This art icle was downloaded by: [ I ngent a Cont ent Dist ribut ion ( Publishing Technology) ] On: 13 Oct ober 2014, At : 19: 06 Publisher: Taylor & Francis I nform a Lt d Regist ered in England and Wales Regist ered Num ber: 1072954 Regist ered office: Mort im er House, 37- 41 Mort im er St reet , London W1T 3JH, UK Grana Publicat ion det ails, including inst ruct ions f or aut hors and subscript ion inf ormat ion: ht t p: / / www. t andf online. com/ loi/ sgra20 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 To link to this article: ht t p: / / dx. doi. org/ 10. 1080/ 00173130152987535 PLEASE SCROLL DOWN FOR ARTI CLE Taylor & Francis m akes every effort t o ensure t he accuracy of all t he inform at ion ( t he “ Cont ent ” ) cont ained in t he publicat ions on our plat form . However, Taylor & Francis, our agent s, and our licensors m ake no represent at ions or warrant ies what soever as t o t he accuracy, com plet eness, or suit abilit y for any purpose of t he Cont ent . Any opinions and views expressed in t his publicat ion are t he opinions and views of t he aut hors, and are not t he views of or endorsed by Taylor & Francis. The accuracy of t he Cont ent should not be relied upon and should be independent ly verified wit h prim ary sources of inform at ion. 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Pollen and spore keys for Quaternary deposits in the northern Pindos Mountains, Greece. ± Grana 40: 299±387. ISSN 0017-3134. . Keys for identiŽcation 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 4 302 302 302 302 304 304 305 306 306 306 306 309 313 313 313 313 314 317 317 319 321 323 323 325 327 327 327 331 332 333 337 342 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 identiŽcation 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, SŽkas 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 identiŽed 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 identiŽed 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 identiŽed speciŽcally, but pollen of Isatis tinctoria (woad ) and Camelina sativa (gold-ofpleasure), and most other crucifers and legumes cannot be speciŽcally 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. Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Pollen and spore keys, Pindos Mts, Greece Thus Grevena can be divided into Žve 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 inuenced 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 Grevena landscape is intensively used and little, if any, unmodiŽed 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 Grevena 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 Žr 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 deŽnition 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 deŽned 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) 302 P. I. Chester and J. I. Raine Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 signiŽes 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 Žgure numbers is provided in the body of the key. Note that some Žgures 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 Grana 40 (2001) 303 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 uniora 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 Žlix-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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žlix-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. UnidentiŽable 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-Žmbriate, 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 signiŽcantly 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 Grana 40 (2001) 306 P. I. Chester and J. I. Raine 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žxed 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 conŽned 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 Žxed 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 Grana 40 (2001) 307 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Grana 40 (2001) 308 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 conŽned 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 3. (For legend see page 310). Pollen and spore keys, Pindos Mts, Greece 309 Grana 40 (2001) 310 P. I. Chester and J. I. Raine Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žstulosus 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). Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žstulosus type (e.g. A. Žstulosus); E, Galanthus type (e.g. G. nivalis); F, Convallaria majalis; G, Polygonatum type (e.g. P. odoratum ); H, Fritillaria (e.g. F. lusitanica). Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. Grana 40 (2001) 314 P. I. Chester and J. I. Raine Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 superŽcially 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. Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece 315 C2 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žne. 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). Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 316 Grana 40 (2001) P. I. Chester and J. I. Raine Fig. 5. (For legend see page 315). Pollen and spore keys, Pindos Mts, Greece M2 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žne. 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). Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 6. 318 (For legend see page 317). P. I. Chester and J. I. Raine Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece C3 C2 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 C1 319 Very Žnely 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 Žne, 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 Žne ........................................................ 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 Želd 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. Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 7. 320 (For legend see page 319). P. I. Chester and J. I. Raine Grana 40 (2001) 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 (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 Grana 40 (2001) P. I. Chester and J. I. Raine Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 322 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). Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece 323 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žne, 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, Žne, 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 Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 324 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 Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 10. 326 (For legend see page 325). P. I. Chester and J. I. Raine Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece 327 D2 Psilate-scabrate with uniform minute spinules and perforations ............................................................... Spergula CARYOPHYLLACEAE D1 Psilate, spinules absent ....................................................................................................................... Spergularia t. CARYOPHYLLACEAE Key E10. Dicolporate Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 deŽne 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 Žner. 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 Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 328 P. I. Chester and J. I. Raine 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 Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece 329 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 11. 330 (For legend see page 329). P. I. Chester and J. I. Raine Grana 40 (2001) 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. Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 conuent 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). Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 12. 332 (For legend see page 331). P. I. Chester and J. I. Raine Grana 40 (2001) 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 A2 A1 Operculate. See Special key ............................................................................................................................... Rosaceae No operculum. B2 Endoaperture a transversal colpus. Costae transversales present. Anacardiaceae. C2 Colpus long, transverse colpus well deŽned. 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 deŽned, 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 Žne. 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 Žne. 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. Grana 40 (2001) 334 P. I. Chester and J. I. Raine 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 Žliformis 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 Žliformis 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Pollen and spore keys, Pindos Mts, Greece 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. Grana 40 (2001) 336 P. I. Chester and J. I. Raine Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 G2 E1 Grana 40 (2001) 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 Žne 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. Pollen and spore keys, Pindos Mts, Greece 337 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žner and more irregular. Reticulum in polar area not diVerent, but Žner 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). Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 14. 338 (For legend see page 337). P. I. Chester and J. I. Raine Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Pollen and spore keys, Pindos Mts, Greece E1 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, Žne. 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. Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 15. 340 (For legend see page 339). P. I. Chester and J. I. Raine Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Pollen and spore keys, Pindos Mts, Greece 341 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 Grana 40 (2001) 342 P. I. Chester and J. I. Raine 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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, superŽcially scabrate except in polar area which has an intectate Želd 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. Grana 40 (2001) 343 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Pollen and spore keys, Pindos Mts, Greece 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. Grana 40 (2001) 344 P. I. Chester and J. I. Raine Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Q2 Grain depressed oval (barrel-shaped) to polyhedric. Polar area with an intectate Želd 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. Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece 345 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Pollen grains of Ficus Žt 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 magniŽcation; 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 Grana 40 (2001) P. I. Chester and J. I. Raine Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece 347 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Grana 40 (2001) 348 P. I. Chester and J. I. Raine C1 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece 349 C1 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žne. 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 Žne 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). Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 18. 350 (For legend see page 349). P. I. Chester and J. I. Raine Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece 351 F1 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 deŽned, regular . T. lucidum RANUNCULACEAE L2 Grain >17 mm. Columellae distinct. Porus margins poorly deŽned, 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 Žne and short. Fumaria. S2 Usually 12 pores .............................................................. Fumaria parviora PAPAVERACEAE S1 Usually 6 pores ............................................................. Fumaria densiora 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. Grana 40 (2001) 352 P. I. Chester and J. I. Raine B2 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 A1 Tricolporate with large pores, the three colpi joining at one pole only. C2 Coarsely reticulate ....................................................................................................... Thesium divaricatum t. SANTALACEAE C1 Psilate to Žnely 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 Želd. 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 Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece 353 D1 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 19. 354 (For legend see page 353). P. I. Chester and J. I. Raine Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žne, 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 Žne 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 Žne 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). Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 20. 356 (For legend see page 355). P. I. Chester and J. I. Raine Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece 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. Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žnger ................................................................................................. 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) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 21. 358 (For legend see page 357). P. I. Chester and J. I. Raine Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 360 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Fig. 25. 362 (For legend see page 361). P. I. Chester and J. I. Raine Grana 40 (2001) Pollen and spore keys, Pindos Mts, Greece Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 deŽned, 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 Žne. Tectum perforate. H2 Vallae transversal, extremely Žne. 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 Žne. L2 Vallae paired. M3 Striation weak. Size 29±42 mm. Fig. 25R±T ..................................................................... Crataegus M2 Striation deŽnite. 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 Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 364 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 identiŽed 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 identiŽed to genus; a species name signiŽes 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žlix-femina Athyrium distentifolium Cystopteris fragilis ASPIDIACEAE Polystichum P. aculeatum, P. lonchitis, P. setiferum ?Polystichum x illyricum Dryopteris Žlix-mas type Dryopteris Žlix-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. Žssum, 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 Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 366 P. I. Chester and J. I. Raine 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. pedunculiora, 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. thyrsiorus ?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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 ?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. Žcifolium, 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. deexus, A. hybridus, A. lividus, A. retroexus 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 biora, A. conferta, A. cretica, A. Žlicaulis, 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. minutiorus, D. myrtinervius, D. pallens, D. petraeus, D. pinifolius, D. pontederae, 367 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. densiora, 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. viridiora, 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) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 368 P. I. Chester and J. I. Raine RANUNCULACEAE Helleborus H. odorus ?Helleborus cyclophyllus Nigella N. arvensis, N. damascena, N. sativa Trollius europaeus Actaea spicata Consolida type Consolida orientalis, C. regalis, Delphinium Žssum, 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 lateriorus 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 Žcaria 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 densiora type Fumaria densiora, F. kralikii, F. oYcinalis, F. rostellata ?Fumaria petteri, F. schleicheri Fumaria parviora 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. doereri, 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 grandiora, 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, Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. subava, 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. reexum, 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 369 PARNASSIACEAE Parnassia palustris GROSSULARIACEAE Ribes R. alpinum, R. multiorum 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. thyrsiorus, 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) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 370 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 parviora 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. parviorus, Grana 40 (2001) L. stenodon, L. tenuis, L. uliginosus, Tetragonolobus biorus, 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. laxiorus, 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. grandiora, 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. pubiorus, 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. reexum 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. nodiorum, 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. deexa, 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) 372 P. I. Chester and J. I. Raine BUXACEAE Buxus sempervirens Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. parviorus Cistus salvifolius Fumana F. aciphylla, F. arabica, F. bonapartei TAMARICACEAE Myricaria germanica Tamarix T. dalmatica, T. hampeana , T. parviora, T. smyrnensis, T. tetrandra ELATINACEAE Elatine alsinastrum CUCURBITACEAE Cucumis melo ?Citrullus lanatus Bryonia B. alba, B. cretica Lagenaria siceraria Cucurbita C. Žcifolia, 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 Epilobium angustifolium type Epilobium angustifolium, Oenothera biennis Epilobium hirsutum type Epilobium hirsutum, E. montanum, E. palustre, E. parviorum, 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. multiorum, 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. Žstulosa 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 uniora 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 Žliformis Centaurium C. erythraea, C. pulchellum, C. tenuiorum 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. planiora ?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. sparsiora, M. suaveolens, Rochelia disperma Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 tenuiora 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 biŽdum, 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. grandiora, 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 grandiora, 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) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. densiorum, V. nigrum, V. phlomoides , V. pulverulentum, V. sinuatum, V. thapsus ?Digitalis ferruginea, D. grandiora, D. laevigata, D. lanata, D. viridiora, 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. doereri, M. Žmbriatum, 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. doereri, 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. hirtiora Utricularia U. intermedia, U. minor, U. vulgaris Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 longiorus 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. magniŽca, 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 377 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 (`Tubuliorae 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, LogŽa 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, Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 378 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 uniosculosa ??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. lactiora, C. melitensis, C. nicopolitana, C. niederi, C. orphanidea, C. pallidior, C. pannosa, C. pelia, C. solstitialis, C. tymphaea, C. uniora, 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 (`Liguliorae 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. parviora 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 densiorum, 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 Ždum group (H. caesiiorum, 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, Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 , 379 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 pauciora, 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. Žbrosa, G. Žstulosa, 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. parviorum, M. tenuiorum 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 Žstulosus type Allium paniculatum type Asphodelus aestivus, A. Žstulosus Asphodelus albus Allium carinatum, A. cepa, A. chamaemoly, Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 380 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 multiorum, 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 colchiciora 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. biorus, 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. inexus, J. minutulus, J. sphaerocarpus, J. striatus, J. subnodulosus , J. tenuis, J. triŽdus, Luzula campestris, L. forsteri, L. luzulina, L. multiora, 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 multiorum, 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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. quinqueora, 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. laxiora, 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. fuciora, O. spruneri, Orchis simia, O. spitzelii, Pseudorchis frivaldii ??Corallorhiza triŽda Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žstulosus 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 Žlix-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 Žliformis 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Žlix-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 densiora t. E17.3:S1 Fumaria parviora 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Grana 40 (2001) Downloaded by [Ingenta Content Distribution (Publishing Technology)] at 19:07 13 October 2014 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 Želd 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 Želd was partly Žnanced by the Grevena Archaeological Project (director Nancy Wilkie), and the National Science Foundation of the United States of America. REFERENCES Andersen, S. T. 1979. IdentiŽcation of wild grass and cereal pollen. Ê rbog, 1978: 69±92. ± Danmarks Geologiske Undersùgelse A Andersen, S. T. & Bertelsen, F. 1972. Scanning electronmicroscope studies of pollen of cereals and other grasses. ± Grana 12: 79±86. Ayala-Nieto, M. L., Lira Saade, R. & Alvarado, J. L. 1988. Morfologia polinica de las Cucurbitaceae de la Peninsula de Yucatan, Mexico. ± Pollen et Spores 30(1): 5±28. Belmonte, J., PeÂrez-Obiol, R. & Roure, J. M. 1986. 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