Nuclear DNA Amounts in Angiosperms: Progress, Problems and Prospects

M D BENNETT; I J LEITCH

doi:10.1093/aob/mci003

. 2005 Jan;95(1):45–90. doi: 10.1093/aob/mci003

Nuclear DNA Amounts in Angiosperms: Progress, Problems and Prospects

M D BENNETT ^1,^*, I J LEITCH ¹

PMCID: PMC4246708 PMID: 15596457

Abstract

CONTENTS

INTRODUCTION	45
PROGRESS	46
Improved systematic representation (species and families)	46
(i) First estimates for species	46
(ii) First estimates for families	47
PROBLEMS	48
Geographical representation and distribution	48
Plant life form	48
Obsolescence time bomb	49
Errors and inexactitudes	49
Genome size, ‘complete’ genome sequencing, and, the euchromatic genome	50
The completely sequenced genome	50
Weeding out erroneous data	52
What is the smallest reliable C-value for an angiosperm?	52
What is the minimum C-value for a free-living angiosperm and other free-living organisms?	53
PROSPECTS FOR THE NEXT TEN YEARS	54
Holistic genomics	55
LITERATURE CITED	56
APPENDIX	59
Notes to the Appendix	59
Original references for DNA values	89

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• Background The nuclear DNA amount in an unreplicated haploid chromosome complement (1C-value) is a key diversity character with many uses. Angiosperm C-values have been listed for reference purposes since 1976, and pooled in an electronic database since 1997 (http://www.kew.org/cval/homepage). Such lists are cited frequently and provide data for many comparative studies. The last compilation was published in 2000, so a further supplementary list is timely to monitor progress against targets set at the first plant genome size workshop in 1997 and to facilitate new goal setting.

• Scope The present work lists DNA C-values for 804 species including first values for 628 species from 88 original sources, not included in any previous compilation, plus additional values for 176 species included in a previous compilation.

• Conclusions 1998–2002 saw striking progress in our knowledge of angiosperm C-values. At least 1700 first values for species were measured (the most in any five-year period) and familial representation rose from 30 % to 50 %. The loss of many densitometers used to measure DNA C-values proved less serious than feared, owing to the development of relatively inexpensive flow cytometers and computer-based image analysis systems. New uses of the term genome (e.g. in ‘complete’ genome sequencing) can cause confusion. The Arabidopsis Genome Initiative C-value for Arabidopsis thaliana (125 Mb) was a gross underestimate, and an exact C-value based on genome sequencing alone is unlikely to be obtained soon for any angiosperm. Lack of this expected benchmark poses a quandary as to what to use as the basal calibration standard for angiosperms. The next decade offers exciting prospects for angiosperm genome size research. The database (http://www.kew.org/cval/homepage) should become sufficiently representative of the global flora to answer most questions without needing new estimations. DNA amount variation will remain a key interest as an integrated strand of holistic genomics.

Keywords: Angiosperm DNA amounts, DNA C-values, nuclear genome size, plant DNA C-values database

INTRODUCTION

It has been possible to estimate the amount of DNA in plant nuclei for over 50 years, and since the key role of DNA in biology was discovered in 1953, such research has increased in each successive decade. Work on plants has played a leading part in research to describe and understand the origin, extent and effects of variation in the DNA amount in the unreplicated haploid nuclear chromosome complement (defined by Swift, 1950, as the 1C-value) of different taxa. Indeed, angiosperms are probably the most intensively studied major taxonomic ‘group’ of organisms, with published C-values for over 4100 species.

Early research to address questions such as possible relationships between DNA C-value and the rate of cell development (e.g. Van't Hof, 1965) usually required work to estimate C-values for most of the taxa concerned, as these were unavailable. Later, as taxa with ‘known’ C-values increased, it was possible to use such data in new comparisons (supplemented by further first estimates made for sample taxa). However, it was often difficult to know whether a C-value existed for a particular taxon, and if so, where to find it. Such estimates were widely scattered in the literature or even unpublished. Small lists of nuclear DNA amounts were published in reviews and research papers, but the first large list of DNA amounts for angiosperms, compiled primarily as a reference source was published in 1976. This contained data for over 750 species from 54 original sources (Bennett and Smith, 1976), and noted an intention to publish supplementary lists for reference purposes at intervals. Five such lists, together giving pooled data for over 2900 species from 323 original sources, have followed (Bennett et al., 1982, 2000; Bennett and Smith, 1991; Bennett and Leitch, 1995, 1997). Data from the first five publications were pooled in an electronic form – the Angiosperm DNA C-values database, which went live in April 1997. This was updated as release 3.1 and incorporated, with databases for gymnosperms, pteridophytes and bryophytes, into the Plant DNA C-values database (release 1.0) in 2001.

These data are clearly much used, as the published lists have been cited over 1400 times, including over 700 times since 1997, whilst the electronic database has received over 50000 hits. Recently they have provided the large samples of data needed for many diverse comparative studies, such as testing for possible relationships between nuclear DNA amount and risk of extinction (Vinogradov, 2003), ecological factors in California (Knight and Ackerly, 2002), lead pollution in Slovenia (B. Vilhar, University of Ljubljana, Slovenia, pers. comm.); ploidy level (Leitch and Bennett, 2004), and land plant evolution (Leitch et al., 2005).

Given their ongoing use as reference sources, publication of a sixth supplementary list of angiosperm C-values is timely, if not overdue. The present work lists DNA C-values for 804 species from 88 original sources, including first estimates for 628 species not included in any previous compilation, plus additional estimates for 176 species already included in one or more previous compilation. Data in the Appendix table were prepared for analysis at the second Plant Genome Size Discussion Meeting in September 2003, so it is fitting that they are included in this special supplement. Whilst they represent most of the new C-value data published or estimated in 2000–2002, we are already aware of a further large sample estimated but unpublished either by late 2002, or subsequently. Thus, despite its large size, the present list will soon be followed by a seventh supplement.

Appendix.

Chromosome number, ploidy level, life-cycle type, and nuclear DNA content in 804 angiosperm species (the superscript letters refer to notes preceding this table)

								DNA amount
Entry number^g	Species	Voucher	Family	Higher group^#	2n^‡	Ploidylevel (x)	Life cycletype^§	1C (Mbp^s)	1C (pg)	2C (pg)	4C (pg)	Original ref.^a	Present amount^†	Standard species^*^b1	Method^††
1a	Acacia dealbata Link.	No	Leguminosae	E	26	2	P	853	0·9	1·7	3·5	441^bb	O	J	FC:PI
2	Acacia dealbata Link.	No	Leguminosae	E	39°	3	P	—^t	—^t	2·5	5·1	441^bb	O	J	FC:PI
3	Acacia dealbata Link.	No	Leguminosae	E	52	4	P	1,671	1·7	3·4	6·8	441^bb	O	J	FC:PI
4a	Acacia mangium Willd.	No	Leguminosae	E	26	2	P	637	0·7	1·3	2·6	441^bb	O	J	FC:PI
5	Acridocarpus natalitius A. Juss.	No	Malpighiaceae	E	c. 216	24	P	1,490	1·5	3·0	6·1	379	O	J	Fe
6	Adenanthera microsperma Teijsm & Binn.	No	Leguminosae	E	—ⁿ	—^p	P	681	0·7	1·4	2·8	454	O	B^c	Fe
7	Adenanthera pavonina L.	No	Leguminosae	E	26°	—^p	P	666	0·7	1·4	2·7	454	O	B^c	Fe
8	Adina cordifolia (Roxb.) Hook. f.	No	Rubiaceae	E	22°	2	P	816	0·8	1·7	3·3	454	O	B^c	Fe
9	Aeonium haworthii Webb & Berth.	No	Crassulaceae	E	72°	4 or 8	P	760	0·8	1·6	3·1	378	O	J	Fe
10a	Aesculus hippocastanum L.	No	Sapindaceae	E	40	2	P	588	0·6	1·2	2·4	465	O	Lycopers.^c	FC:PI
11b	Agave americana L.	—^m	Asparagaceae	M	120	4	P	7,791	8·0	15·9	31·8	384^aa	O	Homo^f	FC:PI
12d	Agave sisalana Perr.	—^m	Asparagaceae	M	150	5	P	—^t	—^t	20·0	40·0	384^aa	O	Homo^f	FC:PI
13	Agave stricta Salm.	—^m	Asparagaceae	M	60	2	P	3,822	3·9	7·8	15·6	384^aa	O	Homo^f	FC:PI
14	Agrostis palustris Huds.	No	Gramineae	M	28	4	P	2,769	2·8	5·7	11·3	417	O	Gallus^f	FC:PI
15	Ailanthus grandis Prain	No	Simaroubaceae	E	64°	—^p	P	2,134	2·2	4·4	8·7	454	O	B^c	Fe
16a	Albuca pendula B.Mathew	No	Asparagaceae	M	16	2	P	2,967	3·0	6·1	12·1	465	O	G	Fe
16b	Albuca pendula B.Mathew	No	Asparagaceae	M	14	2	P	3,033	3·1	6·2	12·4	465	O	G	Fe
17k	Allium cepa L.	No	Alliaceae^k	M	16°	2	P	16,415	16·8	33·5	67·0	457^bm	O	B^d	Fe
18	Allocasuarina verticillata (Lam.) L.Johnson	No	Casuarinaceae	E	20-28°	2	P	931	1·0	1·9	3·8	452	O	Petunia^e	FC:PI
19	Alocasia cucullata (Lour) Schott	No	Araceae	M	98	—^p	AP	8,200	8·4	16·7	33·5	411	O	B^c	Fe
20	Alocasia hilobeauty Host.	No	Araceae	M	32	—^p	A	3,680	3·8	7·5	15·0	411	O	B^c	Fe
21	Aloe albiflora Guillaumin	No	Xanthorrhoeaceae^k	M	14°	2	P	15,337	15·7	31·3	62·6	440	O	Agave sp.^ab	FC:PI
22	Aloe alooides (Bolus) Druten	No	Xanthorrhoeaceae^k	M	14°	2	P	13,083	13·4	26·7	53·4	440	O	Agave sp.^ab	FC:PI
23	Aloe antandroi (Decary) H.Perrier	No	Xanthorrhoeaceae^k	M	14°	2	P	17,199	17·6	35·1	70·2	440	O	Agave sp.^ab	FC:PI
24	Aloe arborescens (yellow flowers) Mill.ⁱ	No	Xanthorrhoeaceae^k	M	14°	2	P	13,671	14·0	27·9	55·8	440	O	Agave sp.^ab	FC:PI
25a	Aloe aristata Haw.	No	Xanthorrhoeaceae^k	M	14°	2	P	15,729	16·1	32·1	64·2	440	O	Agave sp.^ab	FC:PI
25b	Aloe aristata var. parvifolia Baker Haw.	No	Xanthorrhoeaceae^k	M	14°	2	P	16,023	16·4	32·7	65·4	440	O	Agave sp.^ab	FC:PI
26	Aloe bakeri Scott-Elliot	No	Xanthorrhoeaceae^k	M	14°	2	P	15,925	16·3	32·5	65·0	440	O	Agave sp.^ab	FC:PI
27	Aloe barberae Dyer	No	Xanthorrhoeaceae^k	M	14°	2	P	15,043	15·4	30·7	61·4	440	O	Agave sp.^ab	FC:PI
28	Aloe bellatula Reynolds	No	Xanthorrhoeaceae^k	M	14°	2	P	16,268	16·6	33·2	66·4	440	O	Agave sp.^ab	FC:PI
29	Aloe boiteaui Guillaumin	No	Xanthorrhoeaceae^k	M	14°	2	P	16,023	16·4	32·7	65·4	440	O	Agave sp.^ab	FC:PI
30	Aloe bowiea Schult. & Schult.f.	No	Xanthorrhoeaceae^k	M	14°	2	P	16,268	16·6	33·2	66·4	440	O	Agave sp.^ab	FC:PI
31	Aloe brevifolia Mill.	No	Xanthorrhoeaceae^k	M	14°	2	P	14,553	14·9	29·7	59·4	440	O	Agave sp.^ab	FC:PI
32b	Aloe cameronii Hemsl.	No	Xanthorrhoeaceae^k	M	14°	2	P	17,052	17·4	34·8	69·6	440	O	Agave sp.^ab	FC:PI
33	Aloe capitata Baker	No	Xanthorrhoeaceae^k	M	14°	2	P	15,386	15·7	31·4	62·8	440	O	Agave sp.^ab	FC:PI
34	Aloe chabaudii Schonland	No	Xanthorrhoeaceae^k	M	14°	2	P	17,934	18·3	36·6	73·2	440	O	Agave sp.^ab	FC:PI
35	Aloe ciliaris var. tidmarshii Schonland Haw.	No	Xanthorrhoeaceae^k	M	14°	2	P	10,535	10·8	21·5	43·0	440	O	Agave sp.^ab	FC:PI
36	Aloe ciliaris Haw.	No	Xanthorrhoeaceae^k	M	35°	5	P	—^t	—^t	53·3	106·6	440	O	Agave sp.^ab	FC:PI
37	Aloe ciliaris Haw.	No	Xanthorrhoeaceae^k	M	42°	6	P	30,723	31·4	62·7	125·4	440	O	Agave sp.^ab	FC:PI
38	Aloe comptonii Reynolds	No	Xanthorrhoeaceae^k	M	14°	2	P	13,426	13·7	27·4	54·8	440	O	Agave sp.^ab	FC:PI
39a	Aloe cryptopoda Baker	No	Xanthorrhoeaceae^k	M	14°	2	P	14,161	14·5	28·9	57·8	440	O	Agave sp.^ab	FC:PI
39b	Aloe cryptopoda “Wickensii” Baker	No	Xanthorrhoeaceae^k	M	14°	2	P	14,357	14·7	29·3	58·6	440	O	Agave sp.^ab	FC:PI
40	Aloe dawei Berger	No	Xanthorrhoeaceae^k	M	28°	4	P	35,231	36·0	71·9	143·8	440	O	Agave sp.^ab	FC:PI
41a	Aloe descoingsii Reynolds	No	Xanthorrhoeaceae^k	M	14°	2	P	15,974	16·3	32·6	65·2	440	O	Agave sp.^ab	FC:PI
41b	Aloe descoingsii Reynolds ssp. augustina Lavranos	No	Xanthorrhoeaceae^k	M	14°	2	P	16,219	16·6	33·1	66·2	440	O	Agave sp.^ab	FC:PI
42a	Aloe dichotoma Masson var. ramosissima (Pillans) Glen & D.S.Hardy	No	Xanthorrhoeaceae^k	M	14°	2	P	12,005	12·3	24·5	49·0	440	O	Agave sp.^ab	FC:PI
42b	Aloe dichotoma Masson	No	Xanthorrhoeaceae^k	M	14°	2	P	12,103	12·4	24·7	49·4	440	O	Agave sp.^ab	FC:PI
43	Aloe dinteri A.Berger	No	Xanthorrhoeaceae^k	M	14°	2	P	16,366	16·7	33·4	66·8	440	O	Agave sp.^ab	FC:PI
44b	Aloe distans Haw.	No	Xanthorrhoeaceae^k	M	14°	2	P	13,622	13·9	27·8	55·6	440	O	Agave sp.^ab	FC:PI
45	Aloe dorotheae A.Berger	No	Xanthorrhoeaceae^k	M	14°	2	P	15,288	15·6	31·2	62·4	440	O	Agave sp.^ab	FC:PI
46	Aloe elegans Tod.	No	Xanthorrhoeaceae^k	M	14°	2	P	17,346	17·7	35·4	70·8	440	O	Agave sp.^ab	FC:PI
47	Aloe erinacea D.S.Hardy	No	Xanthorrhoeaceae^k	M	14°	2	P	12,103	12·4	24·7	49·4	440	O	Agave sp.^ab	FC:PI
48	Aloe ferox Mill.	No	Xanthorrhoeaceae^k	M	14°	2	P	14,896	15·2	30·4	60·8	440	O	Agave sp.^ab	FC:PI
49	Aloe fleurentiniorum Lavranos & L.E.Newton	No	Xanthorrhoeaceae^k	M	14°	2	P	18,179	18·6	37·1	74·2	440	O	Agave sp.^ab	FC:PI
50	Aloe gariepensis (?) Pillans	No	Xanthorrhoeaceae^k	M	14°	2	P	15,729	16·1	32·1	64·2	440	O	Agave sp.^ab	FC:PI
51	Aloe glauca Mill.	No	Xanthorrhoeaceae^k	M	14°	2	P	15,680	16·0	32·0	64·0	440	O	Agave sp.^ab	FC:PI
52	Aloe globuligemma Pole-Evans	No	Xanthorrhoeaceae^k	M	14°	2	P	16,611	17·0	33·9	67·8	440	O	Agave sp.^ab	FC:PI
53	Aloe haemanthifolia A.Berger & Marloth	No	Xanthorrhoeaceae^k	M	14°	2	P	7,938	8·1	16·2	32·4	440	O	Agave sp.^ab	FC:PI

								DNA amount
Entry number^g	Species	Voucher	Family	Higher group^#	2n^‡	Ploidylevel (x)	Life cycletype^§	1C (Mbp^s)	1C (pg)	2C (pg)	4C (pg)	Original ref.^a	Present amount^†	Standard species^*^b1	Method^††
54	Aloe haworthioides Baker	No	Xanthorrhoeaceae^k	M	14°	2	P	14,749	15·1	30·1	60·2	440	O	Agave sp.^ab	FC:PI
55	Aloe hereroensis Engl.	No	Xanthorrhoeaceae^k	M	14°	2	P	18,130	18·5	37·0	74·0	440	O	Agave sp.^ab	FC:PI
56	Aloe humilis (small form) (L.) Mill.ⁱ	No	Xanthorrhoeaceae^k	M	14°	2	P	16,562	16·9	33·8	67·6	440	O	Agave sp.^ab	FC:PI
57	Aloe jacksonii Reynolds	No	Xanthorrhoeaceae^k	M	28°	4	P	32,487	33·2	66·3	132·6	440	O	Agave sp.^ab	FC:PI
58	Aloe jucunda Reynolds	No	Xanthorrhoeaceae^k	M	14°	2	P	17,591	18·0	35·9	71·8	440	O	Agave sp.^ab	FC:PI
59b	Aloe juvenna Brandham & Carter	No	Xanthorrhoeaceae^k	M	28°	4	P	34,790	35·5	71·0	142·0	440	O	Agave sp.^ab	FC:PI
60	Aloe krapohliana var. dumoulinii Lavranos	No	Xanthorrhoeaceae^k	M	14°	2	P	17,346	17·7	35·4	70·8	440	O	Agave sp.^ab	FC:PI
61	Aloe linearifolia A.Berger	No	Xanthorrhoeaceae^k	M	14°	2	P	12,936	13·2	26·4	52·8	440	O	Agave sp.^ab	FC:PI
62	Aloe lomatophylloides Balf.f.	No	Xanthorrhoeaceae^k	M	14°	2	P	17,248	17·6	35·2	70·4	440	O	Agave sp.^ab	FC:PI
63	Aloe longistyla Baker	No	Xanthorrhoeaceae^k	M	14°	2	P	15,582	15·9	31·8	63·6	440	O	Agave sp.^ab	FC:PI
64	Aloe macrosiphon Bak.	No	Xanthorrhoeaceae^k	M	14°	2	P	17,934	18·3	36·6	73·2	440	O	Agave sp.^ab	FC:PI
65	Aloe maculata Allionii	No	Xanthorrhoeaceae^k	M	14°	2	P	18,620	19·0	38·0	76·0	440	O	Agave sp.^ab	FC:PI
66a	Aloe marlothii A.Berger “Spectabilis”	No	Xanthorrhoeaceae^k	M	14°	2	P	15,435	15·8	31·5	63·0	440	O	Agave sp.^ab	FC:PI
66b	Aloe marlothii A.Berger var. bicolor Reynolds	No	Xanthorrhoeaceae^k	M	14°	2	P	15,631	16·0	31·9	63·8	440	O	Agave sp.^ab	FC:PI
67b	Aloe mcloughlinii Christian	No	Xanthorrhoeaceae^k	M	14°	2	P	16,219	16·6	33·1	66·2	440	O	Agave sp.^ab	FC:PI
68	Aloe melanacantha A.Berger	No	Xanthorrhoeaceae^k	M	14°	2	P	12,299	12·6	25·1	50·2	440	O	Agave sp.^ab	FC:PI
69	Aloe microstigma Salm-Dyck	No	Xanthorrhoeaceae^k	M	14°	2	P	15,092	15·4	30·8	61·6	440	O	Agave sp.^ab	FC:PI
70	Aloe mitriformis Mill.	No	Xanthorrhoeaceae^k	M	14°	2	P	13,475	13·8	27·5	55·0	440	O	Agave sp.^ab	FC:PI
71b	Aloe ngobitensis Reynolds	No	Xanthorrhoeaceae^k	M	28°	4	P	28,420	29·0	58·0	116·0	440	O	Agave sp.^ab	FC:PI
72	Aloe occidentalis (H.Perrier) L.E.Newton & G.D.Rowley	No	Xanthorrhoeaceae^k	M	14°	2	P	20,286	20·7	41·4	82·8	440	O	Agave sp.^ab	FC:PI
73	Aloe parvula A.Berger	No	Xanthorrhoeaceae^k	M	14°	2	P	16,562	16·9	33·8	67·6	440	O	Agave sp.^ab	FC:PI
74	Aloe pearsonii Schonland	No	Xanthorrhoeaceae^k	M	14°	2	P	12,348	12·6	25·2	50·4	440	O	Agave sp.^ab	FC:PI
75b	Aloe peckii Bally & Verdoorn	No	Xanthorrhoeaceae^k	M	14°	2	P	17,444	17·8	35·6	71·2	440	O	Agave sp.^ab	FC:PI
76	Aloe peglerae Schonland	No	Xanthorrhoeaceae^k	M	14°	2	P	15,729	16·1	32·1	64·2	440	O	Agave sp.^ab	FC:PI
77	Aloe petricola Pole-Evans	No	Xanthorrhoeaceae^k	M	14°	2	P	15,092	15·4	30·8	61·6	440	O	Agave sp.^ab	FC:PI
78	Aloe pillansii L.Guthrie	No	Xanthorrhoeaceae^k	M	14°	2	P	12,593	12·9	25·7	51·4	440	O	Agave sp.^ab	FC:PI
79	Aloe plicatilis (L.) Mill.	No	Xanthorrhoeaceae^k	M	14°	2	P	8,624	8·8	17·6	35·2	440	O	Agave sp.^ab	FC:PI
80	Aloe pluridens Haworth	No	Xanthorrhoeaceae^k	M	14°	2	P	14,161	14·5	28·9	57·8	440	O	Agave sp.^ab	FC:PI
81	Aloe polyphylla Schonland	No	Xanthorrhoeaceae^k	M	14°	2	P	13,377	13·7	27·3	54·6	440	O	Agave sp.^ab	FC:PI
82	Aloe prinslooi Verdoorn & Hardy	No	Xanthorrhoeaceae^k	M	14°	2	P	17,444	17·8	35·6	71·2	440	O	Agave sp.^ab	FC:PI
83	Aloe prostrata (H.Perrier) L.E.Newton & G.D.Rowley	No	Xanthorrhoeaceae^k	M	14°	2	P	20,139	20·6	41·1	82·2	440	O	Agave sp.^ab	FC:PI
84	Aloe rauhii Reynolds	No	Xanthorrhoeaceae^k	M	14°	2	P	15,337	15·7	31·3	62·6	440	O	Agave sp.^ab	FC:PI
85	Aloe richardsiae Reynolds	No	Xanthorrhoeaceae^k	M	14°	2	P	21,756	22·2	44·4	88·8	440	O	Agave sp.^ab	FC:PI
86	Aloe rivierei Lavranos & L.E. Newton	No	Xanthorrhoeaceae^k	M	14°	2	P	16,562	16·9	33·8	67·6	440	O	Agave sp.^ab	FC:PI
87	Aloe secundiflora Engl.	No	Xanthorrhoeaceae^k	M	14°	2	P	17,591	18·0	35·9	71·8	440	O	Agave sp.^ab	FC:PI
88	Aloe sinkatana Reynolds (red flowers)ⁱ	No	Xanthorrhoeaceae^k	M	14°	2	P	17,542	17·9	35·8	71·6	440	O	Agave sp.^ab	FC:PI
89	Aloe sladeniana Pole-Evans	No	Xanthorrhoeaceae^k	M	14°	2	P	15,974	16·3	32·6	65·2	440	O	Agave sp.^ab	FC:PI
90	Aloe speciosa Baker	No	Xanthorrhoeaceae^k	M	14°	2	P	14,112	14·4	28·8	57·6	440	O	Agave sp.^ab	FC:PI
91	Aloe spicata L.f.	No	Xanthorrhoeaceae^k	M	14°	2	P	14,259	14·6	29·1	58·2	440	O	Agave sp.^ab	FC:PI
92	Aloe striata Haw.	No	Xanthorrhoeaceae^k	M	14°	2	P	18,914	19·3	38·6	77·2	440	O	Agave sp.^ab	FC:PI
93	Aloe suprafoliata Pole-Evans	No	Xanthorrhoeaceae^k	M	14°	2	P	14,014	14·3	28·6	57·2	440	O	Agave sp.^ab	FC:PI
94	Aloe suzannae Decary	No	Xanthorrhoeaceae^k	M	14°	2	P	16,317	16·7	33·3	66·6	440	O	Agave sp.^ab	FC:PI
95b	Aloe tenuior Haw.	No	Xanthorrhoeaceae^k	M	14°	2	P	10,633	10·9	21·7	43·4	440	O	Agave sp.^ab	FC:PI
96	Aloe trichosantha Berger	No	Xanthorrhoeaceae^k	M	14°	2	P	18,179	18·6	37·1	74·2	440	O	Agave sp.^ab	FC:PI
97	Aloe vanbalenii Pillans	No	Xanthorrhoeaceae^k	M	14°	2	P	13,965	14·3	28·5	57·0	440	O	Agave sp.^ab	FC:PI
98a	Aloe variegata L. “Ausana”	No	Xanthorrhoeaceae^k	M	14°	2	P	16,268	16·6	33·2	66·4	440	O	Agave sp.^ab	FC:PI
98b	Aloe variegata L.	No	Xanthorrhoeaceae^k	M	14°	2	P	16,562	16·9	33·8	67·6	440	O	Agave sp.^ab	FC:PI
99	Aloe vera (L.) Burm.f.	No	Xanthorrhoeaceae^k	M	14°	2	P	16,072	16·4	32·8	65·6	440	O	Agave sp.^ab	FC:PI
100	Alstonia macrophylla Wall. ex G.Don.	No	Apocynaceae	E	—ⁿ	—^p	P	718	0·7	1·5	2·9	454	O	B^c	Fe
101c	Alstroemeria aurea Graham^h	No	Alstroemeriaceae	M	16	2	P	24,843	25·4	50·7	101·4	436	O	B	FC:PI
101d	Alstroemeria aurea Graham^h	No	Alstroemeriaceae	M	16	2	P	27,097	27·7	55·3	110·6	436	O	B	FC:PI
102	Alstroemeria aurea Graham^h	No	Alstroemeriaceae	M	24	3	P	—^t	—^t	80·9	161·8	436	O	B	FC:PI
103d	Alstroemeria ligtu L. ssp. incarnata L.^h	No	Alstroemeriaceae	M	16	2	P	34,300	35·0	70·0	140·0	436	O	B	FC:PI
103e	Alstroemeria ligtu L. ssp. simsii^h	No	Alstroemeriaceae	M	16	2	P	31,948	32·6	65·2	130·4	436	O	B	FC:PI
103f	Alstroemeria ligtu L. ssp. simsii^h	No	Alstroemeriaceae	M	16	2	P	38,661	39·5	78·9	157·8	436	O	B	FC:PI
103g	Alstroemeria ligtu L. ssp. ligtu^h	No	Alstroemeriaceae	M	16	2	P	34,006	34·7	69·4	138·8	436	O	B	FC:PI
103h	Alstroemeria ligtu L. ssp. ligtu^h	No	Alstroemeriaceae	M	16	2	P	33,369	34·1	68·1	136·2	436	O	B	FC:PI
104c	Alstroemeria magnifica Herb. ssp. magnifica^h	No	Alstroemeriaceae	M	16	2	P	17,885	18·3	36·5	73·0	436	O	B	FC:PI
104d	Alstroemeria magnifica Herb. ssp. magnifica^h	No	Alstroemeriaceae	M	16	2	P	20,531	21·0	41·9	83·8	436	O	B	FC:PI
105	Alstroemeria magnifica Herb. ssp. magnifica^h	No	Alstroemeriaceae	M	24	3	P	—^t	—^t	61·6	123·2	436	O	B	FC:PI
106	Amborella trichopoda Baill.	No	Amborellaceae	BA	26°	—^p	P	870	0·9	1·8	3·6	381	O	K	FC:PI
107	Amoreuxia wrightii A.Gray	No	Cochlospermaceae	E	c.12-14	2	P	164	0·2	0·3	0·7	378	O	J	Fe
108	Anthemis altissima^l	No	Compositae^j	E	—ⁿ	—^p	—^q	7,742	7·9	15·8	31·6	457^bm	O	B^d	Fe
109	Anthemis montana^l	No	Compositae^j	E	—ⁿ	—^p	—^q	8,281	8·5	16·9	33·8	457^bm	O	B^d	Fe
110	Anthurium grande Host.	No	Araceae	M	28	—^p	P	13,252	13·5	27·0	54·1	411	O	B^c	Fe
111	Anthurium tetragonum Schott	No	Araceae	M	30	—^p	P	7,485	7·6	15·3	30·6	411	O	B^c	Fe
112	Aphyllanthes monspeliensis L.	No	Asparagaceae	M	c. 32	—^p	P	635	0·6	1·3	2·6	380	O	J	FC:PI
113	Arabidopsis korshynskyi^l	No	Cruciferae^j	E	—ⁿ	—^p	—^q	245	0·3	0·5	1·0	457^bm	O	B^d	Fe
114a	Arabidopsis thaliana (L.) Heynh. ecotype Columbia	No	Cruciferae	E	10	2	A	157	0·2	0·3	0·6	461	O	Caeno.^f	FC:PI
114g	Arabidopsis thaliana (L.) Heynh. ecotype Columbia	No	Cruciferae	E	10	2	A	164	0·2	0·3	0·7	461	O	Gallus^f	FC:PI
114h	Arabidopsis thaliana (L.) Heynh. ecotype Columbia	No	Cruciferae	E	10	2	A	150	0·2	0·3	0·6	461	O	Dros.^f	FC:PI
114i	Arabidopsis thaliana (L.) Heynh.	—^m	Cruciferae	E	10	2	A	125^be	0·1^be	0·3^be	0·5^be	448^be	O	—	GS
114j	Arabidopsis thaliana (L.) Heynh. ecotype Columbia	No	Cruciferae	E	10	2	A	167	0·2	0·3	0·7	463	O	Gallus^f	FC:DAPI
114k	Arabidopsis thaliana (L.) Heynh. ecotype Columbia	No	Cruciferae	E	10	2	A	162	0·2	0·3	0·7	463	O	Gallus^f	FC:HO
114l	Arabidopsis thaliana (L.) Heynh. ecotype Columbia	No	Cruciferae	E	10	2	A	157	0·2	0·3	0·6	463	O	Gallus^f	FC:MI
114m	Arabidopsis thaliana (L.) Heynh. line Landsberg erecta	No	Cruciferae	E	10	2	A	51	0·05	0·10	0·21	464	O	—	RK
115a	Arachis duranensis Krapov. & W.C.Gregory^h	No	Leguminosae^j	E	20	2	A	1,243	1·3	2·5	5·1	396	O	G^c	FC:PI
115b	Arachis duranensis Krapov. & W.C.Gregory^h	No	Leguminosae^j	E	20	2	A	1,324	1·4	2·7	5·4	396	O	G^c	FC:PI
115c	Arachis duranensis Krapov. & W.C.Gregory^h	No	Leguminosae^j	E	20	2	A	1,333	1·4	2·7	5·4	396	O	G^c	Fe
116a	Arachis hypogaea L.	No	Leguminosae	E	40	4	A	2,813	2·9	5·7	11·5	395^aj	O	G^c	Fe
116b	Arachis hypogaea L.	No	Leguminosae	E	40	4	A	2,898	3·0	5·9	11·8	395^aj	O	G^c	FC:PI
116r	Arachis hypogaea L.	No	Leguminosae^j	E	40°	4	A	1,568	1·6	3·2	6·4	457^bm	O	B^d	Fe
117a	Arachis monticola Krapov. & Rigoni	No	Leguminosae	E	40	4	A	2,891	3·0	5·9	11·8	395^aj	O	G^c	Fe
117b	Arachis monticola Krapov. & Rigoni	No	Leguminosae	E	40	4	A	2,930	3·0	6·0	12·0	395^aj	O	G^c	FC:PI
118	Archidendron monadelphum (Roxb.) I.C. Neilsen	No	Leguminosae	E	—ⁿ	—^p	P	1,470	1·5	3·0	6·0	454	O	B^c	Fe
119c	Artemisia absinthium L.	No	Compositae^j	E	18	2	P	4,175	4·3	8·5	17·0	386	O	G-120d	FC:PI
120d	Artemisia annua L.	No	Compositae^j	E	18	2	A	1,715	1·8	3·5	7·0	386	O	G^c	FC:PI
121	Artemisia barrelieri Besser	No	Compositae^j	E	36	4	P	6,350	6·5	13·0	25·9	386	O	G^c	FC:PI
122	Artemisia caerulescens L. ssp. gallica (Willd.) K.Persson	No	Compositae^j	E	18	2	P	3,263	3·3	6·7	13·3	386	O	G^c	FC:PI
123	Artemisia campestris L.	No	Compositae^j	E	18	2	P	2,876	2·9	5·9	11·7	386	O	G^c	FC:PI
124	Artemisia campestris L.	No	Compositae^j	E	36	4	P	5,390	5·5	11·0	22·0	386	O	G^c	FC:PI
125	Artemisia cana Pursh.	No	Compositae^j	E	72	8	P	12,569	12·8	25·7	51·3	386	O	G^c	FC:PI
126	Artemisia chamaemelifolia Vill.	No	Compositae^j	E	18	2	P	2,960	3·0	6·0	12·1	386	O	G^c	FC:PI
127	Artemisia crithmifolia L.	No	Compositae^j	E	54	6	P	7,644	7·8	15·6	31·2	386	O	G^c	FC:PI
128	Artemisia dracunculus L.	No	Compositae^j	E	90	10	P	11,378	11·6	23·2	46·4	386	O	G^c	FC:PI
129	Artemisia fragrans Willd.	No	Compositae^j	E	18	2	P	2,622	2·7	5·4	10·7	386	O	G^c	FC:PI
130	Artemisia herba-alba Asso ssp. valentina (Lam.) Mascl.	No	Compositae^j	E	18	2	P	3,219	3·3	6·6	13·1	386	O	G^c	FC:PI
131	Artemisia herba-alba Asso ssp. herba-alba	No	Compositae^j	E	36	4	P	6,115	6·2	12·5	25·0	386	O	G^c	FC:PI
132c	Artemisia judaica L.	No	Compositae^j	E	16	2	P	5,645	5·8	11·5	23·0	386	O	G^c	FC:PI
133	Artemisia lucentica O.Bolos, Valles & Vigo in O.Bolos & Vigo	No	Compositae^j	E	16	2	P	3,763	3·8	7·7	15·4	386	O	G-120d	FC:PI
134	Artemisia molinieri Quezel, Barbero & R.Loisel	No	Compositae^j	E	18	2	P	2,920	3·0	6·0	11·9	386	O	G^c	FC:PI
135	Artemisia monosperma Delile	No	Compositae^j	E	36	4	P	5,400	5·5	11·0	22·0	386	O	G^c	FC:PI
136	Artemisia splendens Willd.	No	Compositae^j	E	32	4	P	6,659	6·8	13·6	27·2	386	O	G^c	FC:PI
137	Artemisia thuscula Cav.	No	Compositae^j	E	18	2	P	5,155	5·3	10·5	21·0	386	O	G^c	FC:PI
138	Artemisia tournefortiana Reichenb.	No	Compositae^j	E	18	2	AB	3,278	3·3	6·7	13·4	386	O	G^c	FC:PI
139	Artemisia tridentata Nutt. ssp. spiciformis Kartesz & Gandhi	No	Compositae^j	E	18	2	P	4,008	4·1	8·2	16·4	386	O	G-120d	FC:PI
140	Artemisia umbelliformis Lam. ssp. umbelliformis	No	Compositae^j	E	34	4	P	6,081	6·2	12·4	24·8	386	O	G^c	FC:PI
141b	Artemisia vulgaris L.	No	Compositae^j	E	16	2	P	2,979	3·0	6·1	12·2	386	O	G^c	FC:PI
142	Artemisia vulgaris L.	No	Compositae^j	E	34	4	P	4,773	4·9	9·7	19·5	386	O	G^c	FC:PI
143b	Arum maculatum L.	No	Araceae	M	56°	8	P	10,682	10·9	21·8	43·6	457^bm	O	B^d	Fe
144	Asarum europaeum L.	No	Aristolochiaceae	BA	—ⁿ	—^p	P	4,753	4·9	9·7	19·4	457^bm	O	B^d	Fe
145	Astelia fragrans Colenso	No	Asteliaceae	M	c. 60	8	P	1,240	1·3	2·5	5·1	380	O	K	FC:PI
146	Atalantia ceylanica (Arn.) Oliv.)	No	Rutaceae	E	18°	2	P	515	0·5	1·1	2·1	426	O	Gallus^f	FC:PI
147	Austrobaileya scandens C.T.White	No	Austrobaileyaceae	BA	44°	—^p	P	9,327	9·5	19·0	38·1	381	O	G	FC:PI
148	Averrhoa carambola L.	No	Oxalidaceae	E	—ⁿ	—^p	P	235	0·2	0·5	1·0	454	O	B^c	Fe
149	Azadirachta indica A.Juss	No	Meliaceae	E	28°	—^p	P	385	0·4	0·8	1·6	454	O	B^c	Fe
150	Bauhinia hookeri (F.Muell.) Pedley	No	Leguminosae	E	26°	—^p	P	620	0·6	1·3	2·5	454	O	B^c	Fe
151b	Bauhinia purpurea L.	No	Leguminosae	E	28°	2	P	573	0·6	1·2	2·3	454	O	B^c	Fe
152	Bauhinia tomentosa L.	No	Leguminosae	E	28°	2	P	613	0·6	1·3	2·5	454	O	B^c	Fe
153	Bellevalia rixii P.Wendalbo	No	Asparagaceae	M	8	2	P	9,102	9·3	18·6	37·2	465	O	B	Fe
154	Berberidopsis corallina Hook. f.	No	Berberidopsidaceae	E	c. 42	6	P	252	0·3	0·5	1·0	379	O	J	Fe
155	Berrya cordifolia (Willd.) Burret	No	Malvaceae	E	—ⁿ	—^p	P	549	0·6	1·1	2·2	454	O	B^c	Fe
156a	Bixa orellana L.	No	Bixaceae	E	14	2	P	191	0·2	0·4	0·8	379	O	J	Fe
156b	Bixa orellana L.	No	Bixaceae	E	14°	2	P	203	0·2	0·4	0·8	454	O	B^c	Fe
157	Blandfordia punicea Sweet.	No	Blandfordiaceae	M	68	4	P	7,970	8·1	16·3	32·5	380	O	G	Fe
158	Bombax ceiba L.	No	Malvaceae	E	92°	—^p	P	1,590	1·6	3·2	6·5	454	O	B^c	Fe
159	Boswellia serrata Roxb.	No	Burseraceae	E	22°	—^p	P	684	0·7	1·4	2·8	454	O	B^c	Fe
160	Brachychiton discolor F.Muell	No	Malvaceae	E	40°	—^p	P	1,132	1·2	2·3	4·6	454	O	B^c	Fe
161a	Brachypodium distachyon (L.) P.Beauv.	No	Gramineae	M	10	2	A	355	0·4	0·7	1·5	465	O	K	FC:PI
161b	Brachypodium distachyon (L.) P.Beauv.	—^m	Gramineae	M	10	2	A	294	0·3	0·6	1·2	460^bh	O	J	Fe
162e	Brassica napus L.	No	Cruciferae^j	E	38°	4	AB	1,568	1·6	3·2	6·4	457^bm	O	B^d	Fe
163b	Bromus arvensis L.	No	Gramineae	M	14	2	A	5,699	5·8	11·6	23·3	391^af	O	H-164b	FC:PI
164b	Bromus carinatus Hooker & Arnott cv. Broma	No	Gramineae	M	56	8	P	11,241	11·5	22·9	45·9	391^af	O	H^c	FC:PI
165c	Bromus erectus Hudson	No	Gramineae	M	56	8	P	12,079	12·3	24·7	49·3	391^af	O	H^c	FC:PI
166c	Bromus hordeaceus L.	No	Gramineae	M	28	4	A	11,285	11·5	23·0	46·1	391^af	O	H^c	FC:PI
167b	Bromus inermis Leysser	No	Gramineae	M	56	8	P	12,025	12·3	24·5	49·1	391^af	O	H^c	FC:PI
168	Bromus willdenowii Knuth	No	Gramineae	M	42	6	P	6,365	6·5	13·0	26·0	391^af	O	H^c	FC:PI
169	Buchloe dactyloides (Nutt.) Engelm.	No	Gramineae	M	40	4	P	779	0·8	1·6	3·2	417	O	Gallus^f	FC:PI
170	Buddleja globosa Hope	No	Buddlejaceae	E	38	2	P	840	0·9	1·7	3·4	378	O	J	Fe
171	Bulbine alooides Willd.	No	Xanthorrhoeaceae	M	14	2	P	10,601	10·8	21·6	43·3	465	O	B	Fe
172	Bulbine fallax Poelln.	No	Xanthorrhoeaceae	M	14	2	P	11,201	11·4	22·9	45·7	465	O	B	Fe
173	Bulbine lagopus (Thunb.) N.E.Brown	No	Xanthorrhoeaceae	M	—ⁿ	—^p	P	7,938	8·1	16·2	32·4	465	O	B	Fe
174	Bulbine praemorsa Spreng.	No	Xanthorrhoeaceae	M	14	2	P	12,213	12·5	24·9	49·9	465	O	B	Fe
175a	Bunias erucago L.	No	Cruciferae	E	14	2	A	2,029	2·1	4·1	8·3	393	O	G^c	Fe
175b	Bunias erucago L.	No	Cruciferae	E	14	2	A	2,136	2·2	4·4	8·7	393	O	Glycine^e	FC:PI
176a	Bunias orientalis L.	No	Cruciferae	E	14	2	P	2,538	2·6	5·2	10·4	393	O	G^c	Fe
176b	Bunias orientalis L.	No	Cruciferae	E	14	2	P	2,636	2·7	5·4	10·8	393	O	Glycine^e	FC:PI
177	Buxus papillosa C.K. Schneid.	No	Buxaceae	E	—ⁿ	—^p	P	1,389	1·4	2·8	5·7	454	O	B^c	Fe
178	Buxus sempervirens^l	No	Buxaceae	E	28	2 or 4	P	794	0·8	1·6	3·2	380	O	K	FC:PI
179	Byblis liniflora Salisb.	No	Byblidaceae	E	32	2	A	870	0·9	1·8	3·6	378	O	J	Fe
180	Cajanus albicans (Wight. & Am.) Maesen	No	Leguminosae	E	22	2	A	1,259	1·3	2·6	5·1	443^bc	O	B^c	Fe
181	Cajanus mollis (Benth.) Maesen	No	Leguminosae	E	22	2	B	804	0·8	1·6	3·3	443^bc	O	B^c	Fe
182	Cajanus sericeus (Benth. ex Bak.) Maesen	No	Leguminosae	E	22	2	P	1,414	1·4	2·9	5·8	443^bc	O	B^c	Fe
183	Caladium bicolor Vent. var. red polka	No	Araceae	M	32	—^p	A	5,407	5·5	11·0	22·1	411	O	B^c	Fe
184	Caladium bicolor Vent. var. red polka large	No	Araceae	M	66	—^p	A	9,927	10·1	20·3	40·5	411	O	B^c	Fe
185	Calceolaria acutifolia Witasek	No	Scrophulariaceae	E	—ⁿ	—^p	—^q	1,348	1·4	2·8	5·5	465	O	J	Fe
186	Calceolaria gracilis^l	No	Scrophulariaceae	E	—ⁿ	—^p	—^q	1,335	1·4	2·7	5·5	465	O	J	Fe
187	Calibrachoa calycina (Sendtn.) Wijsman	No	Solanaceae	E	18	2	P	1,504	1·5	3·1	6·1	387^ae	O	Gallus-398p	FC:PI
188	Calibrachoa dusenii (R.E.Fr.) Stehmann & Semir.	No	Solanaceae	E	18	2	P	1,401	1·4	2·9	5·7	387^ae	O	Gallus-398p	FC:PI
189	Calibrachoa eglandulata Stehmann & Semir.	No	Solanaceae	E	18°	2	P	1,411	1·4	2·9	5·8	387^ae	O	Gallus-398p	FC:PI
190	Calibrachoa elegans (Miers) Stehmann & Semir.	No	Solanaceae	E	18	2	P	1,563	1·6	3·2	6·4	387^ae	O	Gallus-398p	FC:PI
191	Calibrachoa ericaefolia (R.E.Fr.) Wijsman	No	Solanaceae	E	18	2	P	1,436	1·5	2·9	5·9	387^ae	O	Gallus-398p	FC:PI
192	Calibrachoa heterophylla (Sendtn.) Wijsman	No	Solanaceae	E	18°	2	P	1,455	1·5	3·0	5·9	387^ae	O	Gallus-398p	FC:PI
193	Calibrachoa linearis (Hook.) Wijsman	No	Solanaceae	E	18	2	P	1,485	1·5	3·0	6·1	387^ae	O	Gallus-398p	FC:PI
194	Calibrachoa linoides (Sendtn.) Wijsman	No	Solanaceae	E	18	2	P	1,397	1·4	2·9	5·7	387^ae	O	Gallus-398p	FC:PI
195	Calibrachoa macrodactylon (L.B.Sm. & Downs) Wijsman	No	Solanaceae	E	18	2	P	1,480	1·5	3·0	6·0	387^ae	O	Gallus-398p	FC:PI
196	Calibrachoa micrantha (R.E.Fr.) Stehmann & Semir.	No	Solanaceae	E	18	2	P	1,411	1·4	2·9	5·8	387^ae	O	Gallus-398p	FC:PI
197	Calibrachoa parviflora (Juss.) Wijsman	No	Solanaceae	E	18	2	A	936	1·0	1·9	3·8	387^ae	O	Gallus-398p	FC:PI
198	Calibrachoa pygmaea (R.E.Fr.) Wijsman	No	Solanaceae	E	18	2	A	764	0·8	1·6	3·1	387^ae	O	Gallus-398p	FC:PI
199	Calibrachoa rupestris (Dusen) Wijsman	No	Solanaceae	E	18°	2	P	1,597	1·6	3·3	6·5	387^ae	O	Gallus-398p	FC:PI
200	Calibrachoa selloviana (Sendtn.) Wijsman	No	Solanaceae	E	18	2	P	1,455	1·5	3·0	5·9	387^ae	O	Gallus-398p	FC:PI
201	Calibrachoa sendtneriana (R.E.Fr.) Stehmann & Semir.	No	Solanaceae	E	18°	2	P	1,450	1·5	3·0	5·9	387^ae	O	Gallus-398p	FC:PI
202	Calibrachoa serrulata (L.B.Sm. & Downs) Stehmann & Semir.	No	Solanaceae	E	18	2	P	1,446	1·5	3·0	5·9	387^ae	O	Gallus-398p	FC:PI
203	Calibrachoa spathulata (L.B.Sm. & Downs) Stehmann & Semir.	No	Solanaceae	E	18	2	P	1,416	1·4	2·9	5·8	387^ae	O	Gallus-398p	FC:PI
204	Calibrachoa thymifolia (A. St.-Hil.) Stehmann & Semir.	No	Solanaceae	E	18	2	P	1,485	1·5	3·0	6·1	387^ae	O	Gallus-398p	FC:PI
205	Callistemon citrinus (Curtis) Skeels	No	Myrtaceae	E	22°	—^p	P	1,014	1·0	2·1	4·1	454	O	B^c	Fe
206	Callistemon rigidus R.Br.	No	Myrtaceae	E	—ⁿ	—^p	P	1,526	1·6	3·1	6·2	454	O	B^c	Fe
207	Camellia sinensis Kuntze	No	Theaceae	E	30	2	P	3,824	3·9	7·8	15·6	379	O	G	Fe
208	Canna indica^l	No	Cannaceae	M	18	2	P	706	0·7	1·4	2·9	379	O	J	Fe
209a	Cannabis sativa L. (female)	No	Cannabaceae	E	20	2	A	818.^am	0·8^am	1·7^am	3·3^am	414	O	Arab.^e	FC:DAPI
209b	Cannabis sativa L. (male)	No	Cannabaceae	E	20	2	A	843.^am	0·9^am	1·7^am	3·4^am	414	O	Arab.^e	FC:DAPI
210	Canotia holacantha Torr.	No	Celastraceae	E	30	2	P	181	0·2	0·4	0·7	378	O	J	Fe
211b	Capsella bursa-pastoris (L.) Medic.	No	Cruciferae^j	E	32°	4	A	686	0·7	1·4	2·8	457^bm	O	B^d	Fe
212h	Capsicum annuum L. cv. Doux Long des Landes	No	Solanaceae	E	24	2	—^q	3,734	3·8	7·6	15·2	434	O	G^c	FC:PI
213c	Capsicum baccatum L. ssp. pendulum	No	Solanaceae	E	24	2	—^q	4,111	4·2	8·4	16·8	434^ay	O	G-212h	FC:PI
213d	Capsicum baccatum L. ssp. baccatum	No	Solanaceae	E	24	2	—^q	4,131	4·2	8·4	16·9	434^ay	O	G-212h	FC:PI
214	Capsicum cardenasii Heiser & Smith	No	Solanaceae	E	24°	2	—^q	4,395	4·5	9·0	17·9	434	O	G-212h	FC:PI
215	Capsicum chacoense A.T.Hunz.	No	Solanaceae	E	24°	2	—^q	3,753	3·8	7·7	15·3	434	O	G-212h	FC:PI
216b	Capsicum chinense Jacq.	No	Solanaceae	E	24	2	—^q	3,940	4·0	8·0	16·1	434^ay	O	G-212h	FC:PI
217b	Capsicum eximium A.T.Hunz.	No	Solanaceae	E	24°	2	—^q	4,263	4·4	8·7	17·4	434^ay	O	G-212h	FC:PI
218b	Capsicum frutescens L.	No	Solanaceae	E	24°	2	—^q	3,891	4·0	7·9	15·9	434^ay	O	G-212h	FC:PI
219	Capsicum praetermissum Heiser & Smith	No	Solanaceae	E	24°	2	—^q	4,474	4·6	9·1	18·3	434	O	G-212h	FC:PI
220b	Capsicum pubescens R. & P.	No	Solanaceae	E	24°	2	—^q	4,763	4·9	9·7	19·4	434^ay	O	G-212h	FC:PI
221	Capsicum tovarii Eshbaugh, Smith & Nickrent	No	Solanaceae	E	24°	2	—^q	3,886	4·0	7·9	15·9	434	O	G-212h	FC:PI
222a	Cardamine amara L.	No	Cruciferae	E	16	2	P	238	0·2	0·5	1·0	465	O	Lycopers.^c	FC:PI
223	Castanospermum australe A.Cunn. & C.Fraser	No	Leguminosae	E	—ⁿ	—^p	P	554	0·6	1·1	2·3	454	O	B^c	Fe
224	Casuarina glauca Sieb. ex Spring.	No	Casuarinaceae	E	18°	2	P	343	0·4	0·7	1·4	452	O	Petunia^e	FC:PI
225	Catunaregam spinosa (Thunb.) Trivengadum	No	Rubiaceae	E	22°	—^p	P	343	0·4	0·7	1·4	454	O	B^c	Fe
226b	Centaurea scabiosa^l	No	Compositae	E	—ⁿ	—^p	P	1,254	1·3	2·6	5·1	465	O	J	Fe
227	Cephalotus follicularis Labill.	No	Cephalotaceae	E	20°	2	P	625	0·6	1·3	2·6	378	O	J	Fe
228a	Cerastium alpinum L.	No	Caryophyllaceae	E	72	4	P	1,813	1·9	3·7	7·4	427	O	Cerastium^e	FC:DAPI
228b	Cerastium alpinum L.	No	Caryophyllaceae	E	72	4	P	1,970	2·0	4·0	8·0	427	O	B^c	Fe
229	Cerastium arcticum Lange s. str.	No	Caryophyllaceae	E	108	6	P	3,126	3·2	6·4	12·8	427	O	Cerastium^e	FC:DAPI
230	Cerastium arvense L. ssp. glandulosum (Kit.) Soo	No	Caryophyllaceae	E	36	2^au	P	666	0·7	1·4	2·7	427	O	Cerastium^e	FC:DAPI
231	Cerastium arvense L. ssp. arvense	No	Caryophyllaceae	E	72	4	P	1,274	1·3	2·6	5·2	427	O	Cerastium^e	FC:DAPI
232a	Cerastium banaticum (Rochel) Heuff.	No	Caryophyllaceae	E	36	2^au	P	1,470	1·5	3·0	6·0	427	O	B^c	Fe
232b	Cerastium banaticum (Rochel) Heuff.	No	Caryophyllaceae	E	36	2^au	P	1,529	1·6	3·1	6·2	427	O	Cerastium^e	FC:DAPI
233	Cerastium carinthiacum Vest	No	Caryophyllaceae	E	36	2^au	P	1,480	1·5	3·0	6·0	427	O	Cerastium^e	FC:DAPI
234a	Cerastium eriophorum Kit. in Schult.	No	Caryophyllaceae	E	36	2^au	P	1,264	1·3	2·6	5·2	427	O	B^c	Fe
234b	Cerastium eriophorum Kit. in Schult.	No	Caryophyllaceae	E	36	2^au	P	1,274	1·3	2·6	5·2	427	O	Nicot.^e	FC:DAPI
235b	Cerastium fontanum Baumg.	No	Caryophyllaceae	E	144	8	P	3,469	3·5	7·1	14·2	427	O	Cerastium^e	FC:DAPI
236a	Cerastium latifolium L.	No	Caryophyllaceae	E	36	2^au	P	1,421	1·5	2·9	5·8	427	O	B^c	Fe
236b	Cerastium latifolium L.	No	Caryophyllaceae	E	36	2^au	P	1,470	1·5	3·0	6·0	427	O	Cerastium^e	FC:DAPI
237a	Cerastium transsylvanicum Schur ex Griseb. & Schenk	No	Caryophyllaceae	E	108	6	P	3,038	3·1	6·2	12·4	427	O	Cerastium^e	FC:DAPI
237b	Cerastium transsylvanicum Schur ex Griseb. & Schenk	No	Caryophyllaceae	E	108	6	P	3,048	3·1	6·2	12·4	427	O	B^c	Fe
238	Ceratophyllum demersum L.	No	Ceratophyllaceae	BA	c.70	6	P	674	0·7	1·4	2·8	381	O	J	Fe
239	Chenopodium album L.^h	No	Amaranthaceae	E	18°	2	A	750	0·8	1·5	3·1	455	O	B^c	Fe
240	Chenopodium album L.^h	No	Amaranthaceae	E	36°	4	A	1,597	1·6	3·3	6·5	455	O	B^c	Fe
241b	Chenopodium album L.^h	No	Amaranthaceae	E	54°	6	A	2,423	2·5	4·9	9·9	455	O	B^c	Fe
242	Chenopodium berlandieri (Saff.) Wilson & Heiser ssp. nuttalliae^h	No	Amaranthaceae	E	36°	4	A	1,446	1·5	3·0	5·9	455	O	B^c	Fe
243	Chenopodium bushianum Aellen	No	Amaranthaceae	E	36°	4	A	1,558	1·6	3·2	6·4	455	O	B^c	Fe
244	Chenopodium ficifolium Sm.	No	Amaranthaceae	E	18°	2	A	649	0·7	1·3	2·7	455	O	B^c	Fe
245	Chenopodium giganteum D.Don	No	Amaranthaceae	E	54°	6	A	2,151	2·2	4·4	8·8	455	O	B^c	Fe
246	Chenopodium murale L.	No	Amaranthaceae	E	18°	2	A	610	0·6	1·2	2·5	455	O	B^c	Fe
247	Chenopodium opulifolium Schrad. ex Koch & Ziz	No	Amaranthaceae	E	36°	4	A	1,303	1·3	2·7	5·3	455	O	B^c	Fe
248b	Chenopodium pallidicaule Aellen	No	Amaranthaceae	E	18°	2	A	617	0·6	1·3	2·5	455	O	B^c	Fe
249b	Chenopodium quinoa Willd.^h	No	Amaranthaceae	E	36°	4	A	1,585	1·6	3·2	6·5	455	O	B^c	Fe
250	Chenopodium ugandae (Aell.) Aell.	No	Amaranthaceae	E	32°	—^p	A	1,401	1·4	2·9	5·7	455	O	B^c	Fe
251	Chenopodium vulvaria L.	No	Amaranthaceae	E	18°	2	A	622	0·6	1·3	2·5	455	O	B^c	Fe
252	Chloranthus spicatus Mak.	No	Chloranthaceae	BA	30	—^p	P	3,526	3·6	7·2	14·4	381	O	G	FC:PI
253	Chorisia speciosa St. Hill	No	Malvaceae	E	86°	—^p	P	845	0·9	1·7	3·5	454	O	B^c	Fe
254	Cicca acida (L.) Merr.	No	Euphorbiaceae	E	—ⁿ	—^p	P	938	1·0	1·9	3·8	454	O	B^c	Fe
255	Cicer songaricum Steph. ex DC.	No	Leguminosae	E	16	2	P	1,328	1·4	2·7	5·4	435	O	B^c	Fe
256	Cienfuegosia tripartita H.B.K. Gurke	—^m	Malvaceae	E	20	2	—^q	931	1·0	1·9	3·8	444^bd	O	G^b2	FC:PI
257	Cienfuegosia yucatanensis Millspaugh	—^m	Malvaceae	E	22	2	—^q	980	1·0	2·0	4·0	444^bd	O	G^b2	FC:PI
258	Cistus albanicus E.F.Warburg ex Heywood	No	Cistaceae	E	18°	2	P	2,617	2·7	5·3	10·7	404	O	Raphanus^e	FC:PI
259	Cistus albidus L.	No	Cistaceae	E	18°	2	P	2,342	2·4	4·8	9·6	404	O	Raphanus^e	FC:PI
260	Cistus clusii Dunal	No	Cistaceae	E	18°	2	P	2,587	2·6	5·3	10·6	404	O	Raphanus^e	FC:PI
261	Cistus creticus L.	No	Cistaceae	E	18°	2	P	2,127	2·2	4·3	8·7	404	O	Raphanus^e	FC:PI
262	Cistus crispus L.	No	Cistaceae	E	18°	2	P	1,921	2·0	3·9	7·8	404	O	Raphanus^e	FC:PI
263	Cistus heterophyllus Desf. ssp. carthaginensis (Pau) Crespo & Mateo	No	Cistaceae	E	18°	2	P	2,362	2·4	4·8	9·6	404	O	Raphanus^e	FC:PI
264	Cistus ladanifer L.	No	Cistaceae	E	18°	2	P	2,181	2·2	4·5	8·9	404	O	Raphanus^e	FC:PI
265	Cistus laurifolius L.	No	Cistaceae	E	18°	2	P	2,185	2·2	4·5	8·9	404	O	Raphanus^e	FC:PI
266	Cistus libanotis L.	No	Cistaceae	E	18°	2	P	2,827	2·9	5·8	11·5	404	O	Raphanus^e	FC:PI
267	Cistus monspeliensis L.	No	Cistaceae	E	18°	2	P	2,881	2·9	5·9	11·8	404	O	Raphanus^e	FC:PI
268	Cistus osbeckiaefolius Webb ex Pitard & Proust	No	Cistaceae	E	18°	2	P	2,024	2·1	4·1	8·3	404	O	Raphanus^e	FC:PI
269	Cistus parviflorus Lam.	No	Cistaceae	E	18°	2	P	2,430	2·5	5·0	9·9	404	O	Raphanus^e	FC:PI
270	Cistus populifolius L.	No	Cistaceae	E	18°	2	P	2,102	2·1	4·3	8·6	404	O	Raphanus^e	FC:PI
271	Cistus psilosepalus Sweet	No	Cistaceae	E	18°	2	P	2,558	2·6	5·2	10·4	404	O	Raphanus^e	FC:PI
272	Cistus salviifolius L.	No	Cistaceae	E	18°	2	P	2,332	2·4	4·8	9·5	404	O	Raphanus^e	FC:PI
273	Cistus symphytifolius Lam.	No	Cistaceae	E	18°	2	P	2,406	2·5	4·9	9·8	404	O	Raphanus^e	FC:PI
274b	Citrus aurantium L.	No	Rutaceae	E	18°	2	P	431	0·4	0·9	1·8	426	O	Gallus^f	FC:PI
275b	Citrus grandis (L.) Osbeck	No	Rutaceae	E	18°	2	P	377	0·4	0·8	1·5	426	O	Gallus^f	FC:PI
276b	Citrus limon (L.) Burm. f.ⁱ	No	Rutaceae	E	18°	2	P	392	0·4	0·8	1·6	426	O	Gallus^f	FC:PI
277	Citrus limonia Osbeck cv. Brome Rangpur	No	Rutaceae	E	18°	2	P	402	0·4	0·8	1·6	426	O	Gallus^f	FC:PI
278b	Citrus paradisi Macfad.	No	Rutaceae	E	18°	2	P	392	0·4	0·8	1·6	426	O	Gallus^f	FC:PI
279	Citrus reshni Hort. ex Tanaka	No	Rutaceae	E	18°	2	P	402	0·4	0·8	1·6	426	O	Gallus^f	FC:PI
280e	Citrus sinensis (L.) Osbeck cv. Sargoins Grosse Ronde^h	No	Rutaceae	E	18°	2	P	372	0·4	0·8	1·5	426	O	Gallus^f	FC:PI
280f	Citrus sinensis (L.) Osbeck cv. Pineapple^h	No	Rutaceae	E	18°	2	P	417	0·4	0·9	1·7	426	O	Gallus^f	FC:PI
280g	Citrus sinensis (L.) Osbeck	No	Rutaceae	E	18°	2	P	588	0·6	1·2	2·4	457^bm	O	B^d	Fe
281	Citrus volkameriana Ten. & Pasq.	No	Rutaceae	E	18°	2	P	387	0·4	0·8	1·6	426	O	Gallus^f	FC:PI
282	Coccoloba diversifolia Jacq.	No	Polygonaceae	E	—ⁿ	—^p	P	1,127	1·2	2·3	4·6	454	O	B^c	Fe
283h	Coffea arabica L.	—^m	Rubiaceae	E	44	4	P	1,279	1·3	2·6	5·2	424	O	K^c	FC:PI
283i	Coffea arabica L.	No	Rubiaceae	E	44°	4	P	1,122	1·1	2·3	4·6	454	O	B^c	Fe
284c	Coffea brevipes Hiern.	—^m	Rubiaceae	E	22	2	P	760	0·8	1·6	3·1	424	O	K^c	FC:PI
285d	Coffea canephora Pierre. ex Froehn.	—^m	Rubiaceae	E	22	2	P	755	0·8	1·5	3·1	424	O	K^c	FC:PI
286c	Coffea congensis Froehn.	—^m	Rubiaceae	E	22	2	P	794	0·8	1·6	3·2	424	O	K^c	FC:PI
287d	Coffea eugenioides S.Moore	—^m	Rubiaceae	E	22	2	P	681	0·7	1·4	2·8	424	O	K^c	FC:PI
288c	Coffea humilis A.Cheval.	—^m	Rubiaceae	E	22	2	P	872	0·9	1·8	3·6	424	O	K^c	FC:PI
289d	Coffea liberica L. ssp. dewevrei Wild & Dur. Hiern	No	Rubiaceae	E	22	2	P	703	0·7	1·4	2·9	401	O	Petunia^e	FC:PI
289e	Coffea liberica L.	—^m	Rubiaceae	E	22	2	P	823	0·8	1·7	3·4	424	O	K^c	FC:PI
290d	Coffea pseudozanguebariae D.M.Bridson	—^m	Rubiaceae	E	22	2	P	534	0·5	1·1	2·2	424	O	K^c	FC:PI
291c	Coffea racemosa^l	—^m	Rubiaceae	E	22	2	P	466	0·5	1·0	1·9	424	O	K^c	FC:PI
292c	Coffea sessiliflora D.M.Bridson	—^m	Rubiaceae	E	22	2	P	510	0·5	1·0	2·1	424	O	K^c	FC:PI
293	Coffea sp. F. Bridson^y	—^m	Rubiaceae	E	22	2	P	652	0·7	1·3	2·7	424	O	K^c	FC:PI
294	Coffea sp. Moloundou^y	—^m	Rubiaceae	E	22	2	P	828	0·8	1·7	3·4	424	O	K^c	FC:PI
295c	Coffea stenophylla G.Don.	—^m	Rubiaceae	E	22	2	P	662	0·7	1·4	2·7	424	O	K^c	FC:PI
296b	Colocasia antiquorum Schott var. 1ⁱ	No	Araceae	M	32	—^p	AP	4,951	5·1	10·1	20·2	411	O	B^c	Fe
297	Commiphora mossambicensis Engl.	No	Burseraceae	E	26	2	P	613	0·6	1·3	2·5	379	O	J	Fe
298	Coriaria myrtifolia L.	No	Coriariaceae	E	c. 72	8	P	326	0·3	0·7	1·3	378	O	J	Fe
299	Cosmos atrosanguineus^l	No	Compositae	E	48	—^p	P	7,191	7·3	14·7	29·4	465	O	F	Fe
300c	Crepis biennis L.^h	No	Compositae	E	c. 40	10	B	7,448	7·6	15·2	30·4	394^ah	O	G^c	CIA
300d	Crepis biennis L.^h	No	Compositae	E	c. 40	10	B	7,928	8·1	16·2	32·4	394^ah	O	G^c	CIA
300e	Crepis biennis L.^h	No	Compositae	E	c. 40	10	B	8,173	8·3	16·7	33·4	394^ah	O	G^c	Fe
300f	Crepis biennis L.^h	No	Compositae	E	c. 40	10	B	8,555	8·7	17·5	34·9	394^ah	O	G^c	Fe
301a	Crepis bithynica var. pirinica Acht.ⁱ	No	Compositae	E	10	2	P	3,156	3·2	6·4	12·9	394	O	G^c	Fe
301b	Crepis bithynica var. bithynica Boiss.ⁱ	No	Compositae	E	10	2	P	3,244	3·3	6·6	13·2	394	O	G^c	Fe
302h	Crepis capillaris (L.) Wallr.	No	Compositae^j	E	6°	2	A	2,597	2·7	5·3	10·6	457^bm	O	B^d	Fe
303a	Crepis conyzaefolia (Gouan) A.Kernerⁱ	No	Compositae	E	8	2	P	5,400	5·5	11·0	22·0	394	O	G^c	Fe
303b	Crepis conyzaefolia (Gouan) A.Kernerⁱ	No	Compositae	E	8	2	P	5,576	5·7	11·4	22·8	394	O	E^c	FC:PI
304a	Crepis paludosa (L.) Moenchⁱ	No	Compositae	E	12	2	P	4,077	4·2	8·3	16·6	394	O	G^c	Fe
304b	Crepis paludosa (L.) Moenchⁱ	No	Compositae	E	12	2	P	4,361	4·5	8·9	17·8	394	O	E^c	FC:PI
305b	Crepis pulchra L.^h	No	Compositae	E	8	2	A	4,459	4·6	9·1	18·2	394	O	G^c	CIA
305c	Crepis pulchra L.^h	No	Compositae	E	8	2	A	5,449	5·6	11·1	22·2	394^ai	O	G^c	CIA
305d	Crepis pulchra L.^h	No	Compositae	E	8	2	A	4,851	5·0	9·9	19·8	394	O	G^c	Fe
305e	Crepis pulchra L.^h	No	Compositae	E	8	2	A	5,400	5·5	11·0	22·0	394^ai	O	G^c	Fe
305f	Crepis pulchra L.^h	No	Compositae	E	8	2	A	5,380	5·5	11·0	22·0	394	O	E^c	FC:PI
305g	Crepis pulchra L.^h	No	Compositae	E	8	2	A	5,988	6·1	12·2	24·4	394	O	E^c	FC:PI
306a	Crepis sancta (L.) Babc.^h	No	Compositae	E	10	2	A	2,048	2·1	4·2	8·4	394^ah	O	G^c	Fe
306b	Crepis sancta (L.) Babc.^h	No	Compositae	E	10	2	A	2,176	2·2	4·4	8·9	394^ah	O	G^c	CIA
307	Crepis schachtii Babc.ⁱ	No	Compositae	E	10	2	P	2,764	2·8	5·6	11·3	394	O	G^c	Fe
308b	Crepis setosa Haller f.ⁱ	No	Compositae	E	8	2	A	1,695	1·7	3·5	6·9	394	O	G^c	CIA
309a	Crepis viscidula Froel.ⁱ	No	Compositae	E	12	2	P	4,283	4·4	8·7	17·5	394	O	G^c	CIA
309b	Crepis viscidula Froel.ⁱ	No	Compositae	E	12	2	P	4,782	4·9	9·8	19·5	394	O	E^c	FC:PI
310b	Crepis zacintha (L.) Babc.ⁱ	No	Compositae	E	6	2	A	1,058	1·1	2·2	4·3	394	O	G^c	CIA
311	Crocus biflorus Mill.	No	Iridaceae	M	8^ao	2	P	4,253	4·3	8·7	17·4	419	O	Homo^f	FC:PI
312	Crocus cartwrightianus Herb.ⁱ	No	Iridaceae	M	16	2	P	3,886	4·0	7·9	15·9	419	O	Homo^f	FC:PI
313	Crocus etruscus Parl.	No	Iridaceae	M	8	2	P	3,553	3·6	7·3	14·5	419	O	Homo^f	FC:PI
314	Crocus sativus L.ⁱ	No	Iridaceae	M	24	3	P	—^t	—^t	11·8	23·6	419	O	Homo^f	FC:PI
315	Crocus thomasii Ten.	No	Iridaceae	M	16	2	P	4,258	4·3	8·7	17·4	419	O	Homo^f	FC:PI
316	Cyclamen hederifolium^l	No	Primulaceae	E	—ⁿ	—^p	P	2,923	3·0	6·0	11·9	465	O	G	FC:PI
317a	Cyclamen trochopteranthum O.Schwarz^h	No	Primulaceae	E	30	—^p	P	10,373	10·6	21·2	42·3	465	O	G	Fe
317b	Cyclamen trochopteranthum O.Schwarz^h	No	Primulaceae	E	30	—^p	P	13,458	13·7	27·5	54·9	465	O	G	Fe
318b	Cynodon dactylon (L.) Pers.	No	Gramineae	M	36	4	P	956	1·0	2·0	3·9	417	O	Gallus^f	FC:PI
318c	Cynodon dactylon (L.) Pers. var. dactylon	No	Gramineae	M	36	4	P	1,103	1·1	2·3	4·5	438^ba	O	Sus^f	FC:PI
319	Cynodon dactylon (L.) Pers. var. dactylon	No	Gramineae	M	54	6	P	1,436	1·5	2·9	5·9	438^ba	O	Ictal.^f	FC:PI
320a	Cynodon transvaalensis Burtt-Davy	No	Gramineae	M	18	2	P	505	0·5	1·0	2·1	417	O	Gallus^f	FC:PI
320b	Cynodon transvaalensis Burtt-Davy	No	Gramineae	M	18	2	P	544	0·6	1·1	2·2	438^ba	O	Ictal.^f	FC:PI
321	Dalbergia horrida Dennst.	No	Leguminosae	E	20	2	P	1,928	2·0	3·9	7·9	445	O	G	Fe
322a	Dalbergia lanceolaria Lin. f.	No	Leguminosae	E	20	2	P	1,431	1·5	2·9	5·8	445	O	G	Fe
323a	Dalbergia latifolia Roxb.	No	Leguminosae	E	20	2	P	1,681	1·7	3·4	6·9	445	O	G	Fe
324	Dalbergia malabarica Prain.	No	Leguminosae	E	20	2	P	1,847	1·9	3·8	7·5	445	O	G	Fe
325	Dalbergia melanoxylon Guill & Perr.	No	Leguminosae	E	20	2	P	1,806	1·8	3·7	7·4	445	O	G	Fe
326	Dalbergia paniculata Roxb.	No	Leguminosae	E	20	2	P	1,551	1·6	3·2	6·3	445	O	G	Fe
327	Dalbergia rubiginosa Roxb.	No	Leguminosae	E	20	2	P	1,803	1·8	3·7	7·4	445	O	G	Fe
328	Dalbergia sissoides Grah.	No	Leguminosae	E	20	2	P	1,764	1·8	3·6	7·2	445	O	G	Fe
329a	Dalbergia sissoo Roxb. ex DC.	No	Leguminosae	E	20	2	P	1,585	1·6	3·2	6·5	445	O	G	Fe
329b	Dalbergia sissoo Roxb. ex DC.	No	Leguminosae	E	20°	2	P	691	0·7	1·4	2·8	454	O	B^c	Fe
330	Dalbergia volubilis Roxb.	No	Leguminosae	E	20	2	P	1,901	1·9	3·9	7·8	445	O	G	Fe
331	Damasonium alisma Mill.	No	Alismataceae	M	—ⁿ	—^p	A	23,143	23·6	47·2	94·5	465	O	B	FC:PI
332	Dasypogon hookeri Drumm.	No	Dasypogonaceae	M	14	2	P	426	0·4	0·9	1·7	380	O	J	FC:PI
333	Dasypyrum hordeaceum (Cosson & Durieu) Candargy	No	Gramineae	M	28	4	P	10,288	10·5	21·0	42·0	431	C	—^aw	Fe
334d	Dasypyrum villosum (=Haynaldia villosa) (L.) P. Candargy^h	No	Gramineae	M	14	2	A	10,408	10·6	21·2	42·5	430	O	C^c	Fe
334e	Dasypyrum villosum (=Haynaldia villosa) (L.) P. Candargy^h	No	Gramineae	M	14	2	A	6,262	6·4	12·8	25·6	430	O	C^c	Fe
335b	Decaisnea fargesii Franch.	No	Lardizabalaceae^k	E	40°	2	P	2,450	2·5	5·0	10·0	457^bm	O	B^d	Fe
336	Deutzia prunifolia Rehder	No	Hydrangeaceae	E	52	4	P	1,835	1·9	3·7	7·5	378	O	J	Fe
337	Dictamnus albus L.	No	Rutaceae	E	36°	—^p	P	3,381	3·5	6·9	13·8	457^bm	O	B^d	Fe
338	Dieffenbachia picta Schott	No	Araceae	M	36	—^p	P	12,083	12·3	24·7	49·3	411	O	B^c	Fe
339	Diospyros discolor Willd.	No	Ebenaceae	E	30°	—^p	P	1,174	1·2	2·4	4·8	454	O	B^c	Fe
340	Diospyros malabarica Kost.	No	Ebenaceae	E	30°	—^p	P	1,436	1·5	2·9	5·9	454	O	B^c	Fe
341	Dissotis canescens Hook. f.	No	Melastomataceae	E	c. 28-32	—^p	P	181	0·2	0·4	0·7	378	O	J	Fe
342a	Doritis pulcherrima Lindl.	No	Orchidaceae	M	38	2	P	6,610	6·7	13·5	27·0	447	O	G^b2	FC:PI
343	Doryanthes palmeri W. Hill ex Benth.	No	Doryanthaceae	M	48	—^p	P	3,239	3·3	6·6	13·2	380	O	G	FC:PI
344	Drimys vickeriana A.C. Smith	No	Winteraceae	BA	—ⁿ	—^p	P	1,105	1·1	2·3	4·5	381	O	K	FC:PI
345	Drypetes roxburghii Wall.	No	Putranjavaceae^k	E	42°	—^p	P	1,002	1·0	2·0	4·1	449^bf	O	B^c	Fe
346	Ehretia laevis (Rottler ex G.Don) Roxb.	No	Boraginaceae	E	—ⁿ	—^p	P	3,533	3·6	7·2	14·4	454	O	B^c	Fe
347	Eremochloa ophiuroides (Munro) Hack.	No	Gramineae	M	18	2	P	813	0·8	1·7	3·3	417	O	Gallus^f	FC:PI
348	Eriocaulon aquaticum^l	No	Eriocaulaceae	M	32	4	P	4,101	4·2	8·4	16·7	380	O	G	FC:PI
349	Escallonia rubra^l	No	Escalloniaceae	E	24	2	P	414	0·4	0·8	1·7	380	O	J	Fe
350	Eucnide grandiflora Rose	No	Loasaceae	E	c. 38-40	2 or 6	P	588	0·6	1·2	2·4	378	O	J	Fe
351	Eucommia ulmoides Oliver	No	Eucommiaceae	E	34	2	P	725	0·7	1·5	3·0	379	O	G	Fe
352a	Fagus sylvatica L. var. tortuosa Pepin Willk.	No	Fagaceae	E	24°	2	P	544	0·6	1·1	2·2	433	O	Petunia^e	FC:PI
352b	Fagus sylvatica L.	No	Fagaceae	E	24°	2	P	544	0·6	1·1	2·2	433	O	Petunia^e	FC:PI
352c	Fagus sylvatica L. var. purpurea Ait.	No	Fagaceae	E	24°	2	P	549	0·6	1·1	2·2	433	O	Petunia^e	FC:PI
352d	Fagus sylvatica L. var. pendula Lodd.	No	Fagaceae	E	24°	2	P	554	0·6	1·1	2·3	433	O	Petunia^e	FC:PI
353d	Festuca arundinacea Schreb.	No	Gramineae	M	42	6	P	7,639	7·8	15·6	31·2	417	O	Nicot.^e	FC:PI
354	Festuca longifolia Thuill.	No	Gramineae	M	42	6	P	6,223	6·4	12·7	25·4	417	O	F^c	FC:PI
355	Firmiana colorata (Roxb.) R.Br.	No	Malvaceae	E	40°	—^p	P	1,615	1·6	3·3	6·6	454	O	B^c	Fe
356	Flagellaria guineensis Schum.	No	Flagellariaceae	M	38°	2	P	880	0·9	1·8	3·6	380	O	K	FC:PI
357	Flemingia bracteata Wight	No	Leguminosae	E	22	2	P	1,570	1·6	3·2	6·4	443^bc	O	B^c	Fe
358	Fortunella hindsii Swing.	No	Rutaceae	E	36°	4	P	622	0·6	1·3	2·5	426	O	Gallus^f	FC:PI
359	Fouquieria splendens Engelm.	No	Fouquieriaceae	E	24	4	P	519	0·5	1·1	2·1	378	O	J	Fe
360	Fragaria x ananassa cv. Redcoat Duch.	—^m	Rosaceae	E	56	8	P	598	0·6	1·2	2·4	442	O	Gallus^f	FC:M
361a	Gagea lutea (L.) Ker Gawl.	No	Liliaceae	M	72	6	P	19,355	19·8	39·5	79·0	413	O	B^c	Fe
361b	Gagea lutea (L.) Ker Gawl.	No	Liliaceae	M	72	6	P	19,825	20·2	40·5	80·9	413	O	B^c	FC:EB
362	Gardenia resiniflua Hiern	No	Rubiaceae	E	—ⁿ	—^p	P	1,269	1·3	2·6	5·2	454	O	B^c	Fe
363	Garrya fremontii Torr.	No	Garryaceae	E	c. 20	2	P	1,490	1·5	3·0	6·1	380	O	Lycopers.^c	FC:PI
364j	Glycine max (L.) Merr. strain T215^h	No	Leguminosae	E	40°	2	A	1,161	1·2	2·4	4·7	423^as	O	F^c	FC:PI
364k	Glycine max (L.) Merr. strain PI423·894^h	No	Leguminosae	E	40°	2	A	1,215	1·2	2·5	5·0	423^as	O	F^c	FC:PI
364l	Glycine max (L.) Merr.^h	—^m	Leguminosae	E	40°	2	A	1,250	1·3	2·6	5·1	432^ax	C	Glycine^e	FC:PI
364m	Glycine max (L.) Merr.^h	—^m	Leguminosae	E	40°	2	A	1,401	1·4	2·9	5·7	432^ax	C	Glycine^e	FC:PI
365	Goodenia mimuloides S.Moore	No	Goodeniaceae	E	16	2	A	507	0·5	1·0	2·1	379	O	G	Fe
366	Gossypioides herbaceum L.	—^m	Malvaceae	E	26°	2	—^q	1,813	1·9	3·7	7·4	444^bd	O	G^b2	FC:PI
367	Gossypioides raimondii Ulbrich	—^m	Malvaceae	E	26°	2	—^q	980	1·0	2·0	4·0	444^bd	O	G^b2	FC:PI
368	Gunnera manicata Linden	No	Gunneraceae	E	34	2	P	7,286	7·4	14·9	29·7	379	O	F	Fe
369	Gymnostoma deplancheana (Miq.) L.Johnson	No	Moraceae^k	E	16°	2	P	368	0·4	0·8	1·5	452	O	Petunia^e	FC:PI
370	Haldina cordifolia (Roxb.) Ridsdale	No	Rubiaceae	E	44°	4	P	1,296	1·3	2·6	5·3	454	O	B^c	Fe
371	Hampea appendiculata (J. Donnell-Smith) Standley	—^m	Malvaceae	E	26°	2	—^q	2,891	3·0	5·9	11·8	444^bd	O	G^b2	FC:PI
372	Hanguana malayana Merrill	No	Hanguanaceae	M	c. 170	—^p	P	1,612	1·6	3·3	6·6	380	O	F	Fe
373a	Hedera canariensis Willd.	No	Araliaceae	E	48°	2	P	1,372	1·4	2·8	5·6	429	O	Glycine^e	FC:PI
373b	Hedera canariensis Willd.	No	Araliaceae	E	48°	2	P	1,509	1·5	3·1	6·2	429	O	G^c	CIA
374a	Hedera colchica C.Koch.	No	Araliaceae	E	192	8	P	5,341	5·5	10·9	21·8	429	O	Glycine^e	FC:PI
374b	Hedera colchica C.Koch.	No	Araliaceae	E	192	8	P	5,586	5·7	11·4	22·8	429	O	G^c	CIA
375e	Hedera helix L.	No	Araliaceae	E	48	2	P	1,460	1·5	3·0	6·0	429^av	O	G^c	CIA
375f	Hedera helix L.	No	Araliaceae	E	48	2	P	1,372	1·4	2·8	5·6	429^av	O	Glycine^e	FC:PI
375g	Hedera helix L. f. arborescens C.K. Schneider	No	Araliaceae	E	48	2	P	1,509	1·5	3·1	6·2	429	O	G^c	CIA
375h	Hedera helix L. f. arborescens C.K. Schneider	No	Araliaceae	E	48	2	P	1,382	1·4	2·8	5·6	429	O	Glycine^e	FC:PI
376r	Helianthus annuus L.	No	Compositae	E	34	2	A	3,577	3·7	7·3	14·6	403	O	G	FC:PI
377	Heliconia rostrata Ruiz & Pav.	No	Heliconiaceae	M	24	2	P	441	0·5	0·9	1·8	379	O	J	Fe
378	Helleborus argutifolius Viv.	No	Ranunculaceae	E	32	2	P	9,261	9·5	18·9	37·8	383^z	O	Agave^e	FC:PI
379a	Helleborus atrorubens Waldst. & Kit. ‘Cupreus’ⁱ	No	Ranunculaceae	E	32	2	P	14,504	14·8	29·6	59·2	383^z	O	Agave^e	FC:PI
379b	Helleborus atrorubens Waldst. & Kit.ⁱ	No	Ranunculaceae	E	32	2	P	15,092	15·4	30·8	61·6	383^z	O	Agave^e	FC:PI
380	Helleborus cyclophyllus (A.Br.) Boiss.	No	Ranunculaceae	E	32	2	P	14,651	15·0	29·9	59·8	383^z	O	Agave^e	FC:PI
381	Helleborus dumetorum Waldst. & Kit.	No	Ranunculaceae	E	32	2	P	15,876	16·2	32·4	64·8	383^z	O	Agave^e	FC:PI
382a	Helleborus foetidus L. ‘Wester Flisk’	No	Ranunculaceae	E	32	2	P	11,417	11·7	23·3	46·6	383^z	O	Agave^e	FC:PI
382b	Helleborus foetidus L.ⁱ	No	Ranunculaceae	E	32	2	P	11,466	11·7	23·4	46·8	383^z	O	Agave^e	FC:PI
383	Helleborus lividus Aiton	No	Ranunculaceae	E	32	2	P	9,310	9·5	19·0	38·0	383^z	O	Agave^e	FC:PI
384a	Helleborus multifidus Vis. ssp. hercegovinus (Martinis) B.Mathew	No	Ranunculaceae	E	32	2	P	14,504	14·8	29·6	59·2	383^z	O	Agave^e	FC:PI
384b	Helleborus multifidus Vis. ssp. istriacus (Schiffner) Merxm & Podl.	No	Ranunculaceae	E	32	2	P	14,749	15·1	30·1	60·2	383^z	O	Agave^e	FC:PI
384c	Helleborus multifidus Vis. ssp. multifidus	No	Ranunculaceae	E	32	2	P	14,798	15·1	30·2	60·4	383^z	O	Agave^e	FC:PI
384d	Helleborus multifidus Vis. ssp. bocconei siculus	No	Ranunculaceae	E	32	2	P	15,043	15·4	30·7	61·4	383^z	O	Agave^e	FC:PI
384e	Helleborus multifidus Vis.ssp. bocconei (Tenore) B.Mathew	No	Ranunculaceae	E	32	2	P	15,092	15·4	30·8	61·6	383^z	O	Agave^e	FC:PI
385a	Helleborus niger L. (double flower)ⁱ	No	Ranunculaceae	E	32	2	P	13,720	14·0	28·0	56·0	383^z	O	Agave^e	FC:PI
385b	Helleborus niger L.ⁱ	No	Ranunculaceae	E	32	2	P	13,867	14·2	28·3	56·6	383^z	O	Agave^e	FC:PI
385c	Helleborus niger L. ssp. macranthus (Freyn) Schiffnerⁱ	No	Ranunculaceae	E	32	2	P	14,406	14·7	29·4	58·8	383^z	O	Agave^e	FC:PI
386	Helleborus odorus Waldst. & Kit.ⁱ	No	Ranunculaceae	E	32	2	P	15,043	15·4	30·7	61·4	383^z	O	Agave^e	FC:PI
387a	Helleborus orientalis Lamarck ssp. orientalisⁱ	No	Ranunculaceae	E	32	2	P	14,553	14·9	29·7	59·4	383^z	O	Agave^e	FC:PI
387b	Helleborus orientalis Lamarckⁱ	No	Ranunculaceae	E	32	2	P	14,725	15·0	30·1	60·1	383^z	O	Agave^e	FC:PI
387c	Helleborus orientalis Lamarck ssp. guttatus (A.Br. & Sauer) B.Mathewⁱ	No	Ranunculaceae	E	32	2	P	14,749	15·1	30·1	60·2	383^z	O	Agave^e	FC:PI
387d	Helleborus orientalis Lamarck ssp. abchasicus (A.Br.) B.Mathewⁱ	No	Ranunculaceae	E	32	2	P	14,798	15·1	30·2	60·4	383^z	O	Agave^e	FC:PI
387e	Helleborus orientalis Lamarck ‘Kochii’ⁱ	No	Ranunculaceae	E	32	2	P	14,994	15·3	30·6	61·2	383^z	O	Agave^e	FC:PI
388	Helleborus purpurascens Waldst. & Kit.	No	Ranunculaceae	E	32	2	P	14,945	15·3	30·5	61·0	383^z	O	Agave^e	FC:PI
389	Helleborus thibetanus Franchet	No	Ranunculaceae	E	32	2	P	17,493	17·9	35·7	71·4	383^z	O	Agave^e	FC:PI
390a	Helleborus torquatusArcher Hind ‘Dido’ (double flowers)ⁱ	No	Ranunculaceae	E	32	2	P	14,602	14·9	29·8	59·6	383^z	O	Agave^e	FC:PI
390b	Helleborus torquatus Archer Hindⁱ	No	Ranunculaceae	E	32	2	P	14,749	15·1	30·1	60·2	383^z	O	Agave^e	FC:PI
390c	Helleborus torquatus Archer Hind ‘Croaticus’ⁱ	No	Ranunculaceae	E	32	2	P	14,700	15·0	30·0	60·0	383^z	O	Agave^e	FC:PI
391	Helleborus vesicarius Aucher	No	Ranunculaceae	E	32	2	P	13,867	14·2	28·3	56·6	383^z	O	Agave^e	FC:PI
392a	Helleborus viridis L. ssp. viridis	No	Ranunculaceae	E	32	2	P	14,896	15·2	30·4	60·8	383^z	O	Agave^e	FC:PI
392b	Helleborus viridis L. ssp. occidentalis (Reut.) Schiffner	No	Ranunculaceae	E	32	2	P	15,092	15·4	30·8	61·6	383^z	O	Agave^e	FC:PI
393	Hernandia nymphaeifolia (C.Presl.) Kubitzki	No	Hernandiaceae	BA	—ⁿ	—^p	P	2,340	2·4	4·8	9·6	454	O	B^c	Fe
394	Herniaria glabra Linn.	No	Caryophyllaceae	E	18	2	AP	515	0·5	1·1	2·1	465	O	J	Fe
395b	Hieracium aurantiacum L.	No	Compositae^j	E	36°	4	P	3,626	3·7	7·4	14·8	457^bm	O	B^d	Fe
396	Holoptelea integrifolia Planch.	No	Ulmaceae	E	28°	—^p	P	666	0·7	1·4	2·7	454	O	B^c	Fe
397	Homalomena rubescens Kunth	No	Araceae	M	34	—^p	AP	8,955	9·1	18·3	36·6	411	O	B^c	Fe
398p	Hordeum vulgare L. cv. New Golden	No	Gramineae	M	14°	2	A	5,096	5·2	10·4	20·8	387^ae	O	Gallus^f	FC:PI
399	Hosta capitata (Koidzumi) Nakai	No	Asparagaceae^k	M	60	2	P	9,457	9·7	19·3	38·6	384^aa	O	Agave sp.^ab	FC:PI
400	Hosta clausa var. normalis F.Maekawa	No	Asparagaceae^k	M	60	2	P	9,408	9·6	19·2	38·4	384^aa	O	Agave sp.^ab	FC:PI
401	Hosta clausa Nakai var. clausa	No	Asparagaceae^k	M	90	3	P	—^t	—^t	28·5	57·0	384^aa	O	Agave sp.^ab	FC:PI
402	Hosta gracillima F.Maekawa	No	Asparagaceae^k	M	60	2	P	10,829	11·1	22·1	44·2	384^aa	O	Agave sp.^ab	FC:PI
403	Hosta hypoleuca Murata	No	Asparagaceae^k	M	60	2	P	12,495	12·8	25·5	51·0	384^aa	O	Agave sp.^ab	FC:PI
404	Hosta jonesii M.Chung	No	Asparagaceae^k	M	60	2	P	8,575	8·8	17·5	35·0	384^aa	O	Agave sp.^ab	FC:PI
405	Hosta kikutii F.Maekawa	No	Asparagaceae^k	M	60	2	P	11,172	11·4	22·8	45·6	384^aa	O	Agave sp.^ab	FC:PI
406	Hosta kiyosumiensis F.Maekawa	No	Asparagaceae^k	M	60	2	P	11,907	12·2	24·3	48·6	384^aa	O	Agave sp.^ab	FC:PI
407	Hosta longipes var. longipes Matsumura	No	Asparagaceae^k	M	60	2	P	12,740	13·0	26·0	52·0	384^aa	O	Agave sp.^ab	FC:PI
408	Hosta longissima Honda	No	Asparagaceae^k	M	60	2	P	9,604	9·8	19·6	39·2	384^aa	O	Agave sp.^ab	FC:PI
409	Hosta minor Nakai ‘Gosan’	No	Asparagaceae^k	M	60	2	P	8,428	8·6	17·2	34·4	384^aa	O	Agave sp.^ab	FC:PI
410	Hosta plantaginea (Lamarck) Ascherson	No	Asparagaceae^k	M	60	2	P	12,103	12·4	24·7	49·4	384^aa	O	Agave sp.^ab	FC:PI
411	Hosta pulchella N.Fujita	No	Asparagaceae^k	M	60	2	P	10,633	10·9	21·7	43·4	384^aa	O	Agave sp.^ab	FC:PI
412	Hosta pycnophylla F.Maekawa	No	Asparagaceae^k	M	60	2	P	10,878	11·1	22·2	44·4	384^aa	O	Agave sp.^ab	FC:PI
413b	Hosta rectifolia Nakai	No	Asparagaceae^k	M	60	2	P	10,437	10·7	21·3	42·6	384^aa	O	Agave sp.^ab	FC:PI
414	Hosta rupifraga Nakai	No	Asparagaceae^k	M	60	2	P	12,985	13·3	26·5	53·0	384^aa	O	Agave sp.^ab	FC:PI
415	Hosta shikokiana N.Fujita	No	Asparagaceae^k	M	60	2	P	11,221	11·5	22·9	45·8	384^aa	O	Agave sp.^ab	FC:PI
416	Hosta sieboldiana var. sieboldiana (Hooker) Engler	No	Asparagaceae^k	M	60	2	P	11,564	11·8	23·6	47·2	384^aa	O	Agave sp.^ab	FC:PI
417	Hosta sieboldii P.O.(Paxton) Ingram	No	Asparagaceae^k	M	60	2	P	11,025	11·3	22·5	45·0	384^aa	O	Agave sp.^ab	FC:PI
418	Hosta tibae F.Maekawa	No	Asparagaceae^k	M	60	2	P	8,624	8·8	17·6	35·2	384^aa	O	Agave sp.^ab	FC:PI
419	Hosta tsushimensis N.Fujita	No	Asparagaceae^k	M	60	2	P	8,477	8·7	17·3	34·6	384^aa	O	Agave sp.^ab	FC:PI
420	Hosta ventricosa Stearn	No	Asparagaceae^k	M	120	4	P	19,208	19·6	39·2	78·4	384^aa	O	Agave sp.^ab	FC:PI
421	Hosta venusta F.Maekawa	No	Asparagaceae^k	M	60	2	P	8,477	8·7	17·3	34·6	384^aa	O	Agave sp.^ab	FC:PI
422	Hosta yingeri S.B.Jones	No	Asparagaceae^k	M	60	2	P	9,359	9·6	19·1	38·2	384^aa	O	Agave sp.^ab	FC:PI
423a	Hydrangea anomala D.Don ssp. petiolaris Sieb. & Zucc.	No	Hydrangeaceae	E	36	2	P	1,328	1·4	2·7	5·4	397	O	G^c	FC:EB
423b	Hydrangea anomala D.Don ssp. anomala McClint.	No	Hydrangeaceae	E	36	2	P	1,534	1·6	3·1	6·3	397	O	G^c	FC:EB
424	Hydrangea arborescens L.	No	Hydrangeaceae	E	36	2	P	1,132	1·2	2·3	4·6	397	O	G^c	FC:EB
425a	Hydrangea aspera Don. ssp. robusta McClint. (=H. longipes Franch.)	No	Hydrangeaceae	E	34	2	P	1,480	1·5	3·0	6·0	397	O	G^c	FC:EB
425b	Hydrangea aspera Don. ssp. sargentiana (Rehder) McClint.	No	Hydrangeaceae	E	34	2	P	1,529	1·6	3·1	6·2	397	O	G^c	FC:EB
425c	Hydrangea aspera Don. ssp. strigosa McClint.	No	Hydrangeaceae	E	34	2	P	1,700	1·7	3·5	6·9	397	O	G^c	FC:EB
425d	Hydrangea aspera Don. ssp. aspera McClint.	No	Hydrangeaceae	E	36	2	P	2,323	2·4	4·7	9·5	397	O	G^c	FC:EB
426	Hydrangea heteromalla D.Don	No	Hydrangeaceae	E	36	2	P	1,446	1·5	3·0	5·9	397	O	G^c	FC:EB
427	Hydrangea involucrata Sieb.	No	Hydrangeaceae	E	30	2	P	2,450	2·5	5·0	10·0	397	O	G^c	FC:EB
428a	Hydrangea macrophylla (Thunb.) Ser. ssp. serrata (Thunb.) Makino	No	Hydrangeaceae	E	36	2	P	1,887	1·9	3·9	7·7	397	O	G^c	FC:EB
428b	Hydrangea macrophylla (Thunb.) Ser. ssp. macrophylla McClint.	No	Hydrangeaceae	E	36	2	P	2,107	2·2	4·3	8·6	397	O	G^c	FC:EB
429	Hydrangea paniculata Sieb.	No	Hydrangeaceae	E	36	2	P	1,847	1·9	3·8	7·5	397	O	G^c	FC:EB
430	Hydrangea quercifolia Bartr.	No	Hydrangeaceae	E	36	2	P	956	1·0	2·0	3·9	397	O	G^c	FC:EB
431a	Hydrangea scandens (L.f.) ssp. scandens McClint.	No	Hydrangeaceae	E	36	2	P	1,803	1·8	3·7	7·4	397	O	G^c	FC:EB
431b	Hydrangea scandens (L.f.) ssp. luikinensis (Nakai) McClint.	No	Hydrangeaceae	E	36	2	P	1,872	1·9	3·8	7·6	397	O	G^c	FC:EB
432	Hydrangea seemannii Riley	No	Hydrangeaceae	E	36	2	P	1,024	1·0	2·1	4·2	397	O	G^c	FC:EB
433	Hypericum hirsutum^l	No	Hypericaceae	E	—ⁿ	—^p	P	147	0·2	0·3	0·6	465	O	J	Fe
434	Inga dulcis (Roxb.) Willd.	No	Leguminosae	E	—ⁿ	—^p	P	402	0·4	0·8	1·6	454	O	B^c	Fe
435a	Iris stenophylla Hausskn. ex Baker^h	No	Iridaceae	M	24	2	P	8,359	8·5	17·1	34·1	465	O	B	Fe
435b	Iris stenophylla Hausskn. ex Baker^h	No	Iridaceae	M	26	2	P	10,743	11·0	21·9	43·9	465	O	B	Fe
435c	Iris stenophylla Hausskn. ex Baker^h	No	Iridaceae	M	26	2	P	11,439	11·7	23·3	46·7	465	O	B	Fe
436	Ixiolirion ledebourii Fisch. & Mey.	No	Ixioliriaceae	M	c. 24	2	P	995	1·0	2·0	4·1	380	O	J	Fe
437	Ixora arborea Roxb. ex Sm.	No	Rubiaceae	E	—ⁿ	—^p	P	1,365	1·4	2·8	5·6	454	O	B^c	Fe
438	Jacquinia aristata Jacq.	No	Theophrastaceae	E	38	2	P	593	0·6	1·2	2·4	378	O	J	Fe
439	Kentranthus ruber Druce	No	Valerianaceae	E	32	4	P	407	0·4	0·8	1·7	379	O	G	Fe
440	Khaya senegalensis (Desr.) A.Juss.	No	Meliaceae	E	—ⁿ	—^p	P	853	0·9	1·7	3·5	454	O	B^c	Fe
441	Kigelia africana (Lam.) Benth.	No	Bignoniaceae	E	—ⁿ	—^p	P	1,700	1·7	3·5	6·9	454	O	B^c	Fe
442	Kirkia acuminata Oliver	No	Kirkiaceae	E	c. 30	2	P	319	0·3	0·7	1·3	378	O	J	Fe
443	Lagerstroemia tomentosa C.Presl. Presl.	No	Lythraceae	E	—ⁿ	—^p	P	965	1·0	2·0	3·9	454	O	B^c	Fe
444	Lantana camara L.	No	Verbenaceae	E	22°	—^p	P	2,697	2·8	5·5	11·0	454	O	B^c	Fe
445	Lapageria rosea Ruiz & Pav.	No	Philesiaceae	M	30 + 1B	2	P	6,644	6·8	13·6	27·1	380	O	G	Fe
446b	Lathyrus amphicarpos L.	—^m	Leguminosae	E	14	2	A	5,123	5·2	10·5	20·9	412	O	—^v	Fe
447e	Lathyrus annuus L.	—^m	Leguminosae	E	14	2	A	6,429	6·6	13·1	26·2	412	O	—^v	Fe
447f	Lathyrus annuus L.	—^m	Leguminosae	E	14	2	A	6,282	6·4	12·8	25·6	418	O	G^c	FC:PI
448j	Lathyrus aphaca L.	—^m	Leguminosae	E	14	2	A	4,528	4·6	9·2	18·5	418	O	C & E^c	FC:PI
449b	Lathyrus chloranthus Boiss.	—^m	Leguminosae	E	14	2	A	5,944	6·1	12·1	24·3	412	O	—^v	Fe
450f	Lathyrus cicera L.	—^m	Leguminosae	E	14	2	A	5,194	5·3	10·6	21·2	418	O	G^c	FC:PI
451e	Lathyrus clymenum L.	—^m	Leguminosae	E	14	2	A	4,297	4·4	8·8	17·5	418	O	C & E^c	FC:PI
451f	Lathyrus clymenum L.	—^m	Leguminosae	E	14	2	A	6,399	6·5	13·1	26·1	412	O	—^v	Fe
452	Lathyrus gmelinii Fritsch	—^m	Leguminosae	E	14	2	P	8,791	9·0	17·9	35·9	412	O	—^v	Fe
453b	Lathyrus grandiflorus Sibth. & Sm.	—^m	Leguminosae	E	14	2	P	8,724	8·9	17·8	35·6	412	O	—^v	Fe
454b	Lathyrus heterophyllus L.	—^m	Leguminosae	E	14	2	P	8,639	8·8	17·6	35·3	412	O	—^v	Fe
455	Lathyrus laevigatus (Waldst. & Kit.)	—^m	Leguminosae	E	14	2	P	11,008	11·2	22·5	44·9	412	O	—^v	Fe
456c	Lathyrus maritimus Bigelow	—^m	Leguminosae	E	14	2	P	6,777	6·9	13·8	27·7	412	O	—^v	Fe
457e	Lathyrus nissolia L.	—^m	Leguminosae	E	14	2	A	4,853	5·0	9·9	19·8	412	O	—^v	Fe
458e	Lathyrus ochrus (L.) DC	—^m	Leguminosae	E	14	2	A	4,542	4·6	9·3	18·5	418	O	C & E^c	FC:PI
459i	Lathyrus odoratus L.	No	Leguminosae^j	E	14°	2	A	8,134	8·3	16·6	33·2	457^bm	O	B^d	Fe
460f	Lathyrus sativus L.	—^m	Leguminosae	E	14	2	A	6,551	6·7	13·4	26·7	418	O	G^c	FC:PI
460g	Lathyrus sativus L.	—^m	Leguminosae	E	14	2	A	6,899	7·0	14·1	28·2	412	O	—^v	Fe
461f	Lathyrus sylvestris L.	—^m	Leguminosae	E	14	2	P	10,104	10·3	20·6	41·2	412	O	—^v	Fe
462g	Lathyrus tingitanus L.	—^m	Leguminosae	E	14	2	A	7,693	7·9	15·7	31·4	418	O	G^c	FC:PI
462h	Lathyrus tingitanus L.	—^m	Leguminosae	E	14	2	A	7,691	7·8	15·7	31·4	412	O	—^v	Fe
465	Lawsonia inermis L.	No	Lythraceae	E	c. 30-34	4	P	333	0·3	0·7	1·4	378	O	J	Fe
464	Lebronnecia kokioides Fosberg	—^m	Malvaceae	E	26	2	—^q	1,764	1·8	3·6	7·2	444^bd	O	G^b2	FC:PI
465	Lemna minor L.	No	Araceae	M	126	6	P	1,426	1·5	2·9	5·8	400	O	G	Fe
466	Leucaena collinsii Britton & Rose	No	Leguminosae	E	52, ?56°	2	P	529	0·5	1·1	2·2	425	O	—^at	FC:EB
467b	Leucaena confertiflora S.Zarate	No	Leguminosae	E	112°	4	P	828	0·8	1·7	3·4	425	O	—^at	FC:EB
468	Leucaena cuspidata Standley	No	Leguminosae	E	—ⁿ	—^p	P	686	0·7	1·4	2·8	425	O	—^at	FC:EB
469	Leucaena diversifolia (Schltdl. Benth.	No	Leguminosae	E	104	4	P	1,328	1·4	2·7	5·4	425	O	—^at	FC:EB
470c	Leucaena esculenta (Sesse & Moc. ex DC) Benth.	No	Leguminosae	E	52 (?56, ?112)	—^p	P	706	0·7	1·4	2·9	425	O	—^at	FC:EB
471	Leucaena greggii S.Watson	No	Leguminosae	E	56°	2	P	887	0·9	1·8	3·6	425	O	—^at	FC:EB
472	Leucaena involucrata S.Zarate	No	Leguminosae	E	—ⁿ	—^p	P	1,122	1·1	2·3	4·6	425	O	—^at	FC:EB
473c	Leucaena lanceolata S.Watson	No	Leguminosae	E	52	2	P	706	0·7	1·4	2·9	425	O	—^at	FC:EB
474	Leucaena lempirana C.E.Hughes	No	Leguminosae	E	—ⁿ	—^p	P	426	0·4	0·9	1·7	425	O	—^at	FC:EB
475b	Leucaena leucocephala (Lam.) De Wit.	No	Leguminosae	E	104	4	P	1,455	1·5	3·0	5·9	425	O	—^at	FC:EB
476	Leucaena macrophylla Benth.	No	Leguminosae	E	?52°	—^p	P	299	0·3	0·6	1·2	425	O	—^at	FC:EB
477	Leucaena magnifica (C.E.Hughes) C.E.Hughes	No	Leguminosae	E	?52°	—^p	P	500	0·5	1·0	2·0	425	O	—^at	FC:EB
478	Leucaena matudae (S.Zarate) C.E.Hughes	No	Leguminosae	E	?52°	—^p	P	519	0·5	1·1	2·1	425	O	—^at	FC:EB
479	Leucaena multicapitula Schery	No	Leguminosae	E	?52°	—^p	P	470	0·5	1·0	1·9	425	O	—^at	FC:EB
480	Leucaena pallida Britton & Rose	No	Leguminosae	E	104 (?110/112)°	4	P	774	0·8	1·6	3·2	425	O	—^at	FC:EB
481	Leucaena pueblana Britton & Rose	No	Leguminosae	E	—ⁿ	—^p	P	490	0·5	1·0	2·0	425	O	—^at	FC:EB
482	Leucaena pulverulenta (Schltdl.) Benth.	No	Leguminosae	E	56	2	P	686	0·7	1·4	2·8	425	O	—^at	FC:EB
483	Leucaena retusa Benth.	No	Leguminosae	E	56°	2	P	764	0·8	1·6	3·1	425	O	—^at	FC:EB
484	Leucaena salvadorensis Standley ex Britton & Rose	No	Leguminosae	E	?56°	—^p	P	887	0·9	1·8	3·6	425	O	—^at	FC:EB
485	Leucaena shannonii J.D.Smith	No	Leguminosae	E	52°	2	P	691	0·7	1·4	2·8	425	O	—^at	FC:EB
486	Leucaena trichandra (Zucc.) Urban	No	Leguminosae	E	52 (?56)°	2	P	764	0·8	1·6	3·1	425	O	—^at	FC:EB
487	Leucaena trichodes (Jacq.) Benth.	No	Leguminosae	E	52°	2	P	539	0·6	1·1	2·2	425	O	—^at	FC:EB
488	Limnanthes douglasii R. Br.	No	Limnanthaceae	E	10	2	A	1,362	1·4	2·8	5·6	379	O	J	Fe
489	Litsea glutinosa (Lour.) C.B. Robinson	No	Lauraceae	BA	48°	—^p	P	2,766	2·8	5·6	11·3	454	O	B^c	Fe
490c	Lolium perenne L.	No	Gramineae	M	14	2	P	2,773	2·8	5·7	11·3	417	O	Gallus^f	FC:PI
491	Loranthus europaeus^l	No	Loranthaceae^k	E	—ⁿ	—^p	—^q	8,085	8·3	16·5	33·0	457^bm	O	B^d	Fe
492b	Lupinus angustifolius L.	No	Leguminosae	E	38, 42, 44	—^p	A	752.^an	0·8^an	1·5^an	3·1^an	416	C	—^an	FC:PI
493	Lupinus atlanticus Gladst.	No	Leguminosae	E	38	—^p	A	1,458.^an	1·5^an	3·0^an	6·0^an	416	C	—^an	FC:PI
494	Lupinus cosentinii Guss.	No	Leguminosae	E	32	—^p	A	1,126.^an	1·1^an	2·3^an	4·6^an	416	C	—^an	FC:PI
495	Lupinus digitatus Forssk.	No	Leguminosae	E	36	—^p	A	1,286.^an	1·3^an	2·6^an	5·3^an	416	C	—^an	FC:PI
496	Lupinus micranthus Guss.	No	Leguminosae	E	52	—^p	A	461.^an	0·5^an	0·9^an	1·9^an	416	C	—^an	FC:PI
497	Lupinus palaestinus Boiss.	No	Leguminosae	E	42	—^p	A	1,201.^an	1·2^an	2·5^an	4·9^an	416	C	—^an	FC:PI
498c	Lupinus pilosus Murr.	No	Leguminosae	E	42	—^p	A	1,201.^an	1·2^an	2·5^an	4·9^an	416	C	—^an	FC:PI
499c	Luzula campestris (L.) DC.	No	Juncaceae	M	12	—^u	P	1,443	1·5	2·9	5·9	420	O	B^c	Fe
500c	Luzula elegans Guthnick	No	Juncaceae	M	6	—^u	P	1,512	1·5	3·1	6·2	420	O	B^c	Fe
501d	Luzula luzuloides (Lam.) Dandy & Wilmott	No	Juncaceae	M	12	—^u	P	1,722.^ap	1·8^ap	3·5^ap	7·0^ap	420	O	B^c	Fe
502d	Luzula nivea Lam. & DC.	No	Juncaceae	M	12	—^u	P	1,482.^ap	1·5^ap	3·0^ap	6·1^ap	420	O	B^c	Fe
503c	Luzula pedemontana Boiss & Reut.	No	Juncaceae	M	30	—^u	P	1,717	1·8	3·5	7·0	420	O	B^c	Fe
504c	Luzula pediformis DC.	No	Juncaceae	M	12	—^u	P	1,583	1·6	3·2	6·5	420	O	B^c	Fe
505	Luzula spicata DC.	No	Juncaceae	M	24	—^u	P	1,904	1·9	3·9	7·8	420	O	B^c	Fe
506	Luzula sudetica DC.	No	Juncaceae	M	48	—^u	P	1,686	1·7	3·4	6·9	420	O	B^c	Fe
507k	Lycopersicon esculentum Mill. cv. Gardener's Delight	No	Solanaceae	E	—ⁿ	—^p	A	980	1·0	2·0	4·0	382	O	H^c	FC:PI
508b	Malva sylvestris L.	—^m	Malvaceae	E	42°	6	P	1,470	1·5	3·0	6·0	444^bd	O	G^b2	FC:PI
509b	Mangifera indica L.	No	Anacardiaceae	E	40°	4	P	882	0·9	1·8	3·6	454	O	B^c	Fe
510	Matthiola incana^l	No	Cruciferae^j	E	—ⁿ	—^p	—^q	2,597	2·7	5·3	10·6	457^bm	O	B^d	Fe
511a	Melaleuca leucadendra L.	No	Myrtaceae	E	22°	—^p	P	1,110	1·1	2·3	4·5	449bf	O	B^c	Fe
512	Melampyrum arvense Linn.	No	Orobanchaceae	E	—ⁿ	—^p	P	8,073	8·2	16·5	33·0	465	O	G	FC:PI
513	Melia azedarach L.	No	Meliaceae	E	28°	—^p	P	421	0·4	0·9	1·7	454	O	B^c	Fe
514	Melianthus major L.	No	Melianthaceae	E	36, 38°	2 or 4	P	627	0·6	1·3	2·6	378	O	J	Fe
515	Mentha longifolia^l	No	Labiatae	E	24	2	P	385	0·4	0·8	1·6	465	O	J	Fe
516	Merrilliodendron megacarpum (Hemsl.) Sleum.	No	Icacinaceae	E	30	2	P	1,071	1·1	2·2	4·4	380	O	K	FC:PI
517	Mimusops elengi L.	No	Sapotaceae	E	24°	—^p	P	274	0·3	0·6	1·1	454	O	B^c	Fe
518a	Monstera deliciosa Liebm.	No	Araceae	M	50	—^p	P	9,384	9·6	19·2	38·3	411	O	B^c	Fe
519	Monstera obliqua Miq.	No	Araceae	M	44	—^p	P	8,822	9·0	18·0	36·0	411	O	B^c	Fe
520	Montinia caryophyllacea Thunb.	No	Montiniaceae	E	24	2	P	554	0·6	1·1	2·3	380	O	J	Fe
521d	Musa acuminata Colla ssp. banksii	—^m	Musaceae	M	22	2	P	600	0·6	1·2	2·5	402	O	Glycine^e	FC:PI
521e	Musa acuminata Colla ssp. siamea	—^m	Musaceae	M	22	2	P	618	0·6	1·3	2·5	402	O	Glycine^e	FC:PI
521f	Musa acuminata Colla ssp. banksii^h	No	Musaceae	M	22°	2	P	588	0·6	1·2	2·4	410	O	Petunia^e	FC:EB
521g	Musa acuminata Colla ssp. malaccensis Accession Selangor^h	No	Musaceae	M	22°	2	P	598	0·6	1·2	2·4	410	O	Petunia^e	FC:EB
521h	Musa acuminata Colla ssp. banksii^h	No	Musaceae	M	22°	2	P	637	0·7	1·3	2·6	410	O	Petunia^e	FC:EB
521i	Musa acuminata Colla ssp. siamea Accession Siam^h	No	Musaceae	M	22	2	P	652	0·7	1·3	2·7	410	O	Petunia^e	FC:EB
521j	Musa acuminata Colla ssp. truncata	—^m	Musaceae	M	22	2	P	626	0·6	1·3	2·6	421^aq	O	Glycine^e	FC:PI
521k	Musa acuminata Colla genotype PPC	—^m	Musaceae	M	22	2	P	610	0·6	1·2	2·5	421^aq	O	Glycine^e	FC:PI
522c	Musa balbisiana Colla	—^m	Musaceae	M	22	2	P	556	0·6	1·1	2·3	421	O	Glycine^e	FC:PI
522d	Musa balbisiana Colla	—^m	Musaceae	M	22	2	P	549	0·6	1·1	2·2	402	O	Glycine^e	FC:PI
522e	Musa balbisiana Colla Accession Tani (1120)	No	Musaceae	M	22°	2	P	568	0·6	1·2	2·3	410	O	Petunia^e	FC:EB
523	Musa ornata Roxb.	No	Musaceae	M	22°	2	P	603	0·6	1·2	2·5	410	O	Petunia^e	FC:EB
524	Musa violascens^l	—^m	Musaceae	M	20	2	P	691	0·7	1·4	2·8	421	O	Glycine^e	FC:PI
525	Muscari adilii M.B. Guner & H. Duman	No	Asparagaceae	M	18	2	P	2,550	2·6	5·2	10·4	465	O	G	Fe
526	Muscari mcbeathianum^l	No	Asparagaceae	M	18	2	P	2,989	3·1	6·1	12·2	465	O	B	Fe
527	Myoporum mauritianum A. DC.	No	Myoporaceae	E	72	8	P	1,904	1·9	3·9	7·8	379	O	G	Fe
528	Myrica gale Linn.	No	Myricaceae	E	16, 48, 80, 96°	—^p	P	409	0·4	0·8	1·7	379	O	J	Fe
529	Myrsine africana L.	No	Myrsinaceae	E	46	—^p	P	1,205	1·2	2·5	4·9	380	O	J	Fe
530	Narthecium ossifragum Huds.	No	Nartheciaceae	M	26°	2	P	404	0·4	0·8	1·7	380	O	J	FC:PI
531	Navarretia squarrosa Hook. & Arn.	No	Polemoniaceae	E	18	2	A	1,289	1·3	2·6	5·3	378	O	J	Fe
532	Nelumbo nucifera Gaertn.	No	Nelumbonaceae	E	16	2	P	238	0·2	0·5	1·0	379	O	J	Fe
533	Nemophila menziesii Hook. & Arn.	No	Boraginaceae	E	18	2	A	1,220	1·2	2·5	5·0	465	O	J	Fe
534	Nepenthes pervillei Blume	No	Nepenthaceae	E	—ⁿ	—^p	P	274	0·3	0·6	1·1	378	O	J	Fe
535k	Nicotiana tabacum L.	No	Solanaceae	E	48°	4	A	7,546	7·7	15·4	30·8	457^bm	O	B^d	Fe
536	Nyctanthes arbor-tristis L.	No	Oleaceae	E	44°	—^p	P	1,203	1·2	2·5	4·9	454	O	B^c	Fe
537	Odontites lutea^l	No	Orobanchaceae	E	—ⁿ	—^p	P	554	0·6	1·1	2·3	465	O	J	Fe
538	Odontites verna Dum.	No	Orobanchaceae	E	c. 18	2	P	559	0·6	1·1	2·3	465	O	J	Fe
539	Odontostomum hartwegii Torr.	No	Techophilaeaceae	M	20	2	P	2,506	2·6	5·1	10·2	380	O	G	FC:PI
540	Oenothera ammophila Focke	No	Onagraceae	E	14	2	BP	1,142	1·2	2·3	4·7	439	O	Sorghum^e	FC:PI
541	Oenothera biennis L.	No	Onagraceae	E	14	2	BP	1,196	1·2	2·4	4·9	439	O	Sorghum^e	FC:PI
542	Olea africana Mill.	No	Oleaceae	E	46	2	P	1,548	1·6	3·2	6·3	462	O	Sorghum^e	Fe
543	Olea cuspidata Wall.	No	Oleaceae	E	46	2	P	2,029	2·1	4·1	8·3	462	O	Sorghum^e	Fe
544a	Olea europaea L. cv. Dolce Agogia^h	No	Oleaceae	E	46	2	P	1,911	2·0	3·9	7·8	462	O	Sorghum^e	Fe
544b	Olea europaea L. cv. Pendolino^h	No	Oleaceae	E	46	2	P	2,283	2·3	4·7	9·3	462	O	Sorghum^e	Fe
545	Olea ferruginea Royale	No	Oleaceae	E	46	2	P	1,813	1·9	3·7	7·4	462	O	Sorghum^e	Fe
546	Olea indica Klein	No	Oleaceae	E	46	2	P	1,646	1·7	3·4	6·7	462	O	Sorghum^e	Fe
547r	Oryza sativa L. ssp. indica	No	Gramineae	M	24	2	A	466	0·5	1·0	1·9	450^bg	O	—	GS
547s	Oryza sativa L. ssp. indica cv. IR8	—^m	Gramineae	M	24	2	A	440	0·4	0·9	1·8	389	O	Arab.^e	FC:PI
547t	Oryza sativa L. ssp. japonica cv. Nipponbare	—^m	Gramineae	M	24	2	A	401	0·4	0·8	1·6	389	O	Arab.^e	FC:PI
547u	Oryza sativa L. ssp. japonica cv. Nipponbare	No	Gramineae	M	24	2	A	420	0·4	0·9	1·7	451^bh	O	—	GS
548	Oxalis boliviana Britton	No	Oxalidaceae	E	—ⁿ	—^p	P	469	0·5	1·0	1·9	456^bk	O	Gallus^f	FC:PI
549	Oxalis coralleoides R.Knuth	No	Oxalidaceae	E	16°	2	P	521	0·5	1·1	2·1	456^bk	O	Gallus^f	FC:PI
550	Oxalis cuzcensis R.Knuth	No	Oxalidaceae	E	16°	2	AP	575	0·6	1·2	2·3	456^bk	O	Gallus^f	FC:PI
551	Oxalis herrerae R.Knuth	No	Oxalidaceae	E	16°	2	P	454	0·5	0·9	1·9	456^bk	O	Gallus^f	FC:PI
552	Oxalis humbertii R.Knuth	No	Oxalidaceae	E	16°	2	P	510	0·5	1·0	2·1	456^bk	O	Gallus^f	FC:PI
553	Oxalis lotoides Kunth	No	Oxalidaceae	E	16°	2	P	432	0·4	0·9	1·8	456^bk	O	Gallus^f	FC:PI
554a	Oxalis lucumayensis R.Knuth ssp. lucumayensis	No	Oxalidaceae	E	16°	2	P	455	0·5	0·9	1·9	456^bk	O	Gallus^f	FC:PI
554b	Oxalis lucumayensis R.Knuth ssp. subiens Lourteig	No	Oxalidaceae	E	16°	2	P	483	0·5	1·0	2·0	456^bk	O	Gallus^f	FC:PI
555	Oxalis marcapatensis R.Knuth	No	Oxalidaceae	E	16°	2	P	440	0·4	0·9	1·8	456^bk	O	Gallus^f	FC:PI
556	Oxalis medicaginea Kunth	No	Oxalidaceae	E	16°	2	P	417	0·4	0·9	1·7	456^bk	O	Gallus^f	FC:PI
557	Oxalis megalorrhiza Jacquin	No	Oxalidaceae	E	14 or 18°	2	P	399	0·4	0·8	1·6	456^bk	O	Gallus^f	FC:PI
558	Oxalis mollis Kunth	No	Oxalidaceae	E	16°	2	P	437	0·4	0·9	1·8	456^bk	O	Gallus^f	FC:PI
559	Oxalis oulophora Lourteig	No	Oxalidaceae	E	16°	2	P	433	0·4	0·9	1·8	456^bk	O	Gallus^f	FC:PI
560	Oxalis paucartambensis R.Knuth	No	Oxalidaceae	E	16°	2	P	470	0·5	1·0	1·9	456^bk	O	Gallus^f	FC:PI
561a	Oxalis peduncularis Kunth^h	No	Oxalidaceae	E	16°	2	P	454	0·5	0·9	1·9	456^bk	O	Gallus^f	FC:PI
561b	Oxalis peduncularis Kunth var. pilosa^h	No	Oxalidaceae	E	16°	2	P	570	0·6	1·2	2·3	456^bk	O	Gallus^f	FC:PI
562	Oxalis petrophila R.Knuth	No	Oxalidaceae	E	16°	2	P	482	0·5	1·0	2·0	456^bk	O	Gallus^f	FC:PI
563	Oxalis phaeotricha Diels	No	Oxalidaceae	E	32°	4	P	820	0·8	1·7	3·3	456^bk	O	Gallus^f	FC:PI
564	Oxalis picchensis R.Knuth	No	Oxalidaceae	E	32°	4	P	821	0·8	1·7	3·4	456^bk	O	Gallus^f	FC:PI
565a	Oxalis ptychoclada Diels	No	Oxalidaceae	E	16°	2	P	434	0·4	0·9	1·8	456^bk	O	Gallus^f	FC:PI
565b	Oxalis ptychoclada Diels var. trichocarpa Lourteig	No	Oxalidaceae	E	16°	2	P	466	0·5	1·0	1·9	456^bk	O	Gallus^f	FC:PI
566	Oxalis san-miguelii R.Knuth	No	Oxalidaceae	E	16°	2	P	428	0·4	0·9	1·7	456^bk	O	Gallus^f	FC:PI
567	Oxalis sp. cfr. melilotoides Zuccarini^y	No	Oxalidaceae	E	16°	2	P	456	0·5	0·9	1·9	456^bk	O	Gallus^f	FC:PI
568	Oxalis sp. cfr. teneriensis R. Knuth^y	No	Oxalidaceae	E	16°	2	P	473	0·5	1·0	1·9	456^bk	O	Gallus^f	FC:PI
569a	Oxalis spiralis R. & P. ex G.Don^h	No	Oxalidaceae	E	16°	2	AP	520	0·5	1·1	2·1	456^bk	O	Gallus^f	FC:PI
569b	Oxalis spiralis R. & P. ex G.Don^h	No	Oxalidaceae	E	16°	2	AP	656	0·7	1·3	2·7	456^bk	O	Gallus^f	FC:PI
570	Oxalis tabaconasensis R.Knuth	No	Oxalidaceae	E	16°	2	P	515	0·5	1·1	2·1	456^bk	O	Gallus^f	FC:PI
571	Oxalis tuberosa Molina	No	Oxalidaceae	E	64°	8	P	1,722	1·8	3·5	7·0	456^bl	O	Gallus^f	FC:PI
572	Oxalis unduavensis (Rusby) R.Knuth	No	Oxalidaceae	E	16°	2	P	505	0·5	1·0	2·1	456^bk	O	Gallus^f	FC:PI
573	Oxalis urubambensis R.Knuth	No	Oxalidaceae	E	16°	2	P	431	0·4	0·9	1·8	456^bk	O	Gallus^f	FC:PI
574	Oxalis vulcanicola Donn. Sm.	No	Oxalidaceae	E	16°	2	P	434	0·4	0·9	1·8	456^bk	O	Gallus^f	FC:PI
575a	Paeonia caucasica (Schipcz.) Schipcz.^h	—^m	Paeoniaceae	E	10	2	P	15,592	15·9	31·8	63·6	459	C	—^bq	Fe
575b	Paeonia caucasica (Schipcz.) Schipcz.^h	—^m	Paeoniaceae	E	10	2	P	16,087	16·4	32·8	65·7	459	C	—^bq	Fe
576a	Paeonia daurica Andr.^h	—^m	Paeoniaceae	E	10	2	P	11,804	12·0	24·1	48·2	459	C	—^bq	Fe
576b	Paeonia daurica Andr.^h	—^m	Paeoniaceae	E	10	2	P	12,970	13·2	26·5	52·9	459	C	—^bq	Fe
577a	Paeonia lagodechiana Kem.-Nath.^h	—^m	Paeoniaceae	E	10	2	P	11,981	12·2	24·5	48·9	459	C	—^bq	Fe
577b	Paeonia lagodechiana Kem.-Nath.^h	—^m	Paeoniaceae	E	10	2	P	14,009	14·3	28·6	57·2	459	C	—^bq	Fe
578a	Paeonia macrophylla (Albov) Lomak.^h	—^m	Paeoniaceae	E	20	4	P	29,449	30·1	60·1	120·2	459	C	—^bq	Fe
578b	Paeonia macrophylla (Albov) Lomak.^h	—^m	Paeoniaceae	E	20	4	P	30,086	30·7	61·4	122·8	459	C	—^bq	Fe
579b	Paeonia mlokosewitschi Lomak.^h	—^m	Paeoniaceae	E	10	2	P	16,790	17·1	34·3	68·5	459	C	—^bq	Fe
579c	Paeonia mlokosewitschi Lomak.^h	—^m	Paeoniaceae	E	10	2	P	17,571	17·9	35·9	71·7	459	C	—^bq	Fe
580b	Paeonia officinalis L.^h	—^m	Paeoniaceae	E	20	4	P	25,995	26·5	53·1	106·1	459	C	—^bq	Fe
581a	Paeonia ruprechtiana Kem.-Nath.^h	—^m	Paeoniaceae	E	10	2	P	15,224	15·5	31·1	62·1	459	C	—^bq	Fe
581b	Paeonia ruprechtiana Kem.-Nath.^h	—^m	Paeoniaceae	E	10	2	P	17,331	17·7	35·4	70·7	459	C	—^bq	Fe
582a	Paeonia steveniana Kem.-Nath.^h	—^m	Paeoniaceae	E	20	4	P	27,955	28·5	57·1	114·1	459	C	—^bq	Fe
582b	Paeonia steveniana Kem.-Nath.^h	—^m	Paeoniaceae	E	20	4	P	29,939	30·6	61·1	122·2	459	C	—^bq	Fe
583b	Paeonia tenuifolia L.^h	—^m	Paeoniaceae	E	10	2	P	7,700	7·9	15·7	31·4	459	C	—^bq	Fe
583c	Paeonia tenuifolia L.^h	—^m	Paeoniaceae	E	10	2	P	11,598	11·8	23·7	47·3	459	C	—^bq	Fe
584a	Paeonia tomentosa (Lomak.) N.Busch^h	—^m	Paeoniaceae	E	20	4	P	25,260	25·8	51·6	103·1	459	C	—^bq	Fe
584b	Paeonia tomentosa (Lomak.) N.Busch^h	—^m	Paeoniaceae	E	20	4	P	27,391	28·0	55·9	111·8	459	C	—^bq	Fe
585a	Paeonia wittmanniana Hartwiss ex Lindl.^h	—^m	Paeoniaceae	E	20	4	P	27,514	28·1	56·2	112·3	459	C	—^bq	Fe
585b	Paeonia wittmanniana Hartwiss ex Lindl.^h	—^m	Paeoniaceae	E	20	4	P	31,042	31·7	63·4	126·7	459	C	—^bq	Fe
586d	Papaver rhoeas L.	No	Papaveraceae	E	14°	2	A	2,548	2·6	5·2	10·4	457^bm	O	B^d	Fe
587	Parmentiera cereifera Seem.	No	Bignoniaceae	E	—ⁿ	—^p	P	647	0·7	1·3	2·6	454	O	B^c	Fe
588	Paspalum notatum Flugge.	No	Gramineae	M	20	2	P	706	0·7	1·4	2·9	417	O	Gallus^f	FC:PI
589	Peltophorum pterocarpum (DC.) Baker ex K.Heyne	No	Leguminosae	E	26°	—^p	P	777	0·8	1·6	3·2	454	O	B^c	Fe
590a	Petroselinum crispum cv. Champion Moss Curled^l	No	Umbelliferae	E	—ⁿ	—^p	P	2,205	2·3	4·5	9·0	382	O	Lycopers.^c	FC:PI
	Petunia – Some taxa once included in Petunia are now included in Calibrachoa (see footnote ae)
591	Petunia alpicola L.B.Sm. & Downs	No	Solanaceae	E	18	2	P	1,450	1·5	3·0	5·9	387^ae	O	Gallus-398p	FC:PI
592	Petunia altiplana T.Ando & Hashim.	No	Solanaceae	E	14	2	P	1,274	1·3	2·6	5·2	387^ae	O	Gallus-398p	FC:PI
593b	Petunia axillaris (Lam.) Britton, Sterns & Poggenb. ssp. axillaris	No	Solanaceae	E	14	2	P	1,436	1·5	2·9	5·9	387^ae	O	Gallus-398p	FC:PI
593c	Petunia axillaris (Lam.) Britton, Sterns & Poggenb. ssp. subandina	No	Solanaceae	E	14°	2	P	1,465	1·5	3·0	6·0	387^ae	O	Gallus-398p	FC:PI
593d	Petunia axillaris (Lam.) Britton, Sterns & Poggenb. ssp. parodii	No	Solanaceae	E	14	2	P	1,470	1·5	3·0	6·0	387^ae	O	Gallus-398p	FC:PI
594	Petunia bajeensis T.Ando & Hashim.	No	Solanaceae	E	14°	2	P	1,450	1·5	3·0	5·9	387^ae	O	Gallus-398p	FC:PI
595	Petunia bonjardinensis T.Ando & Hashim.	No	Solanaceae	E	14	2	P	1,421	1·5	2·9	5·8	387^ae	O	Gallus-398p	FC:PI
596	Petunia exserta Stehmann.	No	Solanaceae	E	14°	2	P	1,539	1·6	3·1	6·3	387^ae	O	Gallus-398p	FC:PI
597	Petunia guarapuavensis T.Ando & Hashim.	No	Solanaceae	E	14	2	P	1,499	1·5	3·1	6·1	387^ae	O	Gallus-398p	FC:PI
598	Petunia helianthemoides Sendtn.	No	Solanaceae	E	18	2	P	1,436	1·5	2·9	5·9	387^ae	O	Gallus-398p	FC:PI
599e	Petunia hybrida Vilm. cv. Pearl Sky Blue	No	Solanaceae	E	14°	2	P	1,441	1·5	2·9	5·9	387^ae	O	Gallus-398p	FC:PI
600a	Petunia integrifolia (Hook.) Schinz & Thell. ssp. inflata (R.E.Fr.)	No	Solanaceae	E	14	2	A	1,333	1·4	2·7	5·4	387^ae	O	Gallus-398p	FC:PI
600b	Petunia integrifolia (Hook.) Schinz & Thell. ssp. integrifolia var. integrifolia	No	Solanaceae	E	14	2	P	1,436	1·5	2·9	5·9	387^ae	O	Gallus-398p	FC:PI
600c	Petunia integrifolia (Hook.) Schinz & Thell. ssp. integrifolia var. depauperata (R.E.Fr.)	No	Solanaceae	E	14	2	P	1,490	1·5	3·0	6·1	387^ae	O	Gallus-398p	FC:PI
601	Petunia interior T.Ando & Hashim.	No	Solanaceae	E	14	2	P	1,455	1·5	3·0	5·9	387^ae	O	Gallus-398p	FC:PI
602	Petunia kleinii L.B.Sm. & Downs	No	Solanaceae	E	18	2	P	1,436	1·5	2·9	5·9	387^ae	O	Gallus-398p	FC:PI
603	Petunia littoralis L.B. Sm. & Downs	No	Solanaceae	E	14	2	P	1,455	1·5	3·0	5·9	387^ae	O	Gallus-398p	FC:PI
604	Petunia mantiqueirensis T.Ando & Hashim.	No	Solanaceae	E	14°	2	P	1,524	1·6	3·1	6·2	387^ae	O	Gallus-398p	FC:PI
605	Petunia occidentalis R.E.Fr.	No	Solanaceae	E	14	2	A	1,362	1·4	2·8	5·6	387^ae	O	Gallus-398p	FC:PI
606	Petunia pubescens (Spreng.) R.E.Fr.	No	Solanaceae	E	18	2	P	1,446	1·5	3·0	5·9	387^ae	O	Gallus-398p	FC:PI
607	Petunia reitzii L.B.Sm. & Downs.	No	Solanaceae	E	14	2	P	1,406	1·4	2·9	5·7	387^ae	O	Gallus-398p	FC:PI
608	Petunia riograndensis T.Ando & Hashim.	No	Solanaceae	E	14°	2	P	1,460	1·5	3·0	6·0	387^ae	O	Gallus-398p	FC:PI
609	Petunia saxicola L.B.Sm. & Downs	No	Solanaceae	E	14	2	P	1,411	1·4	2·9	5·8	387^ae	O	Gallus-398p	FC:PI
610	Petunia scheideana L.B.Sm. & Downs	No	Solanaceae	E	14	2	P	1,436	1·5	2·9	5·9	387^ae	O	Gallus-398p	FC:PI
611	Petunia variabilisR.E.Fr.	No	Solanaceae	E	18	2	P	1,441	1·5	2·9	5·9	387^ae	O	Gallus-398p	FC:PI
612	Phalaenopsis amboinensis J. J. Smith	No	Orchidaceae	M	38	2	P	7,036	7·2	14·4	28·7	447	O	G^b2	FC:PI
613	Phalaenopsis aphrodite Rchb.f.	No	Orchidaceae	M	38	2	P	1,372	1·4	2·8	5·6	447	O	G^b2	FC:PI
614	Phalaenopsis bellina (Rchb. f.) Cristenson	No	Orchidaceae	M	38	2	P	7,365	7·5	15·0	30·1	447	O	G^b2	FC:PI
615	Phalaenopsis cornu-cervi (Breda) Bl & Rchb.f.	No	Orchidaceae	M	38	2	P	3,156	3·2	6·4	12·9	447	O	G^b2	FC:PI
616a	Phalaenopsis equestris (Schauer) Rchb.f.	No	Orchidaceae	M	38	2	P	1,651	1·7	3·4	6·7	447	O	G^b2	FC:PI
617	Phalaenopsis fasciata Rchb.f.	No	Orchidaceae	M	38	2	P	3,214	3·3	6·6	13·1	447	O	G^b2	FC:PI
618	Phalaenopsis gigantea J.J.Smith	No	Orchidaceae	M	38	2	P	2,587	2·6	5·3	10·6	447	O	G^b2	FC:PI
619a	Phalaenopsis lueddemanniana Rchb.f.	No	Orchidaceae	M	38	2	P	3,180	3·2	6·5	13·0	447	O	G^b2	FC:PI
620	Phalaenopsis mannii Rchb.f.	No	Orchidaceae	M	38	2	P	6,615	6·8	13·5	27·0	447	O	G^b2	FC:PI
621	Phalaenopsis mariae Burb. ex Warn. & Wms.	No	Orchidaceae	M	38	2	P	3,175	3·2	6·5	13·0	447	O	G^b2	FC:PI
622	Phalaenopsis micholitzii Rolfe	No	Orchidaceae	M	38	2	P	3,180	3·2	6·5	13·0	447	O	G^b2	FC:PI
623	Phalaenopsis modesta J.J.Smith	No	Orchidaceae	M	38	2	P	2,524	2·6	5·2	10·3	447	O	G^b2	FC:PI
624	Phalaenopsis parishii Rchb.f.	No	Orchidaceae	M	38	2	P	8,139	8·3	16·6	33·2	447	O	G^b2	FC:PI
625	Phalaenopsis pulchra (Rchb.f.) Sweet	No	Orchidaceae	M	38	2	P	3,121	3·2	6·4	12·7	447	O	G^b2	FC:PI
626	Phalaenopsis sanderiana Rchb.f.ⁱ	No	Orchidaceae	M	38	2	P	1,372	1·4	2·7	5·6	447	O	G^b2	FC:PI
627	Phalaenopsis stuartiana Rchb.f.	No	Orchidaceae	M	38	2	P	1,534	1·6	3·1	6·3	447	O	G^b2	FC:PI
628	Phalaenopsis sumatrana Korth. & Rchb.f.	No	Orchidaceae	M	38	2	P	3,244	3·3	6·6	13·2	447	O	G^b2	FC:PI
629	Phalaenopsis venosa Shim & Fowl.	No	Orchidaceae	M	38	2	P	4,665	4·8	9·5	19·0	447	O	G^b2	FC:PI
630c	Phaseolus acutifolius var. latifolius G.Freeman	—^m	Leguminosae	E	22°	2	A	794	0·8	1·6	3·2	390	O	Petunia^e	FC:DAPI
630d	Phaseolus acutifolius var. tenuifolius (Wood & Standl) A.Gray	—^m	Leguminosae	E	22°	2	A	799	0·8	1·6	3·3	390	O	Petunia^e	FC:DAPI
630e	Phaseolus acutifolius var. acutifolius A.Gray	—^m	Leguminosae	E	22°	2	A	862	0·9	1·8	3·5	390	O	Petunia^e	FC:DAPI
631	Phaseolus angustissimus A.Gray	—^m	Leguminosae	E	22°	2	—^q	647	0·7	1·3	2·6	390	O	Petunia^e	FC:DAPI
632e	Phaseolus coccineus L.	—^m	Leguminosae	E	22°	2	P	784	0·8	1·6	3·2	390	O	Petunia^e	FC:DAPI
632f	Phaseolus coccineus L. ssp. Purpurascens	—^m	Leguminosae	E	22°	2	P	794	0·8	1·6	3·2	390	O	Petunia^e	FC:DAPI
632g	Phaseolus coccineus L. ssp. coccineus cv. Hammond's Dwarf Scarlet^h	—^m	Leguminosae	E	22°	2	P	794	0·8	1·6	3·2	390	O	Petunia^e	FC:DAPI
632h	Phaseolus coccineus L. ssp. coccineus cv. Preisgewinner^h	—^m	Leguminosae	E	22°	2	P	809	0·8	1·7	3·3	390	O	Petunia^e	FC:DAPI
632i	Phaseolus coccineus L.	No	Leguminosae^j	E	22°	2	P	1,715	1·8	3·5	7·0	457^bm	O	B^d	Fe
633b	Phaseolus filiformis Benth.	—^m	Leguminosae	E	22°	2	P	691	0·7	1·4	2·8	390	O	Petunia^e	FC:DAPI
634b	Phaseolus glabellus Piper	—^m	Leguminosae	E	22°	2	P	1,024	1·0	2·1	4·2	390	O	Petunia^e	FC:DAPI
635	Phaseolus grayanus Wood. & Standl	—^m	Leguminosae	E	22°	2	—^q	931	1·0	1·9	3·8	390	O	Petunia^e	FC:DAPI
636b	Phaseolus hintonii Delgado	—^m	Leguminosae	E	22°	2	P	715	0·7	1·5	2·9	390	O	Petunia^e	FC:DAPI
637	Phaseolus leptostachys var. leptostachys Benth.	—^m	Leguminosae	E	22°	2	—^q	613	0·6	1·3	2·5	390	O	Petunia^e	FC:DAPI
638d	Phaseolus lunatus L. var. lunatus cv. Early Thorogreen^h	—^m	Leguminosae	E	22°	2	P	691	0·7	1·4	2·8	390	O	Petunia^e	FC:DAPI
638e	Phaseolus lunatus L. var. silvester Baudet	—^m	Leguminosae	E	22°	2	P	696	0·7	1·4	2·8	390	O	Petunia^e	FC:DAPI
638f	Phaseolus lunatus L. var. lunatus cv. Henderson Bush^h	—^m	Leguminosae	E	22°	2	P	701	0·7	1·4	2·9	390	O	Petunia^e	FC:DAPI
639b	Phaseolus marechalli Delgado	—^m	Leguminosae	E	22°	2	P	784	0·8	1·6	3·2	390	O	Petunia^e	FC:DAPI
640	Phaseolus micranthus Hook. & Arn.	—^m	Leguminosae	E	22°	2	—^q	588	0·6	1·2	2·4	390	O	Petunia^e	FC:DAPI
641	Phaseolus microcarpus Mart.	—^m	Leguminosae	E	22°	2	—^q	505	0·5	1·0	2·1	390	O	Petunia^e	FC:DAPI
642b	Phaseolus neglectus Herm.	—^m	Leguminosae	E	22°	2	—^q	941	1·0	1·9	3·8	390	O	Petunia^e	FC:DAPI
643	Phaseolus parviflorus G.Freytag	—^m	Leguminosae	E	22°	2	—^q	637	0·7	1·3	2·6	390	O	Petunia^e	FC:DAPI
644b	Phaseolus pluriflorus Marechal	—^m	Leguminosae	E	22°	2	—^q	1,068	1·1	2·2	4·4	390	O	Petunia^e	FC:DAPI
645b	Phaseolus polyanthus Greenm.	—^m	Leguminosae	E	22°	2	P	799	0·8	1·6	3·3	390	O	Petunia^e	FC:DAPI
646g	Phaseolus vulgaris L. cv. Kentucky Wonder^h	—^m	Leguminosae	E	22°	2	A	686	0·7	1·4	2·8	390	O	Petunia^e	FC:DAPI
646h	Phaseolus vulgaris L. var. aborigineus (Burk.) Baudet	—^m	Leguminosae	E	22°	2	A	720	0·7	1·5	2·9	390	O	Petunia^e	FC:DAPI
646i	Phaseolus vulgaris L. var. mexicanus	—^m	Leguminosae	E	22°	2	A	735	0·8	1·5	3·0	390	O	Petunia^e	FC:DAPI
646j	Phaseolus vulgaris L. cv. Sanilac	—^m	Leguminosae	E	22°	2	A	750	0·8	1·5	3·1	390	O	Petunia^e	FC:DAPI
646k	Phaseolus vulgaris L.	No	Leguminosae^j	E	22°	2	A	1,666.^bn	1·7^bn	3·4^bn	6·8^bn	457^bm	O	B^d	Fe
647b	Phaseolus xanthotrichus Piper var. xanthotrichus	—^m	Leguminosae	E	22°	2	P	662	0·7	1·4	2·7	390	O	Petunia^e	FC:DAPI
647c	Phaseolus xanthotrichus Piper	—^m	Leguminosae	E	22°	2	P	848	0·9	1·7	3·5	390	O	Petunia^e	FC:DAPI
648	Philodendron erubescens C.Koch & Bousche	No	Araceae	M	42	—^p	P	5,174	5·3	10·6	21·1	411	O	B^c	Fe
649	Philodendron selloum C.Koch	No	Araceae	M	36	—^p	P	4,895	5·0	10·0	20·0	411	O	B^c	Fe
650	Philodendron squamiferum Poepp. & Endl.	No	Araceae	M	30	—^p	P	4,557	4·7	9·3	18·6	411	O	B^c	Fe
651	Phormium tenax^l	No	Hemerocallidaceae	M	32	2	P	740	0·8	1·5	3·0	379	O	J	Fe
652	Pinguicula primuliflora C.E.Wood & Godfrey	No	Lentibulariaceae	E	22	2	P	669	0·7	1·4	2·7	378	O	J	Fe
653	Piptocalyx moorei Oliver	No	Trimeniaceae	BA	16°	2	P	4,001	4·1	8·2	16·3	381	O	G	FC:PI
654b	Pistia stratiotes L.	No	Araceae^k	M	28	2	P	250	0·3	0·5	1·0	400	O	G	Fe
655c	Pisum abyssinicum A.Braun^h	No	Leguminosae	E	14	2	A	4,371	4·5	8·9	17·8	458^bp	C^bp	G^c	FC:EB
655d	Pisum abyssinicum A.Braun^h	No	Leguminosae	E	14	2	A	4,751	4·8	9·7	19·4	458^bp	C^bp	G^c	FC:EB
656c	Pisum elatius Steven ex M.Bieb.^h	No	Leguminosae	E	14	2	A	4,231	4·3	8·6	17·3	458^bp	C^bp	G^c	FC:EB
656d	Pisum elatius Steven ex M.Bieb.^h	No	Leguminosae	E	14	2	A	4,978	5·1	10·2	20·3	458^bp	C^bp	G^c	FC:EB
657b	Pisum fulvum Sibth. & Smith	No	Leguminosae	E	14	2	A	4,716	4·8	9·6	19·3	458^bp	C^bp	G^c	FC:EB
658c	Pisum humile Boiss & Noë^h	No	Leguminosae	E	14	2	A	4,258	4·3	8·7	17·4	458^bp	C^bp	G^c	FC:EB
658d	Pisum humile Boiss & Noë^h	No	Leguminosae	E	14	2	A	4,809	4·9	9·8	19·6	458^bp	C^bp	G^c	FC:EB
659	Pittosporum tenuifolium Gaertn.	No	Pittosporaceae	E	24	2	P	453	0·5	0·9	1·9	379	O	G	Fe
660	Planchonella eerwah (F.M. Bailey) van Royen	No	Sapotaceae	E	c. 24	2	P	527	0·5	1·1	2·2	380	O	J	Fe
661	Plantago afra L.	No	Plantaginaceae	E	12	2	A	1,129	1·2	2·3	4·6	388	O	B^c	Fe
662	Plantago arenaria W. & K.	No	Plantaginaceae	E	12	2	A	1,115	1·1	2·3	4·6	388	O	B^c	Fe
663	Plantago coronopus L.	No	Plantaginaceae	E	10	2	AP	845	0·9	1·7	3·5	388	O	B^c	Fe
664	Plantago indica L.	No	Plantaginaceae	E	12	2	—^q	1,088	1·1	2·2	4·4	388	O	B^c	Fe
665b	Plantago lagopus L.	No	Plantaginaceae	E	12	2	—^q	1,046	1·1	2·1	4·3	388	O	B^c	Fe
666c	Plantago lanceolata L.	No	Plantaginaceae	E	12	2	P	1,299	1·3	2·7	5·3	388	O	B^c	Fe
667c	Plantago major L.	No	Plantaginaceae	E	12	2	P	867	0·9	1·8	3·5	388	O	B^c	Fe
668	Plantago psyllium L.	No	Plantaginaceae	E	12	2	—^q	1,142	1·2	2·3	4·7	388	O	B^c	Fe
669	Plantago serraria L.	No	Plantaginaceae	E	10	2	—^q	882	0·9	1·8	3·6	388	O	B^c	Fe
670	Plantago stepposa K.	No	Plantaginaceae	E	24	4	—^q	1,593	1·6	3·3	6·5	388	O	B^c	Fe
671	Platanus orientalis L.	No	Platanaceae	E	42	2	P	1,274	1·3	2·6	5·2	379	O	J	Fe
672	Poa pratensis L.	No	Gramineae	M	58-62	—^p	P	4,155	4·2	8·5	17·0	417	O	Gallus^f	FC:PI
673	Poncirus trifoliata (L.) Raf.	No	Rutaceae	E	18°	2	P	377	0·4	0·8	1·5	426	O	Gallus^f	FC:PI
674	Prosopis cineraria (L.) Druce	No	Leguminosae	E	52°	—^p	P	1,252	1·3	2·6	5·1	454	O	B^c	Fe
675	Protium serratum (Wall. ex Colebr.)	No	Burseraceae	E	—ⁿ	—^p	P	924	0·9	1·9	3·8	454	O	B^c	Fe
676	Pterospermum lanceifolium Roxb.	No	Malvaceae	E	38°	—^p	P	786	0·8	1·6	3·2	454	O	B^c	Fe
677	Pterostyrax psilophylla Diels ex Perkins	No	Styracaceae	E	24	2	P	867	0·9	1·8	3·5	380	O	J	Fe
678	Punica granatum L.	No	Sonneratiaceae	E	16°	2	P	706	0·7	1·4	2·9	454	O	B^c	Fe
679	Reseda luteola L.	No	Resedaceae	E	26	2	B	500	0·5	1·0	2·0	378	O	J	Fe
680	Rhapidophora montana Schott	No	Araceae	M	30	—^p	P	9,829	10·0	20·1	40·1	411	O	B^c	Fe
681	Rhapidophora peepla Schott	No	Araceae	M	18	—^p	P	8,984	9·2	18·3	36·7	411	O	B^c	Fe
682	Rhipogonum papuanum C.T. White	No	Rhipogonaceae	M	30	2	P	10,922	11·1	22·3	44·6	380	O	G	Fe
683	Rhodocoma gigantea (Kunth) H.P. Linder	No	Restionaceae	M	—ⁿ	—^p	P	728	0·7	1·5	3·0	380	O	J	FC:PI
684	Rhodohypoxis milloides (Baker) Hilliard & B.L. Burtt	No	Hypoxidaceae	M	24 + 1-2B	4	P	1,394	1·4	2·8	5·7	379	O	J	Fe
685c	Rhoeo discolor Hance	No	Commelinaceae	M	12°	2	P	7,987	8·2	16·3	32·6	457^bm	O	B^d	Fe
686	Rhoiacarpos capensis A. DC.	No	Santalaceae	E	—ⁿ	—^p	P	304	0·3	0·6	1·2	379	O	J	Fe
687	Rhynchosia cyanosperma Benth. Ex Baker	No	Leguminosae	E	22	2	B	2,727	2·8	5·6	11·1	443^bc	O	B^c	Fe
688	Rhynchosia minima (L.) DC.	No	Leguminosae	E	22	2	P	1,227	1·3	2·5	5·0	443^bc	O	B^c	Fe
689	Ribes glutinosum^l	No	Grossulariaceae	E	16	2	P	534	0·5	1·1	2·2	379	O	J	Fe
690	Roridula gorgonias Planch.	No	Roridulaceae	E	12	2	P	186	0·2	0·4	0·8	379	O	G	Fe
691c	Ruta graveolens L.	No	Rutaceae	E	—ⁿ	—^p	P	735	0·8	1·5	3·0	457^bm	O	B^d	Fe
692	Salix alba L.^h	—^m	Salicaceae	E	76^ad	4^ad	P	809	0·8	1·7	3·3	385^ac	O	Lycopers.^c	FC:PI
693	Salix atrocinerea Brot.^h	—^m	Salicaceae	E	76^ad	4^ad	P	804	0·8	1·6	3·3	385^ac	O	Lycopers.^c	FC:PI
694b	Salix caprea L.^h	—^m	Salicaceae	E	38^ad	2^ad	P	470	0·5	1·0	1·9	385^ac	O	Lycopers.^c	FC:PI
695	Salix cinerea L.^h	—^m	Salicaceae	E	76^ad	4^ad	P	828	0·8	1·7	3·4	385^ac	O	Lycopers.^c	FC:PI
696	Salix elaeagnos Scop.^h	—^m	Salicaceae	E	38^ad	2^ad	P	417	0·4	0·9	1·7	385^ac	O	Lycopers.^c	FC:PI
697	Salix fragilis L.^h	—^m	Salicaceae	E	76^ad	4^ad	P	843	0·9	1·7	3·4	385^ac	O	Lycopers.^c	FC:PI
698	Salix purpurea L.^h	—^m	Salicaceae	E	38^ad	2^ad	P	461	0·5	0·9	1·9	385^ac	O	Lycopers.^c	FC:PI
699	Salix pyrenaica Gouan^h	—^m	Salicaceae	E	38^ad	2^ad	P	470	0·5	1·0	1·9	385^ac	O	Lycopers.^c	FC:PI
700	Salix triandra L.^h	—^m	Salicaceae	E	38^ad	2^ad	P	387	0·4	0·8	1·6	385^ac	O	Lycopers.^c	FC:PI
701	Salix viminalis L.^h	—^m	Salicaceae	E	38^ad	2^ad	P	402	0·4	0·8	1·6	385^ac	O	Lycopers.^c	FC:PI
702	Salix viminalis L.^h	—^m	Salicaceae	E	76^ad	4^ad	P	794	0·8	1·6	3·2	385^ac	O	Lycopers.^c	FC:PI
703a	Sambucus nigra L.	No	Adoxaceae^k	E	36°	2	P	14,945.^bo	15·3^bo	30·5^bo	61·0^bo	457^bm	O	B^d	Fe
704	Santalum album L.	No	Santalaceae	E	20°	—^p	P	282	0·3	0·6	1·2	454	O	B^c	Fe
705b	Saxifraga granulata L. ssp. granulata	No	Saxifragaceae	E	22	—^p	P	662	0·7	1·4	2·7	453	O	G^c	FC:EB
706	Saxifraga granulata L.	No	Saxifragaceae	E	—^bj	—^p	P	1,122	1·1	2·3	4·6	453	O	G^c	FC:EB
707a	Saxifraga granulata L.	No	Saxifragaceae	E	52	—^p	P	2,332	2·4	4·8	9·5	453	O	G^c	FC:EB
707b	Saxifraga granulata L. ssp. fernandesii Redondo & Horjalesⁱ	No	Saxifragaceae	E	44-56	—^p	P	1,735	1·8	3·5	7·1	453^bi	O	G^c	FC:EB
708	Schisandra rubriflora^l	No	Schisandraceae	BA	—ⁿ	—^p	P	8,938	9·1	18·2	36·5	381	O	G	FC:PI
709	Schleichera oleosa (Lour.) Oken	No	Sapindaceae	E	32°	—^p	P	1,142	1·2	2·3	4·7	454	O	B^c	Fe
710b	Scilla indica (Roxb.) Baker (cytotype II)	No	Asparagaceae	M	30	—^p	P	3,504	3·6	7·2	14·3	422^ar	O	B^c	Fe
710c	Scilla indica (Roxb.) Baker (cytotype I)^h	No	Asparagaceae	M	30	—^p	P	5,701	5·8	11·6	23·3	422^ar	O	B^c	Fe
711	Scilla nervosa (Burch.) J.P.Jessop	No	Asparagaceae	M	38	—^p	P	3,964	4·0	8·1	16·2	422	O	B^c	Fe
712f	Scilla siberica Haw. in Andr.	No	Asparagaceae	M	12	2	P	30,135	30·8	61·5	123·0	422	O	B^c	Fe
713	Scilla talosii D.Tzanoudakis & Kypriotakis	No	Asparagaceae	M	c. 150	—^p	P	45,840	46·8	93·6	187·1	465	O	B	Fe
714c	Scilla vindobonensis Speta	No	Asparagaceae	M	18	3	P	—^t	—^t	17·9	35·7	422	O	B^c	Fe
715	Scindapsus pictus Hassk	No	Araceae	M	60	—^p	P	11,517	11·8	23·5	47·0	411	O	B^c	Fe
716	Sedum acre L.	No	Crassulaceae	E	—ⁿ	—^p	P	1,225	1·3	2·5	5·0	457^bm	O	B^d	Fe
717	Sedum album L.	—^m	Crassulaceae	E	34	2	P	142	0·1	0·3	0·6	398	O	—^w	—^r
718	Sedum forsterianum Sm.	—^m	Crassulaceae	E	24	2	P	451	0·5	0·9	1·8	398	O	—^w	—^r
719	Sedum montanum Song. & Perrier	—^m	Crassulaceae	E	34	2	P	515	0·5	1·1	2·1	398	O	—^w	—^r
720a	Sedum obtusifolium C.A.Meyer	—^m	Crassulaceae	E	12	2	P	206	0·2	0·4	0·8	398	O	—^w	—^r
720b	Sedum obtusifolium C.A.Meyer	—^m	Crassulaceae	E	12	2	P	206	0·2	0·4	0·8	399	O	B-723b	Fe
721	Sedum obtusifolium C.A.Meyer	—^m	Crassulaceae	E	30	5	P	—^t	—^t	1·7	3·4	399	O	B-723b	Fe
722	Sedum ochroleucum Chaix	—^m	Crassulaceae	E	34	2	P	446	0·5	0·9	1·8	398	O	—^w	—^r
723a	Sedum rupestre L. ssp. erectum	—^m	Crassulaceae	E	64	4	P	1,014	1·0	2·1	4·1	398	O	—^w	—^r
723b	Sedum rupestre L. ssp. rupestre	—^m	Crassulaceae	E	—ⁿ	—^p	P	2,244	2·3	4·6	9·2	399	O	B	Fe
724	Sedum sediforme (Jacq.) Pau	—^m	Crassulaceae	E	32	2	P	568	0·6	1·2	2·3	398	O	—^w	—^r
725	Sedum spurium Bieb.	—^m	Crassulaceae	E	28	4	P	1,735	1·8	3·5	7·1	399	O	B-723b	Fe
726	Sedum spurium Bieb.	—^m	Crassulaceae	E	42	6	P	2,764	2·8	5·6	11·3	399	O	B-723b	Fe
727a	Sedum stellatum L.	—^m	Crassulaceae	E	10	2	P	289	0·3	0·6	1·2	399	O	B-723b	Fe
727b	Sedum stellatum L.	—^m	Crassulaceae	E	10	2	P	289	0·3	0·6	1·2	398	O	—^w	—^r
728a	Sedum stoloniferum S.G.Gmelin	—^m	Crassulaceae	E	14	2	P	309	0·3	0·6	1·3	399	O	B-723b	Fe
728b	Sedum stoloniferum S.G.Gmelin	—^m	Crassulaceae	E	14	2	P	309	0·3	0·6	1·3	398	O	—^w	—^r
729	Senecio viscosus L.	No	Compositae^j	E	—ⁿ	—^p	A	1,519	1·6	3·1	6·2	457^bm	O	B^d	Fe
730	Sesamum alatum Thonn.	No	Pedaliaceae	E	26	2	A	1,651	1·7	3·4	6·7	446	O	G	Fe
731	Sesamum capense Burm.	No	Pedaliaceae	E	26	2	A	1,188	1·2	2·4	4·9	446	O	G	Fe
732	Sesamum indicum L.	No	Pedaliaceae	E	26	2	A	951	1·0	1·9	3·9	446	O	G	Fe
733	Sesamum laciniatum Klein.	No	Pedaliaceae	E	32	4	A	1,154	1·2	2·4	4·7	446	O	G	Fe
734	Sesamum latifolium Gillett.	No	Pedaliaceae	E	32	4	A	933	1·0	1·9	3·8	446	O	G	Fe
735	Sesamum mulayanum Nair.	No	Pedaliaceae	E	26	2	A	870	0·9	1·8	3·6	446	O	G	Fe
736	Sesamum occidentale Regel.	No	Pedaliaceae	E	64	8	A	1,551	1·6	3·2	6·3	446	O	G	Fe
737	Sesamum radiatum Schumach.	No	Pedaliaceae	E	64	8	A	1,306	1·3	2·7	5·3	446	O	G	Fe
738	Sesamum schinzianum Aschers.	No	Pedaliaceae	E	64	8	A	1,343	1·4	2·7	5·5	446	O	G	Fe
739	Sesamum triphyllum Welw. ex Aschers.	No	Pedaliaceae	E	c. 26	2	A	524	0·5	1·1	2·1	378	O	J	Fe
740b	Sesleria albicans Kit. ex Schult.^h	No	Gramineae^j	M	28°	4	P	4,748	4·8	9·7	19·4	428	O	Homo^f	FC:PI
740c	Sesleria albicans Kit. ex Schult.^h	No	Gramineae^j	M	28°	4	P	4,827	4·9	9·9	19·7	428	O	Homo^f	FC:PI
741	Severinia buxifolia (Poir.) Ten.	No	Rutaceae	E	18°	2	P	328	0·3	0·7	1·3	426	O	Gallus^f	FC:PI
742	Silene chalcedonica L.	No	Caryophyllaceae	E	24	2	P	3,229	3·3	6·6	13·2	437	O	Lycopers.^c	FC:PI
743c	Silene latifolia Poiret (female)	No	Caryophyllaceae	E	24	2	AP	2,808	2·9	5·7	11·5	437^az	O	Lycopers.^c	FC:PI
743d	Silene latifolia Poiret (male)	No	Caryophyllaceae	E	24	2	AP	2,867	2·9	5·9	11·7	437^az	O	Lycopers.^c	FC:PI
744	Silene pendula L.	No	Caryophyllaceae	E	24	2	A	1,152	1·2	2·4	4·7	437	O	H^c	FC:PI
745	Silene vulgaris (Moench) Garcke	No	Caryophyllaceae	E	24	2	P	1,103	1·1	2·3	4·5	437	O	H^c	FC:PI
746a	Simmondsia chinensis (Link) C.K.Schneid.	No	Simmondsiaceae	E	48-50	—^p	P	723	0·7	1·5	3·0	465	O	J	Fe
747	Spirodela polyrrhiza (L.) Schleid.	No	Araceae^k	M	80	4	P	292	0·3	0·6	1·2	400	O	G	Fe
748	Spirodela punctata (G.F.W.Meyer) Thompson	No	Araceae	M	46	2	P	363	0·4	0·7	1·5	400	O	G	Fe
749	Stemona tuberosa Lour.	No	Stemonaceae	M	—ⁿ	—^p	P	715	0·7	1·5	2·9	379	O	J	Fe
750	Stenotaphrum secundatum (Walt.) Kuntze.	No	Gramineae	M	18	2	P	529	0·5	1·1	2·2	417	O	Gallus^f	FC:PI
751	Strelitzia nicolai Regel & C. Koch	No	Strelitziaceae	M	14, 22°	2	P	566	0·6	1·2	2·3	379	O	J	Fe
752	Streptocarpus cyaneus S. Moore	No	Gesneriaceae	E	c. 30	2 or 4	P	662	0·7	1·4	2·7	378	O	J	Fe
753	Stylidium adnatum R. Br.	No	Stylidiaceae	E	30	2	P	1,495	1·5	3·1	6·1	378	O	J	Fe
754	Stylobasium spathulatum Desf.	No	Surianaceae	E	30	2	P	1,284	1·3	2·6	5·2	378	O	J	Fe
755	Syngonium albo-lineatum Bull	No	Araceae	M	22	—^p	AP	4,628	4·7	9·4	18·9	411	O	B^c	Fe
756	Syngonium podophyllum Schott	No	Araceae	M	24	—^p	AP	4,755	4·9	9·7	19·4	411	O	B^c	Fe
757	Tabebuia argentea (Bureau & K.Schum.) Britton	No	Bignoniaceae	E	—ⁿ	—^p	P	782	0·8	1·6	3·2	454	O	B^c	Fe
758	Tamarindus indica L.	No	Leguminosae	E	26°	—^p	P	821	0·8	1·7	3·4	454	O	B^c	Fe
759	Tecoma stans (L.) Juss. ex Kunth	No	Bignoniaceae	E	36°	—^p	P	593	0·6	1·2	2·4	454	O	B^c	Fe
760	Thespesia lampas (Cavanilles) Dalzell ex Dalzell & Gibson	—^m	Malvaceae	E	26°	2	—^q	1,568	1·6	3·2	6·4	444^bd	O	G^b2	FC:PI
761a	Thespesia populnea (L.) Solander ex Correa	—^m	Malvaceae	E	26	2	P	4,018	4·1	8·2	16·4	444^bd	O	G^b2	FC:PI
761b	Thespesia populnea (L.) Solander ex Correa	No	Malvaceae	E	26°	2	P	3,011	3·1	6·1	13·3	454	O	B^c	Fe
762	Thespesia thespesioides (R.Brown ex Bentham) Fryxell	—^m	Malvaceae	E	26	2	—^q	1,568	1·6	3·2	6·4	444^bd	O	G^b2	FC:PI
763	Triteleia laxa Benth.	No	Asparagaceae	M	28°	4	P	10,435	10·6	21·3	42·6	380	O	B	FC:PI
764	Trochodendron aralioides Siebold & Zucc.	No	Trochodendraceae	E	38	2	P	1,872	1·9	3·8	7·6	380	O	Lycopers.^c	FC:PI
765	Typhonium cuspidatum Decne.	No	Araceae	M	16	—^p	A	5,064	5·2	10·3	20·7	411	O	B^c	Fe
766	Typhonium trilobatum Schott	No	Araceae	M	40	—^p	A	6,453	6·6	13·2	26·3	411	O	B^c	Fe
767	Vicia canescens Lab.	No	Leguminosae	E	10	2	P	3,058	3·1	6·2	12·5	409	O	B^c	Fe
768a	Vicia cracca L. ssp. tenuifolia	No	Leguminosae	E	14	2	P	5,797	5·9	11·8	23·7	409	O	B^c	Fe
768b	Vicia cracca L. ssp. cracca	No	Leguminosae	E	14	2	P	6,409.^al	6·5^al	13·1^al	26·2^al	409	O	B^c	Fe
769a	Vicia epetiolaris Burk.^h	No	Leguminosae^j	E	14	2	A	4,067	4·2	8·3	16·6	415	O	B	Fe
769b	Vicia epetiolaris Burk.^h	No	Leguminosae^j	E	14	2	A	4,533	4·6	9·3	18·5	415	O	B	Fe
770b	Vicia eristalioides Maxted	No	Leguminosae	E	14	2	A	9,452	9·6	19·3	38·6	406	O	Vicia narb.^e	Fe
771r	Vicia faba L.	No	Leguminosae^j	E	12°	2	A	12,740	13·0	26·0	52·0	457^bm	O	B^d	Fe
771s	Vicia faba L. var. equina	No	Leguminosae	E	12	2	A	14,377	14·7	29·3	58·7	408	O	C^d	Fe
771t	Vicia faba L. ‘Futura RZ’	—^m	Leguminosae	E	—ⁿ	—^p	A	12,985	13·3	26·5	53·0	384^aa	O	Homo^f	FC:PI
772a	Vicia galilaea Plitm. & Zoh.	No	Leguminosae	E	14	2	A	6,392	6·5	13·0	26·1	408	O	C^d	Fe
772b	Vicia galilaea Plitm. & Zoh.	No	Leguminosae	E	14	2	A	7,904	8·1	16·1	32·3	409	O	B^c	Fe
773d	Vicia graminea Sm.ⁱ	No	Leguminosae^j	E	14	2	AB	4,969	5·1	10·1	20·3	415	O	B	Fe
774b	Vicia hyaeniscyamus Mout.	No	Leguminosae	E	14	2	A	7,654	7·8	15·6	31·2	408	O	C^d	Fe
775e	Vicia hybrida L.	No	Leguminosae	E	12	2	A	8,306	8·5	17·0	33·9	409	O	B^c	Fe
776e	Vicia hyrcanica Fisch. & Mey.	No	Leguminosae	E	12	2	A	7,634	7·8	15·6	31·2	409	O	B^c	Fe
777c	Vicia johannis Tamamsch.	No	Leguminosae	E	14	2	A	6,145	6·3	12·5	25·1	408	O	C^d	Fe
778b	Vicia kalakhensis Khattab, Maxted & Bisby	No	Leguminosae	E	14	2	A	10,344	10·6	21·1	42·2	406	O	Vicia narb.^e	Fe
779	Vicia macrograminea Burk.	No	Leguminosae^j	E	14	2	BP	5,821	5·9	11·9	23·8	415	O	B	Fe
780d	Vicia melanops Sibth. & Sm.	No	Leguminosae	E	10	2	A	6,764.^ak	6·9^ak	13·8^ak	27·6^ak	408	O	C^d	Fe
781	Vicia nana Vog.	No	Leguminosae^j	E	14	2	A	4,376	4·5	8·9	17·9	415	O	B	Fe
782a	Vicia pampicola Burk.^h	No	Leguminosae^j	E	14	2	A	4,640	4·7	9·5	18·9	415	O	B	Fe
782b	Vicia pampicola Burk.^h	No	Leguminosae^j	E	14	2	A	5,223	5·3	10·7	21·3	415	O	B	Fe
783d	Vicia peregrina L.	No	Leguminosae	E	14	2	A	9,540	9·7	19·5	38·9	409	O	B^c	Fe
784d	Vicia pisiformis L.^h	No	Leguminosae	E	12	2	P	6,223	6·4	12·7	25·4	407	O	C^d	Fe
784e	Vicia pisiformis L.^h	No	Leguminosae	E	12	2	P	8,038	8·2	16·4	32·8	407	O	C^d	Fe
785v	Vicia sativa L. line 20·1ⁱ	No	Leguminosae	E	12	2	A	1,882	1·9	3·8	7·7	409	O	B^c	Fe
785w	Vicia sativa L. line 31ⁱ	No	Leguminosae	E	12	2	A	2,445	2·5	5·0	10·0	409	O	B^c	Fe
785x	Vicia sativa L. ssp. amphicarpa	No	Leguminosae	E	12	2	A	2,102	2·1	4·3	8·6	409	O	B^c	Fe
785y	Vicia sativa L. ssp. nigra var. nigra	No	Leguminosae	E	12	2	A	2,337	2·4	4·8	9·5	409	O	B^c	Fe
786e	Vicia sepium L.	No	Leguminosae	E	14	2	P	4,719	4·8	9·6	19·3	409	O	B^c	Fe
787c	Vicia serratifolia Jacq.	No	Leguminosae	E	14	2	A	9,700	9·9	19·8	39·6	405	O	C^c	Fe
788d	Viscum album L.	No	Loranthaceae^k	E	20°	2	P	52,430	53·5	107·0	214·0	457^bm	O	B^d	Fe
789	Vitex negundo L.	No	Lamiaceae	E	34°	—^p	P	1,590	1·6	3·2	6·5	454	O	B^c	Fe
790	Vitex pinnata L.	No	Lamiaceae	E	—ⁿ	—^p	P	1,411	1·4	2·9	5·8	454	O	B^c	Fe
791	Voacanga grandifolia (Miq.) Rolfe	No	Apocynaceae	E	—ⁿ	—^p	P	358	0·4	0·7	1·5	454	O	B^c	Fe
792	Wolffia arrhiza (L.) Horkel ex Wimmer	No	Araceae	M	42	2	P	1,600	1·6	3·3	6·5	400	O	G	Fe
793	Wolffiella oblonga (Phil.) Hegelm.	No	Araceae	M	42	2	P	742	0·8	1·5	3·0	400	O	G	Fe
794	Xanthorrhoea preisii Endl.	No	Xanthorrhoeaceae	M	22	2	P	1,014	1·0	2·1	4·1	380	O	J	Fe
795	Xanthosoma sagittifolium (L.) Schott	No	Araceae	M	38	—^p	P	8,609	8·8	17·6	35·1	411	O	B^c	Fe
796	Xeronema callistemon W.R.B. Oliv.	No	Xeronemataceae	M	34	2 or 4	P	3,210	3·3	6·6	13·1	380	O	G	FC:PI
797	Xerophyta humilis Th. Dur. & Schinz.	No	Velloziaceae	M	48°	4 or 8	P	532	0·5	1·1	2·2	378	O	J	Fe
798	Ximenia americana Linn.	No	Olacaceae	E	26	2	P	1,595	1·6	3·3	6·5	379	O	J	Fe
799	Xiphidium caeruleum Aubl. var. caeruleum	No	Haemodoraceae	M	38°	2	P	767	0·8	1·6	3·1	378	O	J	Fe
800	Xyris gracilis ssp. gracilis^l	No	Xyridaceae	M	26°	2	P	6,867	7·0	14·0	28·0	380	O	B	FC:PI
801bx	Zea mays ssp. mays L. line opaque 2^h	No	Gramineae^j	M	20	4	A	3,262	3·3	6·7	13·3	392	O	B	Fe
801by	Zea mays ssp. mays L. race Altiplano^h	No	Gramineae^j	M	20	4	A	2,454	2·5	5·0	10·0	392^ag	O	B-801bx	Fe
801bz	Zea mays ssp. mays L. race Blanco y ocho rayas^h	No	Gramineae^j	M	20	4	A	3,311	3·4	6·8	13·5	392^ag	O	B-801bx	Fe
801ca	Zea mays L.	No	Gramineae^j	M	20°	4	A	3,283	3·4	6·7	13·4	457^bm	O	B^d	Fe
802	Zizyphus glabrata Heyne	No	Rhamnaceae	E	24°	—^p	P	1,517	1·5	3·1	6·2	454	O	B^c	Fe
803	Zostera marina^l	No	Zosteraceae	M	12°	2	P	309	0·3	0·6	1·3	380	O	J	Fe
804	Zoysia japonica Steud.	No	Gramineae	M	40	4	P	421	0·4	0·9	1·7	417	O	Gallus^f	FC:PI

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^‡

Chromosome number.

^§

E, ephemeral; A, annual; B, biennial; P, perennial.

^†

O, original value; C, calibrated value

The standard species used to calibrate the present amount.

^††

Fe, Feulgen microdensitometry; FC, flow cytometry using one of the following fluorochromes: PI, propidium iodide; DAPI, 4′, 6-diamidinophenylindole; EB, ethidium bromide; MI, mithramycin; HO,Hoechst 33258; GS, genome sequencing; CIA, computer image analysis; RK, reassociation kinetics.

E, eudicot; M, monocot; BA, basal angiosperm.

PROGRESS

Research on DNA C-values in angiosperms is unique in having been subject to detailed analyses of its quantity and quality over a long period (Bennett and Leitch, 1995). The importance of identifying gaps in our knowledge concerning this key biodiversity character, of recommending targets for new work to fill them by collaboration of international partners, and of monitoring progress to ensure that any shortfall is recognized, was confirmed by the first plant genome size workshop in 1997 (http://www.kew.org/cval/conference.html#outline, Bennett et al., 2000) and reviewed by participants at the second plant genome size workshop in 2003. Thus, what follows is mainly a summary of the overall progress for angiosperms against key targets set in 1997 for the following quinquennium (1998–2002). However, it also notes meaningful statistics for the data included in the Appendix table, or known to us from personal communications made after the Appendix table was closed.

In 1997 C-values for 2802 species (approximately 1 %) of angiosperm species had been estimated in the previous 40 years. The 1997 workshop concluded that the ideal of a C-value for all taxa was unrealistic, but long-term, estimates for 10–20 % of angiosperms seemed both ultimately achievable and adequate for all conceivable uses provided they were carefully targeted to be representative of the various taxonomic groups, geographical regions, and life forms in the global flora. So the first recommended target was to estimate first C-values for the next 1 % of angiosperm species (i.e. another 2500 species) by 2003. Many saw this goal as aspirational, as achieving it would mean estimating as many C-values in five years as in the past 40. Others thought that new technology (e.g. flow cytometry) would make it easy to achieve.

Improved systematic representation (species and families)

(i) First estimates for species

In September 1997 the Angiosperm DNA C-values database contained data for 2802 species. By September 2003 C-values were listed for 4119 species, including 689 first values for species listed in Bennett et al. (2000) and 628 such values for species in the Appendix table. Progress toward the first target in the five year period (1998–2002) considerably exceeded the average of ∼110 first values for species per annum in the early 1990s. Clearly, the 1997 workshop stimulated an increase in the total output of first C-values for species to its highest level for any five-year period (almost 200 per annum; Fig. 1A). Moreover, the proportion of newly published C-values that were also first estimates for species, which had previously fallen (Bennett et al., 2000), rose as a result of recent targeting and averaged 72·5 % for values published since 1997 (Fig. 1B). Nevertheless, the total number of published first C-values for species (1032) listed since 1997 was only 41 % of the 1997 target of approx. 2500.

Fig. 1. — (A) Mean number per year of total (open symbols) and ‘first’ (closed symbols) DNA C-value estimates communicated in ten successive 5-year periods and the 3-year period 2000–2002, between 1950 and 2002. Based on analysis of data listed in the present Appendix table, and the Angiosperm DNA C-values database (release 4.0, January 2003). (B) Percentage of C-value estimates published or communicated during 1965–2002 that are first values for species listed in the present Appendix table and the Angiosperm DNA C-values database (release 4.0, January 2003).

The real total of first C-values for angiosperms estimated after 1996 but unpublished by 2003 was much higher, but is difficult to determine exactly. For example, several hundred values were measured by Ben Zonneveld (pers. comm.) using flow cytometry but not published. Listing for the Appendix table closed in August 2002, ready for the workshop; however, we saw 158 first C-values published by other authors later in 2002, and 22 such values were estimated at RBG, Kew. Adding these data to those listed in our compilations suggests that the total number of first C-values for species estimated in 1997–2002 was probably at least 1700 and hence not less than approx. 66 % of the target set in 1997. Analysis shows that this was achieved by international collaboration involving at least 18 research groups in ten countries. Whilst a target of 2500 was aspirational, it seems attainable as a future five-year goal. However, at the present rate achieving 20 % species representation would take 100 years, so an ultimate goal of 10 % (approx. 25 000 angiosperm species) is more sensible.

(ii) First estimates for families

The 1997 workshop noted that a first C-value was available for only 30 % of angiosperm families recognized at that time. Thus, a second recommended target was ‘To obtain at least one C-value estimate for a species in all angiosperm families’. Monitoring first C-values for species listed in Bennett et al. (2000) showed that progress towards this goal was initially very slow. Indeed, ‘since 1997 first C-values had been listed for 691 angiosperm species, but only 12 (1·7 %) were also first estimates for families’. Work to correct this began at RBG, Kew in 1999. In 2001 two papers reported first C-values for 50 families (Hanson et al., 2001a, b), and 30 more followed, including five basal angiosperm families (Leitch and Hanson, 2002; Hanson et al., 2003), all included in the present Appendix table.

Analysis of listed data for 4119 species shows that a first published value is available for at least 217 of the 457 angiosperm families currently recognized by the Angiosperm Phylogeny Group (APG) (APG II, 2003). Together with first estimates for 11 unlisted families (Hanson, RBG, Kew, pers. comm.; Koce et al., 2003) measured or seen after listing for the Appendix table was closed, the total is 228. Thus, since 1997 (after losses owing to new familial circumscriptions—APG II, 2003; Hanson et al., 2003) first values for at least 85 such families have been measured, so good progress has been made. However, the proportion of families represented rose only from 30 % to 49·9 % (Fig. 2), which is less than one third of the target (100 %) set in 1997. Major factors limiting progress were discussed previously (Hanson et al., 2003). Unlike progress towards the species target, which involved many research groups, movement towards the goal for families since 1997 has depended mainly on work by one institution, as RBG Kew estimated 65 of the 74 (87 %) first values for families listed in the Appendix table.

Fig. 2. — Cumulative percentage of angiosperm families recognized by the Angiosperm Phylogeny Group (APG) (APGII, 2003) with a first C-value represented in the present Appendix table, the Angiosperm DNA C-values database (release 4.0, January 2003), plus eleven known to the present authors in September 2003.

The Plant Genome Size Workshop in 2003 confirmed that global capacity for estimating DNA C-values (determined by available equipment, funding and trained operators) remains very limited. Consequently, any increased focus on targets for other plant groups (e.g. bryophytes, pteridophytes, gymnosperms—reviewed in Leitch and Bennett, 2002b) inevitably reduces progress to improve representation for angiosperm targets, a problem discussed below. However, it should not detract from the highly successful progress to make C-values more representative of the global flora described above.

PROBLEMS

Geographical representation and distribution

We first noted the need to improve geographical representation for angiosperm C-values in Bennett and Leitch (1995). This was confirmed by the 1997 plant genome size workshop, although no specific regional targets were recommended. Perhaps, in consequence, progress in this area has never been monitored in detail, although we have been at pains to advertise the problem in a general way and to provoke action to rectify it in particular regions, such as southern Africa (Leitch and Bennett, 2002a).

There are two critical concerns regarding the geographical distribution of angiosperm C-value work. (i) The first concerns the small number of publications with original C-values by first authors in many regions (Table 1). This reflects a serious imbalance between the geographical distribution of research scientists working on genome size and of taxa whose C-values are unknown. Bennett et al. (2000) noted that ‘Africa remains an unexplored continent’ and that ‘Whereas six out of 377 original sources have first authors with addresses in Africa, still none has an angiosperm C-value estimated in Africa, as all six reported work done in Europe or the USA.’ Analysis of the 88 original sources in the present work shows no improvement, as the number of original sources from Africa (2), China (2), and South America (2) remains low (Table 1). (ii) The second concerns the small number of first C-values by any authors for species endemic to several large geographical regions. With some exceptions, the sample is still dominated by crops and their wild relatives, model species grown for experimental use, and other species growing near laboratories in temperate regions, mainly in Western Europe and North America. Analysis of data in the Appendix table shows that none presented data for other taxa endemic to China, Japan, Brazil, Mexico or Central Africa. Similarly, although island floras are known to be rich in endemics, no original source has reported C-values for any large islands such as Borneo, New Guinea or Madagascar, where 80 % of the 12000 described plant species are endemic (Robinson, 2004).

Table 1.

The number and percentage (in brackets) of original references with first authors from various geographical areas among the total of 465 sources contributing to the present Appendix table and the six lists of angiosperm DNA amounts previously compiled for reference purposes that were pooled in the Angiosperm DNA C-values database (release 4.0, January 2003)

	DNA C-value compilation
Area	1976¹	1982²	1991³	1995⁴	1997⁵	2000⁶	Present Appendix	Total
Europe	34 (63·0)	38 (71·7)	30 (53·6)	43 (40·6)	18 (48·6)	38 (51·4)	54 (61·4)	255 (54·8)
UK	28 (51·9)	13 (24·5)	22 (39·3)	23 (21·7)	5 (13·5)	8 (10·8)	10 (11·4)	109 (23·4)
North America	14 (25·9)	11 (20·8)	16 (28·6)	19 (17·9)	5 (13·5)	11 (14·9)	13 (14·8)	89 (19·1)
South and Meso America	0 (0·0)	0 (0·0)	3 (5·4)	9 (8·5)	1 (2·7)	6 (8·1)	2 (2·3)	21 (4·5)
Africa	1 (1·9)	0 (0·0)	0 (0·0)	1 (0·9)	2 (5·4)	1 (1·4)	2 (2·3)	7 (1·5)
Asia	1 (1·9)	3 (5·7)	4 (7·1)	30 (28·3)	11 (29·7)	8 (10·8)	17 (19·3)	74 (15·9)
India	1 (1·9)	1 (1·9)	2 (3·6)	28 (26·4)	11 (29·7)	4 (5·4)	11 (12·5)	58 (12·5)
China	0 (0·0)	0 (0·0)	0 (0·0)	0 (0·0)	0 (0·0)	0 (0·0)	2 (2·3)	2 (0·4)
Australasia	4 (7·4)	1 (1·9)	3 (5·4)	4 (3·8)	0 (0·0)	7 (9·5)	0 (0·0)	19 (4·1)
Australia	4 (7·4)	1 (1·9)	3 (5·4)	1 (0·9)	0 (0·0)	1 (1·4)	0 (0·0)	10 (2·2)
Total	54	53	56	106	37	71	88	465 (100)

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Bennett and Smith (1976)

Bennett et al. (1982)

Bennett and Smith (1991)

⁴

Bennett and Leitch (1995)

⁵

Bennett and Leitch (1997)

⁶

Bennett et al. (2000).

Plant life form

There is also a need for the overall sample to represent better the full range of plant types and life forms. We previously identified several associations and life forms as being poorly represented in the database (Bennett and Leitch, 1995), yet taxa from bog, fen, tundra, alpine and desert environments, and halophytic, insectivorous, parasitic, saprophytic and epiphytic species and their associated taxa are all still under-represented.

Solving this problem needs a proactive approach, as recent experience with first C-values for angiosperm families shows. First, a target must be set for each gap. Second, monitoring newly published data against targets must begin. Third, if work on poorly-represented floras or life forms does not increase, then established research groups must re-focus on target material available in existing collections. Unless global capability for estimating plant DNA C-values is significantly increased by new technology, funds or skilled operators, then this change in strategy will reduce progress towards achieving other targets. However, the prime objective remains: to generate a sample representative of the global flora that is able to support most comparative studies. Managers of the limited global capacity for estimating genome size should keep this firmly in mind when targeting taxa for new work.

Obsolescence time bomb

Several methods have been used to measure plant DNA C-values, but most values have been estimated by Feulgen microdensitometry (Fe), both overall and since 1997. In 1997 we identified an imminent problem, likely to limit future estimations. This was the failure and non-replacement of densitometers long used by many groups to estimate DNA C-values. Manufacturers were ending their support for such equipment, and users faced difficulty in funding new equipment for this purpose. Moreover, this problem was likely to be most acute in regions where some of the greatest gaps in our knowledge lay. Reviewing the position at the second Plant Genome Size Workshop confirmed that, as predicted, the ‘obsolescence time bomb’ had exploded. By 2003 several laboratories that had long published C-values listed in the Angiosperm DNA C-values database were now unable to estimate C-values by this (e.g. in Mexico, the USA), or any method (e.g. in Argentina). Vickers Instruments no longer supports their M85 microdensitometer, and spare parts for it are unobtainable. A few laboratories, including ours, can still use such machines, but now without servicing and only until they fail catastrophically.

As expected, one response to this problem was the increased use of flow cytometry (FC) to replace Fe. Analysis of data in the Appendix table shows a higher proportion of C-values obtained by FC (58·4 %), mostly since 1997, than noted previously (48·6 %) for data listed in Bennett and Leitch (2000), whilst in Bennett and Leitch (1995, 1997) FC averaged 26·7 %. Several groups have undertaken careful studies to compare DNA estimates made by FC and Fe, to define best practice for FC, or to show that FC can be applied widely to most plants across the full range of known DNA C-values (e.g. see review by Doležel and Bartos, 2005, this volume). Fortunately, the cost of a basic flow cytometer for such work has fallen, and suitable models (e.g. Partec PAII) have recently been set up for this use for approx. £20K (US$30K). If this technology continues to improve, and its costs continue to fall, FC should be more easily available. However, FC easily yields poor data in unskilled hands and by itself does not provide the cytological view of test material(s) that is essential to count chromosome number(s). Its use in some less-developed countries (where the greatest gaps in our knowledge still remain) will depend on training local operators, but such capacity-building may be thwarted by a lack of in-country support by the suppliers of flow cytometers.

A second solution to the problem is a new availability of relatively inexpensive computer-based image analysis (CIA) systems, which can estimate DNA amounts using Feulgen-stained cytological preparations in place of a microdensitometer. Although proprietary hard-wired CIA systems have been available since the 1970s (e.g. Zeiss Quantimet system), they cost much more than microdensitometers, and analysis of the literature shows they have not been used to estimate plant C-values. However, in the 1990s, with advances in computer technology, less expensive systems were developed (e.g. CIRES system) primarily for medical use, and these have also been used to good effect for plant C-value estimations (e.g. Temsch et al., 1998; Greilhuber et al., 2000).

Sadly, the CIRES system that adapted well for this purpose is no longer available, as the software is incompatible with the operating system used on modern PCs. However, computer-literate groups can assemble the kit needed to estimate C-values using CIA, and several software packages written specifically for this purpose are available (Vilhar et al., 2001; Hardie et al., 2002). Hardie et al. (2002) give an excellent review of this technique and practical issues concerned with its use for animal materials, and Vilhar et al. (2001) have compared CIA, Fe and FC, to help define best practice for CIA, demonstrating that CIA can be applied to plants with an approx. 100-fold range of C-values. Vilhar et al. (2001) concluded that ‘DNA image photometry gives accurate and reproducible results, and may be used as an alternative to photometric cytometry in plant nuclear DNA measurements’. CIA can use an existing microscope, costs less than FC to set up, and is easier to service in countries that lack FC manufacturers' support. The field would benefit from development of a standard inexpensive CIA ‘kit’, an agreed best practice CIA technique, and easy access to leading laboratories for training and technology transfer. Given this, CIA could soon become the method of choice for estimating C-values in angiosperms, replacing Fe as a method of choice along with FC, but with the advantage that, unlike FC, it uses microscope slide preparations, allowing users to make cytological observations.

Errors and inexactitudes

Swift (1950) defined the DNA content of an unreplicated haploid complement as its 1C-value (C standing for ‘constant’). Thus, replicated diplophase nuclei have a 4C DNA amount and produce two unreplicated 2C nuclei by mitotic division, and four 1C gametic nuclei after meiosis, irrespective of the organism's ploidy level. This convention applies well to polyploid taxa with diploidized meiotic chromosome pairing such as hexaploid breadwheat, which produce mainly functional, balanced polyhaploid gametes with 1C DNA amounts at meiosis (Rees and Walters, 1965). Consequently, for several previous reference lists, 4C DNA estimates for all taxa were divided by 2 and 4 to generate 2C- and 1C-values respectively (e.g. Bennett et al., 2000). However, a problem with this practice was identified for the few taxa with odd ploidy levels in release 3.1 of the Angiosperm DNA C-values database (namely 45 out of 3493 listed taxa, ∼1·3 %), as the resulting 1C-values are not biologically meaningful. For example, triploids with a 4C amount in their fully replicated metaphase nuclei do regularly produce two 2C nuclei at mitosis, but do not regularly produce four 1C products at meiosis. The authors are grateful to several colleagues who noted this problem and suggested solutions.

This problem has several practical consequences. (i) Regrettably, researchers who use 1C data from the literature or downloaded from the Angiosperm DNA C-values database may have included this error in the samples that they used for comparative analyses. However, this is unlikely to have influenced their conclusions significantly, since the magnitude of the error is relatively small (ranging between −0·25C for a triploid, to +0·25C for a pentaploid—which tend to cancel out), and affects only 1·3 % of all taxa listed. Overall, errors in mean DNA amounts for samples are probably less than 0·5 %. Studies that used data from the 2C or 4C columns for samples of odd-ploid taxa are unaffected by the error. (ii) To ensure that researchers are aware of the problem and do not generate 1C data for taxa with odd ploidy levels in the future, release 4.0 of the Angiosperm DNA C-values database gives 2C- and 4C-values for the 45 out of 3493 entries with odd ploidy levels, plus a warning note in response to any queries for 1C-values. This approach is also followed in the present Appendix table (see footnote t). (iii) This problem also highlights a general need to re-assess definitions of ‘C-value’ and ‘genome size’ in light of recent usage and new theoretical understanding, a topic explored by Greilhuber et al. (2005). Indeed, the above problem shows the need for care when handling data, and the danger of using computer-generated numbers uncritically. It is clearly perilous to ignore basic biology or the literature, as the recent history of genome size, ‘complete’ genome sequencing, and interest in the smallest angiosperm genome clearly shows.

Genome size, ‘complete’ genome sequencing, and, the euchromatic genome

A growing semantic problem concerns different uses of the term ‘genome’ (Greilhuber et al., 2005). As originally defined by Winkler (1920), genome referred to a monoploid chromosome complement. Since a monoploid is defined as ‘having one chromosome set with the basic (x) number of chromosomes’ (Rieger et al., 1991), it followed by definition that any polyploid taxon had three or more genomes. However, an alternative meaning, now in common usage, uses genome as an interchangeable alternative for the 1C-value to refer to the DNA content of an unreplicated gametic nuclear complement, irrespective of ploidy level. Unless the meaning intended is clearly defined on each occasion, this can be confusing, especially when authors use both meanings for a polyploid taxon in the same paper. For example, Devos and Gale (1997) used the term ‘genome’ to refer to both the entire complement of nuclear DNA in a hexaploid wheat nucleus and to the individual A, B and D ‘genomes’.

Further potential for confusion comes from new uses of the term ‘genome’ recently spawned by genome sequencers. These concern the counter-intuitive meaning of a ‘wholly’, ‘completely’ or ‘entirely’ sequenced genome, or of equating ‘genome’ with ‘euchromatic genome’—a confusing concept in which ‘genome’ equals the parts which could be cloned and sequenced, but not the rest (see below). None of these qualitative new uses of genome equates to its quantitative use to mean either a 1C-value, or one monoploid parental genome in a polyploid.

The completely sequenced genome

Since 2000 the scientific and popular press has reported and celebrated the ‘complete’ sequencing of the first insect (Drosophila melanogaster) and plant genome (Arabidopsis thaliana) and the human genome (in 2001). For example, a title in Nature reported: ‘The sequencing of an entire plant genome is now complete.’ Readers could be forgiven for assuming this meant the entire linear sequence of the nuclear DNA had been sequenced and assembled, so that the total size of the nuclear genome in these organisms was now known with certainty, and hence much more accurately than any previous estimate based on other methods subject to various experimental errors. The popular and scientific literature easily gives that impression, and unfortunately that is what many, incorrectly, understood. The truth is otherwise, as a ‘completely sequenced’ genome is a very relative concept. In the same issue of Science where Brenner (2000) wrote ‘We have the complete sequence of the 125-megabase genome of the fruit-fly Drosophila’, Pennisi (2000) noted that ‘the fly sequence still has c. 1000 small gaps’—referring only to the sequenced euchromatin part. But what of the rest? Speaking of heterochromatin, Adams et al. (2000) explained that the ‘genomes of eukaryotes generally contain heterochromatic regions surrounding the centromeres that are intractable to all current sequencing methods’ and that ‘Because of the unclonable repetitive DNA surrounding the centromeres it is highly unlikely that the genomic sequence of chromosomes from eukaryotes such as Drosophila or human will ever be ‘complete’. Moreover, Adams et al. (2000) stated that the unsequenced centric heterochromatin regions comprised ‘one third’ of the approx.180 Mb genome of Drosophila. But how was its size determined? Careful reading revealed that the Mb size of these unsequenced centromeric heterochromatic segments was measured not by any modern molecular method, but by using a ruler on one cell of a plate in a paper by Yamamoto et al. (1990). This important detail is not stated in the main text, but in the legend to fig. 1 in Adams et al. (2000). As Bork and Copley (2001) clearly explain, ‘There are regions, often highly repetitive, that are difficult or impossible to clone (one of the initial steps in a sequencing project) or sequence with current technology.… The extent of these regions varies widely in different species. So, rather than applying a universal gold standard, each sequencing project has made pragmatic decisions as to what constitutes a sufficient level of coverage for a particular genome. For example, as much as one-third of the sequence of the fruitfly Drosophila melanogaster was not stable in the cloning systems used, and so was not sequenced.’

Thus, workers interested in C-values should clearly understand that a ‘completely’, ‘entirely’ or ‘wholly’ sequenced genome is not what those words might imply if taken at face value, and the size given for such a genome may indicate either the amount of DNA sequenced, or the size of that euchromatic genome sequenced plus a best-guess estimate of a lot of unsequenced heterochromatin. Further, it can mean that every type of sequence in an organism has been sequenced, but it need not mean that all copies of all types have been sequenced, or that their copy numbers are known. Without this information total genome size (the DNA C-value) cannot be determined based on genome sequencing (Bennett et al., 2003).

Swift (1953) stated that, ‘in general estimates of the nucleic acids in cells are at present accurate to 10 or 20 %’. Later, Bennett and Smith (1976) concluded that ‘While a few estimates are not accurate even to within 20 %, careful measurements of 4C DNA amounts in species with 0·5–2·0 times that of a standard species are probably accurate to within 5–10 %’. Greilhuber (1998) noted ‘much suspect or demonstrably wrong data have accumulated and continue to be accumulated in the literature’. Sadly, the ‘complete’ genome sequencing of Arabidopsis (Arabidopsis Genome Initiative, 2000), which was expected to provide a new baseline, only added to this phenomenon.

Plant genome size researchers have long recognized the need for an exact calibration standard, whose C-value is not subject to technical errors. Thus, the publication of a precise C-value for the first plant to have its genome completely sequenced was eagerly awaited, as it was expected to provide a baseline, gold-standard reference point, against which all other plants could be compared and expressed. Arabidopsis thaliana ecotype ‘Columbia’ was chosen for complete genome sequencing, partly because its tiny genome should be less costly to sequence than larger genomes in other species.

In 2000 the Arabidopsis Genome Initiative (AGI) published the genome size of Arabidopsis thaliana as 125 Mb, comprising 115·4 Mb in the sequenced regions plus a rough estimate of 10 Mb in unsequenced centromere and ribosomal DNA regions. The accuracy of this estimate was set not by the precision of sequencing and assembling contigs, but by the total inaccuracy in the sizes assumed for the unsequenced gaps (Bennett et al., 2003) and hence was no more accurate than many estimates in the range 150–180 Mb made by other methods. Further analysis showed that the AGI's rough estimate of 10 Mb in the unsequenced gaps was highly inaccurate. Thus, new comparisons using flow cytometry, which co-ran A. thaliana ecotype ‘Columbia’ with three animal species including Caenorhabditis elegans Bristol N2 (whose genome size is accurately established by genome sequencing as just over 100 Mb), gave C-value estimates for A. thaliana in the range 154–162 Mb (with 157 Mb when C. elegans was used as the standard) (Bennett et al., 2003). This value is about 25 % larger than the AGI estimate of 125 Mb which was clearly a gross underestimate, and hence is not the long-awaited first benchmark C-value for a completely sequenced plant genome—giving those words their natural meaning. Other molecular work has confirmed this conclusion (e.g. Hosouchi et al., 2002).

More recently, the draft DNA sequence of the rice (Oryza sativa) genome was published (O. sativa ssp. japonica, Goff et al., 2002; O. sativa ssp. indica, Yu et al., 2002). However, while the estimated genome sizes based on DNA sequencing did not suffer from the serious shortcomings of the Arabidopsis estimate, neither did they fulfil the criteria essential for a new benchmark calibration standard. Yu et al. (2002) gave a new C-value of 466 Mb for O. sativa ssp. indica calculated by adding up the DNA sequencing data for 362 Mb of sequenced scaffolds and 104 Mb of ‘unassembled data’. In contrast Goff et al. (2002) reported the sequencing of DNA which covered a total of 389 809 244 bp of the O. sativa ssp. japonica genome. They stated that this represented 93 % of the 420 Mb rice genome but did not give a reference to the source of 420 Mb. It is therefore unclear whether the C-value of 420 Mb given by Goff et al. (2002) represents a new C-value based on genome sequencing alone. The 1C-value for rice may yet prove to be slightly higher than the values assumed by Goff et al. (2002) and Yu et al. (2002), and approach 490 Mb, equivalent to the 0·5 pg estimated by Bennett and Smith (1991).

Exact C-values based on complete genome sequences would be invaluable (Bennett et al., 2003). The need to complete sequencing gaps in Arabidopsis remains technically difficult, and it is unclear how, when, or if it will be achieved. Genome sequencing becomes more difficult as genome size increases, and experience with Arabidopsis implies that exact C-values are unlikely to be obtained in this way soon for any larger plant genomes, including the established plant C-value standard Oryza sativa.

The current situation poses a quandary for the plant genome size community, who have long paid serious attention to trying to maximize the accuracy and comparability of plant DNA C-values by using agreed calibration standards (both materials and assumed values; e.g. see http://www.rbgkew.org.uk/cval/conference.html#outline, Bennett et al., 2000), while eagerly awaiting the first absolute measurement for a plant obtained by really complete DNA sequencing. Current options include: (i) continue to use the existing small group of plant calibration standards until a plant C-value which meets the required criteria becomes available; (ii) adopt an animal C-value which meets these criteria as the baseline reference for expressing all other plant species values, e.g. Caenorhabditis elegans Bristol N2, whose C-value is known with confidence to within 1 % from genome sequencing to be just above 100 Mb (or roughly 0·1 pg); (iii) adopt a plant value based on direct comparisons with C. elegans, as the base calibration standard for plants, and create a ladder of secondary calibration standards all measured against it in a study replicated between several groups able to use best practice. The C-value for Arabidopsis thaliana (1C = 157 Mb or 0·16 pg), recently measured against C. elegans (Bennett et al., 2003), could be adopted as the basal plant calibration standard. Seed is readily available from stock centres and gives small, easily grown plants. Moreover, the ladder of values for its many endopolyploid nuclei would also provide convenient calibration reference points for higher values up to approx. 2500 Mb or 2·5 pg (i.e. 0·64 − 4C, 1·28 − 8C, and 2·56 − 16C).

Weeding out erroneous data

The value of the database is determined by the accuracy of the data it contains. Ideally, values should be exact, but in reality they are all subject to various technical and other errors, as noted above. This raises questions as to how accurate data are, and what level of error is acceptable in practice, or makes a datum valueless for a particular use or study.

The existence of a database itself is a valuable means of identifying real or potential errors, and hence of improving the accuracy and quality of the whole body of data. For example, where estimates for the same taxon (with the same chromosome number) disagree greatly this suggests an error. Further, where a body of data for a taxon shows close agreement except for one major departure, this identifies the outlier as almost certainly incorrect. For example, in the Appendix table, the 2C-value for diploid Acacia dealbata (1·7 pg) reported by Blakesley et al. (2002) is similar to that reported by Bukhari (1997) of 2C = 1·6 pg (listed in Bennett et al., 2000), but both values differ considerably from the 2C-value of 2·9 pg reported by Mukerjee and Sharma (1993b) (see Notes to the Appendix bb). Another example concerns Brachypodium distachyon. In 1991, the PhD thesis of Shi reported a 1C-value of 0·15 pg, but later Shi et al. (1993) gave its 1C-value as 0·3 pg. To resolve this discrepancy, RBG, Kew obtained some original material studied by Shi and estimated its 1C-value to be 0·36 pg, confirming the larger C-value for this species (see also footnote br). Thus, real errors can be identified with certainty, and potential errors flagged up for users in cautionary footnotes following Appendix tables.

DNA C-values in angiosperms vary approx. 1000-fold (over three orders of magnitude) from approx. 0·1 pg to over 100 pg. It is, therefore, often useful to know whether a species' 1C DNA amount has approximately 0·1, 1, 10 or 100 pg, even if there is still uncertainty regarding whether a species with approximately 1 pg is really closer to 0·8 pg than to 1·2 pg (an error ±20 %). In terms of its predictive value in nucleotypic correlations, such an error still permits useful conclusions to be drawn. The Arabidopsis community laboured long under the misapprehension that its 1C DNA amount was approx. 70 Mb (Leutwiler et al., 1984), and later approx. 100 Mb (Meyerowitz, 1994), when in reality it is much higher (about 157 Mb, Bennett et al., 2003). The level of inaccuracy involved (approx. 50–100 %) was considerable, yet it did not prevent the selection of Arabidopsis as the model plant for first complete genome sequencing, in no small part on the basis of its ‘small genome size’ (NSF, 1990; Somerville and Somerville, 1999). The Convention on Biological Diversity (United Nations Environment Programme, 1992) noted the need to make biodiversity data available, despite imperfections; a view which merits support (Bennett, 1998). Thus, it is better to list available C-value data subject to errors, until improved data with fewer errors become available. The body of data is needed by the scientific community and can clearly already be used to draw important conclusions, to make valuable predictions, and as a basis for necessary planning.

What is the smallest reliable C-value for an angiosperm?

The above examples show how seeing data in the comparative context of the database can help to identify real or potential errors in particular species. It can also facilitate broader enquiries such as ‘what is the smallest reliable C-value for an angiosperm?’. Again, the comparative approach has enabled researchers to be active in identifying potential errors in species with the smallest reported C-values, and to be transparent in correcting mistakes.

Because of error variation, a population of 1C-value estimates for one taxon should vary according to a normal curve, so those in the lower tail are all too low (Fig. 3A). Such values are lost in the frequency histogram for all angiosperm C-value estimates except at its lowest tail where some of the lowest C-values claimed are expected to be too low. This expectation is strongly supported in practice, as shown below. There are 53 1C estimates in the Angiosperm DNA C-values database or the present Appendix with 0·21–0·30 pg, 29 with 0·11–0·20 pg, but only three with 0·10 pg or below (Fig. 3B). Table 2 lists the 24 lowest estimates listed with 0·175 pg or less, but how robust are they?

Fig. 3. — (A) Expected error variation in a large population of DNA C-value estimates for one genotype as underestimates (in the lower tail) and overestimates (in the upper tail) surround more accurate, intermediate, genome size estimates. (B) Histogram showing frequency of C-values for the 85 smallest species in the database or Appendix.

Table 2.

The 24 lowest angiosperm 1C DNA estimates among data listed in the present Appendix table and the Angiosperm DNA C-values database (release 4.0, January 2003)

Taxon	1C (pg)	Original reference
Arabidopsis thaliana	0·051	Francis et al. (1990)
Cardamine amara	0·055	Band SR (pers. comm. 1984)
Arabidopsis thaliana	0·073	Leutwiler et al. (1984)
Fragaria viridis	0·108	Antonius and Ahokas (1996)
Rosa wichuriana	0·125	Bennett and Smith (1991)
Aesculus hippocastanum	0·125	Bennett et al. (1982)
Arabidopsis thaliana	0·128	Arabidopsis Genome Initiative (2000)
Sedum album	0·145	Hart (1991)
Arabidopsis thaliana	0·150	Arumaganathan and Earle (1991)
Carex nubigera	0·150	Nishikawa et al. (1984)
Carex paxii	0·150	Nishikawa et al. (1984)
Epilobium palustre	0·150	Band SR (pers. comm. 1984)
Hypericum hirsutum	0·150	Hanson, Leitch and Bennett (pers. comm. 2002)
Thlaspi alpestre	0·150	Band SR (pers. comm. 1984)
Arabidopsis thaliana	0·153	Bennett et al. (2003)
Arabidopsis thaliana	0·160	Bennett et al. (2003)
Arabidopsis thaliana	0·160	Galbraith et al. (1991)
Arabidopsis thaliana	0·165	Galbraith et al. (1991)
Arabidopsis thaliana	0·167	Krisai and Greilhuber (1997)
Arabidopsis thaliana	0·167	Bennett et al. (2003)
Amoreuxia wrightii	0·168	Hanson et al. (2001a)
Arabidopsis thaliana	0·170	Galbraith et al. (1991)
Arabidopsis thaliana	0·175	Bennett and Smith (1991)
Arabidopsis thaliana	0·175	Marie and Brown (1993)

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Thirteen of the 24 estimates in Table 2 are for Arabidopsis thaliana. A comparative approach suggests that some, which featured among the lowest C-values reported for angiosperms, are too low. Thus several C-value estimates made by molecular means in the range 0·05–0·125 pg (Leutwiler et al., 1984; Francis et al., 1990; Arabidopsis Genome Initiative, 2000) are now seen as gross underestimates, while many others in the range 0·15–0·18 pg are shown to span the true value of about 0·16 pg (Bennett et al., 2003).

After discounting the 1C estimate for Arabidopsis thaliana of 0·051 pg by Francis et al. (1990), the next smallest estimate listed is 0·055 pg for Cardamine amara (communicated from S. R. Band in 1984). With only a third the DNA amount of its related crucifer A. thaliana (0·16 pg), it seemed suspiciously low. Cardamine amara seed cannot survive drying, so it is unavailable from seed banks. However, we recently used flow cytometry to compare diploid C. amara collected near Sheffield with several calibration standards including A. thaliana ecotype ‘Columbia’. The 1C-value we obtained was around 0·24 pg (almost five-fold the earlier report). This is in close agreement with independent estimates made elsewhere (e.g. see Bennett and Leitch, 2005; Johnston et al., 2005).

Once the underestimates for Arabidopsis thaliana and Cardamine amara are discounted, few 1C-values of 0·125 pg or below remain for other species. One is the 1C estimate of 0·125 pg for Rosa wichuriana, estimated using callus material from Dr Andy Roberts (Bennett and Smith 1991). This value seemed questionably low in the context of the database, especially as it became clear that culturing may induce stain inhibitors. This concern led to a new collaboration with RBG, Kew using non-callous material, and our doubts were confirmed when it was re-estimated as 1C = 0·575 pg (Yokoya et al., 2000).

Perhaps all estimates below 0·125 pg should be doubted until confirmed. Another candidate was Aesculus hippocastanum whose 1C-value was listed as 0·125 pg (Bennett et al., 1982). This material is rich in tannins and a likely candidate for underestimating its DNA amount (Noirot et al., 2000, 2005). Recent work using flow cytometry at RBG, Kew, showed that the 1C-value of 0·125 pg was clearly an underestimate, as the true value is approx. 0·60 pg (L. Hanson, RBG, Kew, pers. comm.). With 0·125 pg for A. hippocastanum rejected, only one estimate below 0·14 pg remains, namely 0·11 pg for the Green strawberry, Fragaria viridis. Since there is considerable interest in knowing the smallest possible angiosperm genome, checks to establish whether this estimate is robust are now urgently required.

What is the minimum C-value for a free-living angiosperm and other free-living organisms?

Such comparative approaches can also facilitate broader questions such as: ‘what is the minimum genome size in angiosperms and other free-living organisms?’. There is a minimum compendium of nuclear genes essential for the life of any organism. This concept was behind Craig Venter's declared intension to synthesize from scratch a minimal bacterial genome (Check, 2002), and a project for a minimal eukaryote genome may eventually follow. Meanwhile we can only speculate on how small the minimum genome is for an angiosperm, and how closely extant species approach the minimum. It is, of course, below the lowest robust C-value for the group, i.e. less than 0·16 pg established for Arabidopsis thaliana. The presence of six other species with C-values of 0·15–0·169 pg in Table 2 strongly supports this conclusion. The estimate(s) of approx. 0·108 pg for Fragaria viridis may indicate a minimum C-value for extant angiosperms of about 100 Mb, but if so, is it a diploid, or a polyploid with three or more even smaller ancestral genomes?

Whilst the robust 1C-value for A. thaliana is 0·16 pg, this includes >25 % of repeated DNA (Bennett et al., 2003) and analysis of sequenced regions shows that >70 % of coding genes are duplicated (Bowers et al., 2003). Thus, in theory, a minimal genome without duplicated coding genes or repetitive DNA should not exceed approx. 50 Mb. Currently, there is no robust 1C estimate below 0·1 pg for an angiosperm, but if any of the seven species with C-values between 0·14–0·15 pg is a tetraploid, this would indicate a minimum genome size in extant taxa of approx. 75 Mb, or approx. 50 Mb if it is a hexaploid.

The comparative approach is usefully extended to include other groups of organisms. Table 3 shows minimum C-value estimates for multicellular organisms in several widely differing groups obtained by genome sequencing or other methods. Such minima for groups as diverse as nematodes, insects, algae and angiosperms range from 59 to 160 Mb. Thus, the minimum C-value known in extant free-living multicellular higher organisms is around 60 Mb. All may be diploidized paleopolyploids (Wendel, 2000), but except for one early and unconfirmed report of a 1C-value of approx. 39 Mb in a most simple multicellular placozoan animal (Ruthmann and Wenderoth, 1975) there is no evidence for extant diploid multicellular eukaryotic life forms with only 40–50 Mb. This tantalizing possibility will be an interesting driver for new work to find a first angiosperm whose 1C-value is <100 Mb, or a first free-living multicellular plant or animal with a robust 1C-value <50 Mb.

Table 3.

Robust minimum 1C-value estimates for several widely different groups of free-living, multicellular, higher organisms obtained by genome sequencing (*), other best practice techniques, or static cytometry using the fluorochrome DAPI for algae¹

Group	Species	Mb	Original reference
ANIMALS
Nematode	Caenorhabditis elegans	100^*	C. elegans sequencing Consortium (1998)
Platyhelminthes (flatworms)	Stenostomum brevipharyngium	59	Gregory et al. (2000)
Crustacea	Scapholeberis kingii (water flea)	157	Beaton (1988)
Annelid	Dinophilus gyrociliatus (polychaete worm)	59	Soldi et al. (1994)
Tardigrades (water bears)	Isohypsibius lunulatus	78	Redi and Garagna (1987)
Insect	Peristenus stygicus	98	TR Gregory (pers. comm.)
Arachnid	Tetranychus urticae (spider mite)	78	TR Gregory (pers. comm.)
Urochordates (tunicates)	Oikopleura dioica	72	Seo et al. (2001)
PLANTS
Chlorophyta (green alga)	Caulerpa paspaloides	88	Kapraun (2005)
Rhodophyta (red algae)	Heydrichia wolkerlingii	69	Kapraun (2005)
Phaeophyta (brown algae)	Stilophora rhizodes	98	Kapraun (2005)
Bryophyte	Holomitrium arboreum	167	Voglmayr (2000)
Lycophyte	Selaginella kraussiana	157	Obermayer et al. (2002)
Angiosperm	Arabidopsis thaliana	157	Bennett et al. (2003)

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The use of the base-specific fluorochrome DAPI for estimating DNA amounts may be less reliable than using intercalating fluorochromes such as propidium iodide (e.g. Doležel et al., 1992).

PROSPECTS FOR THE NEXT TEN YEARS

Apart from better defining the limits of genome size variation, what key developments are targeted, or likely, to occur in angiosperm genome size research in the next decade?

The first concerns the expected progress to increase the total number and representation of angiosperms in the C-values database. As noted above, estimating first values for species reached a historic high during recent years (Fig. 1). At least 1700 such values were added in 1997–2002, and the total number of species' C-value estimates probably reached around 4300. In 2003 the second Plant Genome Size Workshop set a goal of estimating a further 1 % (i.e. approx. 2500 species) in the next five years, and a similar target is likely for the following quinquennium. If so, there is a reasonable prospect that the number of species with a C-value estimate will reach, or significantly exceed 7500 by 2014.

More important than the expected increase in total numbers is the predicted improvement in the spread of new values across taxa, geographical regions and life forms, making the sample more representative of the global angiosperm flora, based on careful targeting to identify and fill knowledge gaps. The next decade should see almost complete representation for families, and a greatly increased representation for genera (especially in monocots), as work focuses increasingly at this taxonomic level. Representation at the generic level is currently approx. 1042 out of an estimated 14000 genera (7·4 %) and is targeted to rise to 10 % by 2009, and might reasonably be expected to approach 15 % within a decade. Moreover, this may approach 100 % for monocots, as they are targeted for holistic genomic studies (including C-values) for the global Monocot Checklist Project (Govaerts, 2004).

Recent experience shows that identifying a gap and setting a target may still not provoke the work needed to fill it. Positive monitoring of trends in published C-value data may also be required to achieve a significant change in research activity (e.g. as with the level of family representation in angiosperms; Hanson et al., 2003). Thus it will be important to monitor by 2009 whether the gaping chasms in the representation of African, South American and Chinese floras noted previously have yet resulted in a significant rise in first estimates for taxa from those regions. If not, then a major effort will be needed to correct this. The same applies to other groups of plants that have been identified as poorly represented in the Angiosperm DNA C-values database (e.g. halophytes, parasitic species and their hosts and tundra species). Whilst less certain, there is a good prospect that this vital process will occur in the next decade, driven by the Genome Size Initiative (GESI: see Bennett and Leitch, 2005).

Hitherto, when a question regarding C-value was framed (e.g. is genome size related to weediness?), it was often necessary to estimate C-values for many species before it could be addressed (Bennett et al., 1998). Clearly, the prime aim is to create a sample of C-values that is sufficiently representative for systematic, regional and life form variation as to allow most questions to be answered with confidence using the available dataset, without recourse to further C-value estimations. This goal is likely to be achieved in the next ten years. Thus, the next decade may be the last to see major efforts devoted to estimating first DNA C-values for taxa. Thereafter, new C-value research will probably concentrate on using and understanding such data, rather than acquiring them.

What important questions regarding genome size in angiosperms are likely to be answered in the next decade? Three closely interrelated issues concerning the possible significance of genome size for extinction, conservation and pollution are worth mentioning here.

The possibility that a large C-value might correlate with an increased risk and rate of extinction was suggested by Rejmanek (1996) and by Bennett et al. (2000). To test this, Vinogradov (2003) identified 3036 diploid species from the Plant DNA C-values database and compared each one against the United Nations Environmental Programme World Conservation Monitoring Centre (UNEP-WCMC) species database to determine its conservation status (i.e. global concern, local concern or no concern). He noted a striking relationship between genome size and conservation status; species with large genomes appeared to be at greater risk of extinction that those with smaller genomes.

Clearly, this was an important finding that now requires independent confirmation, drawn from further independent samples of species in different local regions and environments. Obtaining data for meaningfully large samples of species for such studies will probably be one main driver determining which taxa are targeted for C-value estimates in the future. If so, the next decade offers the prospect of a more definite and detailed understanding of any relationships between C-value and/or genome size and the risk of extinction. This, in turn, may have important practical and theoretical implications for conservation models and strategies. A key question is whether a large nuclear DNA amount gives an increased risk of extinction equally in diploid or polyploid taxa? Vinogradov (2003) tested whether ploidy played a role in increasing a species' risk of extinction, concluding that C-value per se was most important. Polyploidy is supposed to confer many advantages based on increased gene dosage and diversity, but do such advantages overcome the possible risks of a high C-value? If so, the proportion of polyploids should be higher for species with very high C-values than for those with lower C-values. In a test different from that of Vinogradov (2003), we compared the percentage of polyploids in 3400 extant species with known DNA amount and ploidy level in the Angiosperm DNA C-values database, ranked in order of increasing DNA amount and divided into five groups each containing 680 species. We found the percentage of polyploids for species in group 5 with the highest C-values (29·9 %) was actually lower than for species in group 4 (32·1 %) (Fig. 4). This confirms Vinogradov's finding that the prime factor determining increased risk of extinction is high C-value, and that polyploidy does not reduce this risk.

Fig. 4. — The percentage of diploids (open bars) and polyploids (closed bars) among 3400 species of known DNA amount and ploidy level ranked in order of increasing DNA amount and divided into five groups with 680 species per group. Data taken from the Angiosperm DNA C-values database (release 4.0, January 2003) and the present Appendix.

Other enquiries should test whether the risk of extinction in relation to high C-value or genome size varies for different threats and environments. This should compare variation in internal factors affecting the structure and ecology of the genome (e.g. increased ploidy level, and heterochromatin distribution), and in external factors (e.g. pollution and increased competition for space, minerals, light, and pollinators). Vilhar (pers. comm., and Vidic et al., 2003) investigated the effect of genome size on plant survival in lead-polluted soils. With increasing lead concentration in the soil the percentage of species with large genomes decreased significantly, suggesting that species with large genomes were at a selective disadvantage. Similar work on local floras in different areas with various threats is now needed to test whether their results are typical for other pollutants and environments. Understanding which species survive locally is always important, but especially as local loss equals global extinction if a species range is restricted to just that one locality. Such work will increasingly inform local environmental action plans and conservation strategies.

Holistic genomics

Early interest in plant genome size variation (c. 1950s and 1960s) ranged broadly across many fields including its genetic, developmental, ecological and evolutionary implications. However, after the molecular revolution the field fragmented somewhat as interest in DNA sequences was largely separated from more macro interests in C-values. However, given ‘complete sequences’ for genomes and homoeologous segments, and greater computing power, this post-genomic age is seeing a strong convergence of these interests. Thus, leading scientists who work at comparative sequence levels can also work on questions of genome size and evolution (e.g. Zhang and Wessler, 2004; Bennetzen et al., 2005). This is the age of holistic genomics in which knowledge of variation in genome size and C-value can be seamlessly joined up with information at all other levels to embrace information from sequences to ecology and from evolution to the environment. This powerful approach should permit or provoke quantum leaps in understanding the significance of extant variation in C-value and genome size, the processes that produce it, the rate at which it occurs, the factors that limit its extent and the advantages and disadvantages that it confers. Together, such understanding will link across biological fields to explain patterns of genome size variation in development, floras, ecological niches and evolution. The next ten years offer many exciting prospects for angiosperm genome size research. Work on DNA amount will remain a key core interest in biological research, but will increasingly become one integrated strand in holistic genomic studies and understanding, covering its origin(s), mechanisms of change, phenotypic and phenological effects, and its significance for ecological, developmental and environmental issues.

APPENDIX

Notes to the Appendix

The Appendix appears on pp. 59–88.

Named references in the following notes are given above in ‘Literature Cited’, while numbered references are given in ‘Original references for DNA values’ below.

(a) The original references for species DNA amounts in the Appendix are given in a numbered list following the Appendix table. Reference numbers follow on sequentially from those given in ‘Notes to Table 8’ by Bennett and Smith (1976, references 1–54) ‘Notes to Table 1’ by Bennett et al. (1982, references 55–107), Bennett and Smith (1991, references 108–163), ‘Notes to the Appendix’ by Bennett and Leitch (1995, references 164–269), ‘Notes to the Appendix’ by Bennett and Leitch (1997, references 270–306), and ‘Notes to the Appendix’ by Bennett et al. (2000, references 307–377).

(b1) Bennett and Smith (1991) gave absolute 4C DNA values for 11 angiosperm species recommended for use as calibration standards to estimate DNA amounts in other species. These species and their 4C DNA amounts are given in Table 4. If a species was calibrated in direct comparison with any one or more of the 11 standard species then the standard species used is identified in column 15 of the Appendix by the appropriate Key letter given above (e.g. F is Hordeum vulgare, etc.). If a species was first calibrated using a standard species listed above, then the original standard species is identified first and the intermediate standard species used to calibrate those species listed with it is also denoted by its number in column 1 of the Appendix. For instance, standard G (P. sativum) was used to calibrate Capsicum annuum ‘Doux Long des Landes’ (species 212 h in the Appendix), which was then used as an intermediate standard to estimate other Capsicum species given by Belletti et al. (1998, Ref. 434). The calibration standard for such Capsicum species is therefore given as G-212h.

Table 4.

The eleven angiosperm species recommended for use as calibration standards (see Notes to the Appendix, b1)

Key	Standard species	4C DNA amount (pg)
A	Triticum aestivum ‘Chinese Spring’	69·27
B	Allium cepa ‘Ailsa Craig’	67·00
C	Vicia faba PBI, inbred line 6	53·31
D	Anemone virginiana line AV 200	35·67
E	Secale cereale ‘Petkus Spring’	33·14
F	Hordeum vulgare ‘Sultan’	22·24
G	Pisum sativum ‘Minerva Maple’	19·46
H	Zea mays ‘W64A’	10·93
I	Senecio vulgaris (PBI population)	6·33
J	Vigna radiata ‘Berken’	2·12
K	Oryza sativa ‘IR36’	2·02

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(b2) In Refs 444 (Wendel et al., 2002) and 447 (Lin et al., 2001) Pisum sativum ‘Minerva Maple’ was used as the calibration standard but they assumed a 4C DNA value of 19·12 pg (Johnston et al., 1999) instead of 19·46 pg, which is the value given in Bennett and Smith (1976) and listed in Table 4. The 4C-value of P. sativum ‘Minerva Maple’ used in Refs 444 and 447 was estimated using Hordeum vulgare ‘Sultan’ as the calibration standard with an assumed 4C DNA content of 22·24 pg (Johnston et al., 1999).

(c) In several references listed in ‘Original references for DNA values’ the authors used a cultivar of a standard species different from that listed in Table 4, these are listed in Table 5. In some cases the C-value of the cultivar used was assumed to be the same as that of the cultivar given in Table 4. Evidence of intraspecific variation in a number of species suggests that such assumptions may sometimes be incorrect. In other cases the C-value of the cultivar was determined by the authors and was different from that of the standard species listed in Table 4. For example Refs 386, 397, and 453 used the cultivar ‘Express Long’ of Pisum sativum with a 4C DNA value of 16·74 pg. This value is lower than the 4C DNA amount of the cultivar ‘Minerva Maple’ of 19·46 pg given in Table 4.

Table 5.

Cultivars of standard species used that differ from those listed in Table 4

Original reference number	Plant calibration standard used	Assumed 4C DNA amount and reference (pg)
	Allium cepa
413	‘Frühstamm’	67·0 (reference not given)
409	‘Kantar topu’	67·0 (Van't Hof, 1965; Bennett and Smith, 1976)
435, 443, 449, 454, 455	‘Nasik Red’	67·0 (Van't Hof, 1965)
427	‘Stuttgarter Reisen’	67·0 (Bennett and Smith, 1976)
388, 411, 420, 422	var. aggregata	– (amount and reference not given)
	Hordeum vulgare
417	‘Stark’	21·36 (reference not given)
423	strain NE 86954	20·48 (Lee et al., 1997)
	Oryza sativa
424	type japonica	2·20 (Bennett and Smith, 1991)
	Pisum sativum
386, 397, 453, 393, 394, 395, 396, 429, 458	‘Express Long’	16·74 (Marie and Brown, 1993)
	‘Kleine Rheinländerin’	17·68 (Greilhuber and Ebert, 1994)
434	‘Lincoln’	18·14 (Doležel et al., 1992)
418	ssp. sativum convar. sativum var. ponderosum ‘Viktoria’	18·18 (Doležel et al., 1998)
	Secale cereale
418	ssp. cereale	32·38 (Doležel et al., 1998)
394	‘Dankovske’	31·16 (Doležel et al., 1998)
	Vicia faba
418	ssp. minor var. minor subvar. rigida ‘Tinova’	54·00 (Doležel et al., 1998)
405	‘Aquadulce’	53·31 (pers. comm. 2002)
430	‘Superguadulce’	53·30 (Bennett and Smith, 1976)
	Zea mays
391	‘CE-777’	10·86 (Lysák and Doležel, 1998)
437	‘C-777’	10·86 (Lysák and Doležel, 1998)
382	Va35	10·93 (Bennett and Leitch, 1995)

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(d) In References 407, 408 and 457 the cultivar of the calibration standard was not given. Refs 407 and 408 used Vicia faba as a calibration standard, whereas Ref. 457 used Allium cepa. In Ref. 408 Cremonini et al. (1992) assumed the same 4C-value for Vicia faba as for PBI line 6 (i.e. 53·3 pg) given in Table 4. If this species exhibits intraspecific variation then such assumptions may be incorrect.

(e) In a number of original references the authors used a plant species not listed in Table 4 as a calibration standard. These are listed in Table 6.

Table 6.

Plant species used as calibration standards but not listed in Table 4

Original reference number	Plant calibration standard used	Assumed 4C DNA amount and reference (pg)	Abbreviation used in column 15 of Appendix
383	Agave americana	31·80 (Zonneveld and Van Iren, 2001)	Agave
	Arabidopsis thaliana
389	‘Columbia’	– (amount and reference not given)	Arab.
414	‘Columbia’	0·53 (Kaneko et al., 1998)	”
427	Cerastium eriophorum	5·20 (Boscaiu et al., 1999)	Cerastium
	Glycine max
393	‘Ceresia’	4·54 (Greilhuber and Obermayer, 1997)	Glycine
429	‘Ceresia’	4·51 (Obermayer and Greilhuber, 1999)	”
421	var. Palmetto	5·00 (Doležel et al., 1994)	”
432	‘Burlison’	5·56 (Graham et al., 1994)	”
402	‘Polanka’	5·00 (Doležel et al., 1994)	”
	Lycopersicon esculentum
380, 382, 465	‘Gardener's Delight’	4·00 (Obermayer et al., 2002)	Lycopers.
437	‘Stukické’	3·92 (Doležel et al., 1992)	”
385	‘Montfavet’	4·02 (Marie and Brown, 1993)	”
	Nicotiana tabacum
427	‘Petit Havana SR1’	18·00 (Bennett and Leitch, 1995)	Nicot.
417	‘Samsun’	18·15 (reference not given)	”
	Petunia hybrida
390, 410, 433, 452	‘PxPC6’	5·70 (Godelle et al., 1993; Marie and Brown, 1993)	Petunia
390	‘Hit Parade Blau’ R	5·70 (Marie and Brown, 1993)	”
401	No cultivar given	5·70 (Marie and Brown, 1993)	”
	Rhaphanus stativus
404	‘Saxa’	2·20 (Doležel et al., 1992)	Rhaphanus
	Sorghum bicolor
439	Line TX623	3·52 Price and Levin (pers. comm.)	Sorghum
462	No cultivar given	3·20 (Bennett and Smith, 1991)	”
406	Vicia narbonensis	29·10 (Frediani et al., 1992)	Vicia narb.

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(f) Several papers listed in ‘Original references for DNA values’ used animal cells as the calibration standards. Thus Refs 387, 417, 426, 442, 456, 463 used chicken erythrocytes with an assumed 4C DNA value of 4·66 pg (Galbraith et al., 1983). The calibration standard is abbreviated to Gallus in column 15 of the Appendix. In Ref. 438 blood cells from the catfish Ictalurus punctatus were used as a standard with an assumed 4C-value of 4·00 pg (Tiersch et al., 1989), this is abbreviated to Ictal. in the Appendix. In Ref. 438 domestic swine (Sus scrofa) erythrocytes were used as a standard with an assumed 4C-value of 11·34 pg (Taliaferro et al., 1997), and is abbreviated to Sus in the Appendix. Human cells with an assumed 4C DNA amount of 14·00 pg (Tiersch et al., 1989) were used as calibration standards in Refs 384, 428 and 419 (leucocytes, Ref. 384, 428; lymphocytes, Ref. 419) and the abbreviation of Homo is used in the Appendix. Finally, Drosophila melanogaster with an assumed 1C DNA amount of 180 Mb (Adams et al., 2000) and Caenorhabditis elegans with a 1C DNA amount of 100·25 Mb, based on complete genome sequencing (see C. elegans Sequencing Consortium, 1998 and http://wormbase.org), were used as calibration standards for Ref. 461, and the abbreviation of Dros. and Caeno. respectively are used.

If a plant species was calibrated using an animal species and then subsequently used as the calibration species for other plants, then the animal species is identified first, and the intermediate plant species is identified by its entry number given in column 1 of the Appendix. Thus Mishiba et al. (2000, Ref. 387) used Gallus with an assumed 4C DNA amount of 4·66 pg (Galbraith et al., 1983) to calibrate Hordeum vulgare ‘New Golden’ (species 398p in Appendix), this was then used as the calibration standard to estimate DNA C-values of Petunia and Calibrachoa species given by Mishiba et al. (2000). The calibration standard for these Petunia and Calibrachoa species is given as Gallus-398p.

(g) When a new estimate (or estimates) is given for a species or subspecies already listed by Bennett and Smith (1976, 1991), Bennett et al. (1982, 2000) or Bennett and Leitch (1995, 1997), the estimate is given a number and a lower case letter in column 1 of the Appendix. An ‘a’ implies that the value is preferred to any estimate for that species listed previously by the first author. Where several estimates are available for the same species, the ‘a’ value would automatically be chosen in any arithmetical or statistical calculations. In this context, single estimates for species and ‘a’ values are referred to as ‘prime entries’.

(h) Intraspecific variation in nuclear DNA amount is claimed to occur in this species. Consequently the values given in the Appendix should not be assumed to be correct for all accessions of the species. Where several C-values are listed for a single species with the same ploidy level or chromosome number within a taxon, then only the minimum and maximum values reported from a single reference are listed in the Appendix.

(i) A range of DNA amounts was reported for this species in the reference cited in column 13 of the Appendix. Intraspecific variation was not claimed to occur, so the nature of this variation is unclear. Where estimates differed by more than 10 % the minimum and maximum values are given for the same ploidy level or chromosome number in the Appendix, otherwise only the highest value is given.

(j) According to the International Code of Botanical Nomenclature (Greuter et al., 1994), the names of plant families must end in -aceae. However, eight plant families are exceptions in that each has two alternative names, both of which are correct under the Botanical Code. One is a standard name, ending in -aceae, the other is an exception, sanctioned by long usage. These and their alternatives are the following: Palmae (Arecaceae), Gramineae (Poaceae), Cruciferae (Brassicaceae), Leguminosae (Fabaceae), Guttiferae (Clusiaceae), Umbelliferae (Apiaceae), Labiatae (Lamiaceae) and Compositae (Asteraceae). To be consistent with previous DNA lists (Bennett and Smith, 1976, 1991; Bennett et al., 1982, 2000; Bennett and Leitch, 1995, 1997) the ‘non-standard’ plant names are retained in the present work.

(k) Recent cladistic analysis using both molecular and non-molecular phylogenetic data has resulted in a revised classification of families by the Angiosperm Phylogeny Group (APG) (APG II, 2003). Familial names used in the APG classification are followed in the Appendix. Thus, although Zonneveld (2002, Ref. 440) placed Aloe in Aloeaceae, recent molecular and non-molecular phylogenetic data recognizes that this family is embedded within the newly circumscribed Xanthorrhoeaceae (APG II, 2003) so Xanthorrhoeaceae is given in the Appendix. Similarly, the APG II (2003) now recognizes that Hostaceae is embedded within the Asparagaceae, so Hosta, which was placed in Hostaceae in Ref. 384 (Zonneveld and Van Iren, 2000), is listed under Asparagaceae in the Appendix.

(l) The authority for this species is either unknown or unclear to the present authors.

(m) Whether or not voucher specimens exist for this species is unknown to the present authors.

(n) The chromosome number of this species is either unknown or unclear to the present authors.

(o) The chromosome count for this species was taken from the literature and not determined by the authors of the reference cited.

(p) The ploidy level of this species is either uncertain or unclear to the present authors.

(q) The life cycle type of this species is either unknown or unclear to the present authors.

(r) The method used to measure the DNA amount is unclear.

(s) The factor of 1 pg = 980 Mbp was used to convert picograms to Mbp (Cavalier-Smith, 1985; Bennett et al., 2000).

(t) As a rule, replicated diplophase nuclei contain a 4C DNA amount producing two unreplicated 2C nuclei by mitotic division and four 1C gametic nuclei after meiosis (irrespective of ploidy level). This convention applies well to polyploid taxa with diploidized meiotic chromosome pairing which produce functional balanced polyhaploid gametes with 1C DNA amounts at meiosis. Thus 4C estimates were automatically divided by 4 to generate 1C-values given for all taxa of even ploidy level listed in the Appendix. However the resulting ‘1C’ data are not biologically meaningful for taxa with odd ploidies. Consequently the Appendix gives only 2C- and 4C-values for such taxa.

(u) There is no obvious basic number for the genus Luzula due to the presence of holocentric chromosomes. It is therefore impossible to allocate Luzula species with high chromosome numbers to any ploidy level with certainty.

(v) Unal and Callow (1995, Ref. 412) obtained a regression of the nuclear fluorescence of Allium cepa (4C = 67·0 pg), Crepis capillaris (4C = 9·6 pg), Hordeum vulgare (4C = 22·2 pg), Pisum sativum (4C = 20·2 pg), Secale cereale (4C = 33·2 pg) and Vicia faba (4C = 47·9 pg) versus nuclear DNA content, and used this to estimate the DNA C-values of 13 Lathryus species. However, it is noted that the 4C-values for P. sativum, and Vicia faba are non-standard values compared with those given for these species in footnote (b1) above.

(w) The standard species used to convert arbitrary units into absolute DNA amounts is unclear to the present authors.

(x) The DNA value given for this species in the original reference differs considerably (i.e. >100 %) from that given in other original references cited in previous compiled lists of DNA amounts (i.e. Bennett and Smith, 1976, 1991; Bennett et al., 1982, 2000; Bennett and Leitch, 1995, 1997). The reason(s) for this is unknown. This C-value should therefore be used with caution until the question is resolved.

(y) The specific status of the material available for study is unclear. The data are included since information on DNA amounts for this genus is relatively sparse, so an indication of genome size in the genus may be useful.

(z) Zonneveld (2001, Ref. 383) gave C-values for 16 hybrid cultivars which fall within the range that he reported for Helleborus species listed in the Appendix. Our compiled lists have usually been restricted to C-values for species. Following this practice, C-values for Helleborus hybrids were not included in the present Appendix.

(aa) Zonneveld and Van Iren (2000, Ref. 384) gave DNA amounts for 94 accessions of Hosta which were recognised as 23 different species. Their table 1 gives a DNA amount for each accession together with a mean value for each recognised species. Only the later value is given in the present Appendix. They also included C-values for 16 Hosta cultivars (in their table 3). These were once recognized as species, but following pollen viability tests Zonneveld and Van Iren (2000) concluded they were hybrids. Our compiled lists have usually been restricted to C-values for species. Accordingly, C-values for Hosta hybrids were not included in the present Appendix.

(ab) Zonneveld and Van Iren (2000, Ref. 384) and Zonneveld (2002, Ref. 440) used male human leucocytes (2C = 7·0 pg; Tiersch et al., 1989) as their primary standard to estimate the DNA amount of three Agave species, namely: (i) Agave stricta (ii) A. americana and (iii) A. sisalana. A. americana was then used as internal calibration standard for most Hosta (Ref. 384) and Aloe (Ref. 440) taxa. However, in a few cases where the DNA content of Hosta or Aloe coincided with that of A. americana, one of the other two Agave species was used as the internal standard. As neither reference identified which Agave species was used, the calibration standard in column 15 of the present Appendix is given as Agave sp.

(ac) Thibault (1998, Ref. 385) claimed intraspecific variation ranging from 6 to 11 % in the Salix species he studied, but only a mean DNA C-value for each species was given in his table 3. It is these values that are listed in the present Appendix. Thibault (loc. cit.) also included C-values for five hybrids. Our compiled lists have usually been restricted to C-values for species, thus C-values for Salix hybrids are not included in the Appendix. In addition, Thibault (1998, Ref. 385) listed a C-value for ‘S. triandra?’ but concluded its identity was ‘hard to specify’. Consequently, this taxon was not included in the Appendix.

(ad) Thibault (1998, Ref. 385) used DNA C-values to predict the ploidy level of each Salix species given in his table 3, assuming direct proportionality. Moreover, their chromosome numbers were not counted by him, but derived by him assuming a constant basic chromosome number of n = 19 for the genus. These predictions are entered in columns 6 and 7 of the Appendix.

(ae) Some taxa once included in Petunia are now included in Calibrachoa. The taxonomy for most Petunia species listed in Mishiba et al. (2000, Ref. 387) follows that of Wijsman (1990) who split the genus Petunia sensu Jussieu (1803) into two; Petunia sensu Wijsman and Calibrachoa. However, five species listed in Mishiba et al. (2000) were not reclassified by Wijsman and so they were listed under the generic name of Petunia sensu Jussieu in Mishiba et al. (2000) although they ‘were regarded as Calibrachoa’ (see their table 3). By following the taxonomy of Petunia sensu Wijsman, several species originally listed under the genus Petunia, now belong to Calibrachoa (e.g. parviflora was listed in the genus Petunia by White and Rees, 1985, 1987) and this generic name was used in the list of Bennett and Leitch (1995). Yet Mishiba et al. (2000) assigned this species to Calibrachoa. To avoid confusion readers looking under Petunia are referred to Calibrachoa in the Appendix.

(af) Joachimiak et al. (2001, Ref. 391) reported chromosome numbers and C-values for six Bromus species. Chromosome numbers varied considerably in roots of three species, but variation in C-values was ‘virtually absent within leaf mesophyll cells’. The C-values given by Joachimiak et al. (2001) were obtained using leaf mesophyll cells and are listed in the Appendix.

(ag) The study by Rosato et al. (1998, Ref. 392) was primarily concerned with polymorphism in Zea mays ssp. mays races with B-chromosomes, but gave C-values only for plants lacking B-chromosomes. Thus, they listed DNA amounts for 17 populations which differed by 36 % (2C = 5·008–6·757 pg) in plants with 2n = 20. Similar intraspecific variation in this species was reported previously (Laurie and Bennett, 1985; Rayburn et al., 1985). Mean DNA amounts for only the populations with the largest and smallest C-values for A-chromosomes, are listed in the Appendix.

(ah) Dimitrova and Greilhuber (2000, Ref. 394) reported significant intraspecific variation in Crepis biennis (P < 0·05) and C. sancta (P < 0·01), some of which had variable numbers of B-chromosomes. As only means were given for material with 0–2 B-chromosomes, it was impossible to give values (presumably the largest) for the 2B complement. Consequently, the Appendix just lists the smallest and largest C-values for accessions without B-chromosomes.

(ai) Dimitrova and Greilhuber (2000, Ref. 394) reported significant (P < 0·001) intraspecific variation of 11 % for Crepis pulchra. They suggested that the two accessions with the higher C-values may belong to subspecies turkestanica. This is not recognized in the Bulgarian flora (where these accessions were collected), but was described by Babcock (1947). In the Appendix the higher C-values listed for this species (entry numbers 305c and e) may thus correspond to C. pulchra ssp. turkestanica.

(aj) Temsch and Greilhuber (2000, Ref. 395) estimated C-values in 11 accessions of Arachis hypogaea using both Feulgen microdensitometry and flow cytometry. C-values for different accessions showed great stability, so they calculated a mean C-value for each method in the ‘Results and Discussion’ of their paper. Only these mean values are listed in the Appendix.

(ak) Previous estimates for Vicia melanops (2n = 10) (e.g. Chooi, 1971; Raina and Rees, 1983; Raina and Bisht, 1988) all report a 4C-value of approx. 40 pg, which is much higher than the value of 27·6 pg given in Cremonini et al. (1992, Ref. 408, entry number 780d). Thus, this estimate should be viewed with caution until confirmed independently.

(al) Akpinar and Bilaloglu (1997, Ref. 409) gave a 2C-value of 13·1 pg for Vicia cracca ssp. cracca (with 2n = 2x = 14; their original count). However, six previous reports for V. cracca listed in the database (Bennett and Leitch, 2003) gave similar 2C-values (from 10 to 13 pg), but for 2n = 4x = 28. The cause of this discrepancy is unknown, thus the estimate by Akpinar and Bilaloglu (loc. cit.) should be viewed with caution until confirmed independently.

(am) Sakamoto et al. (1998, Ref. 414) estimated the C-value of Cannabis sativa using Arabidopsis thaliana ‘Columbia’ (1C = 130 Mb, Kaneko et al., 1998) as the calibration standard. However, the 1C-value assumed for A. thaliana was low compared with its recently confirmed estimate of 157 Mb (Bennett et al., 2003). If 157 Mb is assumed for A. thaliana, then the 1C-value for C. sativa would be 988 Mb = 1·01 pg (female) and 1016 Mb = 1·04 pg (male).

(an) Gammar et al. (1999, Ref. 416) gave DNA amounts for eight Lupinus species in arbitrary units (a.u.) listed as Mn(x) values in their Figs. 1–4. Bennett and Smith (1976) gave the 4C DNA amount of L. luteus as 4·0 pg (allowing for recalibration of Senecio vulgaris from 5·88 pg to 6·33 pg, see Bennett and Smith, 1991). Gammar et al. (loc. cit.) gave Mn(x) values for three L. luteus populations as 60·4, 58·4, and 63·8 in figure 1A, noting they were not statistically different. The mean of these three values was calculated to be 60·86 a.u. To convert the Mn(x) values for each Lupinus species into absolute DNA amounts, they were multiplied by a conversion factor of 0·07 (i.e. 4·0 pg ÷ 60·86 a.u.).

In some Lupinus species more than one population was studied, and several Mn(x) values were listed. If these did not differ significantly, the average Mn(x) value was calculated and converted into absolute DNA amounts. However, chromosome counts of 2n = 38, 42 and 44 were reported in L. angustifolius, so variation in Mn(x) may correspond to different cytotypes.

Some absolute DNA amounts calculated for Gammar et al. (loc. cit) do differ greatly from those previously reported for the same species (e.g. L. pilosus, 4C = 4·9 pg, is almost double the value of 2·5 pg given by Obermayer et al., 1999). Similarly, the 4C-value of 3·1 pg calculated for L. angustifolius with 2n = 38, 42, or 44 is similar to the estimate (4C = 3·7 pg) by Barlow (pers. comm., listed in Bennett et al., 1982), yet the latter was for material with 2n = 26. Data from Gammer et al. give a useful approximation of C-values in the five species not previously listed, but should be treated with caution unless confirmed independently.

(ao) Brandizzi and Grilli Caiola (1996, Ref. 419) gave 2n = 18 for Crocus biflorus in their table 1, but 2n = 8 in the first paragraph of their text. They also stated in their final paragraph: ‘However, C. biflorus and C. etruscus, having half the chromosome number with respect to C. thomasii and C. cartwrightianus…..’ As C. thomasii and C. cartwrightianus were both recorded with 2n = 16 by Brandizzi and Grilli Ciola (1996), we conclude that C. biflorus had 2n = 8, and so this number is entered in the Appendix.

(ap) The 4C DNA amounts reported by Mukerjee and Sharma (1993a, Ref. 420) for Luzula nivea and L. luzuloides are over 50 % larger than those reported by Barlow (pers. comm. 1976; reference 36 in Bennett and Smith, 1976). The chromosome numbers for each species were the same, so the cause of the discrepancy is unknown. However, Mukerjee and Sharma (1993a) used a single wavelength method, which may suffer from distributional error (Greilhuber, 2005, this volume). Thus, estimates for Luzula in Mukerjee and Sharma (1993a) should be viewed with caution until confirmed independently for these species.

(aq) Asif et al. (2001, Ref. 421) estimated DNA amounts in 14 genotypes of Musa acuminata. Genotype BC3 (belonging to the separate subspecies truncata) had the highest DNA amount and its C-value was shown to be significantly different (P < 0·01) from the other thirteen genotypes. Only the DNA amount of genotype BC3, corresponding to M. acuminata ssp. truncata, and the highest DNA amount out of the 13 other genotypes of M. acuminata are entered in the Appendix.

(ar) Chaudhuri and Sen (2001, Ref. 422) examined two Scilla indica cytotypes (entry numbers 710b and c) which differed considerably in both DNA amount and karyotype structure, although both had 2n = 30. The differences may reflect problems with taxonomy. Studies by Greilhuber and colleagues (Greilhuber, 1979; Greilhuber and Speta, 1985) have shown that large intraspecific differences in C-values in other Scilla species (e.g. S. bifolia) reduce to a level hardly more than methodological error following taxonomic splitting.

(as) Chung et al. (1998, Ref. 423) estimated C-values in 12 soybean (Glycine max) strains varying in seed size. They reported statistically significant differences of 4·6 % in the 2C-values between strains. Only the smallest and largest C-values are entered in the Appendix.

(at) Hartman et al. (2000, Ref. 425) estimated C-values in 22 Leucaena species using flow cytometry. Three species (Pisum sativum 4C = 17·6 pg, Oryza sativa 4C = 1·8 pg and Vicia faba 4C = 53·0 pg) were used as calibration standards at various times, but unfortunately the authors did not state which standard(s) was compared with which Leucaena species.

(au) Boscaiu et al. (1999, Ref. 427) referred to plants of Cerastium with 2n = 36 as diploids in contrast with various other authors who consider them as tetraploids. The assumption was based on Boscaiu et al.'s observations that, while the base chromosome number in Cerastium may be x = 9, no Cerastium species is known with 2n = 18.

(av) The C-values reported for Hedera helix by Obermayer and Greilhuber (1999, Ref. 429) agree well with previous reports by König et al. (1987) of 2C = 3·0 pg, but are only about one third the value reported by Marie and Brown (1993) of 2C = 8·2 pg, which is unsupported.

(aw) Blanco et al. (1996, Ref. 431) gave DNA amounts for Dasypyrum hordaceum and D. villosum in arbitrary units (a.u.), listed as mean values in their fig. 3. The value for D. hordaceum was converted into an absolute DNA amount by multiplying the mean value of 381·7 a.u. by a conversion factor of 0·11. This conversion factor was obtained as the ratio of the 4C estimate for Dasypyrum villosum (listed as the synonym Haynaldia villosa) reported by Bennett (1972) as 21·4 pg, and the estimate of 193·7 a.u. reported by Blanco et al. (1996).

(ax) Rayburn et al. (1997, Ref. 432) estimated C-values in 90 accessions of Glycine max. Accessions showed a 12 % variation in DNA amount and these differences were statistically significant. Only the smallest and largest C-values are listed in the Appendix.

(ay) Comparing C-values given by Belletti et al. (1998, Ref. 434) with those previously published showed DNA amounts for Capsicum baccatum, C. chinese, C. eximium, C. frutescens and C. pubescens were around one third greater than those of Owens (pers. comm.) listed in Bennett and Smith (1976). Belletti et al. (1998) suggested that the cause of the discrepancy could be that Owens used Allium cepa as the calibration standard, whose 2C-value of 33·5 pg differs considerably from those reported in Capsicum species studied.

(az) Široký et al. (2001, Ref. 437) investigated C-values in four Silene species, including S. latifolia, which has previously been listed by Bennett and Leitch (1995) under its synonym Melandrium album.

(ba) Taliaferro et al. (1997, Ref. 438) gave DNA C-values for 18 accessions of Cynodon corresponding to two species: C. transvaalensis (2n = 2x = 18), and C. dactylon var. dactylon (2n = 4x = 36 and 2n = 6x = 54). Only small differences in DNA amounts were noted between five diploid and five tetraploid accessions, and a mean 2C-value for each ploidy level was also given in table 2 of their paper. This mean value is listed in the Appendix. However, the three hexaploid accessions examined comprised one accession of C. dactylon var. dactylon and two hybrids. Thus, only the C-value estimate for hexaploid C. dactylon var. dactylon is entered in the Appendix, rather than the mean for the three hexaploid accessions given in table 2 of Taliaferro et al. (1997).

(bb) Blakesley et al. (2002, Ref. 441) examined seven populations of Acacia dealbata and four of A. mangium to determine ploidy and DNA amount. In A. dealbata, they identified naturally occurring diploid, triploid and tetraploid genotypes. Chromosome numbers were counted in only one diploid and one tetraploid genotype, and C-values for only these populations are given in the Appendix. In naturally occurring A. mangium only diploid populations were found, and the C-value for the only population whose chromosome number was determined is given in the Appendix. C-values for colchicine-induced tetraploid genotypes of A. mangium are not included.

The 2C-value for diploid A. dealbata (1·7 pg) is similar to that reported by Bukhari (1997) as 2C = 1·6 pg. In contrast the 2C-value (2·9 pg) reported by Mukherjee and Sharma (1993b) is nearly twice that of Blakesley et al. (2002). Perhaps this discrepancy reflects the use of Allium cepa (2C = 33·5 pg), whose genome size is over an order of magnitude greater than that of Acacia, as a calibration standard by Mukherjee and Sharma (1993b). Similar discrepancies were noted between DNA estimates for A. mangium reported by Blakesley et al. (loc. cit.) of 2C = 1·3 pg, and those by Mukherjee and Sharma (1995) of 2C = 2·3 pg.

(bc) Ohri and Singh (2002, Ref. 443) listed C-values for 20 wild relatives of cultivated pigeon pea (Cajanus cajan). However, C-values for 14 of these species had already been communicated to MD Bennett in 1996 and listed in the Appendix of Bennett and Leitch (1997) under Original reference number 303. To avoid duplication of data in the database, only C-values for six species not listed previously are included in the Appendix.

(bd) Wendel et al. (2002, Ref. 444) listed DNA amounts for 13 species in the tribe Gossypieae. However, C-values for three of these species had already been communicated to MD Bennett in 1999 and listed in the Appendix of Bennett et al. (2000) under Original reference number 349. To avoid duplication of data in the database, only C-values for ten species not listed previously are included in the Appendix.

(be) The C-value of Arabidopsis thaliana given by the Arabidopsis Genome Initiative (2000, Ref. 448) was based on DNA sequencing data for 115·4 Mb of the genome, plus a guestimate of 10 Mb for several unsequenced gaps in the genome. Recent work places its 1C-value around 157 Mb (Bennett et al., 2003).

(bf) Ohri (2002, Ref. 449) listed DNA amounts for 36 tropical hardwood species belonging to 13 families. However, C-values of 35 of these had already been communicated to MD Bennett in 1996 and listed in the Appendix of Bennett and Leitch (1997) under Original reference number 301. To avoid duplication of data in the database, a C-value for the only species not included in a previous compilation (Drypetes roxburghii) is listed in the present Appendix. A new C-value for Melaleuca leucadendra, double that given in Bennett and Leitch (1997), is also listed in the Appendix, to correct an error in communication which confused the 2C- and 4C-values for this species.

(bg) The C-value of 466 Mb for Oryza sativa ssp. indica given in Yu et al. (2002, Ref. 450) was based on DNA sequencing data for 362 Mb of sequenced scaffolds, and 104 Mb of ‘unassembled data’ subject to numerous assumptions (see Yu et al., loc. cit. – page 80).

(bh) The C-value of 420 Mb for Oryza sativa ssp. japonica given in Goff et al. (2002, Ref. 451) was derived from DNA sequencing data for 389·9 Mb, and their assumption that this equals 93 % of the genome, perhaps using some previously published 1C-value. The source of this assumption, as of any such DNA estimate, and the method by which it was obtained, was not clearly cited by Goff et al. (2002).

(bi) Redondo et al. (1996, Ref. 453) estimated C-values in four populations of Saxifraga granulata. In one population chromosome numbers ranged from 2n = 44 to 56, but 2n = 44 was predominant. They noted that the DNA amount was also variable but gave only one C-value, which is listed in the Appendix. However, intraspecific variation in DNA amount may occur in this species, so the C-value listed may not apply to all members of the population.

(bj) Redondo et al. (1996, Ref. 453) estimated DNA amounts in four populations of Saxifraga granulata. They reported DNA amounts for a population in which they could not obtain a chromosome count (entry number 706), but based on the DNA amount, they suggested this population may have 2n = 30.

(bk) Emshwiller (2002, Ref. 456) estimated C-values in 10 accessions of cultivated oca (2n = 8x = 64; Oxalis tuberosa), two tetraploid wild species, and 78 diploid accessions which were provisionally identified as 35 species. As variation in 2C DNA amounts was usually no more than 0·1 pg, and considered to be technical in nature, only the highest C-value was reported in table 3 of Emshwiller (2002) for most species and is entered in the Appendix. Variation in DNA amounts greater than 0·1 pg was considered real for O. spiralis (2C = 1·062–1·339 pg) and O. peduncularis (2C = 0·927–1·163 pg), so both the lowest and highest values are entered in the Appendix for these species. Emshwiller (2002) noted that this variation may reflect problems of taxonomy and species boundaries.

(bl) Emshwiller (2002, Ref. 456) estimated the DNA amounts in ten accessions of cultivated oca (2n = 8x = 64; Oxalis tuberosa). Variation was noted, even in measurements made for all accessions estimated on one day (see table 3 of Emshwiller, 2002), but she did not consider it to represent intraspecific variation and a mean 2C estimate calculated from all measurements made was given in her ‘Results’ section as 2C = 3·514 pg. It is this value that is entered in the Appendix.

(bm) Nagl et al. (1983, Ref. 457) included DNA amounts for 49 species. However, C-values for 20 of these had been published elsewhere, and already included in previous compilations by Bennett and colleagues (listed under original reference numbers 34, 36, 60, 61, 81, 82, 84, 85, 86). To avoid duplication only C-values for 29 species that had not been listed previously are included in the Appendix.

(bn) Values for Phaseolus coccineus and P. vulgaris given in Nagl et al. (1983, Ref. 457) are around twice those given in another paper by Nagl and Treviranus (1995, Ref. 390), listed in the present Appendix. 2C-values for both species given in Nagl et al. (loc. cit.) agree with those reported by Ayonoadu (1974), but are around twice that reported in Ingle et al. (1975) and Arumuganathan and Earle (1991). The basis of this discrepancy is unclear, so C-values for these Phaseolus taxa should be viewed with caution until confirmed independently.

(bo) The value for Sambucus nigra (2C = 30·5 pg ) given by Nagl et al. (1983, Ref. 457) is similar to the value of 2C = 21·8 pg for a related species, S. racemosa, reported by Nagl et al. (1979) and listed under Ref. 86 in Bennett et al. (1982). However, it is very different from 2C = 3·1 pg reported for S. nigra by Mowforth (1986) and listed under Ref. 158 in Bennett and Smith (1991). The cause for the discrepancy remains unclear, so C-values for S. nigra should be used with caution until confirmed independently.

(bp) Baranyi et al. (1996, Ref. 458) investigated C-values in 75 accessions of four wild Pisum species. Results were given as percentages relative to P. sativum ‘Kleine Rheinländerin’ ( = 100 %) which was used as the calibration standard. To convert these into absolute DNA amounts the 4C-values were multiplied by the value of Pisum sativum ‘Kleine Rheinländerin’ of 17·68 pg (Greilhuber and Ebert, 1994) and then divided by 100.

Pisum fulvum was homogeneous in DNA amount size (4C = approx. 19·3 pg), but wide variation was seen between accessions of the other species studied (P. abyssinicum, P. humile and P. elatius). This variation was interpreted to show that these taxa with variable genome sizes were genetically heterogeneous, suggesting that the current species delimitations did not reflect the true biological species groups adequately. Only the smallest and largest C-values for each of these species are listed in the Appendix.

(bq) Punina and Alexandrova (1992, Ref. 459) estimated DNA amounts in 11 Paeonia species but gave the results as percentage values relative to P. caucasica. Since Mulry and Hanson (pers. comm. 1999) had estimated the 4C DNA value of this species as 65·2 pg (see entry number 602 in Bennett et al., 2000), the relative percentage values given in Punina and Alexandrova (loc. cit.) were converted into absolute DNA amounts by multiplying by 0·652.

(br) A PhD thesis by Shi (1991) gave a 1C-value of 0·15 pg for two accessions of diploid Brachypodium distachyon (2n = 10), plus values for four other Brachypodium species. Later, Shi et al. (1993, Ref. 460) gave the 1C-value for diploid B. distachyon as 1C = 0·3 pg, but cited the PhD thesis (Shi, 1991) as the source for this figure. In order to confirm which was correct, Clive Stace kindly supplied seed of one original accession (B306) and RBG, Kew estimated its DNA amount as 0·36 pg using Oryza sativa ‘IR36’ (4C = 2·02 pg) as a calibration standard (see entry number 161a in the present Appendix). This was much closer to the value in Shi et al. (1993). As C-values for four other Brachypodium species in Shi (1991) may also be under-estimates, they are therefore not included in the present Appendix, and should be viewed with caution until confirmed independently.

Original references for DNA values

Named references in the ‘Notes to the Appendix’ are given in ‘Literature cited’. Only numbered references of original sources of species DNA values in the Appendix (column 13) are given in the Key below.

378. Hanson L, McMahon KA, Johnson MAT, Bennett MD. 2001. First nuclear DNA C-values for 25 angiosperm families. Annals of Botany 87: 251–258.

379. Hanson L, McMahon KA, Johnson MAT, Bennett MD. 2001. First nuclear DNA C-values for another 25 angiosperm families. Annals of Botany 88: 851–858.

380. Hanson L, Brown RL, Boyd A, Johnson MAT, Bennett MD. 2003. First nuclear DNA C-values for 28 angiosperm genera. Annals of Botany 91: 1–8.

381. Leitch IJ, Hanson L. 2002. DNA C-values in seven families fill phylogenetic gaps in the basal angiosperms. Botanical Journal of the Linnean Society 140: 175–179.

382. Obermayer R, Leitch IJ, Hanson L, Bennett MD. 2002. Nuclear DNA C-values in 30 species double the familial representation in pteridophytes. Annals of Botany 90: 209–217.

383. Zonneveld BJM. 2001. Nuclear DNA contents of all species of Helleborus (Ranunculaceae) discriminate between species and sectional divisions. Plant Systematics and Evolution 229: 125–130.

384. Zonneveld BJM, Van Iren F. 2001. Genome size and pollen viability as taxonomic criteria: Application to the genus Hosta Plant Biology 3: 176–185.

385. Thibault J. 1998. Nuclear DNA amount in pure species and hybrid willows (Salix): a flow cytometric investigation. Canadian Journal of Botany 76: 157–165.

386. Torrell M, Valles J. 2001. Genome size in 21 Artemisia L. species (Asteraceae, Anthemideae): Systematic, evolutionary, and ecological implications. Genome 44: 231–238.

387. Mishiba KI, Ando T, Mii M, Watanabe H, Kokubun H, Hashimoto G, Marchesi E. 2000. Nuclear DNA content as an index character discriminating taxa in the genus Petunia sensu Jussieu (Solanaceae). Annals of Botany 85: 665–673.

388. Sen S, Sharma AK. 1990. Chromosome complements, nuclear DNA and genetic distance as measures of interrelationship in Plantago The Nucleus 33: 4–10.

389. Ohmido N, Kijima K, Akiyama Y, de Jong JH, Fukui K. 2000. Quantification of total genomic DNA and selected repetitive sequences reveals concurrent changes in different DNA families in indica and japonica rice. Molecular and General Genetics 263: 388–394.

390. Nagl W, Treviranus A. 1995. A flow cytometric analysis of the nuclear 2C DNA content in 17 Phaseolus species (53 genotypes). Botanica Acta 108: 403–406.

391. Joachimiak A, Kula A, Sliwinska E, Sobieszczanska A. 2001. C-banding and nuclear DNA amount in six Bromus species. Acta Biologica Cracoviensia Series Botanica 43: 105–115.

392. Rosato M, Chiavarino AM, Naranjo CA, Hernandez JC, Poggio L. 1998. Genome size and numerical polymorphism for the B chromosome in races of maize (Zea mays ssp. mays, Poaceae). American Journal of Botany 85: 168–174.

393. Greilhuber J, Obermayer R. 1999. Cryptopolyploidy in Bunias (Brassicaceae) revisited – a flow-cytometric and densitometric study. Plant Systematics and Evolution 218: 1–4.

394. Dimitrova D, Greilhuber J. 2000. Karyotype and DNA-content evolution in ten species of Crepis (Asteraceae) distributed in Bulgaria. Botanical Journal of the Linnean Society 132: 281–297.

395. Temsch EM, Greilhuber J. 2000. Genome size variation in Arachis hypogaea and A. monticola re-evaluated. Genome 43: 449–451.

396. Temsch EM, Greilhuber J. 2001. Genome size in Arachis duranensis: a critical study. Genome 44: 826–830.

397. Cerbah M, Mortreau E, Brown S, Siljak-Yakovlev S, Bertrand H, Lambert C. 2001. Genome size variation and species relationships in the genus Hydrangea Theoretical and Applied Genetics 103: 45–51.

398. Hart H. 1991. Evolution and classification of the European Sedum species (Crassulaceae). Flora Mediterranea 1: 31–61.

399. Hart HT, Tomlik A, Alpinar K. 1993. Biosystematic studies in Sedum (Crassulaceae) fromTurkey. 4. The cytology of Sedum subsect. Spathulata Boriss. Acta Botanica Neerlandica 43: 289–298.

400. Geber G. 1989.Zur Karyosystematik der Lemnaceae. Ph.D. Thesis, University of Vienna, Vienna.

401. Noirot M, Barre P, Louarn J, Duperray C, Hamon S. 2002. Consequences of stoichiometric error on nuclear DNA content evaluation in Coffea liberica var. dewevrei using DAPI and propidium iodide. Annals of Botany 89: 385–389.

402. Lysák MA, Doleželova M, Horry JP, Swennen R, Doležel J. 1999. Flow cytometric analysis of nuclear DNA content in Musa Theoretical and Applied Genetics 98: 1344–1350.

403. Price HJ, Hodnett G, Johnston JS. 2000. Sunflower (Helianthus annuus) leaves contain compounds that reduce nuclear propidium iodide fluorescence. Annals of Botany 86: 929–934.

404. Ellul P, Boscaiu M, Vicente O, Moreno V, Rossello JA. 2002. Intra- and interspecific variation in DNA content in Cistus (Cistaceae). Annals of Botany 90: 345–351.

405. Cremonini R, Castiglione MR, Venora G, Blangiforti S, Losavio FP, Pignone D. 1998. Cytology of Vicia species. VI. Nuclear chromatin organization, karyomorphological analysis and DNA amount in Vicia serratifolia Jacq. Caryologia 51: 195–205.

406. Cremonini R, Miotto D, Ngu MA, Tota D, Pignone D, Blangiforti S, Venora G. 1998. Cytology of Vicia species. 5. Nuclear chromatin structure, karyomorphological analysis and DNA content in newly discovered relatives of Vicia faba L.: Vicia kalakhensis Khattab, Maxted et Bisby and Vicia eristalioides Maxted. Cytologia 63: 371–379.

407. Venora G, Blangiforti S, Castiglione MR, Black-Samuelsson S, Cremonini R. 1999. Cytology of Vicia species. VIII. Nuclear DNA contents, chromatin organization and karyotype evolution in Vicia pisiformis L. populations. Caryologia 52: 105–115.

408. Cremonini R, Funari S, Mazzuca S. 1992. Cytology of Vicia species: nuclear structure, karyological analysis and DNA content. Chromatin 1: 135–146.

409. Akpinar N, Bilaloglu R. 1997. Cytological investigations of certain species of Vicia L. Turkish Journal of Botany 21: 197–207.

410. Kamaté K, Brown S, Durand P, Bureau JM, De Nay D, Trinh TH. 2001. Nuclear DNA content and base composition in 28 taxa of Musa Genome 44: 622–627.

411. Ghosh P, Mukherjee S, Sharma AK. 2001. Cytophotometric estimation of in situ DNA content in several species of Araceae. Cytobios 105: 177–183.

412. Unal F, Callow RS. 1995. Estimation of genome size by Feulgen photometry. Turkish Journal of Botany 19: 523–529.

413. Greilhuber J, Ebert I, Lorenz A, Vyskot B. 2000. Origin of facultative heterochromatin in the endosperm of Gagea lutea (Liliaceae). Protoplasma 212: 217–226.

414. Sakamoto K, Akiyama Y, Fukui K, Kamada H, Satoh S. 1998. Characterization: genome sizes and morphology of sex chromosomes in Hemp (Cannabis sativa L.). Cytologia 63: 459–464.

415. Naranjo CA, Ferrari MR, Palermo AM, Poggio L. 1998. Karyotype, DNA content and meiotic behaviour in five South American species of Vicia (Fabaceae). Annals of Botany 82: 757–764.

416. Gammar ZG, Puech S, Zouaghi M. 1999. Flow cytometry DNA assay of Mediterranean lupins. Candollea 54: 45–56.

417. Arumuganathan K, Tallury SP, Fraser ML, Bruneau AH, Qu R. 1999. Nuclear DNA content of thirteen turfgrass species by flow cytometry. Crop Science 39: 1518–1521.

418. Ali HBM, Meister A, Schubert I. 2000. DNA content, rDNA loci, and DAPI bands reflect the phylogenetic distance between Lathyrus species. Genome 43: 1027–1032.

419. Brandizzi F, Grilli Caiola M. 1996. Quantitative DNA analysis in different Crocus species (Iridaceae) by means of flow cytometry. Giornale Botanico Italiano 130: 643–645.

420. Mukherjee S, Sharma AK. 1993. Cytophotometric DNA estimation in Luzula species. Current Science 65: 987–989.

421. Asif MJ, Mak C, Othman RY. 2001. Characterization of indigenous Musa species based on flow cytometric analysis of ploidy and nuclear DNA content. Caryologia 54: 161–168.

422. Chaudhuri D, Sen S. 2001. Genotypic diversity in species of Scilla Cytobios 104: 75–81.

423. Chung J, Lee JH, Arumuganathan K, Graef GL, Specht JE. 1998. Relationships between nuclear DNA content and seed and leaf size in soybean. Theoretical and Applied Genetics 96: 1064–1068.

424. Cros J, Combes MC, Chabrillange N, Duperray C, Desangles AM, Hamon S. 1995. Nuclear DNA content in the subgenus Coffea (Rubiaceae): inter- and intra-specific variation in African species. Canadian Journal of Botany 73: 14–20.

425. Hartman TPV, Jones J, Blackhall NW, Power JB, Cocking EC, Davey MR. 2000. Cytogenetics, molecular cytogenetics and genome size in Leucaena (Leguminosae, Mimosideae). In: Guttenberger H, Borzan Z, Schlarbaum SE, Hartman TPV, eds. Cytogenetic studies of forest trees and shrubs – review, present status, and outlook on the future. Zvolen, Slovakia: Arbora Publishers, 57–70.

426. Kayim M, Koç NK, Rokka VM. 1998. Variation of the nuclear DNA content of species of subtribe Citrinae (Rutaceae). Hortscience 33: 1247–1250.

427. Boscaiu M, Vicente O, Ehrendorfer F. 1999. Chromosome numbers, karyotypes and nuclear DNA contents from perennial polyploid groups of Cerastium (Caryophyllaceae). Plant Systematics and Evolution 218: 13–21.

428. Lysák MA, Rostková AR, Dixon JM, Rossi G, Doležel J. 2000. Limited genome size variation in Sesleria albicans Annals of Botany 86: 399–403.

429. Obermayer R, Greilhuber J. 2000. Genome size in Hedera helix L.—a clarification. Caryologia 53: 1–4.

430. Caceres ME, De Pace C, Mugnozza GTS, Kotsonis P, Ceccarelli M, Cionini PG. 1998. Genome size variations within Dasypyrum villosum: correlations with chromosomal traits, environmental factors and plant phenotypic characteristics and behaviour in reproduction. Theoretical and Applied Genetics 96: 559–567.

431. Blanco A, Simeone R, Resta P, DePace C, Delre V, Caccia R, Mugnozza GTS, Frediani M, Cremonini R, Cionini PG. 1996. Genomic relationships between Dasypyrum villosum (L) Candargy and D. hordeaceum (Cosson et Durieu) Candargy. Genome 39: 83–92.

432. Rayburn AL, Biradar DP, Bullock DG, Nelson RL, Gourmet C, Wetzel JB. 1997. Nuclear DNA content diversity in Chinese soybean introductions. Annals of Botany 80: 321–325.

433. Gallois A, Burrus M, Brown S. 1999. Evaluation of the nuclear DNA content and GC percent in four varieties of Fagus sylvatica L. Annals of Forest Science 56: 615–618.

434. Belletti P, Marzachi C, Lanteri S. 1998. Flow cytometric measurement of nuclear DNA content in Capsicum (Solanaceae). Plant Systematics and Evolution 209: 85–91.

435. Ohri D. 1999. Cytology of Cicer songaricum Steph ex DC, a wild relative of chickpea. Genetic Resources and Crop Evolution 46: 111–113.

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PERMALINK

Nuclear DNA Amounts in Angiosperms: Progress, Problems and Prospects

M D BENNETT

I J LEITCH

Abstract

INTRODUCTION

Appendix.

PROGRESS

Improved systematic representation (species and families)

(i) First estimates for species

Fig. 1.

(ii) First estimates for families

Fig. 2.

PROBLEMS

Geographical representation and distribution

Table 1.

Plant life form

Obsolescence time bomb

Errors and inexactitudes

Genome size, ‘complete’ genome sequencing, and, the euchromatic genome

The completely sequenced genome

Weeding out erroneous data

What is the smallest reliable C-value for an angiosperm?

Fig. 3.

Table 2.

What is the minimum C-value for a free-living angiosperm and other free-living organisms?

Table 3.

PROSPECTS FOR THE NEXT TEN YEARS

Fig. 4.

Holistic genomics

APPENDIX

Notes to the Appendix

Table 4.

Table 5.

Table 6.

Original references for DNA values

LITERATURE CITED

ACTIONS

PERMALINK

RESOURCES

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