,
T. praecox, T. rivale
Arabis
A. glabra, A. turrita
Sinapis type
Alliaria petiolata, Arabis alpina, Barbarea bracteosa,
B. sicula, B. vulgaris, Brassica napus, B. nigra, B. rapa,
Bunias erucago, Calepina irregularis, Camelina
microcarpa, Cardamine bulbifera, C. hirsuta, C. pratensis,
Diplotaxis muralis, D. viminea, Draba aizoides, D. athoa,
D. lasiocarpa, D. muralis, D. scardica, Erophila minima,
E. verna, Erysimum cephalonicum, E. cheiranthoides,
E. comatum, E. cuspidatum, E. diVusum, E. linariifolium,
E. microstylum, E. odoratum, E. pulchellum, E. pusillum,
E. repandum, E. sylvestre, Iberis saxatilis, I. sempervirens,
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Pollen and spore keys, Pindos Mts, Greece
I. umbellata, Isatis lusitanica, I. tinctoria, I. vermia,
Lepidium latifolium, L. ruderale, Lunaria annua,
L. rediviva, Matthiola fruticulosa, Raphanu s raphanistrum,
Rorippa amphibia, R. prolifera, R. pyrenaica, R. sylvestris,
Sinapis alba, S. arvensis, Sisymbrium altissimum, S. irio,
S. loeselii, S. oYcinale, S. orientale, S. polyceratium,
Teesdalia coronopifolia
?Alyssoides utriculata, Arabis allionii, A. bryoides,
A. caucasica, A. collina, A. laxa, A. muralis,
A. pseudoturritis, A. recta, A. sagittata, A. subava,
Aubrieta deltoidea, A. glabrescens, A. gracilis,
A. intermedia, Bornmuellera baldaccii, B. tymphaea,
Camelina alyssum, C. rumelica, C. sativa, Cardamine
barbaraecoides, C. carnosa, C. glauca, C. graeca,
C. hirsuta, C. pectinata, C. plumieri, C. raphanifolia,
Clypeola jonthlaspi, Conringia austriaca, C. orientalis,
Coronopus squamatus, Eruca vesicaria, Hesperis dinarica,
H. laciniata, H. matronalis, H. rechingeri, H. sylvestris,
H. theophrasti, Hirschfeldia incana, Hutchinsia alpina,
Hymenolobus procumbens, Iberis odorata, I. pinnata,
I. pruitii, I. umbellata, Lobularia maritima, Malcolmia
africana, M. angulifolia, M. bicolor, M. graeca,
M. maritima, M. orsiniana, Myagrum perfoliatum,
Nasturtium oYcinale, Neslia paniculata, Peltaria
emarginata, Ptilotrichum cyclocarpum, Rapistrum rugosum
RESEDACEAE
Reseda
R. lutea, R. luteola, R. phyteuma
CRASSULACEAE
Umbilicus
U. erectus, U. horizontalis, U. rupestris
Jovibarba heuVelii
Crassula
C. tillaea, C. vaillantii
Sedum
S. acre, S. album, S. alpestre, S. annuum, S. athoum,
S. atratum, S. cepaea, S. dasyphyllum , S. exuosum,
S. hispanicum, S. laconicum, S. litoreum, S. magellense,
S. ochroleucum, S. pallidum, S. reexum, S. sartorianum,
S. sediforme, S. telephium, S. tenuifolium
Sempervivum
S. ballsii, S. ciliosum, S. kindingeri, S. marmoreum,
S. reginae-amaliae, S. zeleborii
SAXIFRAGACEAE
Chrysosplenium alternifolium
Saxifraga stellaris
Saxifraga tridactylites type
Saxifraga chrysosplenifolia, S. exarata, S. rotundifolia,
S. tridactylites
Saxifraga oppositifolia type
Saxifraga adscendens, S. glabella, S. marginata,
S. oppositifolia, S. paniculata
Saxifraga bulbifera
Saxifraga sempervivum type
Saxifraga porophylla , S. sempervivum, S. stribrnyi
?Saxifraga graeca, S. grisebachii, S. luteoviridis,
S. scardica, S. spruneri, S. taygetea
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PARNASSIACEAE
Parnassia palustris
GROSSULARIACEAE
Ribes
R. alpinum, R. multiorum
PLATANACEAE
Platanus orientalis
ROSACEAE
Filipendula
F. ulmaria, F. vulgaris
Rubus type
Rubus caesius, R. candicans, R. canescens, R. corylifolius,
R. hirtus, R. idaeus, R. serpens, R. thyrsiorus,
R. ulmifolius
?Rubus thessalus
??Spiraea crenata
Sorbus type
Sorbus aria, S. aucuparia, S. domestica, S. graeca,
S. torminalis, S. umbellata
?Amelanchier ovalis
Rosa
R. agrestis, R. andegavensis, R. arvensis, R. caesia,
R. canina, R. corymbifera, R. elliptica, R. foetida,
R. gallica, R. majalis, R. micrantha, R. montana,
R. moschata, R. nitidula, R. obtusifolia, R. pendulina,
R. pimpinellifolia, R. rubiginosa, R. squarrosa, R. villosa,
R. vosagiaca
?Rosa caryophyllacea, R. glutinosa, R. heckeliana,
R. orientalis, R. sempervirens, R. turcica
Cotoneaster type
Cotoneaster integerrimus, C. nebrodensis
?Pyracantha coccinea
Mespilus germanica
Prunus avium type
Prunus avium, P. mahaleb
Prunus spinosa type
Prunus cerasus, P. domestica, P. persica, P. spinosa
?Prunus armeniaca, P. cerasifera, P. cocomilia, P. dulcis,
P. prostrata
Malus
M. domestica, M. pumila, M. sylvestris
?Malus orentina
Pyrus
P. amygdaliformis , P. communis, P. cordata, P. pyraster
?Pyrus eleagrifolia, P. salvifolia
Crataegus
C. heldreichii, C. laciniata, C. monogyna, C. pentagyna
?Crataegus schraderana
Cydonia oblonga
Agrimonia type
Agrimonia eupatoria, Aremonia agrimonoides
Sanguisorba oYcinalis
Sanguisorba minor
Sanguisorba minor ssp. minor, S. minor ssp. muricata
Geum
G. coccineum, G. molle, G. montanum, G. reptans,
G. urbanum
Grana 40 (2001)
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P. I. Chester and J. I. Raine
Potentilla
P. argentea, P. aurea, P. erecta, P. inclinata,
P. micrantha, P. recta, P. reptans, P. sterilis, P. supina
?Potentilla astracanica, P. australis, P. detommasii,
P. geoides, P. pedata, P. speciosa
Fragaria viridis type
Fragaria viridis
?Fragaria x ananassa
Fragaria vesca type
Fragaria vesca
?Sibbaldia parviora
Alchemilla type
Alchemilla xanthochlora, Aphanes arvensis, A. microcarpa
?Alchemilla acutiloba, A. cinerea, A. fallax, A. gorcensis,
A. heterotricha, A. incisa, A. plicatula, A. straminea,
A. subglabra group
LEGUMINOSAE
Genista type
Cytisus decumbens, C. procumbens, C. villosus, Genista
acanthoclada , G. albida, G. carinalis, G. hassertiana,
G. januensis, G. radiata, G. tinctoria
?Chamaecytisus austriacus, C. ciliatus, C. heuVelii,
C. hirsutus, C. polytrichus, C. spinescens, C. supinus,
Lembotropis nigricans, Podocytisus caramanicus
Onobrychis
O. aequidentata, O. alba, O. arenaria, O. caput-galli,
O. ebenoides, O. hypargyrea, O. montana, O. oxyodonta,
O. pindicola
Hedysarum coronarium
Ononis type
Colutea arborescens, Melilotus alba, M. altissima,
M. indica, M. neapolitana, M. oYcinalis, M. segetalis,
Ononis pusilla, O. reclinata, O. spinosa
Trifolium dubium type
Medicago lupulina, Trifolium angustifolium, T. arvense,
T. aureum, T. badium, T. campestre, T. dubium,
T. lappaceum, T. micranthum, T. nigrescens, T. pallidum,
T. resupinatum
?Medicago aculeata, M. ciliaris, M. coronata,
M. disciformis, M. orbicularis, M. praecox, M. prostrata,
M. rigidula, M. scutellata, M. turbinata, Trifolium
alpestre, T. aurantiacum, T. bocconei, T. cherieri,
T. dalmaticum, T. diVusum, T. dolopium, T. echinatum,
T. glomeratum, T. heldreichianum, T. hirtum, T. latinum,
T. leucanthum, T. noricum, T. ochroleucon,
T. pannonicum , T. parnassi, T. patens, T. patulum,
T. phleoides, T. physodes, T. pignantii, T. pilczii,
T. purpureum, T. sebastianii, T. smyrnaeum, T. strictum,
T. suVocatum, T. tenuifolium, T. trichopterum,
T. vesiculosum
Trifolium repens type
Medicago arabica, M. minima, M. polymorpha ,
Scorpiurus muricatus, Trifolium fragiferum, T. globosum,
T. hybridum, T. incarnatum, T. medium, T. pratense,
T. repens, T. scabrum, T. squarrosum, T. striatum,
T. subterraneum
Lotus type
Lotus aegaeus, L. angustissimus, L. corniculatus,
L. ornithopodioides , L. palustris, L. parviorus,
Grana 40 (2001)
L. stenodon, L. tenuis, L. uliginosus, Tetragonolobus
biorus, T. purpureus, Trigonella monspeliaca
?Glycyrrhiza echinata, G. glabra, Trigonella caerulea,
T. foenum-graecum, T. procumbens, T. spicata, T. striata
??Hymenocarpus circinnatus, Securigera securidaca
Lathyrus pratensis type
Lathyrus alpestris, L. aphaca, L. cicera, L. digitatus,
L. grandi orus, L. hallersteinii, L. hirsutus, L. latifolius,
L. laxiorus, L. niger (mostly), L. nissolia, L. pallescens,
L. pratensis, L. setifolius, L. sphaericus, L. sylvestris,
L. tuberosus, L. venetus, L. vernus, Lupinus albus,
Trigonella gladiata
?Lathyrus ochrus
Lens
L. culinaris
?Lens ervoides, L. nigricans
Vicia sativa type
Vicia cracca (sometimes), V. hirsuta (mostly), V. hybrida,
V. lutea, V. peregrina, V. sativa, V. sepium (mostly),
V. tetrasperma (mostly)
?Vicia articulata, V. barbazitae, V. canescens,
V. cassubica, V. dumetorum, V. ervilia, V. grandiora,
V. incana, V. melanops, V. narbonensis, V. onobrychioides,
V. pannonica , V. pubescens, V. sibthorpii, V. villosa
Vicia lathyroides
Vicia faba
Vicia cracca type
Lathyrus niger (sometimes), Vicia cracca (mostly),
V. hirsuta (sometimes), V. sepium (sometimes),
V. tenuifolia, V. tetrasperma (sometimes)
Astragalus
A. depressus, A. exscapus, A. glycyphyllos, A. hamosus,
A. monspessulanus, A. onobrychis, A. purpureus,
A. sesameus, A. vesicarius
?Astragalus angustifolius, A. baldaccii, A. creticus,
A. glycyphylloides , A. mayeri, A. parnassi, A. pubiorus,
A. sinaicus, A. sirinicus, A. subuliformis
Galega oYcinalis
Coronilla scorpioides type
Coronilla scorpioides, C. valentina, C. varia
?Coronilla coronata, C. cretica, C. elegans, C. rostrata
Medicago sativa
Medicago sativa p.p.
Ornithopus compressus
Dorycnium
D. graecum, D. hirsutum, D. pentaphyllum, D. rectum
Trifolium montanum type
Medicago sativa p.p., Trifolium montanum
Anthyllis
A. aurea, A. montana, A. vulneraria
Coronilla emerus
Spartium junceum
Psoralea type
Psoralea bituminosa, Vigna unguiculat a
Phaseolus
P. vulgaris
?Phaseolus coccineus
??Cicer arietinum, Glycine max
Pisum sativum
Lathyrus sativus
Pollen and spore keys, Pindos Mts, Greece
Ceratonia siliqua
Cercis siliquastrum
Hippocrepis
H. ciliata, H. comosa, H. unisiliquosa
Robinia pseudoacacia
Arachis hypogaea
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OXALIDACEAE
Oxalis
O. acetosella, O. corniculata, O. pes-caprae
GERANIACEAE
Geranium
Geranium robertianum group
G. purpureum, G. pusillum, G. pyrenaicum,
G. robertianum, G. rotundifolium
Geranium bohemicum group
G. bohemicum, G. macrorrhizum
?Geranium aristatum, G. cinereum, G. reexum
Geranium pratense group
G. pratense, G. sanguineum, G. sylvaticum
?Geranium asphodeloides, G. macrostylum, G. tuberosum
Geranium molle group
G. columbinum, G. dissectum, G. divaricatum, G. lucidum,
G. molle, G. versicolor
?Geranium brutium
Erodium
E. botrys, E. ciconium, E. cicutarium, E. hoeVtianum,
E. malacoides, E. moschatum
ZYGOPHYLLACEAE
Peganum harmala
Tribulus terrestris
LINACEAE
Linum usitatissimum type
Linum bienne, L. usitatissimum
Linum austriacum type
Linum austriacum, L. perenne, L. punctatum
Radiola linoides
Linum catharticum type
Linum catharticum
Linum tenuifolium
Linum trigynum
Linum strictum group
Linum hirsutum, L strictum
?Linum pubescens, L. spathulatum
Linum hologynu m type
Linum hologynu m
?Linum aroanium, L. capitatum, L. elegans, L. nodiorum,
L. tauricum, L. thracicum
EUPHORBIACEAE
Andrachne telephioides
Mercurialis perennis
Mercurialis annua type
Mercurialis annua
?Mercurialis ovata
Euphorbia
E. amygdaloides , E. brittingeri, E. chamaesyce, E. esula,
371
E. exigua, E. falcata, E. helioscopia, E. lathyris,
E. nicaeensis, E. palustris, E. peplus, E. platyphyllos ,
E. pubescens, E. segetalis, E. seguierana, E. serrulata,
E. spinosa, E. villosa
?Euphorbia agraria, E. aleppica, E. barrelieri,
E. cyparissias, E. deexa, E. epithymoides, E. avicoma,
E. glabri ora, E. heldreichii, E. herniariifolia, E. lathyris,
E. lingulata, E. lucida, E. myrsinites, E. oblongata,
E. phymatosperma, E. salicifolia, E. taurinensis
Ricinus communis
RUTACEAE
Dictamnus albus
Ruta chalepensis
Ruta montana type
Ruta montana
?Ruta graveolens
Haplophyllum
?Haplophyllum boissieranum, H. coronatum, H. patavinum
Citrus
C. aurantium, C. limon, C. sinensis
SIMAROUBACEAE
Ailanthus altissima
POLYGALACEAE
Polygal a monspeliaca type
Polygala monspeliaca
?Polygala alpestris, P. major, P. nicaeensis, P. supina
Polygal a vulgaris type
Polygala vulgaris
?Polygala comosa
ANACARDIACEAE
Rhus coriaria
Schinus molle
Pistacia
P. lentiscus, P. terebinthus
Cotinus coggygria
ACERACEAE
Acer
A. campestre, A. monspessulanum , A. opalus,
A. platanoides, A. pseudoplatanu s
?Acer heldreichii, A. hyrcanum, A. obtusatum,
A. sempervirens, A. tataricum
HIPPOCASTANACEAE
Aesculus hippocastanum
BALSAMINACEAE
Impatiens noli-tangere
AQUIFOLIACEAE
Ilex aquifolium
CELASTRACEAE
Euonymu s europaeus type
Euonymus europaeus
?Euonymus latifolius, E. verrucosus
Grana 40 (2001)
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BUXACEAE
Buxus sempervirens
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RHAMNACEAE
Frangula
F. alnus
?Frangula rupestris
Rhamnus type
Paliurus spina-christi, Rhamnus alpinus, R. saxatilis,
Ziziphus jujuba, Z. lotus
?Paliurus microcarpus, Rhamnus prunifolius
VITACEAE
Vitis vinifera
Vitis vinifera ssp. sylvestris, V. vinifera ssp. vinifera
TILIACEAE
Tilia platyphyllos
?Tilia rubra
Tilia cordata type
Tilia cordata, T. tomentosa
MALVACEAE
Abutilon type
Abutilon theophrasti, Malvella sherardiana
Alcea type
Alcea pallida, Althaea cannabina
?Alcea heldreichii
Althaea hirsuta
Althaea oYcinalis
Malva moschata
Malva pusilla
Malva pusilla p.p.
Malva sylvestris type
Lavatera thuringiaca, Malva neglecta, M. nicaeensis,
M. pusilla p.p., M. sylvestris, M. verticillata
Hibiscus
H. trionum
?Hibiscus syriacus
Abelmoschus esculentus
Gossypium
G. herbaceum, G. hirsutum
THYMELAEACEAE
Daphne
D. blagayana, D. gnidium, D. laureola, D. mezereum,
D. oleoides
ELAEAGNACEAE
Elaeagnus angustifolia
GUTTIFERAE
Hypericum hircinum type
Hypericum hircinum
?Hypericum empetrifolium
Hypericum perforatum type
Hypericum hirsutum, H. perforatum, H. tetrapterum
?Hypericum annulatum, H. barbatum, H. linarioides,
H. olympicum, H. rumeliacum, H. spruneri, H. thasium
VIOLACEAE
Viola hirta type
Grana 40 (2001)
Viola alba, V. elatior, V. hirta, V. odorata, V. riviniana
p.p.
Viola riviniana type
Viola reichenbachiana, V. riviniana p.p.
Viola arvensis type
Viola arvensis, V. kitaibeliana, V. tricolor
?Viola aetolica, V. allchariensis, V. brachyphylla ,
V. chelmea, V. dukadjinica, V. epirota, V. eximia,
V. gracilis, V. heterophylla, V. magellensis, V. orphanidis,
V. pyrenaica, V. rupestris, V. sieheana
CISTACEAE
Helianthemum
H. apenninum , H. canum, H. ledifolium, H. nummularium ,
H. oelandicum, H. salicifolium
Tuberaria guttata
Cistus monspeliensis type
Cistus monspeliensis
?Cistus incanus, C. parviorus
Cistus salvifolius
Fumana
F. aciphylla, F. arabica, F. bonapartei
TAMARICACEAE
Myricaria germanica
Tamarix
T. dalmatica, T. hampeana , T. parviora, T. smyrnensis,
T. tetrandra
ELATINACEAE
Elatine alsinastrum
CUCURBITACEAE
Cucumis melo
?Citrullus lanatus
Bryonia
B. alba, B. cretica
Lagenaria siceraria
Cucurbita
C. cifolia, C. maxima, C. mixta, C. moschata, C. pepo
LYTHRACEAE
Lythrum salicaria type
Lythrum hyssopifolia, L. junceum, L. salicaria,
L. tribracteatum
?Lythrum borysthenicum, L. virgatum
Lythrum portula
TRAPACEAE
Trapa natans
MYRTACEAE
Myrtus communis
PUNICACEAE
Punica granatum
ONAGRACEAE
Circaea lutetiana
Ludwigia palustris
Pollen and spore keys, Pindos Mts, Greece
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Epilobium angustifolium type
Epilobium angustifolium, Oenothera biennis
Epilobium hirsutum type
Epilobium hirsutum, E. montanum, E. palustre,
E. parviorum, E. tetragonum
?Epilobium adenocaulon , E. alsinifolium,
E. anagallidifolium , E. dodonaei , E. gemmascens,
E. lanceolatum, E. obscurum, E. roseum
HALORAGACEAE
Myriophyllum verticillatum
Myriophyllum spicatum
THELIGONACEAE
Theligonum cynocrambe
CORNACEAE
Cornus sanguinea
Cornus mas
ARALIACEAE
Hedera helix
UMBELLIFERAE
Hydrocotyle vulgaris
Anthriscus caucalis
Torilis nodosa
Physospermum cornubiense
Smyrnium
S .lusatrum, S. perfoliatum
Ligusticum
?Ligusticum lucidum, L. rhizomaticum
Bupleurum
B. asperuloides, B. falcatum, B. avicans, B. fruticosum,
B. gerardi, B. glumaceum, B. karglii, B. lancifolium,
B. praealtum, B. rotundifolium, B. tenuissimum
Sison amomum
Carum
?Carum heldreichii, C. multiorum, C. rigidulum
??StefanoYa daucoides
Berula erecta
Trinia
T. dalechampii, T. glauca
Sanicula europaea
Conium maculatum
Cachrys
?Cachrys cristata, C. ferulacea
Cnidium silaifolium
Seseli type
Opopanax chironium, Seseli libanotis, S. pallasii,
S. parnassicum, S. peucedanoides, S. rigidum,
S. tortuosum
?Opopana x hispidus
Oenanthe type
Oenanthe aquatica group
O. aquatica, O. lachenalii
Oenanthe pimpinelloides group
O. pimpinelloides
?Oenanthe tenuifolia
373
Oenanthe silaifolia group
O. silaifolia
?Athamanta macedonica, Oenanthe banatica
O. stulosa
Cicuta virosa
Chaerophyllum hirsutum type
Chaerophyllum hirsutum
?Chaerophyllum aromaticum
Chaerophyllum temulentum type
Chaerophyllum temulentum
??Myrrhoides nodosa
Ammi
A. majus, A. visnaga
Pimpinella saxifraga
Torilis arvensis
Anethum graveolens
Falcaria vulgaris
Peucedanum austriacum type
Peucedanum austriacum
?Peucedanum aegopodioides , P. longifolium,
P. oligophyllum , P. stridii, P. vittijugum, P. vourinense
Angelica sylvestris
Petroselinum crispum
Peucedanum schotti
Bunium
B. ferulaceum
?Bunium tenerum
??Huetia cynapioides
Foeniculum vulgare ssp. vulgare
F. vulgare ssp. vulgare
??Ferulago campestris, F. nodosa, F. sylvatica
Anthriscus sylvestris type
Anthriscus sylvestris
?Anthriscus nemorosa
Orlaya
O. grandi ora
?Orlaya daucorlaya, O. kochii, O. topaliana
Pastinaca sativa
Eryngium
E. campestre
?Eryngium amethystinum, E. creticum, E. palmatum
Scandix type
Caucalis platycarpos, Scandix australis, S. pecten-veneris,
Turgenia latifolia
Heracleum sphondyliu m
Pimpinella anisum type
Pimpinella anisum
?Pimpinella peregrina, P. tragium
Anthriscus cerefolium
Laserpitium siler
Torilis japonica type
Torilis japonica
?Torilis leptophylla, T. ucranica
Daucus
D. broteri, D. carota
?Daucus guttatus
Coriandrum type
Bifora testiculata, Coriandrum sativum
Aegopodium podagraria
Chaerophyllum aureum
Grana 40 (2001)
374
P. I. Chester and J. I. Raine
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Malabaila
M. aurea, M. involucrata
Tordylium oYcinale
Echinophora type
Echinophora tenuifolia, Tordylium maximum
Conopodiu m capillifolium (= Conopodiu m majus type in
NW4)
?Conopodium capillifolium
PYROLACEAE
Monotropa hypopitys
Pyrola
P. chlorantha, P. minor
Moneses uniora
ERICACEAE
Erica herbacea
Vaccinium myrtillus
PRIMULACEAE
Soldanella
?Soldanella hungarica, S. pindicola
Cyclamen
C. hederifolium
?Cyclamen graecum, C. repandum
Androsace villosa
Androsace maxima
Primula
P. elatior, P. veris, P. vulgaris
?Primula halleri
Lysimachia
L. atropurpurea, L. dubia, L. nummularia, L. punctata,
L. vulgaris
Anagallis arvensis type
Anagallis arvensis, A. foemina
Anagallis tenella type
Anagallis minima, A. tenella
Asterolinon linum-stellatum
Samolus valerandi
PLUMBAGINACEAE
Armeria maritima type
Acantholimon androsaceum, Armeria canescens,
A. maritima, A. rumelica, A. sancta, A. undulata,
Goniolimon tataricum
Plumbago europaea
EBENACEAE
Diospyros lotus
STYRACACEAE
Styrax oYcinalis
OLEACEAE
Fraxinus excelsior type
Fraxinus angustifolia ssp. oxycarpa, F. excelsior
Fraxinus angustifolia
Fraxinus ornus
Ligustrum vulgare
Grana 40 (2001)
Phillyrea
P. latifolia, P. media
Jasminum
J. fruticans
?Jasminum humile
GENTIANACEAE
Gentiana verna
Gentianella ciliata
Cicendia liformis
Centaurium
C. erythraea, C. pulchellum, C. tenuiorum
Blackstonia perfoliata
Blackstonia perfoliata ssp. perfoliata, B. perfoliata ssp.
serotina
Gentiana lutea type
Gentiana asclepiadea, G. cruciata, G. lutea
MENYANTHACEAE
Nymphoide s peltata
APOCYNACEAE
Vinca herbacea
ASCLEPIADACEAE
Periploca graeca
Vincetoxicum
V. hirundinaria
?Vincetoxicum fuscatum, V. speciosum
CONVOLVULACEAE
Convolvulus
C. arvensis, C. betonicifolius, C. boissieri, C. cantabrica,
C. holosericeus
Cuscuta epithymum type
Cuscuta epithymum, C. planiora
?Cuscuta approximata, C. australis, C. suaveolens
Cuscuta europaea type
Cuscuta campestris, C. europaea, C. palaestina
Cuscuta monogyn a
Calystegia
C. sepium, C. silvatica
Ipomoea
I. batatas, I. purpurea
BORAGINACEAE
Cynoglossum type
Cynoglossum creticum, C. oYcinale, Rindera graeca
?Cynoglossum hungaricum, C. nebrodense, Solenanthus
albanicus
Heliotropium
H. europaeum, H. suaveolens, H. supinum
?Heliotropium hirsutissimum
Myosotis type
Asperugo procumbens, Myosotis arvensis, M. laxa ssp.
caespitosa, M. nemorosa, M. ramosissima, M. sicula,
M. stricta, M. sylvatica, Omphalode s verna
?Myosotis incrassata, M. refracta, M. sparsiora,
M. suaveolens, Rochelia disperma
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Pollen and spore keys, Pindos Mts, Greece
Lappula
L. squarrosa
?Lappula marginata
Echium
E. italicum, E. plantagineum, E. vulgare
Alkanna type
Alkanna tinctoria
?Alkanna corcyrensis, A. noneiformis, A. pindicola,
Halacsya sendtneri
Borago oYcinalis
Cerinthe type
Cerinthe minor, C. retorta
??Moltkia petraea, Onosma elegantissima, O. graeca,
O. helvetica, O. heterophylla, O. mattirolii, O. montana,
O. pygmaeum , O. rhodopea, O. taurica, O. visianii
Symphytum
S. bulbosum, S. ottomanum
Buglossoides arvensis type
Buglossoides arvensis
?Buglossoides tenuiora
Neatostema apulum
Anchusa arvensis type
Anchusa arvensis, A. azurea
?Anchusa cretica
Nonea
?Nonea pallens, N. pulla, N. ventricosa
Pulmonaria mollis
Anchusa oYcinalis type
Anchusa oYcinalis, A. stylosa
?Anchusa macedonica, A. serpentinicola, A. thessala
Lithospermum
L. oYcinale
?Lithospermum goulandriorum
Buglossoides purpurocaerulea
VERBENACEAE
Verbena
V. oYcinalis, V. supina
Vitex agnus-castus
CALLITRICHACEAE
Callitriche obtusangula
Callitriche stagnalis type
Callitriche stagnalis
?Callitriche truncata
LABIATAE
Teucrium
T. chamaedrys, T. halacsyanum , T. montanum, T. polium,
T. scordium
Stachys sylvatica type
Lamium album, L. amplexicaule, L. maculatum,
L. purpureum, Leonurus cardiaca, L. marrubiastrum,
Melittis melissophyllum, Stachys alopecuros, S. alpina,
S. annua, S. cretica, S. germanica, S. goulimyi,
S. menthifolia, S. oYcinalis, S. palustris, S. sylvatica
?Lamiastrum galeobdolon , Lamium bidum,
L. garganicum, Stachys atherocalyx, S. balcanica,
S. cassia, S. decumbens, S. iva, S. leucoglossa, S. obliqua,
375
S. parolini, S. plumosa, S. scardica, S. serbica,
S. spinulosa, S. thirkei, S. tymphaea
Stachys arvensis
Stachys recta
Stachys angustifolia
Scutellaria type
Ajuga chamaepitys, A. genevensis, A. iva, A. reptans,
Scutellaria alpina, S. galericulata, S. orientalis
?Ajuga pseudochia, Ballota macedonica, B. nigra,
Scutellaria altissima, S. columnae, S. rubicunda
Marrubium
M. vulgare
?Marrubium peregrinum, M. pestalozzae, M. thessalum,
M. velutinum
Phlomis
P. herba-venti, P. tuberosa
?Phlomis samia
Sideritis
S. montana
?Sideritis lanata, S. perfoliata, S. scardica, S. syriaca
Prunella type
Glechoma hederacea, Nepeta cataria, N. nuda, Prunella
grandi ora, P. laciniata, P. vulgaris
?Glechoma hirsuta, Nepeta sibthorpii, N. spruneri
Salvia oYcinalis type
Salvia aethiopis, S. amplexicaulis, S. argentea,
S. candidissima, S. eichlerana, S. glutinosa, S. grandiora,
S. nemorosa, S. oYcinalis, S. ringens, S. sclarea,
S. verbenaca, S. viridis
Salvia verticillata
Mentha type
Acinos arvensis, Clinopodium vulgare, Lycopus europaeus,
Melissa oYcinalis, Mentha aquatica, M. arvensis,
M. longifolia, M. x piperita, M. pulegium, M. spicata,
M. suaveolens, Origanum vulgare, Thymus praecox,
T. pulegioides
?Acinos alpinus, A. rotundifolius, A. suaveolens,
Calamintha grandiora, C. nepeta, C. sylvatica, Lycopus
exaltatus, Mentha microphylla, Micromeria cremnophila,
M. cristata, M. juliana, M. myrtifolia, Origanum
heracleoticum, Satureja horvatii, S. montana, Thymus
cherlerioides, T. dolopicus, T. leucospermus, T.
leucotrichus, T. longicaulis, T. ocheus, T. sibthorpii,
T. stojanovii, T. striatus, T. teucrioides, T. thracicus,
Ziziphora capitata
SOLANACEAE
Solanum dulcamara
Solanum nigrum type
Capsicum annuum, Lycopersicon esculentum, Physalis
alkekengi, Solanum elaeagnifolium, S. luteum, S. nigrum,
S. tuberosum
?Solanum cornutum, S. laciniatum
Solanum melongena
Atropa type
Atropa bella-donna, Hyoscyamus albus
Hyoscyamus niger
Datura stramonium
Nicotiana
N. rustica, N. tabacum
Grana 40 (2001)
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376
P. I. Chester and J. I. Raine
SCROPHULARIACEAE
Scrophularia ttype
Chaenorhinu m minus, Cymbalaria microcalyx, C. muralis,
Limosella aquatica, Linaria alpina, L. angustissima,
L. arvensis, L. genistifolia, L. simplex, L. vulgaris,
Misopates orontium, Scrophularia nodosa, S. scopolii,
S. umbrosa, Sibthorpia europaea, Verbascum blattaria,
V. chaixii, V. densiorum, V. nigrum, V. phlomoides ,
V. pulverulentum, V. sinuatum, V. thapsus
?Digitalis ferruginea, D. grandiora, D. laevigata,
D. lanata, D. viridiora, Limosella tenella, Linaria
peloponnesiaca, Scrophularia aestivalis, S. canina,
S. heterophylla, S. lucida, S. peregrina, Verbascum
adenanthum , V. baldaccii, V. banaticum, V. botuliforme,
V. dieckianum, V. epixanthinum, V. eriophorum,
V. glabratum, V. glandulosum , V. graecum, V. guicciardii,
V. leucophyllum, V. longifolium, V. macrurum,
V. mallophorum , V. orientale, V. ovalifolium,
V. phoeniceum, V. samniticum, V. speciosum,
V. undulatum
Gratiola oYcinalis
Kickxia
K. elatine, K. spuria
Rhinanthus type
Orobanchaceae: Orobanche purpurea, Rhinanthus minor
?Rhinanthus mediterraneus, R. melampyroides , R. pindicus,
R. pubescens, R. sintenisii
Odontites type
Odontites glutinosa, Parentucellia latifolia, P. viscosa
Euphrasia
E. minima, E. pectinata, E. salisburgensis, E. stricta
?Euphrasia liburnica
Veronica anagallis-aquatica type
Odontites verna, Veronica anagallis-aquatica ,
V. beccabunga, V. persica, V. polita
?Odontites lutea, Veronica acinifolia, V. agrestis,
V. anagalloides , V. aphylla, V. argute-serrata, V. arvensis,
V. austriaca, V. bozakmanii , V. catenata, V. dillenii,
V. glauca, V. hederifolia, V. montana, V. oYcinalis,
V. praecox, V. prostrata, V. rhodopaea, V. scardica,
V. scutellata, V. thymifolia, V. urticifolia, V. verna
Veronica triphyllos type
Veronica chamaedrys, V. cymbalaria, V. serpyllifolia,
V. triphyllos
Melampyrum
M. sylvaticum
?Melampyrum barbatum, M. ciliatum, M. doereri,
M. mbriatum, M. heracleoticum, M. hoermannianum ,
M. scardicum
Bellardia trixago
Pedicularis
P. comosa, P. verticillata
?Pedicularis brachyodonta , P. graeca, P. hoermanniana ,
P. leucodon, P. limnogena, P. petiolaris
RUBIACEAE
Putoria calabrica
Crucianella
C. angustifolia
?Crucianella latifolia
Sherardia arvensis
Galium type
Asperula aristata, A. arvensis, A. cynanchica, A. laevigata,
A. taurina, Cruciata laevipes, Galium album, G. aparine,
G. corrudifolium, G. debile, G. divaricatum, G. elongatum,
G. lucidum, G. odoratum, G. palustre, Galium x
pomeranicum, G. rotundifolium, G. setaceum, G. spurium,
G. tricornutum, G. verrucosum, G. verticillatum, G. verum,
Rubia peregrina, R. tinctorum
?Asperula chlorantha, A. doereri, A. involucrata, A. lutea,
A. purpurea, A. rumelica, A. tenella, Cruciata glabra,
C. pedemontana, Galium absurdum, G. advenum,
G. anisophyllon , G. asparagifolium, G. breviramosum,
G. circae, G. degenii, G. hellenicum, G. humifusum,
G. incanum, G. intricatum, G. laconicum, G. monasterium,
G. octonarium, G. ophiolithicum, G. oreophilum,
G. pisoderium, G. procurrens, G. pseudaristatum,
G. rhodopeum, G. rigidifolium, G. rivale, G. sacrorum,
G. scabrifolium, G. speciosum, G. tenuissimum
Valantia
?Valantia aprica, V. hispida
GLOBULARIACEAE
Globularia
G. cordifolia, G. meridionalis, G. punctata
PLANTAGINACEAE
Plantago major
Plantago major ssp. intermedia, P. major ssp. major
Plantago afra
Grana 40 (2001)
ACANTHACEAE
Acanthus
?Acanthus balcanicus, A. spinosus
PEDALIACEAE
Sesamum indicum
GESNERIACEAE
?Ramonda
?Ramonda nathaliae, R. serbica
OROBANCHACEAE
Orobanche minor type
Orobanche amethystea, O. crenata, O. gracilis, O. minor,
O. reticulata
?Orobanche alba, O. cernua, O. elatior, O. grisebachii,
O. hederae, O. loricata, O. lutea, O. oxyloba,
O. pubescens, O. rechingeri, Scrophulariaceae:
Rhynchocorys elephas
Orobanche ramosa type
Orobanche ramosa, Scrophulariaceae: Tozzia alpina
LENTIBULARIACEAE
Pinguicula
P. alpina, P. vulgaris
?Pinguicula balcanica, P. hirtiora
Utricularia
U. intermedia, U. minor, U. vulgaris
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Pollen and spore keys, Pindos Mts, Greece
Plantago alpina type
Plantago alpina, P. serraria, P. subulata
Plantago lanceolata type
Plantago altissima, P. lagopus, P. lanceolata
?Plantago cretica, P. gentianoides, P. holosteum,
P. reniformis, P. squarrosa
Plantago coronopus
Plantago bellardii
Plantago media
Plantago arenaria
Plantago argentea
Plantago albicans
Plantago amplexicaulis
Plantago atrata
CAPRIFOLIACEAE
Sambucus nigra
Sambucus ebulus
Lonicera alpigena
Lonicera periclymenum type
Lonicera periclymenum
?Lonicera etrusca
Lonicera xylosteum type
Lonicera nigra, L. xylosteum
Viburnum
V. lantana, V. opulus, V. tinus
VALERIANACEAE
Valerianella rimosa type
Valerianella coronata, V. eriocarpa, V. rimosa
?Valerianella microcarpa
Valeriana tuberosa
Valeriana oYcinalis type
Valeriana oYcinalis
?Valeriana crinii, V. dioscoridis
Centranthus
C. calcitrapae
?Centranthus longiorus
Valerianella locusta type
Valerianella carinata, V. locusta
?Valerianella costata, V. echinata, V. pumila, V. turgida
DIPSACACEAE
Succisa pratensis
Scabiosa columbaria type
Scabiosa atropurpurea, S. columbaria
?Scabiosa taygetea, S. tenuis, S. triniifolia, S. webbiana
Dipsacus type
Dipsacus fullonum, D. laciniatus
?Cephalaria ambrosioides, C. ava, C. transylvanica
Knautia
itK. integrifolia
?Knautia ambigua, K. drymeia, K. longifolia,
K. macedonica, K. magnica, K. orientalis
Scabiosa sicula type
Scabiosa graminifolia, S. sicula
?Scabiosa argentea, S. crenata, S. epirota
Pterocephalus type
Pterocephalus papposus
?Pterocephalus perennis, Tremastelma palaestinum
Morina persica
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CAMPANULACEAE
Jasione
?Jasione heldreichii, J. laevis
Campanul a erinus type
Campanul a erinus, C. glomerata, C. persicifolia,
C. rapunculus, C. trachelium
?Campanula albanica, C. cervicaria, C. foliosa,
C. formanekiana, C. hawkinsiana, C. lingulata,
C. macrostachya, C. patula, C. phrygia, C. ramosissima,
C. scutellata, C. sparsa, C. spatulata, C. trichocalycina,
C. tymphaea, C. versicolor, Edraianthus graminifolius,
E. tenuifolius, Legousia speculum-veneris, Trachelium
jacquinii
Campanul a rotundifolia
Asyneuma type
Asyneuma canescens, Campanul a p.p., Legousia hybrida
?Asyneuma limonifolium
Phyteuma
P. orbiculare
?Phyteuma confusum
COMPOSITAE
SUBFAMILY ASTEROIDEAE (`Tubuliorae type’)
Solidago type
Tribe Heliantheae: Bidens cernua, B. tripartita,
Helianthus annuus
Tribe Astereae: Aster alpinus, A. amellus, A. linosyris,
A. novi-belgii, A. tripolium, Bellis annua, B. perennis,
B. sylvestris, Conyza canadensis, Erigeron acer, E. alpinus,
E. epiroticus, E. glabratus, Solidago virgaurea
Tribe Inuleae: Antennaria dioica, Dittrichia graveolens,
D. viscosa, Filago pyramidata, F. vulgaris, Gnaphalium
luteo-album, Inula britannica, I. conyza, I. germanica,
I. helenium, I. salicina, Pallenis spinosa, Pulicaria
dysenterica, P. vulgaris
Tribe Calendula: Calendula arvensis
Tribe Senecioneae: Petasites hybridus, Senecio aquaticus,
S. doronicum, S. erucifolius, S. uviatilis, S. gallicus,
S. integrifolius, S. jacobaea, S. macedonicus, S. othonnae ,
S. papposus, S. scopolii, S. squalidus, S. thapsoides,
S. vernalis, S. viscosus, S. vulgaris, Tussilago farfara
Tribe Eupatorieae: Eupatorium cannabinu m
Tribe Lactuceae (Subfamily Cichorioideae): Scorzonera
austriaca
?Tribe Senecioneae: Adenostyles alliariae, Doronicum
austriacum, D. clusii, D. columnae, D. orientale
Tribe Inuleae: Evax pygmaea, Filaginella uliginosa,
Helichrysum italicum, H. plicatum, H. stoechas, Inula
ensifolia, I. hirta, I. oculus-christi, I. verbascifolia, Loga
arvensis, L. gallica, L. minima, Omalotheca hoppeana,
O. roeseri, O. supina, O. sylvatica, Phagnalo n graecum
Echinops
Tribe Cardueae: Echinops bannaticus, E. mirocephalus,
E. ritro, E. sphaerocephalus
Artemisia
Tribe Anthemideae: Artemisia abrotanum, A. absinthium,
A. alba, A. campestris, A. eriantha, A. vulgaris
Carlina type
Tribe Cardueae: Arctium lappa, A. minus, A. tomentosum,
Carlina acanthifolia, C. acaulis, C. corymbosa, C. frigida,
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P. I. Chester and J. I. Raine
C. vulgaris, Carthamus dentatus, C. lanatus, Onopordu m
acanthium, O. bracteatum, O. tauricum
?Jurinea glycacantha, J. mollis, Staehelina uniosculosa
??Tribe Cardueae: Cnicus benedictus
Picnomon acarna
Tribe Cardueae: Picnomon acarna
Achillea type
Tribe Anthemideae: Achillea clypeolata, A. coarctata,
A. millefolium, A. nobilis, A. ptarmica, A. setacea,
Anthemis altissima, A. arvensis, A. carpatica, A. cotula,
A. cretica, A. tinctoria, A. triumfetti, Chamomilla recutita,
Coleostephus myconis, Matricaria caucasica, M. perforata,
M. tempskyana, M. trichophylla, Tanacetum corymbosum,
T. vulgare
Tribe Cardueae: Serratula tinctoria
?Tribe Anthemideae: Achillea abrotanoides ,
A. absinthoides, A. ageratifolia, A. chrysocoma,
A. clavennae, A. clusiana, A. collina, A. crithmifolia,
A. depressa, A. distans, A. fraasii, A. grandifolia,
A. holosericea, A. lingulata, A. pannonica, A. pindicola,
Anthemis auriculata, A. coelopoda, A. meteorica,
A. parnassica, A. ruthenica, A. segetalis, A. tenuiloba,
A. tomentosa, Leucanthemum praecox, L. vulgare,
Tanacetum macrophyllum
Tribe Cardueae: Xeranthemum annuum, X. cylindraceum,
X. inapertum
Centaurea cyanus type
Tribe Cardueae: Centaurea cyanus, C. depressa,
C. pinardii
Centaurea napulifera type
Tribe Cardueae: Centaurea napulifera, C. pindicola,
C. triumfetti
Cirsium type
Tribe Cardueae: Carduus acanthoides, C. acicularis,
C. candicans, C. cronius, C. hamulosus, C. kerneri,
C. pycnocephalus , C. thessalus, C. tmoleus, Cirsium
appendiculatum , C. arvense, C. candelabrum, C. creticum,
C. eriophorum, C. erisithales, C. heldreichii, C. ligulare,
C. mairei, C. tymphaeum, C. vulgare
?Tribe Cardueae: Carduus macrocephalus, C. taygeteus,
C. thoermeri
Xanthium
Tribe Heliantheae: Xanthium spinosum, X. strumarium
Centaurea solstitialis type
Tribe Cardueae: Centaurea aYnis, C. alba, C. attica,
C. calcitrapa, C. charrelii, C. cuneifolia, C. diVusa,
C. grisebachii, C. iberica, C. jacea, C. kalambakensis ,
C. lactiora, C. melitensis, C. nicopolitana, C. niederi,
C. orphanidea, C. pallidior, C. pannosa, C. pelia,
C. solstitialis, C. tymphaea, C. uniora, C. zuccariniana,
Cynara scolymus
?Tribe Cardueae: Amphoricarpos neumayeri, Centaurea
deustiformis, C. epirota, C. immanuelis-loewii,
C. macedonica, C. pannonica, C. pawlowski, C. stenolepis,
C. stereophylla, C. triamularia, C. vlachorum,
C. weldeniana, Ptilostemon afer, P. chamaepeuce,
P. gnaphaloides , P. strictus
Centaurea graeca type
Tribe Cardueae: Centaurea graeca, C. psilacantha,
C. rupestris, C. salonitana, C. spruneri
Grana 40 (2001)
Centaurea amplifolia
Tribe Cardueae: Centaurea amplifolia
Crupina
Tribe Cardueae: Crupina crupinastrum, C. vulgaris
SUBFAMILY CICHORIOIDEAE (`Liguliorae type’,
`Taraxacum type’)
Tragopogon
T. dubius, T. porrifolius, T. pratensis
?Tragopogon balcanicus, T. crocifolius, T. hybridus,
T. tommasinii
Scorzonera laciniata type
Scorzonera doria, S. laciniata, S. purpurea
Scorzonera hispanica type
Scorzonera hispanica
?Scorzonera cana, S. mollis, S. parviora
Sonchus type
Reichardia picroides, Sonchus arvensis, S. asper,
S. oleraceus, S. tenerrimus
??Cephalorrhynchus tuberosus
Lactuca type
Lactuca quercina, L. saligna, L. sativa, L. serriola,
L. viminea, L. virosa, Mycelis muralis, Rhagadiolus
stellatus
?Lactuca aurea, L. graeca
Scolymus
S. hispanicus, S. maculatus
Cichorium type
Cichorium intybus, Crepis pulchra
Hieracium type
Hieracium/Pilosella agg., Hieracium amplexicaule,
H. dasycraspedum, H. epirense, H. ferdinandi-coburgi,
H. lactucella, H. naegelianiforme, H. necopinum,
H. pilosella, H. trikalense, Tolpis barbata
?Hieraceum subgenus Pilosella: Hieracium x auriculoides,
H. caespitosum, H. cymosum, H. x densiorum, H. x
fuscatrum, H. x halacsyi, H. hoppeanum , H. x
macrotrichum, H. pavichii, H. piloselloides, H. praealtum,
H. pseudopilosella, H. x ruprechtii, H. x spurium, H. x
zizianum;
Hieraceum subgenus Hieraceum: H. amplexicaule group
(H. petraeum), H. bi dum group (H. caesiiorum,
H. stenolepis), H. bracteolatum group (H. bracteolatum),
H. dolopicum group (H. dolopicum), H. eriobasis group,
H. gaudryi group (H. gaudryi), H. graecum group
(H. graecum), H. gymnocephalu m group
(H. gymnocephalum) , H. heldreichii group,
H. hypochoeroides group, H. latifolium group
(H. brachyphyllum) , H. lazistanum group (H. leithneri),
H. marmoreum group (H. megalothecum), H. murorum
group (H. grandidens, H. integratum, H. semisilvaticum),
H. naegelianum group (H. naegelianum) , H. pannosum
group (H. friwaldii, H. pannosum) , H. parnassi group
(H. parnassi, H. pseudobracteolatum) , H. pilosissimum
group, H. pseudorieni group (H. scardicum),
H. racemosum group (H. barbatum, H. crinitum,
H. italicum, H. racemosum), H. sabaudum group
(H. obliquum, H. sabaudum) , H. sartorianum group,
H. scapigerum group, H. schmidtii group (H. pallidum),
H. sericophyllum group, H. silesiacum group,
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Pollen and spore keys, Pindos Mts, Greece
H. umbrosum group (H. umbrosum), H. waldsteinii group
(H. delpinoi, H. waldsteinii)
Taraxacum type
Crepis capillaris, C. foetida, C. vesicaria, Hypochoeris
glabra, H. maculata, H. radicata, Lapsana communis,
Leontodon autumnalis , L. crispus, L. hispidus,
L. taraxacoides, Picris echioides, P. hieracioides,
Taraxacum apenninum group, Taraxacum bithynicum
group (T. bithynicum, T. megalorrhizon, T. minimum),
Taraxacum erythrospermum group (T. pindicola),
Taraxacum fulvum group, Taraxacum gasparrinii group
(T. dorchocarpum, T. epirense, T. xanthiense),
Taraxacum oYcinale group (T. copidophyllum ,
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T. oYcinale), Taraxacum serotinum group (Taraxacum
serotinum)
?Crepis aurea, C. baldaccii, C. dioscoridis, C. fraasii,
C. geracioides, C. guioliana, C. merxmuelleri, C. neglecta,
C. rubra, C. sancta, C. setosa, C. turcica, C. viscidula,
Leontodon cichoraceus, Picris pauciora, P. sprengerana,
Taraxacum aganippeum , T. bulgaricum, T. gionense,
T. glaciale, T. graecofontanum, Taraxacum hoppeanu m
group (T. amborum, T. hoppeanum, T. poliochlorum),
T. nudum, Taraxacum palustre group (Taraxacum
scaturiginosum), T. subolivaceum
??Calycocorsus stipitatus, Chondrilla juncea,
C. ramosissima, Hedypnois cretica, Prenanthes purpurea
Monocotyledones
Ornithogalum type
ALISMATACEAE
Asphodeline liburnica, A. taurica, Ornithogalum arabicum,
Sagittaria sagittifolia
O. collinum, O. narbonense, O. nutans, O. orthophyllum,
Baldellia ranunculoides
O. pyrenaicum, O. umbellatum, Scilla hyacinthoide s
Alisma
?Ornithogalum armeniacum, O. comosum, O. divergens,
A. gramineum, A. lanceolatum, A. plantago-aquati c
O. exscapum, O. montanum, O. oligophyllum ,
Damasonium alisma
O. refractum, O. sphaerocarpum
Fritillaria
BUTOMACEAE
F. epirotica, F. gussichiae, F. messanensis, F. mutabilis,
Butomus umbellatus
F. orientalis, F. pontica, F. thessala
HYDROCHARITACEAE
Gagea lutea type
Hydrocharis morsus-ranae
Gagea arvensis, G. bohemica, G. lutea, Paris quadrifolia
Stratiotes aloides
?Gagea amblyopetala, G. dubia, G. brosa, G. stulosa,
Vallisneria spiralis
G. minima, G. pusilla, G. reticulata, G. saxatilis
Gagea pratensis
JUNCAGINACEAE
Tulipa sylvestris
Triglochin bulbosaa
Tulipa praecox type
Tulipa gesnerana, T. praecox
POTAMOGETONACEAE
?Tulipa boeotica, T. clusiana
Potamogeton crispus type
Lilium
Groenlandia densa, Potamogeton acutifolius, P. alpinus,
L. albanicum, L. candidum, L. carniolicum,
P. berchtoldii, P. coloratus, P. compressus, P. crispus,
L. chalcedonicum, L. martagon
P. lucens, P. natans, P. nodosus, P. obtusifolius,
Anthericum liliago
P. perfoliatus, P. polygonifolius , P. trichoides
Scilla type
Potamogeton pectinatus
Polygonatu m verticillatum, Scilla autumnalis, S. bifolia,
ZANNICHELLIACEAE
S. hyacinthoides
Zannichellia palustris
?Scilla albanica
Muscari comosum
NAJADACEAE
Muscari neglectum type
Najas marinaa
Muscari neglectum
?Muscari armeniacum, M. botryoides, M. cummutatum,
LILIACEAE
M.
macrocarpum, M. parviorum, M. tenuiorum
Colchicum
??Bellevalia
ciliata, B. dubia, B. romana, B. trifoliata,
C. autumnale, C. graecum, C. neapolitanum , C. triphyllum
Hyacinthus
orientalis,
Hyacinthella leucophaea,
?Colchicum bivonae, C. callicymbium, C. cupanii,
Strangweia
spicata
C. hungaricum, C. lingulatum, C. turcicum
Allium roseum type
Merendera sobolifera
Allium neapolitanum, A. roseum
Narthecium scardicum
Allium sphaerocephalon type
Veratrum
Allium ampeloprasum , A. atroviolaceum, A. guttatum,
V. album
A. sativum, A. scorodoprasum, A. sphaerocephalon,
?Veratrum nigrum
A. vineale
Asphodelus stulosus type
Allium paniculatum type
Asphodelus aestivus, A. stulosus
Asphodelus albus
Allium carinatum, A. cepa, A. chamaemoly,
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P. I. Chester and J. I. Raine
A. moschatum, A. nigrum, A. pallens, A. paniculatum,
A. schoenoprasum, A. subhirsutum, A. ursinum
?Allium amethystinum, A amphipulchellum ,
A. bornmuelleri, A. breviradium, A. chamaespathum ,
A. favosum, A. avum, A. frigidum, A. heldreichii,
A. hirtovaginum, A. meteoricum, A. phthioticum,
A. sipyleum, A. stamineum, A. suaveolens
Polygonatum type
Polygonatum multiorum, P. odoratum, Ruscus aculeatus,
R. hypoglossum
?Polygonatu m latifolium
Convallaria majalis
Asparagus
A. acutifolius, A. oYcinalis
?Asparagus stipularis, A. tenuifolius, A. verticillatus
Urginea maritima
Smilax
S. aspera
?Smilax excelsa
AMARYLLIDACEAE
Narcissus papyraceus type
Narcissus papyraceus, N. poeticus, N. serotinus
Narcissus tazetta
Sternbergia
S. lutea
?Sternbergia colchiciora
Galanthus type
Galanthus nivalis, Leucojum aestivum
?Leucojum valentinum
DIOSCOREACEAE
Tamus communis
IRIDACEAE
Hermodactylus tuberosus
Iris pseudacorus
Iris spuria type
Iris spuria
?Iris reichenbachii, I. sintenisii, I. suaveolens
Iris pumila
Iris germanica
Gladiolus type
Gladiolus communis, G. illyricus, G. italicus, Romulea
bulbocodium , R. linaresii
?Gladiolus imbricatus
Gynandriris sisyrinchium
Crocus
C. biorus, C. chrysanthus, C. avus
?Crocus boryi, C. cvijicii, C. hadriaticus, C. olivieri,
C. pulchellus, C. robertianus, C. sieberi, C. veluchensis
Iris planifolia
JUNCACEAE
Juncaceae
Juncus alpinus, J. anceps, J. articulatus, J. bufonius,
J. compressus, J. conglomeratus, J. eVusus, J. fontanesii,
J. hybridus, J. inexus, J. minutulus, J. sphaerocarpus,
J. striatus, J. subnodulosus , J. tenuis, J. tridus, Luzula
campestris, L. forsteri, L. luzulina, L. multiora, L. pilosa,
L. spicata, L. sylvatica
?Juncus atratus, J. heldreichianus, J. thomasii, Luzula
italica, L. pindica, L. sudetica
Grana 40 (2001)
GRAMINEAE
Festuca type
Bellardiochloa violacea, Desmazeria rigida, Festuca alpina,
F. altissima, F. amethystina, F. arundinacea, F. callieri,
F. circummediterranea, F. cyllenica, F. dalmatica,
F. drymeja, F. glauca, F. graeca, F. grandiaristata,
F. hercegovinica, F. heterophylla, F. hirtovaginata,
F. horvatiana, F. jeanpertii, F. korabensis, F. koritnicensis,
F. loliacea, F. nigrescens, F. ovina, F. paniculata,
F. penzesii, F. peristerea, F. pindica, F. polita,
F. pratensis, F. rechingeri, F. rubra, F. spectabilis,
F. thracica, F. valesiaca, F. valida, F. varia, F. violacea,
F. vizzavonae, Lolium multiorum, L. perenne,
L. remotum, L. rigidum, L. temulentum, Micropyrum
tenellum, Poa alpina, P. angustifolia, P. annua,
P. bulbosa, P. cenisia, P. chaixii, P. glauca,
P. macedonica, P. nemoralis, P. ophiolithica, P. palustris,
P. perconcinna, P. pratensis, P. pumila, P. timoleontis,
P. trivialis, P. versicolor, Puccinellia distans, Sclerochloa
dura, Vulpia bromoides, V. ciliata, V. muralis, V. myuros
?Eleusine indica, Eragrostis cilianensis, E. minor, E. pilosa
??Aristida adscensionis, Cleistogenes serotina, Crypsis
aculeata, C. alopecuroides, C. schoenoides, Danthonia
alpina, D. decumbens, Molinia caerulea, Tragus racemosus
Glyceria type
Bromus alopecuros, B. arvensis, B. benekenii,
B. cappadocicus, B. commutatus, B. diandrus, B. erectus,
B. hordeaceus, B. intermedius, B. japonicus,
B. lanceolatus, B. madritensis, B. pindicus, B. ramosus,
B. rigidus, B. riparius, B. rubens, B. scoparius,
B. secalinus, B. squarrosus, B. sterilis, B. tectorum,
Glyceria maxima, G. spicata
Dactylis type
Apera spica-venti, Beckmannia eruciformis, Brachypodiu m
distachyon, B. phoenicoides, B. pinnatum, B. retusum,
B. sylvaticum, Briza humilis, B. maxima, B. media,
B. minor, Catabrosa aquatica, Cynodon dactylon,
Cynosurus cristatus, C. echinatus, C. elegans, Dactylis
glomerata, Echinaria capitata, Festucopsis sancta,
Lamarckia aurea, Melica altissima, M. ciliata, M. minuta,
M. nutans, M. transsilvanica, Mibora minima, Nardus
stricta, Psilurus incurvus, Sesleria bielzii, S. coerulans,
S. korabensis, S. robusta, S. sadlerana, S. tenerrima,
S. vaginalis
??Achnatherum calamagrostis, Agrostis canina,
A. capillaris, A. castellana, A. gigantea, A. stolonifera,
Aira caryophyllea, A. cupaniana , A. elegantissima,
A. tenorii, Alopecurus aequalis, A. arundinaceus ,
A. creticus, A. myosuroides, A. rendlei, A. setarioides,
Ampelodesmos mauritanica, Andropogo n distachyos,
Anthoxanthum gracile, A. odoratum, A. ovatum,
Arrhenatherum elatius, Arundo donax, Avellinia michelii,
Avenula cincinnata, A. compressa, A. pubescens,
Brachiaria eruciformis, Calamagrostis arundinacea,
C. varia, Chrysopogon gryllus, Corynephorus divaricatus,
Danthoniastrum compactum, Dasypyrum villosum,
Deschampsia cespitosa, D. exuosa, D. media,
Dichanthium ischaemum, Digitaria ischaemum,
D. sanguinalis , Echinochloa crus-galli, Gastridium
ventricosum, Hainardia cylindrica, Helictotrichon
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Pollen and spore keys, Pindos Mts, Greece
convolutum, Hemarthria altissima, Holcus lanatus,
H. mollis, H. setiglumis, Hyparrhenia hirta, Imperata
cylindrica, Koeleria lobata, K. macrantha, K. pyramidata,
K. splendens, Lagurus ovatus, Lophochloa cristata,
L. hispida, Milium eVusum, M. vernale, Molineriella
minuta, Parapholis pycnantha, Parvotrisetum myrianthum,
Paspalum paspalodes, Phacelurus digitatus, Phalaris
arundinacea, P. brachystachys, P. canariensis,
P. coerulescens, P. minor, P. paradoxa, Phleum alpinum,
P. arenarium, P. echinatum, P. graecum, P. montanum,
P. paniculatum, P. phleoides, P. pratense, P. subulatum,
Pholiurus pannonicus, Piptatherum coerulescens,
P. holciformis, P. miliaceum, Polypogo n maritimus,
P. monspeliensis, P. viridis, Saccharum ravennae,
S. strictum, Setaria italica, S. pumila, S. verticillata,
S. viridis, Sorghum halepense, Stipa bromoides,
S. capensis, S. capillata, S. epilosa, S. fontanesii,
S. joannis, S. lessingiana, S. pennata, S. pulcherrima,
S. rechingeri, S. thessala, S. tirsa, Trisetum avescens,
Trisetum ?avescens, T. tenuiforme, Ventenata dubia
Agropyron type
Aegilops cylindrica, A. dichasians, A. geniculata,
A. lorentii, A. neglecta, A. speltoides, A. triuncialis,
A. uniaristata, Agropyron cristatum
?Panicum miliaceum, P. repens, Secale montanum
Triticum type
Triticum aestivum, T. compactum, T. dicoccon, T. durum,
T. turgidum
Secale cereale
Hordeum type
Elymus caninus, E. elongatus, E. hispidus, E. repens,
Hordeum murinum, H. vulgare, Triticum monococcum
?Hordelymus europaeus, Hordeum bulbosum, H. distichon,
H. hystrix, H. marinum, Taeniatherum caput-medusae,
Triticum baeoticum
Avena
A.byzantina, A. fatua, A. sativa
?Avena barbata, A. clauda, A. sterilis, A. strigosa
Phragmites australis
Zea mays
ARACEAE
Arum
A. italicum, A. maculatum
?Arum orientale, A. petteri
Dracunculus type
Dracunculus vulgaris
?Biarum tenuifolium
Arisarum vulgare
LEMNACEAE
Lemnaceae
Lemna gibba, L. minor, L. trisulca, Spirodela polyrhiza
SPARGANIACEAE
Sparganium erectum
Sparganium angustifolium
TYPHACEAE
Typha angustifolia type
Typha angustifolia, T. latifolia
381
Typha domingensis type
Typha domingensis
?Typha laxmannii
CYPERACEAE
Scirpus
S. lacustris, S. maritimus
Cladium mariscus
Cyperus michelianus
Cyperus longus type
Cyperus glomeratus, C. laevigatus, C. longus
?Cyperus diVormis, C. esculentus, C. avescens,
C. avidus, C. glaber
Schoenus nigricans
Carex hallerana type
Carex hallerana, Scirpus holoschoenus, S. sylvaticus
?Blysmus compressus, Carex depressa, C. ferruginea,
C. liparocarpos, C. macrolepis, C. melanostachya,
C. sempervirens, Scirpus cernuus, S. cespitosus,
S. litoralis, S. mucronatus, S. setaceus, S. supinus
Carex acca type
Carex acuta, C. acutiformis, C. appropinquata , C. atrata,
C. caryophyllea, C. chordorrhiza, C. depauperata,
C. digitata, C. distachya, C. distans, C. disticha, C. divisa,
C. divulsa, C. echinata, C. elata, C. elongata,
C. ericetorum, C. acca, C. hirta, C. hostiana, C. humilis,
C. lepidocarpa, C. muricata, C. nigra, C. otrubae,
C. ovalis, C. pallescens, C. panicea, C. paniculata,
C. pendula, C. punctata, C. remota, C. riparia,
C. rostrata, C. serotina, C. spicata, C. sylvatica,
C. tomentosa, C. umbrosa, C. vulpina, Cyperus fuscus,
C. rotundus, Eleocharis multicaulis, E. palustris,
Eriophorum angustifolium, E. gracile, E. latifolium,
E. vaginatum
?Carex davalliana, C. hispida, C. kitaibeliana, Eleocharis
acicularis, E. quinqueora, E. uniglumis, Fimbristylis
bisumbellata
ORCHIDACEAE
Epipactis type
Subfamily Neottioideae: Epipactis atrorubens,
E. helleborine, E. leptochila, E. microphylla, E. palustris,
E. purpurata, Listera ovata, Neottia nidus-avis
?Cypripedium calceolus, Epipogium aphyllum
Limodorum abortivum
Subfamily Neottioideae: Limodorum abortivum
Cephalanthera
Subfamily Neottioideae: Cephalanthera damasonium,
C. rubra
Orchis type
Subfamily Orchidoideae: Aceras anthropophorum ,
Anacamptis pyramidalis, Barlia robertiana, Coeloglossum
viride, Dactylorhiza fuchsii, D. incarnata, D. sambucina,
D. sulphurea, Gymnadenia conopsea, Himantoglossum
hircinum, Neotinea maculata, Ophrys apifera,
O. bombyli ora, O. fusca, O. reinholdii, O. scolopax,
Grana 40 (2001)
382
P. I. Chester and J. I. Raine
O. sphegodes, O. tenthredinifera, Orchis coriophora,
O. italica, O. laxiora, O. mascula, O. pallens,
O. papilionacea, O. provincialis, O. purpurea,
O. quadripunctata, O. tridentata, O. ustulata, Platanthera
bifolia, P. chlorantha, Serapias cordigera, S. lingua,
S. vomeracea;
Subfamily Neottioideae: Spiranthes aestivalis, S. spiralis
?Subfamily Orchidoideae: Dactylorhiza baumanniana ,
D. cordigera, D. iberica, D. kalopissii, D. saccifera,
Ophrys carmeli, O. ferrum-equinum, O. fuciora,
O. spruneri, Orchis simia, O. spitzelii, Pseudorchis frivaldii
??Corallorhiza trida
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INDEX
Abelmoschus esculentus Malvaceae special key:E1
Abies A:C2; Fig. 1A
Abutilon t. E11.2:B1 and Malvaceae special key: A3
Acanthus E7.5:C3 and E7.6:D1
Acer E7.4:E3; Fig. 8L±M
Achillea t. E11.2:I1; Fig. 12G±H
Actaea spicata E7.3:F2
Adiantum capillus-veneris D:E3
Adonis E7.2:I3
Aesculus hippocastanu m E11.3:U2
Agrimonia t. Rosaceae special key:H2; Fig. 25Q
Agropyron t. Gramineae special key:F1
Agrostemma githago Caryophyllaceae special key:D2; Fig. 21A
Ailanthus altissima E11.3:J2
Alcea t. Malvaceae special key:G2
Alchemilla t. Rosaceae special key:J1; Fig. 25M±P
Alisma E17.3:K2 and E17.3:T1; Fig. 19E
Alkanna t. E11.1:F2
Allium paniculatum t. E4:C1; Fig. 3K
Allium roseum t. E4:D2; Fig. 3M
Allium sphaerocephalo n t. E4:D1; Fig. 3L
Alnus glutinosa E16:I2; Fig. 18M
Althaea hirsuta Malvaceae special key:F2
Althaea oYcinalis Malvaceae special key:C2
Anagallis arvensis t. E11.5:R2 and E11.5:X2
Anagallis tenella t. E11.5:U1 and E11.5:W1
Anchusa arvensis t. E11.1:B1 and E12:P2; Fig. 17D±E
Anchusa oYcinalis t. E12:W2; Fig. 17F±G
Andrachne telephioides E11.5:A¾4
Androsac e maxima E11.3:A¾1
Androsac e villosa E11.1:I1
Anemone nemorosa group E7.2:M1
Anogramma leptophylla D:D2
Anthericum liliago E4:F2
Anthyllis E11.1:L3
Arabis E7.6:K3
Arachis hypogae a E18:H4
Arceuthobium oxycedri E7.7:C1
Arenaria serpyllifolia Caryophyllaceae special key:O2; Fig. 21J
Arisarum vulgare E2:A5
Aristolochia E2:D3
Armeria maritima t. E7.6:C2
Artemisia E11.2:J1; Fig. 12N±O
Arum E2:I2
Asparagus E4:S1
Asphodelus albus E4:V2
Asphodelus stulosus t. E4:K1; Fig. 4D
Asplenium t. C:I2; Fig. 2F
Asterolinon linum-stellatum E11.5:A ¾3
Astragalus E11.4:M2, O1, and Q2
Asyneuma t. E16:A4 and B1
Athyrium distentifolium C:H2
Athyrium lix-femina C:K2
Atropa t. E11.3:O2
Avena Gramineae special key:E1; Fig. 23C
Baldellia ranunculoide s E17.3:T2
Bellardia trixago E7.1:L2
Grana 40 (2001)
Berberis vulgaris t. E18:I1
Betula pendula E15:W2; Fig. 18L
Bilderdykia E11.1:B2
Blackstonia perfoliata E11.3:S2 and E12:S2; Fig. 13P±Q
Blechnum spicant C:K1
Boraginaceae E19:A2
Borago oYcinalis E12:E1; Fig. 17K±M
Botrychium D:F2
Bryonia E11.5:V3
Buglossoides arvensis t. E12:M2
Buglossoides purpurocaerule a E12:X2
Butomus umbellatus E4:N2; Fig. 4B
Buxus sempervirens E17.2:A2 and E17.2:D1
Cachrys E11.1:I2
Callitriche obtusangul a E2:K2
Callitriche stagnalis t. E2:L2 and E5:E4
Caltha t. E7.2:I1; Fig. 6H±I
Calystegia E17.3:N2; Fig. 19I
Campanula erinus t. E15:K2 and E16:A4
Campanula rotundifoli a E15:J2 and E16:A4; Fig. 18C
Cannabi s sativa E15:U1 and E16:F2
Carex hallerana t. Cyperaceae special key:E2
Carex acca t. Cyperaceae special key:E1; Fig. 22C
Carlina t. E11.2:I2; Fig. 12C±D
Carpinus t. E15:W1 and E16:M2
Caryophyllaceae E17.1:E2, E17.2:B2, E17.3:G2, and E17.3:R1;
Fig. 21
Castanea sativa E11.1:A¾2; Fig. 11X±Z
Cedrus A:C1; Fig. 1C
Celtis E15:N1, E16:E2, and E17.3:F2
Centaurea amplifolia E11.2:G1
Centaurea cyanus t. E11.1:C2; Fig. 11A±C
Centaurea graeca t. E11.2:K2
Centaurea napulifera t. E11.1:C1; Fig. 11D
Centaurea solstitialis t. E11.2:M3; Fig. 12I
Centaurium E11.3:P1 and E11.3:R1; Fig. 13N±O
Centranthus E7.2:F2
Cephalanthera E2:N2 and E5:C2; Fig. 3H, 5D
Cerastium fontanum t. Caryophyllaceae special key:U1; Fig. 21F
Ceratonia siliqua E12:T1
Ceratophyllum C:J2 and E2:F1
Cercis siliquastrum E11.5:F ¾3; Fig. 16Q±S
Cerealia t. Gramineae special key:A2
Cerinthe type E12:H1 and E12:M1
Cheilanthes D:C2
Chelidonium majus E7.6:I1
Chenopodiaceae t. E17.3:Q2 and Caryophyllaceae special key:E1;
Fig. 21G
Chrysospleniu m alternifolium E7.6:L1 and E11.5:H¾1
Cicendia liformis E11.3:T2 and E11.3:Y2
Cichorium t. E20:H2; Fig. 20H±I
Circaea lutetiana E15:B1; Fig. 18B
Cirsium t. E11.2:M1; Fig. 12J±K
Cistus monspeliensis t. E11.5:B3
Cistus salvifolius E11.5:K2; Fig. 16A
Citrus E12:K1 and E12:W1
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Pollen and spore keys, Pindos Mts, Greece
Cladium mariscus Cyperaceae special key:D2
Clematis vitalba group E7.2:M1
Colchicum E14:C2 and E15:D1; Fig. 17N
Comandra elegans E7.6:G1
Compositae Asteroideae E11.2 and E13:B1
Consolida t. E7.2:I2
Convallaria majalis E4:Z1; Fig. 4F
Convolvulus E7.1:C2; Fig. 5I
Cornus mas E7.2:C1 and E11.1:V1
Cornus sanguine a E7.2:C2 and E11.1:V2; Fig. 6A
Coronilla emerus E11.4:O2
Coronilla scorpioides t. E11.3:X2
Corrigiola litoralis E7.1:G1, E15:R1, and Caryophyllaceae special
key:B2
Corydalis ochroleuca E9:E2 and E18:J1
Corydalis solida t. E9:E1
Corylus E15:V1; Fig. 18G
Cotinus coggygria E11.3:C1
Cotoneaster t. Rosaceae special key:J2
Crassula E11.3:Y1
Crataegus Rosaceae special key:M3; Fig. 25R±T
Crocus E2:D5 and E18:I3
Crucianella E8:K3
Crupina E11.2:M2
Cucumis melo E15:D2
Cucurbita E17.1:D2
Cuscuta epithymum t. E7.1:E2; Fig. 5J-K
Cuscuta europaea t. E7.1:E1
Cuscuta monogyna E7.6:I2
Cyclamen E11.1:G1; Fig. 11R±T
Cydonia oblonga Rosaceae special key:M1
Cynoglossum t. E19:B2
Cyperaceae E2:C2, E5:E3, E17.3:B2, and special key; Fig. 22
Cyperus longus t. Cyperaceae special key:C1; Fig. 22B
Cyperus michelianus Cyperaceae special key:A1
Cystopteris fragilis C:M2
Cytinus hypocistis B1:B3 and C:B2
Cytinus ruber E14:B2
Dactylis t. Gramineae special key:H2; Fig. 23G±H
Damasonium alisma E17.3:K1
Daphne E2:P3 and E17.2:D2
Datura stramonium E11.3:F2; Fig. 13A±B
Dianthus deltoides t. Caryophyllaceae special key:H2 and J2;
Fig. 21D
Dictamnus albus E11.5:P2
Diospyros lotus E11.1:J ¾1
Dipsacus t. E7.3:E3 and E15:H2; Fig. 7N±O
Dorycnium E11.1:M2; Fig. 11U±W
Dracunculus t. E2:I1
Dryopteris expansa C:F1
Dryopteris lix-mas t. C:F3
Echinophora t. E10
Echinops E11.2:J2; Fig. 12L±M
Echium E11.1:F1 and E11.4:A2; Fig. 11N±O
Elaeagnus angustifoli a E11.1:D¾1
Elatine alsinastru m E11.5:F¾4
Emex spinosa E11.1:M¾1
Ephedra distachya t. E1:A2
Ephedra fragilis E1:A1; Fig. 3A
Epilobium angustifoliu m t. E15:B2 and E16:F3; Fig. 18A
Epilobium hirsutum t. B1:E2; Fig. 2C
Epipactis t. B1:B2; Fig. 2B
Equisetum D:A2 and C3, and E2:G2; Fig. 3D
Erica herbacea B1:H1; Fig. 2D±E
Erodium E7.4:A2; Fig. 8A±B
Euonymus europaeus t. E11.5:L1
Euphorbia E11.4:G2; Fig. 14C±D
Euphrasia E7.1:K1; Fig. 5N±O
383
Fagus E11.1:M ¾2; Fig. 11J¾±L¾
Festuca t. Gramineae special key:I2; Fig. 23K±L
Ficus carica E14:A1
Filipendula E11.1:A ¾3, E11.1:D ¾2, and Rosaceae special key:I3
Fragaria vesca t. Rosaceae special key:F2
Fragaria viridis t. Rosaceae special key:G2
Frangula E11.1:T2
Fraxinus angustifoli a E7.6:T2 and E8:H2
Fraxinus excelsior t. E7.6:T1 and E8:H1
Fraxinus ornus E11.5:B¾2
Fritillaria E4:O1; Fig. 4H
Fumana E11.5:E¾3
Fumaria densiora t. E17.3:S1
Fumaria parviora E17.3:S2
Gagea lutea t. E4:Z2
Gagea pratensis E4:A6
Galanthus t. E4:T1; Fig. 4E
Galega oYcinalis E11.4:R1
Galium t. E8:D2; Fig. 10Q±R
Genista t. E7.5:H2
Gentiana lutea t. E11.3:P2; Fig. 13I±K
Gentiana verna E11.5:A2 and E11.5:K1; Fig. 15E±F
Gentianella ciliata E11.5:L2
Geranium E7.6:C1; Fig. 9A±B
Geum Rosaceae special key:K2
Gladiolus t. E4:Y3
Glaucium E7.6:R1
Glinus t. E7.2:M3
Globularia E11.1:J ¾2
Glyceria t. Gramineae special key:I1; Fig. 23I±J
Gossypium E13:B2 and Malvaceae special key:A2
Gramineae E5:D2 and Gramineae special key; Fig. 23
Gratiola oYcinalis E11.1:G2; Fig. 11P±Q
Gymnocarpium dryopteris C:F2
Gynandriri s sisyrinchium E4:H2
Haplophyllum E11.3:S3
Hedera helix E11.5:F2; Fig. 15O±Q
Hedysarum coronarium E7.5:C1
Helianthemum E11.3:L2; Fig. 13E±F
Heliotropium E19:C2
Helleborus E7.6:O2; Fig. 9O±P
Hermodactylus tuberosus E4:P3
Herniaria t. E17.3:B1 and Caryophyllaceae special key:B1
Hibiscus Malvaceae special key:E3; Fig. 22D
Hieracium t. E20:G2; Fig. 20D±G
Hippocrepis E11.3:X1
Holosteum umbellatum Caryophyllaceae special key:N2; Fig. 21K
Hordeum t. Gramineae special key:F2; Fig. 23D
Hornungia t. E7.6:K2; Fig. 9C±E
Humulus lupulus E14:D2 and E15:U1
Humulus t. E15:U1 and E16:F2; Fig. 18H
Hydrocharis morsus-rana e E2:B3
Hydrocotyle vulgaris E11.1:O2
Hyoscyamus niger E11.5:G¾2; Fig. 16Y±A¾
Hypecoum E6:B2 and E18:F2
Hypericum E7.5:H1
Hypericum hircinum t. E11.5:G¾1
Hypericum perforatum t. E11.5:F¾1; Fig. 16N±P
Ilex aquifolium E7.7:B1 and E11.6; Fig. 10A±B
Impatiens noli-tangere C:B1 and E8:F3
Ipomoea E17.1:C2; Fig. 19A
Iris germanica E4:G2
Iris planifolia E18:I2
Iris pseudacorus E4:P2
Iris pumila E4:A5
Iris spuria t. E4:P1
Isoetes C:L2
Grana 40 (2001)
384
P. I. Chester and J. I. Raine
Jasione E15:K1
Jasminum E11.5:J2; Fig. 15M±N
Jovibarba heuVelii E11.1:K¾2
Juglans regia E17.3:E2; Fig. 19J
Juncaceae B1:D2 and E2:F1
Juniperus E2:J2, E5:E1 and E18:E2; Fig. 3E±F
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Kickxia E11.5:E¾1; Fig. 16V±X
Knautia E15:F1
Lactuca t. E20:F2; Fig. 20A±C
Lagenaria siceraria E11.1:V3 and E11.4:G1
Lappula E19:D1
Lathyrus pratensis t. E11.4:W2; Fig. 14L
Lathyrus sativus E11.4:T2
Laurus nobilis E2:D4
Lemnaceae E2:B3 and E5:E1
Lens E11.4:X1
Leontice leontopetalum E7.6:D2
Ligustrum vulgare E11.5:G2; Fig. 15A±B
Lilium E4:G1; Fig. 4A
Limodorum abortivum E2:L3 and E5:C1
Linum spp. E7.7 and E11.6
Linum austriacum t. E7.7:F2
Linum catharticum t. E7.7:F1; Fig. 10G±H
Linum hirsutum E7.7:F1
Linum hologynum t. E7.7:F3
Linum strictum group E7.7:F1
Linum tenuifolium group E7.7:F1
Linum trigynum E7.7:F1
Linum usitatissimum t. E7.7:D2; Fig. 10E±F
Lithospermum E12:X1
Lonicera E11.2:B2 and E12:P1
Lonicera alpigena E11.2:D2
Lonicera periclymenum t. E11.2:D1; Fig. 12A±B
Lonicera xylosteum t. E11.2:C2
Loranthu s europaeus E18:H2
Lotus t. E11.1:L1; Fig. 11D¾±E¾
Ludwigia palustris E11.1:C¾2 and E11.3:E3
Lychnis coronaria Caryophyllaceae special key:O1; Fig. 21B
Lychnis os-cuculi Caryophyllaceae special key:I2
Lychnis viscaria Caryophyllaceae special key:G2
Lysimachia E11.5:S2 and U2; Fig. 16B±E
Lythrum portula E19:F2
Lythrum salicaria t. E19:F1; Fig. 18R±T
Mahonia aquifolium E18:J2
Malabaila E11.1:I2
Malus Rosaceae special key:R1; Fig. 26I±L
Malva moschata Malvaceae special key:E2
Malva pusilla Malvaceae special key:G1; Fig. 22E±F
Malva sylvestris t. Malvaceae special key:F1; Fig. 22G±H
Malvaceae E17.1:D1 and Malvaceae special key; Fig. 22
Marrubium E7.5:F2; Fig. 8G±H
Medicago sativa E11.1:A¾1 and E11.4:M3; Fig. 14H
Melampyrum E7.1:B3
Mentha t. E8:K1; Fig. 10I±J
Mercurialis annua t. E11.5:C1
Mercurialis perennis E11.5:C2
Merendera sobolifera E14:C1
Mespilus germanica Rosaceae special key:J3
Minuartia setacea t. Caryophyllaceae special key:K1and L2
Moehringia trinervia t. Caryophyllaceae special key:N1
Moneses uni ora B1:F2
Monotrop a hypopity s E7.1:A3
Montia fontana E9:C2 and E18:J3; Fig. 10S±T
Morina persica E15:T2
Morus alba E14:E1 and E15:O2; Fig. 17O, 18D
Morus nigra E15:O1, E15:U2, and E16:M1
Grana 40 (2001)
Muscari comosum E4:I2
Muscari neglectum t. E4:I1
Myosotis t. E19:D2; Fig. 18U±V
Myricaria germanica E7.1:K2, E7.1:P1, E7.2:B1, and E7.2:L2
Myriophyllu m spicatum E16:K2
Myriophyllu m verticillatum E16:K1
Myrtus communis E18:H1; Fig. 18P±Q
Najas marina C:J3, and E2:A3
Narcissus papyraceus t. E4:L1
Narcissus tazetta E4:V3
Narthecium scardicum E4:M3 and E4:T2
Neatostema apulum E12:L1
Nicotiana E11.3:O1
Nigella E7.2:E3 and E7.3:F1; Fig. 6B±C
Nonea E12:U2
Nuphar lutea E4:B2
Nymphaea alba E5:B2
Nymphoides peltata E18:G2
Odontites t. E7.1:L3; Fig. 5P±Q
Onobrychis E7.5:C2
Ononis t. E11.4:R2; Fig. 14K
Ophioglossu m lusitanicum D:H2
Ophioglossu m vulgatum D:H1
Orchis t. B2:A2
Ornithogalum t. E4:X1
Ornithopus compressus E11.1:M1
Orobanche minor t. E2:D2; Fig. 3B
Orobanche ramosa t. E7.2:K1
Osmunda regalis D:C1
Ostrya t. E15:W1; Fig. 18K
Osyris alba E11.1:Z2
Oxalis E7.4:D2, E7.6:P1, and E8:G2; Fig. 8E±F
Oxyria digyna E11.1:E¾2, E11.5:B¾1, and E13:C1
Papaver argemone t. E17.3:J1
Papaver dubium E7.2:J2 and E8:C1
Papaver hybridum E7.2:J2 and E8:C1
Papaver rhoeas t. E7.2:J2 and E8:C1; Fig. 6D±E
Papaver somniferum E7.2:J2 and E8:C1
Parietaria E15:R2 and E16:H2; Fig. 18I±J
Parnassia palustris E11.5:V1; Fig. 16L±M
Paronychia t. E17.3:V1 and Caryophyllaceae special key:R2
Pedicularis E6:B1 and E18:F1; Fig. 5E±F
Peganum harmala E11.5:C¾2
Periploca graeca B1:B1
Phaseolus E11.4:D2
Phillyrea E11.5:W2 and E12:T2; Fig. 16T±U
Phlomis E7.5:E1
Phragmites australis Gramineae special key:G2; Fig. 23E±F
Phyteuma E15:J1, E16:A4 and B1
Phytolacca americana E7.1:B2
Picea abies A:E2; Fig. 1B
Picnomon acarna E11.2:G2
Pinguicula E12:J2
Pinus (Haploxylon) A:E1; Fig. 1D
Pinus (Diploxylon) A:D1; Fig. 1E±F
Pistacia E17.3:U2; Fig. 19H
Pisum sativum E11.4:V2
Plantaginaceae E17.1:A2, E17.3:H2 and Plantaginaceae special
key; Fig. 24
Plantago afra Plantaginaceae special key:I3; Fig. 24O±Q
Plantago albicans Plantaginaceae special key:E2; Fig. 24E±F
Plantago alpina t. Plantaginaceae special key:C1
Plantago amplexicaulis Plantaginaceae special key:D1; Fig. 24G±H
Plantago arenaria Plantaginaceae special key:H2; Fig. 24I±J
Plantago argentea Plantaginaceae special key:E1
Plantago atrata Plantaginaceae special key:G2
Plantago bellardii Plantaginaceae special key:H1
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Pollen and spore keys, Pindos Mts, Greece
Plantago coronopus Plantaginaceae special key:C2; Fig. 24C±D
Plantago lanceolata t. Plantaginaceae special key:B2; Fig. 24A±B
Plantago major Plantaginaceae special key:I1; Fig. 24K±L
Plantago media Plantaginaceae special key:I2; Fig. 24M±N
Platanus orientalis E7.6:S2; Fig. 9K±L
Plumbago europaea E7.3:A3
Polygala monspeliaca t. E12:G2
Polygala vulgaris t. E12:G1
Polygonatum t. E4:V1; Fig. 4G
Polygonu m amphibium E9:B2
Polygonu m aviculare t. E11.1:B3 and E13:C2; Fig. 11H±J
Polygonu m bistorta t. E11.1:K1 and E11.4:H1
Polygonu m persicaria t. E17.2:C2; Fig. 19C
Polygonu m viviparum E11.1:K2 and E11.4:H2
Polypodium C:J4
Polystichum C:I1
Populus E2:D1; Fig. 3C
Portulaca oleracea E9:C1
Potamogeton crispus t. E2:O1; Fig. 3J
Potamogeton pectinatus E2:O2
Potentilla Rosaceae special key:F1 and G1; Fig. 25F±I
Primula E8:G3
Prunella t. E8:J2; Fig. 10L
Prunus avium t. Rosaceae special key:Q1; Fig. 26D±E
Prunus spinosa t. Rosaceae special key:R2; Fig. 26F±H
Pseudotsuga menziesii E2:G1
Psoralea t. E11.4:D1
Pteridium aquilinum D:E2; Fig. 2G
Pteris cretica D:G2
Pteris vittata D:G1
Pterocephalus t. E15:G1
Pulmonaria mollis E12:V2
Pulsatilla halleri E7.2:M1
Punica granatu m E11.4:L1
Putoria calabrica E7.1:D2
Pyrola B1:G2
Pyrus Rosaceae special key:M2
Quercus frainetto t. E7.2:N1; Fig. 7A±C
Quercus coccifera t. E7.2:N3 and E7.4:D1; Fig. 7D±F
Quercus trojana t. E7.2:N2; Fig. 6M±N
Radiola linoides E7.7:E2
Ranunculus E7.2:M1
Ranunculus acris t. E7.2:M1; Fig. 6F±G
Ranunculus arvensis t. E17.1:F2
Reseda E7.6:G2
Rhamnus t. E11.4:I2; Fig. 14A±B
Rhinanthus t. E7.1:L1
Rhus coriaria E11.1:R2 and E11.3:D1
Ribes E17.3:M2
Ricinus communis E11.1:Z1 and E11.5:A¾2
Robinia pseudoacacia E11.1:L¾2
Rosa Rosaceae special key:H1 and O2; Fig. 25D±E
Rosaceae E7.1:M2, E7.4:B2, E11.3:A2 and H2, and Rosaceae
special key; Figs. 25 & 26
Rubus t. Rosaceae special key:N2; Fig. 25J±L
Rumex t. E11.1:F¾2, E11.5:B¾1, and E13:C1; Fig. 11H¾±I¾
Ruta chalepensis E11.3:T1
Ruta montana t. E11.5:A¾1
Sagittaria sagittifolia E17.1:F1; Fig. 19D
Salix fragilis t. E7.6:Q1 and E11.5:N1; Fig. 9M±N
Salix triandra E7.6:Q2 and E11.5:N2; Fig. 9I±J
Salvia oYcinalis t. E8:L2; Fig. 10M±N
Salvia verticillata E8:L1; Fig. 10O±P
Salvinia natans D:E1
Sambucus ebulus E11.5:H2; Fig. 15J±L
Sambucus nigra E11.5:P1; Fig. 16F±H
Samolus valerandi E11.1:G¾1
385
Sanguisorba minor E11.1:B¾2 and Rosaceae special key:C2;
Fig. 25A±C
Sanguisorba oYcinalis E12:J1 and Rosaceae special key:C1
Saxifraga bulbifera E7.1:P2
Saxifraga oppositifoli a t. E7.4:F1 and E11.3:J1; Fig. 8J±K
Saxifraga sempervivum t. E7.4:G1
Saxifraga stellaris E7.1:N2, E7.6:L2, and E11.5:H¾2; Fig. 5L±M
Saxifraga tridactylites t. E7.4:G2 and E11.3:I1
Scabiosa columbaria t. E7.3:E2; Fig. 7J±K
Scabiosa sicula t. E15:H1
Schinus molle E11.3:D2; Fig. 13C±D
Schoenus nigricans Cyperaceae special key:C2
Scilla t. E4:L2; Fig. 3N
Scirpus Cyperaceae special key:B1 and E2; Fig. 22A
Scleranthus annuus t. Caryophyllaceae special key:T2; Fig. 21I
Scleranthus perennis Caryophyllaceae special key:T1
Scolymus E20:A2
Scorzonera hispanica t. E20:D1
Scorzonera laciniata t. E20:D2
Scrophularia t. E11.5:F ¾2 and V2; Fig. 16I±K
Scutellaria t. E7.5:D1; Fig. 8P
Secale cereale Gramineae special key:C2
Sedum E11.3:A ¾2; Fig. 13L±M
Selaginella D:F1
Sempervivum E11.3:Z2
Sesamum indicum E8:B3
Sherardia arvensis E8:C2; Fig. 10K
Sideritis E8:F4
Silene alba t. Caryophyllaceae special key:G3; Fig. 21C
Silene vulgaris t. Caryophyllaceae special key:I1 and K2; Fig. 21E
Sinapis t. E7.6:R2; Fig. 9F±H
Smilax E5:B1 and E18:B1; Fig. 5A
Solanum dulcamara E11.1:S1; Fig. 11F¾±G¾
Solanum melongena E11.4:N1
Solanum nigrum t. E11.1:T1; Fig. 11A¾
Soldanella E7.2:D3 and E18:H3
Solidago t. E11.2:E2; Fig. 12E±F
Sonchus t. E20:E1; Fig. 20L±M
Sorbus t. Rosaceae special key:Q2; Fig. 26A±C
Sparganium angustifolium E5:F3
Sparganium erectum E5:F4; Fig. 5B
Spartium junceum E11.1:G ¾2
Spergula E7.1:I2 and E9:D2
Spergularia t. E7.1:I1 and E9:D1
Sperguleae Caryophyllaceae special key
Stachys angustifoli a E8:J1
Stachys arvensis E7.5:G2; Fig. 8O
Stachys recta E7.5:G1; Fig. 8N
Stachys sylvatica t. E7.5:E3; Fig. 8I
Stellaria holostea Caryophyllaceae special key:Q1; Fig. 21H
Sternbergia E4:V4
Stratiotes aloides E4:B1
Styrax oYcinalis E11.1:R1
Succisa pratensis E7.3:E1
Symphytum E12:H2
Tamarix E7.6:K1
Tamus communis E6:A2
Taraxacum t. E20:H1; Fig. 20J±K
Taxus baccata E2:J1; Fig. 3G
Teucrium E7.2:H2; Fig. 6L
Thalictrum E17.3:I1 and V2
Thalictrum aquilegifoliu m E17.3:L1; Fig. 19F±G
Thalictrum lucidum E17.3:L3
Thalictrum minus E17.3:L2
Theligonum cynocrambe E16:A3
Thelypteris palustris C:M1
Thelypteris phegopteris C:G2
Thesium alpinum E18:C1
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386
P. I. Chester and J. I. Raine
Thesium divaricatum t. E18:C2
Tilia cordata t. E11.4:C1; Fig. 14I
Tilia platyphyllos E11.4:C2
Tordylium E10 and E11.1:I2
Tragopogon E20:C2
Trapa natans A:A2 and E3
Tribulus terrestris E17.2:C3; Fig. 19B
Trifolium dubium t. E11.4:Y1; Fig. 14G
Trifolium montanum t. E11.1:B¾1 and E11.4:N2; Fig. 14M±N
Trifolium repens t. E11.4:Y2; Fig. 14J
Triglochin bulbosa E2:P2; Fig. 3I
Triticum t. Gramineae special key:E2; Fig. 23B
Trollius europaeus E7.4:F2
Tuberaria guttata E11.3:R2
Tulipa praecox t. E7.6:A3
Tulipa sylvestris E4:X2
Typha angustifoli a t. B1:C2 and E5:F1; Fig. 2A, 5C
Typha domingensis t. B1:C1 and E5:F2
Ulmus glabra t. E16:C2; Fig. 18N±O
Ulmus minor t. E16:C1
Ulmus procera E16:C3
Umbelliferae E11.1:I2; Fig. 11E±G, K±M
Umbilicus E11.1:K¾1
Urginea maritima E4:Y2
Urtica E16:G2
Urtica dioica E14:E2 and E15:Q2; Fig. 18E
Urtica pilulifera t. E16:H1 and E17.3:F1
Urtica urens t. E15:Q1; Fig. 18F
Utricularia E12:F1; Fig. 17A±C
Vaccaria pyramidata Caryophyllaceae special key:U2
Vaccinium myrtillus B1:H2
Valantia E8:D1
Valeriana oYcinalis t. E7.3:D2; Fig. 7G±I
Valeriana tuberosa E7.3:D1
Valerianella locusta t. E7.2:G1; Fig. 6J±K
Valerianella rimosa t. E7.2:G2
Vallisneria spiralis E2:B2
Veratrum E4:N1; Fig. 4C
Verbena E11.1:T2; Fig. 11B¾±C¾
Veronica anagallis-aquatic a t. E7.4:C3; Fig. 8C±D
Veronica triphyllos t. E7.1:H2 and E7.2:K2; Fig. 5G±H
Viburnum E11.5:H1; Fig. 15C±D
Vicia cracca t. E11.4:L2; Fig. 14E±F
Vicia faba E11.4:V1
Vicia lathyroide s E15:S3 and E16:L2
Vicia sativa t. E11.4:X2; Fig. 14O
Vinca herbacea E19:E2
Vincetoxicum B2:A1
Viola arvensis t. E12:K2 and E12:Q2; Fig. 17H±J
Viola hirta t. E7.1:O2
Viola riviniana t. E12:Q1
Viscum album E7.7:C2; Fig. 10C±D
Vitex agnus-castu s E11.3:F1; Fig. 13G±H
Vitis vinifera E11.5:B1; Fig. 15G±I
Xanthium E7.3:F3 and E11.2:L2; Fig. 7L±M
Zannichellia palustris E2:P1
Zea mays Gramineae special key:B2; Fig. 23A
ACKNOWLEDGMENTS
We would like to thank the following people for their assistance: Dr
Oliver Rackham, Corpus Christi College, Cambridge University,
U.K., for identifying plants collected in GrevenaÂ; Drs Nancy Wilkie
and Mary Savina, Carleton College, Minnesota, U.S.A., for assistance with eld facilities; Peter Tijsen, Wellington Botanic Gardens,
Grana 40 (2001)
for permitting collection of pollen from plants in the gardens; Drs
Patrick Brownsey, Natural History Unit, Museum of New Zealand,
and Barry Sneddon, Victoria University of Wellington, for giving
permission to take pollen from their herbarium collections; Dr
Dallas Mildenhall, Institute of Geological and Nuclear Sciences,
New Zealand, for permitting the use of laboratory facilities and
providing materials for making reference slides.
The research was done while P. Chester held a Postgraduate
Scholarship from Victoria University of Wellington, New Zealand,
and a Postgraduate Fellowship from the New Zealand Federation
of University Women, and formed part of the work towards a
doctoral dissertation. Subsistence in the eld was partly nanced by
the Grevena Archaeological Project (director Nancy Wilkie), and
the National Science Foundation of the United States of America.
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