The Families of Flowering Plants
The Families of Flowering Plants |
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#1. Alternatively <alternative family names>/
An attempt has been made via this character to distinguish synonyms denoting (more or less) the same taxonomic concept as the name used here. It is also intended to encompass spelling variants of the ‘accepted’ name. Family names constituting nomenclaturally valid alternatives to those used are indicated (‘nom. altern.’). Combine characters 1 and 3 to list all the synonyms entered for a family.
‘Synonym’: a name rejected because of different taxonomic interpretation, or under the rules of nomenclature.
#2. ~ <Sensu lato family to which sometimes referred (see Notes)>/
~ (‘alternatively’) is here used to indicate ‘sometimes not unreasonably referred to’. The sensu lato interpretations indicated often correspond with Cronquist’s (1981) treatment, but to an increasing extent refer to more recent cladistic pronouncements (few of which are accompanied by any serious attempt to provide revised family descriptions). Sound practical reasons for adopting tenable sensu stricto interpretations in the present context are given in Watson and Dallwitz (1991).
#3. Including <family names treated here as synonyms>/
These nomenclatural data are a compilation, containing no original research. The main source of synonyms has been Airy Shaw (1973), who conducted genuine nomenclatural research and aimed to include ‘every published family name from the appearance of the Genera Plantarum of Jussieu in 1789’; and the data have subsequently been checked against Gunn et al. (1992 onwards). There no doubt remain numerous omissions, and it would be impossible to obtain comprehensive coverage of variant spellings. The latter, including misprints, insertion or omission of accents, deliberate corrections and intended etymological ‘improvements’, inevitably pose problems in computer searches. Nevertheless, if used properly this package should permit referal to appropriate descriptions of most of the family names used (as opposed to appearing in lists of synonyms) in twentieth century floras and research publications. Combine characters 1 and 3 to list all the synonyms entered for a family.
‘Synonym’: a name rejected because of different taxonomic interpretation, or under the rules of nomenclature.
#4. Excluding <families excluded - intended cross-referencing extensive but incomplete>/
#5. <Habit:>/
1. trees/
2. ‘arborescent’/
3. shrubs <currently including ‘subshrubs’>/
4. lianas/
5. herbs/
‘Tree’: a relatively large, perennial woody plant with an evident trunk (bole), nearly always producing bark (the visual syndrome nearly always reflecting persistent vascular and cork cambia).
‘Arborescent’: perennial, attaining more or less treelike aspect without exhibiting all the ‘tree’ characteristics.
‘Shrub’: a woody, branching perennial with persistent above-ground parts, of smaller stature than a tree and without a well expressed trunk.
‘Subshrub’: an under-shrub, or small shrub which may have partially herbaceous stems.
‘Liana’ or ‘liane’: a woody climber.
‘Herb’: a plant without persistent above-ground stems.
#6. <Stems and/or leaves, whether noticeably laticiferous etc. (a ‘casual’ character - see ‘laticifers’):>/
1. laticiferous/
2. with coloured juice/
3. non-laticiferous and without coloured juice <<implicit>>/
#7. <Essential oils in vegetative parts:>/
1. bearing essential oils/
2. without essential oils/
#8. <Vegetative parts, whether resinous:>/
1. resinous/
2. not resinous/
#9. <Plant form: whether ‘normal’, ‘switch’ or very peculiar:>/
1. ‘normal’ plants <with more or less normal stems, leaves and roots> <<implicit>>/
2. switch-plants <with leaves more or less reduced, the leaf blade functions transferred to specialized stems and/or petioles>/
3. plants of very peculiar vegetative form <i.e., form virtually unrecognisable as that of an Angiosperm>/
‘Switch-plants’: plants with the functions normally fulfilled by leaves transferred to chlorophyllous, more or less modified shoots and/or petioles (the true leaves more or less reduced or lacking).
‘Cladode’: a phylloclade of one internode.
‘Phyllode’: a flat, dilated petiole fulfilling the functions of a leaf.
‘Phylloclade’: a stem modified to function as a leaf or leaves.
#10. <Switch-plant type:>/
1. with the principal photosynthesizing function transferred to <non-succulent> stems <including phylloclades and cladodes>/
2. phyllodineous <with flattened petioles>/
3. ‘cactoid’ <with succulent, photosynthetic stems>/
#11. <Nature of ‘very peculiar vegetative morphology’:>/
1. thalloid/
2. filamentous/
3. fungoid/
4. no clear distinction between stems, roots and leaves, the stem bearing photosynthetic appendages equipped with animal-trapping bladders <Utricularia>/
5. neotenic/
#12. Leaves <presence>/
1. well developed <<implicit>>/
2. much reduced/
3. absent/
‘Reduced leaves’ denotes leaves present and identifiable as such, but more or less vestigial. Not applied to phyllodes and cladodes (though they commonly occur in association with the latter — e.g. in Ruscus).
#13. Plants <mature, whether with roots>/
1. with roots <<implicit>>/
2. rootless/
#14. Plants <succulence>/
1. succulent/
2. non-succulent <<implicit>>/
This composite character was introduced (a), as a ‘colloquial’ improver in natural language descriptions, and (b) as an aid in seeking likely additional records for CAM photosynthesis.
#15. Plants <nutrition>/
1. autotrophic <<implicit>>/
2. saprophytic/
3. partially parasitic/
4. totally parasitic/
5. ‘carnivorous’ <including insectivorous>/
#16. Parasitic on <host parts>/
1. roots of the host/
2. aerial parts of the host/
#17. Trapping mechanism <whether active or passive>/
1. active/
2. passive/
#18. The traps <type>/
1. in the form of minute (submerged) bladders/
2. consisting of ‘pitchers’/
3. constituted by sticky leaf glands, associated with subsequent, slow enclosure of the prey by movement of the blade <and/or ‘tentacles’>/
4. consisting of the curiously modified, steel-trap-like leaf blades, which spring shut when adaxial receptors are touched/
5. consisting of the sticky-glandular, non-irritable (flypaper-like) leaves/
#19. <Herbaceous plants, life form:>/
1. annual/
2. biennial/
3. perennial/
‘Annual’: plants progressing from germination to seed production and perishing, within that period.
‘Biennial’: a plant requiring two years to complete its life-cycle, growing vegetatively in the first, then flowering, fruiting and perishing in the second.
‘Perennial’: a plant which persists and continues growth for several to many years.
#20. <Plants, presence of leaf rosettes:>/
1. with a basal <rosette or> aggregation of leaves <‘rosulate’, not necessarily ‘acaulescent’>/
2. with neither basal nor terminal aggregations <rosettes> of leaves/
3. with terminal aggregations <rosettes> of leaves/
#21. Young stems <whether breaking easily at nodes>/
1. breaking easily at the nodes/
2. not breaking easily at the nodes <<implicit>>/
#22. <Height of plants:>/
m high/
#23. <Perennial herbs, mode of perennation:>/
1. bulbaceous/
2. cormous/
3. rhizomatous/
4. tuberous/
‘Bulb’: underground bud, modified for vegetative reproduction/perennation.
‘Corm’: bulblike, fleshy, rooted stem or stem base, associated with vegetative reproduction/perennation.
‘Rhizome’: perennial, prostrate or oblique, more or less subterranean, more or less dorsiventral stem, producing roots below and leaves and/or stems apically or above and apically.
‘Tuber’: a thick, short, swollen vegetative part concerned with food storage (may represent either root or stem).
Unsatisfactory character state definitions, which in practice are not properly exclusive. ‘Tuberous or not’ and specification of kinds of tubers await resolving into separate characters.
#24. Rhizome and root tissues <red pigment, presence>/
1. brightly red-pigmented/
2. not red-pigmented <<implicit>>/
#25. <Whether epiphytic or climbing:>/
1. self supporting <<implicit>>/
2. epiphytic/
3. climbing/
#26. <Types of climbers:>/
1. stem twiners/
2. petiole twiners/
3. tendril climbers/
4. sucker climbers/
5. root climbers/
6. scrambling/
#27. <Direction of twining, as viewed from above (see Notes):>/
1. twining clockwise/
2. twining anticlockwise/
Preliminary data only, from Darwin (1882).
Clockwise (viewed from above and/or as the stem approaches you) = ‘ascending a vertical support from right to left when viewed from the side’ = Darwin’s ‘following the sun’. ‘Dextrorotatory’ = anticlockwise.
#28. Stem growth <whether conspicuously sympodial>/
1. conspicuously sympodial/
2. not conspicuously sympodial <<implicit>>/
#29. <Trees/shrubs/arborescents, form:>/
1. leptocaul/
2. pachycaul/
‘Pachycaul’: thick-stemmed, not ‘twiggy’ (unbranched, or the branches few and thick).
‘Leptocaul’ (= leptocladous): twiggy, with abundant, relatively slender branches.
#30. <Ecological type:>/
1. hydrophytic/
2. helophytic/
3. mesophytic/
4. xerophytic/
‘Hydrophytes’: plants normally living with the vegetative parts submerged or floating in water, or only partially emergent.
‘Helophytes’: marsh plants.
‘Mesophytes’: plants avoiding extremes of moisture and drought — in habitats intermediate between those of hydrophytes and xerophytes.
‘Xerophytes’: plants which normally subsist in habitats affording relatively little moisture. Usually exhibiting one or more recognisable ‘xeromorphic’ features, which include extreme hairiness; thick cuticles; harsh, pungent or rolled leaves with much sclerenchyma; spines, prickles and switch-plant properties (q.v.); succulence.
#31. <Hydrophytic habitat:>/
1. marine/
2. non-marine <<implicit>>/
#32. <Hydrophytes, plant habit:>/
1. free floating/
2. rooted <to the substrate> <<to be implicit>>/
#33. Leaves <of rooted hydrophytes, habit>/
1. submerged/
2. emergent/
3. floating/
#34. <Whether conspicuously heterophyllous:>/
1. heterophyllous/
2. not heterophyllous <<to become implicit?>>/
‘Heterophyllous’: having leaves of two (or more) conspicuously different forms. Exclusive of prophylls, bud-scales etc., and of the relatively minor variations associated with normal annual-incremental leaf spectra.
#35. Leaves <longevity>/
1. evergreen/
2. deciduous/
Of trees and shrubs: a loose but meaningful character, imprecisely expressing differences regarding presence/absence and/or performance of leaf abscission zones.
‘Deciduous’ implies seasonally synchronized leaf fall resulting in a leafless condition in winter, and the leaves are usually non-leathery.
‘Evergreens’ are leafy all year round, and the leaves are often more or less leathery.
#36. Leaves <maximum size as reflected by their greatest dimension: a ‘casual’ character (see Notes)>/
1. minute <5mm or less>/
2. small <3mm to 3cm>/
3. medium-sized <2 to 30cm>/
4. large <15 to 50cm>/
5. very large <greater than 30cm>/
‘Size’ here denotes a ‘colloquial’ character, contributing helpful but imprecisely defined adjectives to some of the natural language descriptions. It is frequently left uncoded. Reliable data on dimensions (lengths, widths, diameters) are not readily available for the larger families, and their practical value would scarcely justify the effort required to obtain them. On the other hand, it is absurd to ignore the identificatory potential of extreme size differences, especially in connection with small families. Liberal overlaps in the attempted state definitions indicate the need for extreme caution in their application during identifications, and the advisability of entering broadly inclusive ranges.
#37. Leaves <insertion>/
1. alternate/
2. opposite/
3. whorled/
‘Opposite’: the leaves in pairs, the members of each pair on the same level as one another and on opposite sides of the stem.
‘Alternate’: the leaves neither manifestly ‘opposite’ nor genuinely whorled. A loose term in widespread taxonomic usage, reflecting failure to describe phyllotaxy more precisely. Implies ‘spiral or distichous’.
‘Whorled’: the leaves (or structures reasonably interpretable as such) disposed in rings of three or more per node.
#38. Leaves <phyllotaxy, when alternate>/
1. spiral/
2. distichous/
3. four-ranked/
4. tristichous/
‘Spiral’: disposed on the axis such that an imaginary line joining the bases of successive, adjacent members constitutes a spiral.
‘Distichous’: borne in two longitudinal ranks, these on opposite sides of the axis.
‘Tristichous’: borne in three longitudinal ranks.
#39. Leaves <whorled, number per whorl>/
per whorl/
#40. Leaves <mature form of lamina>/
1. flat/
2. folded/
3. rolled/
4. terete/
#41. Leaves <texture>/
1. ‘herbaceous’ <<to become implicit?>>/
2. leathery/
3. fleshy/
4. membranous/
5. modified into spines/
#42. Leaves <insertion, distance>/
1. imbricate/
2. not imbricate/
#43. Leaves <whether petiolate>/
1. petiolate <including ‘pseudopetiolate’>/
2. subsessile/
3. sessile/
4. perfoliate/
Here applied indiscriminately to Monocots and Dicots; i.e. ignoring for practical purposes the rather compelling morphological proposition that typical Monocot leaves represent phyllodes, and treating the ‘pseudopetioles’ of bamboos, palms, etc. as petioles.
‘Perfoliate’: when the stem appears to pass through the blade of the leaf.
#44. Leaves <bases>/
1. connate/
2. not connate <<implicit>>/
‘Connate’: united, congenitally or subsequently. Here, referring to opposite or whorled leaves having their bases joined around the supporting axis.
#45. Leaves <insertion: states and data need attention, especially in relation to monocotyledons - in forms with secondary thickening, currently refers to the primary insertion only>/
1. <more or less> sheathing/
2. non-sheathing/
#46. Leaf sheaths <form>/
1. tubular/
2. not tubular/
#47. Leaf sheaths <whether margins joined>/
1. with free margins <<implicit>>/
2. <at least partially tubular> with <at least partially> joined margins/
#48. Leaves <whether gland-dotted (pellucid-punctate)>/
1. gland-dotted/
2. not gland-dotted <<implicit>>/
This refers to internal, translucent glands, readily observable in anatomical sections or when the blade is viewed against a strong light. It does not refer to glandular trichomes (‘leaves gland-dotted below’, etc.).
#49. Leaves <odour>/
1. aromatic/
2. without marked odour <<implicit>>/
3. foetid/
#50. Leaves <primary orientation, plane of flattening>/
1. edgewise to the stem <flattened at right angles to the ‘normal’ plane, vertical or ‘ensiform’ - e.g. Iris, Eucalyptus (exclusive of reorientation by twisting)>/
2. with ‘normal’ orientation <flattened in the ‘normal’ plane> <<implicit>>/
#51. Leaves <simple or compound>/
1. simple <without subdivision into leaflets; excluding ‘unifoliolate’>/
2. compound <comprising leaflets, or exhibiting evidence of representing a single leaflet>/
#52. Leaves <whether peltate>/
1. peltate/
2. not peltate <<unreliably implicit>>/
‘Peltate’: having the petiole attached to the lower surface of the blade, instead of to its end or margin.
#53. Leaves <whether pulvinate>/
1. pulvinate/
2. epulvinate/
Leaf ‘pulvinate’: having a swelling on the petiole (usually at its base), of the kind often associated with movement. Pulvinate/epulvinate is a distinction of demonstrated taxonomic utility (and classificatory value), for example in Leguminosae; but it has been under-utilized, and is rarely mentioned in published descriptions.
#54. Leaves <compound form>/
1. unifoliolate <see Notes>/
2. bifoliolate/
3. ternate <trifoliolate>/
4. pinnate/
5. palmate/
6. bipinnate/
7. multiply compound <e.g. tripinnate, digitate with pinnate divisions>/
Leaf ‘unifoliolate’: presumed (by inference from morphological peculiarities and/or inference from taxonomic relationships) to represent a compound leaf ‘reduced’ to a single leaflet (plus petiole). Typically indicated by a distinct joint at junction of blade and petiole, or by a pulvinus at the base of the blade (i.e. instead of, or in addition to, the more usual basal one).
#55. Leaflets <whether pulvinate>/
1. pulvinate/
2. epulvinate/
Leaflet ‘pulvinate’: having a swelling on the petiolule, of the kind often associated with movement. Little information seems to be available on this potentially interesting character.
#56. <Leaf> lamina <inversion>/
1. <consistently> inverted/
2. <consistently> twisted through 90 degrees/
3. neither <consistently> inverted nor twisted through 90 degrees <<implicit>>/
#57. <Leaf> lamina <simple, dissection>/
1. dissected <-fid, -sect, lobed etc.>/
2. entire <i.e. not dissected - exclusive of the margin>/
#58. <Leaf> lamina <symmetry>/
1. conspicuously asymmetric/
2. not <generally> conspicuously asymmetric <<implicit>>/
#59. <Leaf> lamina <simple, undissected, form>/
1. setaceous/
2. acicular/
3. linear/
4. lanceolate/
5. oblanceolate/
6. oblong/
7. ovate/
8. obovate/
9. orbicular/
10. subulate/
Unsatisfactory character delimitation and inadequate character state definitions in relation to leaf morphology reflect the fact that it is easier to locate elegant glossologies than to obtain useable comparative data.
#60. <Leaf> lamina <simple, dissected, form>/
1. <once> pinnatifid/
2. palmatifid/
3. runcinate/
4. spinose/
5. <more or less> finely dichotomously dissected/
6. much-divided <e.g. bipinnately, tripinnately or digitate-pinnately dissected>/
#61. <Leaf> lamina <basic (primary) vein layout: in compound leaves, refers to the entire leaf, not individual leaflets>/
1. one-veined/
2. pinnately veined/
3. palmately veined/
4. parallel-veined/
Precisely defined terms abound, offering the choice of Greek or Latin roots, but their practical implementation is impeded by absence of comparative data.
#62. <Leaf> lamina <venation system, closed or open>/
1. <conspicuously> cross-venulate <currently very inclusive - including ‘reticulate’>/
2. without <conspicuous> cross-venules/
#63. <Leaf> lamina <form, base>/
1. auriculate at the base/
2. cordate/
3. hastate/
4. sagittate/
5. attenuate at the base/
6. cuneate at the base/
7. oblique at the base/
8. rounded at the base/
#64. Leaves <whether ligulate>/
1. ligulate/
2. eligulate <<implicit>>/
‘Ligule’: a collarlike or tonguelike projection near the junction of sheath or petiole and blade.
#65. Leaves <whether stipulate: hardly ever applied to Monocots>/
1. stipulate/
2. exstipulate <including ‘opposing leaves joined by a raised line’>/
‘Stipules’: appendages of a leaf, borne one on each side of its insertion on the stem (usually in association with a petiole).
‘Stipulate’: having stipules, or structures recognizable as modifications (spines, glands etc.), fusion products or vestiges of them.
Stipules have protective (sometimes including secretory) functions during shoot and leaf development. They are commonly associated with trilacunar nodes, deriving their vascular supply via branches from the lateral leaf traces. In mature material, stipules even when adnate to the petiole are frequently readily recognisable as such (and distinguishable from mere winging of the petiole) through possessing separate vascular supplies, different textures, or abscission zones distinct from those of the associated leaves. However, small stipules often abscise early, so that detecting their ‘presence’ in mature material often depends on recogning their scars. Furthermore, is sometimes hard to distinguish stipules from leaflets or leaf sheaths, or even in some families (notably Rubiaceae) from leaves; and interpretive problems are posed by other structures (e.g. glands) which may or may not be legitimately considered stipular. Convincing solutions to questionable cases require detailed comparative studies, and are inaccessible for routine identifications.
The present compilation no doubt contains errors reflecting failure to distinguish ‘absent’ from ‘caducous’. These can be corrected as they come to light. Meanwhile, ‘presence or absence of stipules’ is a perfectly good character in the context of INTKEY identifications, provided intelligent use is made of the TOLERANCE facility.
#66. Stipules <type>/
1. interpetiolar <implies association/concrescence of stipules from the different members of a leaf pair or whorl>/
2. intrapetiolar <axillary or on the petiole>/
#67. Stipules <intrapetiolar, freedom>/
1. free of one another/
2. concrescent/
#68. Stipules <concrescent, whether sheathing>/
1. ochreate <joined to form a cup-shaped structure round the stem>/
2. not ochreate <<implicit>>/
#69. Stipules <with or without colleters>/
1. with colleters <secreting mucilage>/
2. without colleters <<implicit>>/
‘Colleters’. Glands secreting mucilage, which commonly occur on buds. Their occurrence on stipules is supposed to have great diagnostic value among families around Rubiaceae, but the data are unreliable. As usual, there is a dearth of information on observed absence.
#70. Stipules <form>/
1. scaly/
2. leafy/
3. spiny/
4. represented by glands/
#71. Stipules <persistence>/
1. caducous/
2. persistent/
‘Stipules’: appendages of a leaf, borne one on each side of its insertion.
‘Caducous’: falling off early.
#72. Axillary scales <intravaginal squamules, presence>/
1. present/
2. absent <<implicit>>/
#73. Lamina margins/
1. entire/
2. crenate/
3. serrate/
4. dentate/
#74. Lamina margins/
1. flat/
2. revolute/
3. involute/
‘Revolute’: curled, rolled or bent outwards, to enclose or partially enclose (part of) the outer (abaxial) surface.
‘Involute’: curled, rolled or bent inwards, to enclose or partially enclose (part of) the inner (adaxial) surface.
#75. Vegetative buds <investment>/
1. scaly/
2. not scaly/
#76. <Vegetative> prophylls <number - no data entered>/
#77. <Vegetative> prophylls <orientation - no data entered>/
1. adaxial/
2. lateral/
#78. Leaves <basal meristem>/
1. with a persistent basal meristem, and basipetal development/
2. without a persistent basal meristem/
There seems to be little hard comparative information on occurrence and location of ‘intercalary meristems’ (i.e. as exemplified and clearly demonstrated in Gramineae). Judging from external appearances, however, their presence may represent an important ontogenetic difference between typical Dicots and most Monocots.
#79. Vernation <(ptyxis) of leaves>/
1. conduplicate <folded lengthwise>/
2. plicate <folded like a fan>/
3. involute <margins rolled inwards>/
4. revolute <margins rolled outwards>/
5. reclinate <apex bent to base>/
6. convolute <rolled spirally>/
7. circinnate <coiled inwards from the tip>/
#80. Vernation <(ptyxis) of leaves: a redundant character, covering shortage of data for the other states in the preceding one>/
1. circinnate <coiled inwards from the tip>/
2. not circinnate <<implicit>>/
‘Circinnate’: coiled basipetally, with the apex near the centre of the coil, as in fern fronds.
#81. Leaves <compound, developmental type>/
1. becoming compound by necrosis/
2. becoming compound by ontogenetically predetermined splitting/
3. becoming compound from primordial lobes <<implicit>>/
#82. Domatia <records: from Brewer, Clifford and Gregory 1990 (no explicit negatives)>/
1. occurring in <at least some representatives of> the family/
2. never explicitly mentioned for the family <<implicit>>/
‘Domatia’: structures occurring in the vein axils on the abaxial (under) surface of leaves, frequently inhabited by mites or ants. Data from Brouwer, Clifford and Gregory (1990: no explicit negatives).
#83. Domatia manifested as <types>/
1. pits/
2. pockets/
3. hair tufts/
#84. Plants <presence of laticifers comprising cells or coenocytes>/
1. with laticifers/
2. without laticifers <implicit when anatomy ‘known’>/
Information mainly from Metcalfe and Chalk (1950), Metcalfe (1979). Data on this character are unreliable through lack of direct information on families observed to exhibit ‘negatives’. When unmentioned in anatomical family descriptions, it has perforce been treated as implicitly exhibiting the negative state. Families undescribed or poorly known anatomically have been tagged as ‘unknown’.
#85. The laticifers <location: very incompletely recorded>/
1. in leaves/
2. in stems/
3. in roots/
4. in flowers/
5. in the fruits/
#86. Plants <presence of ‘crystal sand’>/
1. with ‘crystal sand’/
2. without ‘crystal sand’ <implicit when anatomy ‘known’>/
Information mainly from Metcalfe and Chalk (1950) and (1979). Data on this character are unreliable through lack of direct information on families observed to exhibit ‘negatives’. When unmentioned in anatomical family descriptions, it has perforce been treated as implicitly exhibiting the negative state. Families undescribed or poorly known anatomically have been tagged as ‘unknown’.
#87. Plants <whether exhibiting silica bodies: recorded only for Monocots>/
1. with silica bodies/
2. without silica bodies <implicit for Monocots, where anatomy known>/
#88. Chlorenchyma <of leaves and/or stems, presence of ‘peg cells’>/
1. including ‘peg cells’/
2. without ‘peg cells’ <<to become implicit for Monocots>>/
#89. Accumulated starch <type, in seeds and/or vegetative parts: Czaja 1978 Taxon summary (book not yet used)>/
1. exclusively ‘pteridophyte type’/
2. other than exclusively ‘pteridophyte type’/
Leaf
#90. Leaves <presence of ‘pearl glands’>/
1. with ‘pearl glands’/
2. without ‘pearl glands’ <implicit when anatomy ‘known’>/
#91. Extra-floral nectaries <presence on leaves>/
1. present <<specify>>/
2. absent/
Data as yet very incompletely extracted from Metcalfe and Chalk (1950, 1979).
#92. Hydathodes <presence on leaves>/
1. present/
2. absent/
#93. <Lamina> epidermis <differentiation>/
1. conspicuously differentiated into ‘long’ and ‘short’ cells/
2. without differentiation into ‘long’ and ‘short’ cells <<implicit>>/
#94. <Lamina> epidermis <of plants exhibiting silica bodies, whether containing silica bodies>/
1. containing silica bodies/
2. without silica bodies <<implicit>>/
#95. <Lamina> epidermis <presence of crystal idioblasts>/
1. with crystal idioblasts/
2. without crystal idioblasts/
#96. Abaxial epidermis <of the lamina, whether papillose>/
1. papillose/
2. not papillose/
#97. Mucilaginous epidermis <of the lamina, presence>/
1. present/
2. absent/
#98. <Foliar> stomata <presence>/
1. present <<to become implicit>>/
2. absent/
#99. <Foliar> stomata <distribution on the lamina>/
1. mainly confined to one surface/
2. on both surfaces/
#100. <Foliar> stomata <form>/
1. anomocytic <ranunculaceous>/
2. anisocytic <cruciferous>/
3. paracytic <rubiaceous>/
4. tetracytic/
5. diacytic <caryophyllaceous>/
6. cyclocytic/
7. actinocytic/
8. axillocytic <surrounded by a single, doughnut-shaped subsidiary cell>/
Data mainly from Metcalfe and Chalk (1950, for Dicots); Dahlgren, Clifford and Yeoh (1985, for Monocots).
#101. <Stomatal> guard-cells <of leaves, shape>/
1. ‘grass type’ <dumb-bell shaped>/
2. not ‘grass type’ <<implicit>>/
#102. Hairs <foliar, presence>/
1. present/
2. absent/
#103. <Leaf> hairs <type>/
1. eglandular/
2. glandular/
#104. <Leaf> hairs <form>/
1. unicellular/
2. multicellular/
#105. Unicellular <leaf> hairs <whether branched>/
1. branched/
2. unbranched/
#106. Multicellular <leaf> hairs <uni- or multiseriate>/
1. uniseriate/
2. multiseriate/
#107. Multicellular <leaf> hairs <whether branched>/
1. branched/
2. unbranched/
#108. Complex <leaf> hairs <presence>/
1. present/
2. absent/
#109. Complex <leaf> hairs <specialised forms>/
1. peltate/
2. stellate/
3. clavate/
4. capitate/
#110. Urticating hairs <presence on leaves>/
1. present/
2. absent <<implicit>>/
#111. Adaxial hypodermis <presence in lamina>/
1. present <or the epidermis ‘horizontally divided’>/
2. absent/
#112. <Leaf> lamina <general layout in transverse section>/
1. dorsiventral/
2. isobilateral/
3. centric/
‘Dorsiventral’: more or less flattened , and anatomically different adaxially and abaxially. Commonly manifested via conspicuously different epidermes and/or adaxial palisade versus abaxial spongy mesophyll.
‘Isobilateral’: capable of being divided into two similar halves; i.e., more or less flattened (adaxially/abaxially, or laterally), without noticeable adaxial/abaxial or lateral anatomical distinction. Perhaps a new term, ‘isobifacial’, should be introduced.
‘Centric’: with more or less radial anatomical organization.
The leaves characteristic of many Monocots which are sometimes described as ‘unifacial’ here fall under ‘isobilateral’ or ‘centric’, according to the extent of flattening.
#113. <Leaf> lamina <whether with secretory cavities, as distinct from laticifers or secretory cells>/
1. with secretory cavities/
2. without secretory cavities <implicit when leaf anatomy ‘known’>/
#114. Secretory cavities <of leaves>/
1. containing oil/
2. containing mucilage/
3. containing resin/
4. containing latex/
#115. Secretory cavities <of leaves, development>/
1. schizogenous/
2. lysigenous/
#116. Cystoliths <presence in leaves>/
1. present/
2. absent <<implicit>>/
#117. The mesophyll <whether with etherial oil cells>/
1. with spherical etherial oil cells/
2. without etherial oil cells <implicit when anatomy ‘known’>/
#118. The mesophyll <whether with mucilage cells: few negatives coded>/
1. containing mucilage cells/
2. not containing mucilage cells <implicit for Dicots only, when anatomy ‘known’>/
#119. The mesophyll <sclerenchymatous idioblasts: few negatives coded>/
1. with sclerencymatous idioblasts/
2. without sclerenchymatous idioblasts <implicit when anatomy ‘known’>/
#120. The mesophyll <presence of calcium oxalate crystals>/
1. containing calcium oxalate crystals/
2. without calcium oxalate crystals/
Data mainly from Metcalfe and Chalk (1950, for Dicots — incompletely encoded); Dahlgren, Clifford and Yeoh (1985, for Monocots). References to crystal forms which fail to specify location (‘plants with raphides’, etc.) are commonly encountered in taxonomic and related literature: they are useless for practical purposes, and have been ignored.
#121. The mesophyll <calcium oxalate> crystals <form>/
1. raphides <i.e in bundles>/
2. druses <cluster crystals>/
3. solitary-prismatic <whatever the shape, including styloids>/
Data mainly from Metcalfe and Chalk (1950, for Dicots — incompletely encoded); Dahlgren, Clifford and Yeoh (1985, for Monocots). References to crystal forms which fail to specify location (‘plants with raphides’, etc.) are commonly encountered in taxonomic and related literature: they are useless for practical purposes, and have been ignored.
‘Raphides’: slender, needle-shaped calcium oxalate crystals, arranged parallel to each other in tight bundles, the bundles enclosed in raphide sacs. Unorientated acicular crystals are excluded by this definition.
#122. Midrib <of lamina>/
1. conspicuous/
2. not conspicuous/
#123. Midrib <of lamina, comments on vascularisation>/
#124. Main veins <of leaf lamina, excluding midribs>/
1. vertically transcurrent/
2. embedded/
#125. Minor leaf veins <presence of phloem transfer cells: data provided by B.E.S. Gunning, cf. Pate and Gunning 1969. Generic samples given as text comment>/
1. with phloem transfer cells/
2. without phloem transfer cells/
Most of the data derive from observations by J.S. Pate and B.E.S. Gunning, which were summarized taxonomically in their 1969 paper. The detailed generic figures given here are taken from the unpublished original data, provided by B.E.S. Gunning.
#126. <Foliar> vessels <presence>/
1. present/
2. absent/
Data mainly from the comprehensive compilation of Wagner (1977), and restricted to Monocots. No comparable data on Dicots have yet been located.
#127. <Foliar vessel> end-walls <form of perforation>/
1. reticulately perforated/
2. scalariform/
3. <with> simple <perforations>/
#128. Stems <internodes, solidity>/
1. with solid internodes/
2. with spongy internodes/
3. with hollow internodes/
#129. Young stems <shape in transverse section>/
1. <more or less> cylindrical/
2. oval in section/
3. tetragonal/
4. flattened/
5. triangular in section/
#130. Secretory cavities <in stems, presence (as distinct from laticifers)>/
1. present/
2. absent/
#131. Secretory cavities <of stems, contents>/
1. with oil/
2. with resin/
3. with mucilage/
4. with latex/
#132. Cork cambium <presence in stems>/
1. present/
2. absent/
#133. Cork cambium initially <position of origin in young stems>/
1. deep-seated <in the cortex or pericycle>/
2. superficial <originating subepidermally, or in the epidermis>/
Data mainly from Metcalfe and Chalk (1950 and 1979).
#134. The cortex <presence of cristarque cells, with U-shaped thickenings and containing crystals>/
1. containing cristarque cells/
2. without cristarque cells <implicit when anatomy ‘known’>/
#135. Nodes <of foliage leaves: data mostly from Sinnott 1914>/
1. unilacunar/
2. bilacunar/
3. tri-lacunar/
4. penta-lacunar/
5. multilacunar/
#136. Primary vascular tissue <of stems>/
1. in a cylinder, without separate bundles/
2. comprising a ring of bundles/
3. in two or more rings of bundles/
4. in scattered bundles/
#137. Primary vascular tissue <of stems, layout of rings or cylinders>/
1. centrifugal/
2. bicollateral/
#138. Cortical bundles <presence in stems>/
1. present/
2. absent <implicit when anatomy ‘known’>/
#139. Medullary bundles <presence in stems>/
1. present/
2. absent <implicit when anatomy ‘known’>/
#140. Internal <‘intraxylary’> phloem <presence in stem: not to be confused with ‘included’ (‘interxylary’) phloem>/
1. present/
2. absent <implicit when anatomy ‘known’>/
#141. Secondary thickening <of stems>/
1. absent/
2. <present,> developing from a conventional cambial ring <generating xylem to the inside, phloem to the outside>/
3. <present but> anomalous <concentric cambia, interxylary phloem etc.>/
#142. Secondary thickening <of anomalous stems, pattern: states incomplete>/
1. via concentric cambia/
2. from a single cambial ring/
#143. The secondary phloem <stratification: very few negatives coded>/
1. stratified into hard (fibrous) and soft (parenchymatous) zones/
2. not stratified <into fibrous and parenchymatous zones> <<implicit when applicable and anatomy ‘known’>>/
#144. ‘Included’ <‘interxylary’> phloem <presence, sensu lato (see Notes)>/
1. present/
2. absent <<to become implicit when anatomy ‘known’>>/
‘Included’ (= ‘interxylary’) phloem is here used sensu lato. For the sensu stricto version (‘included phloem derived from a single cambial ring’, cf. Carlquist 1988), apply the single/concentric cambia character to lists derived via this one.
Information mainly from Metcalfe and Chalk (1950), Carlquist (1988). Data on this character are unreliable through lack of direct information on families observed to exhibit ‘negatives’. When unmentioned in anatomical family descriptions, it has perforce been treated as implicitly exhibiting the negative state. Families undescribed or poorly known anatomically have been tagged as ‘unknown’.
#145. <Axial, stem> xylem <presence of tracheids - data from Carlquist 1988, lacking explicit negatives>/
1. with tracheids/
2. without tracheids <implicit when wood characters known>/
In Dicots, refers to secondary xylem (including wood). Refers to ‘true tracheids’, cf. Carlquist (1988); excluding ‘vascular tracheids’ and ‘vasicentric tracheids’. For the present purpose, in utilising the lists given by Carlquist (1988: the principal source of data), families listed under any of the four characters concerning tracheids, fibre tracheids, libriform fibres and ‘tile cells’ have been treated as implicitly exhibiting the negative state of those for which they go unmentioned. Families unmentioned by Carlquist for any of these wood characters have generally been recorded as ‘unknown’ for all of them, along with all families that are ‘unknown’ regarding presence/absence of secondary thickening. Families without secondary xylem (e.g. most monocots) have been treated as ‘inapplicable’ for secondary xylem characters.
#146. <Axial, stem> xylem <presence of fibre tracheids - data from Carlquist 1988, lacking explicit negatives>/
1. with fibre tracheids <= fibres with distinctly bordered pits>/
2. without fibre tracheids <implicit when wood characters known>/
See comments under ‘tracheids’.
#147. <Axial, stem> xylem <presence of libriform fibres - data from Carlquist 1988, lacking explicit negatives>/
1. with libriform fibres <= fibres with simple to minutely bordered pits>/
2. without libriform fibres <implicit when wood characters known>/
See comments under ‘tracheids’.
#148. <Axial, stem> xylem <presence of vessels>/
1. with vessels <implicit for Dicots>/
2. without vessels/
Information mainly from Metcalfe and Chalk (1950), Cheadle (1953), Cronquist (1981), Wagner (1977). In Dicots, the data seem to refer ‘mainly’ (Cheadle 1953) to secondary xylem, in Monocots to late metaxylem. See further comments under ‘end-wall perforations’ and ‘root vessels’.
#149. <Stem> vessel end-walls <form>/
1. horizontal/
2. oblique/
Information mainly from Metcalfe and Chalk (1950), Cheadle (1953), Cronquist (1981), Wagner (1977). In Dicots, the data refer ‘mainly’ (e.g. Cheadle 1953) to secondary xylem, in Monocots to late metaxylem. See further comment under ‘end-wall perforations’ (next character).
#150. <Stem> vessel end-walls <form of perforation>/
1. reticulately perforated/
2. scalariform/
3. <with> simple <perforations>/
Information mainly from Metcalfe and Chalk (1950), Cheadle (1953), Cronquist (1981) and (for Monocots) Wagner (1977). The latter compilation and its source references certainly contain good, taxonomically valid data. The information on Dicots, however, is problematic to say the least. Oft-repeated phylogenetic assertions regarding xylem evolution in Dicots, and especially those encompassing both Dicots and Monocots, derive from inaccessible or unpublished primary data on the former: none of the papers by Bailey, Frost and Cheadle listed here (see References) contain any primary data — just references to impressive slide collections, etc., observations summarized by taxonomic Orders, and phylogenetic conclusions (Bailey and Tupper 1918 yet to be seen). Valid taxonomic comparisons between Monocots and Dicots require data on Dicot primary xylem, which may not exist. Bailey (1944) examined ‘primary and adjacent first-formed secondary xylem of more than 350 representatives of 89 families’, and ‘amplified and verified these data’ by studies ‘of many additional species, genera and families’, but presents no primary data. Cheadle, speculating on ‘Independent origin of vessels’ in Monocots and Dicots, used data ‘derived (and reorganized) from Metcalfe and Chalk (1950), Solereder (1908), Engler and Prantl (1887–1915, 1925-), and numerous papers listed by these authors’. Watson has not consulted the ‘numerous papers listed’. However, Metcalfe and Chalk conscientiously organised their compiled family descriptions of axis anatomy under the headings ‘Young stem’ and ‘Wood’, and generally do not distinguish primary from secondary vessels in the former category. It comes as no surprise, therefore, to find Cheadle (1953) stating of Dicots that ‘the information for woody plants is chiefly concerned with the secondary xylem of the stem; that for most of the herbs probably includes both secondary and primary xylem of stems’. It seems to be generally accepted that in Dicots, vessels evolved first in secondary xylem, and that their further evolution and phylogenetic specialization proceeded centripetally: i.e. the primary xylem often differs from the secondary xylem in the same stem. The ‘common knowledge’ data on Dicot vessels quoted by (for example) Cronquist, and incorporated in this database, evidently derive from extensively non-comparative observations, and are of very dubious taxonomic value. The best that can be said of them is that ‘most’ of the records compiled under Dicot families ‘probably’ represent what is supposed to be the most ‘advanced’ stem xylem components; and if the supposed centripetal progression of evolutionary advancement is real, ‘stem vessels absent’ mostly implies absence from the whole stem. Likewise, ‘stem vessel end-wall perforations scalariform’ mostly implies that simple perforations are totally absent from the stem. It is intended soon to reorganize the data given here, so as to distinguish in the character list between the elements of primary and secondary stem xylem. The Dicot data will then probably be reorganized with reference to Metcalfe and Chalk, assigning any vessel information on forms without secondary thickening to primary xylem.
#151. <Stem wood> vessels <presence of vestured pits: best data (including negatives!) from Brazier and Franklin 1961>/
1. with vestured pits/
2. without vestured pits <<to become implicit when wood characters known?>>/
#152. Primary medullary rays <of young stem>/
1. wide <over 100 microns>/
2. mixed wide and narrow/
3. narrow <less than 50 microns>/
Data mainly from Metcalfe and Chalk (1979).
#153. Tile <ray> cells <presence in stems - data from Carlquist 1988>/
1. present <<specify type>>/
2. absent <implicit when wood characters known>/
‘Tile cells’: seemingly empty, upright or square ray cells of about the same height as the procumbent members, occurring in indeterminate horizontal series which are usually interspersed among the procumbent cells.
See comments under ‘tracheids’.
#154. Wood <of stem, porosity>/
1. ring porous/
2. semi-ring porous/
3. diffuse porous/
#155. Wood <organization>/
1. storied/
2. partially storied/
3. not storied/
Data extensively from Carlquist (1988); the assignments to Type given as text comment are exclusively from that source. See comments under ‘tracheids’.
#156. Wood parenchyma/
1. apotracheal <including subtypes diffuse, banded, marginal-terminal, marginal-initial, confluent>/
2. paratracheal <including subtypes scanty, vasicentric, aliform, confluent>/
Data mainly from Metcalfe and Chalk (1950).
#157. Sieve-tube plastids <type, P or S: data from Behnke, 1972–1977, Behnke and Dahlgren 1976>/
1. P-type <ergasic accumulations consisting of, or containing, proteins>/
2. S-type <ergasic accumulations without proteins, usually with starch>/
Occurrence of S-type plastids lacking starch is indicated by text comment in the descriptions.
#158. Sieve-tube plastids <P-type categories, based on protein accumulation types (Behnke, 1977)>/
1. type I <single crystalloids of various shapes and sizes, and/or irregularly arranged filaments>/
2. type II <many cuneate crystalloids>/
3. type III <one ring-shaped bundle of filaments>/
4. type IV <many, mainly polygonal crystalloids>/
#159. Pith <of stems, whether with diaphragms: few negatives coded>/
1. with diaphragms/
2. without diaphragms <implicit when anatomy ‘known’>/
#160. Pith <whether homogeneous or heterogeneous>/
1. homogeneous/
2. heterogeneous/
#161. Pith <crystalline inclusions>/
1. with crystalline inclusions/
2. without crystalline inclusions/
#162. Roots <velamen>/
1. with velamen/
2. without velamen <<implicit>>/
#163. Lateral roots <endodermis>/
1. with a conspicuous endodermis/
2. without a conspicuous endodermis/
#164. Root xylem <presence of vessels>/
1. with vessels/
2. without vessels/
Data mainly from the comprehensive compilation of Wagner (1977), and restricted to Monocots. No comparable body of data on Dicots has yet been located — do ‘vessel-less Dicots’ really lack them throughout the plant?
#165. Root vessel end-walls <form of perforation>/
1. reticulately perforated/
2. scalariform/
3. <with> simple <perforations>/
Data mainly from the comprehensive compilation of Wagner (1977), and restricted to Monocots. No comparable body of data on Dicots has yet been located, though Bailey (1944) implies that they do not exhibit the phylogenetic lag from roots to vessels that seems to have characterised xylem evolution in Monocots.
#166. Fertile flowers <sexuality, with reference to individual flowers: compiled from the ‘species’ data (see below), to facilitate INTKEY identifications>/
1. hermaphrodite/
2. functionally male/
3. functionally female/
Compiled from ‘species sexuality’ data, for easier and more reliable use in identificatory connections.
#167. Unisexual flowers <presence, with reference to complete individual plants: compiled from the ‘species’ data (see below), to facilitate INTKEY identifications>/
1. present/
2. absent <flowers seemingly all hermaphrodite>/
Compiled from ‘species sexuality’ data, for easier and more reliable use in identificatory connections.
#168. Plants <sexuality with reference to species: i.e not absolutely determinable from a single specimen (see Notes). Data extensively adapted from Yampolsky and Yampolsky 1922>/
1. <plants and fertile flowers more or less exclusively> hermaphrodite/
2. monoecious <separate male and female flowers on the same plant>/
3. andromonoecious <hermaphrodite and male flowers on the same plant, but no females>/
4. gynomonoecious <hermaphrodite and female flowers on the same plant, but no males>/
5. dioecious <plants each with male-only or female-only flowers>/
6. androdioecious <flowers male-only or hermaphrodite, on separate plants>/
7. gynodioecious <flowers female-only or hermaphrodite, on separate plants>/
8. polygamomonoecious <‘polygamous’: males-only, female-only and hermaphrodite flowers, together on the one plant>/
9. polygamodioecious <some plants with the flowers male and bisexual, others having them female and bisexual>/
Assignments to the precise monoecious and dioecious states are impracticable or unreliable from single specimens, and the details are probably extensively unreliable in any case. For INTKEY identifications, cautiously entering the possibilities 1/6/7 or (when something other than state 1 evidently obtains) 2/3/4/5/6/7/8/9 will often prove useful.
However, the preceding characters (sexuality of individual flowers, presence on specimens of unisexual flowers) have been compiled from this one, for easier and more reliable use in identificatory connections.
#169. Female flowers <presence of staminodes>/
1. with staminodes/
2. without staminodes/
#170. Gynoecium of male flowers <presence/status>/
1. pistillodial/
2. vestigial/
3. absent/
‘Gynoecium’: a collective term for the total female component of the flower, including all recognisable derivatives of this.
#171. Plants <whether viviparous>/
1. viviparous/
2. not viviparous/
#172. Plants <whether heterostylous>/
1. homostylous/
2. heterostylous/
#173. Floral nectaries <presence - relatively few data entered for Dicots>/
1. present/
2. absent/
Fairly comprehensive data have been entered for Monocots, mainly from Dahlgren and Clifford (1982); Dahlgren, Clifford and Yeoh (1985). Few entries have been made for Dicots.
#174. <Site of> nectar secretion <relatively few data entered for Dicots>/
1. from the perianth/
2. from the disk/
3. from the gynoecium/
4. from the androecium/
Fairly comprehensive data have been entered for Monocots, mainly from Dahlgren and Clifford (1982); Dahlgren, Clifford and Yeoh (1985). Few entries have been made for Dicots.
‘Disk’: an elevated or appendicular, usually more or less fleshy and nectariferous development on the receptacle internal to the perianth or stamens, which may be receptacular or staminodial (or ambiguous) in origin.
‘Gynoecium’: a collective term for the total female component of the flower, including all recognisable derivatives of this.
‘Androecium’: the total male component of the flower, including all recognisable ‘derivatives’ of this. ‘Androecial members’ thus include the stamens (q.v.), plus any staminodial structures (q.v.).
#175. Pollination <type>/
1. anemophilous/
2. entomophilous/
3. ornithophilous/
4. cheiropterophilous/
5. by water/
6. by unusual means <specified in the family descriptions>/
Many families which are obviously at least sometimes entomophilous have been left uncoded, owing to uncertainty regarding the occurrence of other conditions.
#176. Pollination <entomophilous>/
1. via beetles/
2. via hymenoptera/
3. via lepidoptera/
4. via diptera/
#177. Pollination mechanism/
1. conspicuously specialized <<comment how>>/
2. <relatively> unspecialized <<to become implicit?>>/
This character affords an opportunity to enter descriptive information of general interest, but the states are obviously imprecise and the data are of little use for identification or classification.
#178. Flowers <aggregation>/
1. solitary/
2. aggregated in ‘inflorescences’/
#179. Flowers <when solitary, whether terminal or axillary>/
1. terminal/
2. axillary/
‘Axillary’ implies from a leaf axil, rather than from a bract axil.
#180. Flowers <inflorescence, general description: data in process of development>/
1. in cymes/
2. in racemes/
3. in spikes/
4. in heads/
5. in umbels/
6. in corymbs/
7. in fascicles/
8. in panicles <including thyrses>/
9. in verticils/
10. in glomerules/
11. in catkins/
Unsatisfactory character delimitation, inadequate character state definitions and unreliable data on ‘inflorescence’ features reflect the fact that it is easier to locate erudite morphological discussions and elegant terminologies than to obtain useable comparative data.
This unsatisfactory character will eventually be subsumed into the nearby text character (infloresences — type). The terms in everyday use employed here have adequate definitions for practical application and for resolution into a hierarchy of multistate characters, but their haphazard application in published descriptions (in which, for example, different levels of aggregation are seldom mentioned) precludes effectively using them in that fashion without organizing the data de novo.
See the text character for definitions of terms
#181. The fruiting inflorescences <whether conelike (strobiloid)>/
1. conelike <compact, woody, persistent>/
2. not conelike <<implicit>>/
#182. Flowers <whether in ‘spikelets’>/
1. in ‘spikelets’/
2. not in ‘spikelets’ <<implicit>>/
‘Spikelet’ = a secondary spike (ultimate inflorescence unit), consisting of one or more reduced flowers and subtending bracts, as in grasses. A morphologically loose but very useful descriptive term, unambiguously applicable in relation to most flowering plants.
#183. The terminal inflorescence unit <whether racemose or cymose>/
1. cymose <determinate>/
2. racemose <indeterminate>/
Unsatisfactory character delimitation, inadequate character state definitions and unreliable data on ‘inflorescence’ features reflect the fact that it is easier to locate erudite morphological discussions and elegant terminologies than to obtain useable comparative data.
#184. Inflorescences <whether a scape>/
1. scapiflorous <main axis elongate, leafless>/
2. not scapiflorous <<implicit>>/
#185. Inflorescences <position>/
1. terminal/
2. axillary/
3. leaf-opposed/
4. intercalary/
5. epiphyllous/
6. cauliflorous/
#186. Inflorescences <type>/
Unsatisfactory character delimitation, inadequate character state definitions and unreliable data on ‘inflorescence’ features reflect the fact that it is easier to locate erudite morphological discussions and elegant terminologies than to obtain useable comparative data.
The following terms have been employed here. They have adequate definitions for practical application and for resolution into a hierarchy of multistate characters, but their haphazard application in published descriptions (in which, for example, different levels of aggregation are seldom mentioned) precludes effectively using them in that fashion without organizing the data de novo.
‘Catkin’ (ament): a scaly-bracted, deciduous, often pendulous or flexuous raceme or (more usually) raceme of cymules, with reduced (anemophilous), apetalous, usually unisexual flowers.
‘Corymb’: a relatively broad and short-internoded, flat-topped or convex, open, indeterminate (centripetal) inflorescence, whose outer flowers open first.
‘Cyme’: a determinate flower cluster, with central flowers opening first.
‘Fascicle’: a loose term for a condensed or close cluster, of whatever detailed construction (unresolved/unresolvable into racemose/cymose).
‘Glomerule’: a cluster of dense or compact, short-pedicelled or short-peduncled clusters or heads, with a common involucre.
‘Panicle’: an indeterminate (racemose, corymbose), compound inflorescence (the primary branches racemose, the nature of lower order branching unspecified).
‘Raceme’: a simple, elongate-internoded, indeterminate (centripetal) inflorescence with pedicelled flowers which open acropetally.
‘Thyrse’: a mixed (compound) inflorescence, in the form of a contracted or ovate panicle, the primary branching racemose (indeterminate) but the ultimate branching cymose (determinate).
‘Umbel’: A relatively broad, often flat-topped, indeterminate (centripetal) inflorescence, whose pedicels or peduncles arise from a common point and whose outer flowers open first.
‘Verticil’: a whorl of flowers about an axis (of whatever detailed organization).
#187. Inflorescences <whether involucrate>/
1. with involucral bracts/
2. without involucral bracts <<implicit>>/
‘Involucre’: one or more whorls of modified leaves or bracts subtending a cluster of flowers (or florets) or a cluster of inflorescence branches.
#188. Inflorescences <whether incorporating pseudanthia>/
1. pseudanthial/
2. not pseudanthial <<implicit>>/
‘Pseudanthium’: an aggregation of flowers superficially resembling (and perhaps fuctionally, biologically equivalent to) a single flower.
#189. The involucres <accrescence>/
1. accrescent/
2. non-accrescent <<to become implicit>>/
‘Accrescent’: increasing in size after flowering.
#190. Inflorescences <whether spatheate>/
1. spatheate/
2. espatheate <<implicit>>/
‘Spathe’: a conspicuous bract or specialized leaf, enclosing an inflorescence.
#191. Flowers <whether bracteate>/
1. bracteate/
2. ebracteate/
‘Bract’: a modified foliar structure, subtending an infloresence, inflorescence branch or pedicel. Available descriptions frequently fail to distinguish adequately between (or are confused or ambiguous about) bracts and ‘bracteoles’ (q.v.), so the compiled data are unreliable.
#192. Bracts <whether calyptrate and covering the flower bud: not to be confused with calyptrate K or C>/
1. calyptrate/
2. not calyptrate <<implicit>>/
‘Calyptrate’: forming a hood or lid over the inner parts of the flower.
#193. Flowers <whether bracteolate>/
1. bracteolate/
2. ebracteolate/
‘Bracteoles’: ultimate bracts (q.v.), borne singly, in pairs or in series directly on the pedicel. Frequently interpretable as inflorescence prophylls. Available descriptions frequently fail to distinguish adequately between (or are confused or ambiguous about) bracts and bracteoles, so the compiled data are unreliable.
#194. Flowers <size; a ‘casual’ character reflecting their greatest dimension, whether diameter or length (see Notes)>/
1. minute <3mm or less>/
2. small <2mm to 2cm>/
3. medium-sized <1 to 6cm>/
4. large <4 to 20cm>/
5. very large <greater than 15cm>/
#195. Flowers <whether calyptrate/operculate>/
1. calyptrate <operculate>/
2. not calyptrate <<implicit>>/
#196. Flowers <odour>/
1. fragrant/
2. malodorous/
3. odourless/
#197. Flowers <symmetry of K, C and A, exclusive of G>/
1. regular <actinomorphic, radially symmetric>/
2. somewhat irregular <fairly regular alternation of components but posterior perianth members larger, etc.>/
3. very irregular <including monosymmetric, zygomorphic, bisymmetric, bilaterally symmetric, asymmetric etc.>/
#198. Flowers <irregular, nature of irregularity>/
1. zygomorphic <bisymmetric, monosymmetric, medianly zygomorphic etc. - with one or two planes of symmetry>/
2. asymmetric <no plane of symmetry>/
#199. Flowers <whether resupinate>/
1. resupinate <consistently inverted, with the pedicel twisted through (50-)180 degrees>/
2. not resupinate <<implicit>>/
#200. The floral irregularity <components most conspicuously involved>/
1. involving <all or part of> the perianth/
2. involving the androecium/
#201. Flowers <organization: whether the androecium is external or internal to the gynoecium>/
1. not exhibiting the androecium internal to the gynoecium <<implicit>>/
2. uniquely organized, with the stamens inserted internally to the gynoecium <Lacondoniaceae>/
#202. Flowers <whether papilionaceous>/
1. papilionaceous/
2. ‘pseudo-papilionaceous’/
3. neither papilionaceous nor pseudo-papilionaceous <<implicit>>/
‘Papilionaceous’: denotes possession of an imbricate-descending corolla as in Leguminosae-Papilionoideae, the butterfly shape deriving from an outer, adaxial (posterior) ‘standard’ petal, two lateral ‘wings’ and a ‘keel’ constituted by the anterior (abaxial) pair.
‘Pseudo-papilionaceous’: denotes a floral structure superficially resembling the corolla of Leguminosae-Papilionoideae, but morphologically different.
#203. Flowers <-merous: K, C and A - somewhat interpretive, not for identification>/
merous/
#204. Flowers <whether cyclic>/
1. <strictly> cyclic <<implicit>>/
2. partially acyclic <i.e. in part more or less spiral>/
3. acyclic <i.e. more or less spiral>/
#205. <Distribution of acyclic (i.e. more or less spiral) components:>/
1. the perianth acyclic/
2. the androecium acyclic/
3. the gynoecium acyclic/
#206. Flowers <number of cycles (whorls): applicable only to hermaphrodite flowers>/
1. tricyclic/
2. tetracyclic/
3. pentacyclic/
4. polycyclic/
#207. Floral receptacle <elongation>/
1. developing an androphore <an elongation of the floral axis, above the perianth, supporting or bearing the stamens>/
2. developing a gynophore <an elongation of the floral axis between androecium and gynoecium, a stipitate pistil>/
3. with neither androphore nor gynophore <<implicit>>/
As used here, ‘androphore’ means elongation of the receptacle above the perianth and beneath the stamens, or associated with a column of stamens. A ‘gynophore’ is an elongation immediately beneath the gynoecium, and implies a stipitate pistil. A combination of androphore and gynophore (encoded 1&2) represents an ‘androgynophore’. The term ‘gonophore’ (= an elongation of the floral axis, bearing stamens and carpels) seems to be ambiguous.
#208. Floral receptacle <shape>/
1. markedly hollowed/
2. not markedly hollowed <elongated, domed, flattish to slightly depressed> <<to become implicit>>/
#209. Free hypanthium <presence associated with ‘perigyny’, and implied by ‘corolla inserted on the calyx’. Applied only to Dicotyledons>/
1. present/
2. absent/
‘Hypanthium’: usually ostensibly (and once generally thought morphologically interpretable as) a tubular enlargement or development of the receptacle, supporting the perianth and stamens. Nowadays supposed usually to represent congenital fusion of Dicot perianth and androecium, but arguably at least sometimes (e.g. in Calycanthaceae, Juglandaceae, Santalaceae) partially receptacular. A hypanthium is often implied in older literature by ‘corolla (and/or stamens) inserted on the calyx tube’. In encoding the present descriptions, however, it has been assumed that neither corolla nor stamens can be attached to a true calyx; and any tubular structure bearing calyx lobes plus corolla and/or stamens is regarded as hypanthium, not calyx tube. In the presence of a perianth tube interpreted (or interpretable) as hypanthium, however, there is sometimes no obvious means (textural change, etc.) of deciding precisely where hypanthium ends and ‘true’ calyx begins, hence the distinction between polysepaly and gamosepaly (q.v.) becomes ambiguous. An effort has been made in the descriptions to allow for alternative interpretations, but further adjustments will no doubt be required. For practical purposes, it is fortunate that some families posing severe interpretive difficulties in this context (e.g. Proteaceae) are so variable for all the structural aspects involved that the validity of family descriptions does not depend on satisfactory morphological resolutions for all the forms they cover.
Note that the ‘floral tube’ (hypanthium) of Dicots is generally thought not to be homologous with the ‘perigone tube’ of Monocots, which is usually more obviously perianthial in appearance and texture.
#210. Perigone tube <presence - applied only to Monocotyledons>/
1. present/
2. absent/
See comments under ‘hypanthium’.
#211. Hypogynous <nectariferous> disk <presence>/
1. present/
2. absent/
‘Disk’ = an elevated or appendicular, usually more or less fleshy and nectariferous development on the receptacle internal to the perianth or stamens, which may be receptacular or staminodial (or ambiguous) in origin.
#212. Hypogynous disk <position>/
1. extrastaminal/
2. intrastaminal/
#213. Hypogynous disk <coherence>/
1. of separate members <e.g. ‘scales’>/
2. <more or less> annular/
#214. Perianth <presence/absence, constitution: beware states 1, 3, 4, 5, especially in relation to Monocots (see Notes)>/
1. with distinct calyx and corolla <including ‘petaloid calyces’>/
2. sequentially intergrading from sepals to petals <with intermediates>/
3. <exclusively> sepaline <more or less convincingly interpretable as sepals, though sometimes ‘petaloid’; ‘apetalous’>/
4. <exclusively> petaline <more or less convincingly interpretable as petals; ‘asepalous’>/
5. of ‘tepals’ <not readily resolvable into calyx and/or corolla: the condition awarded here to most Monocots>/
6. vestigial/
7. absent <flower both asepalous and apetalous, achlamydeous>/
‘Calyx’: the outer component of the floral envelope, represented by sepals or equivalent structure(s).
‘Corolla’: the inner component of the floral envelope, represented by petals or equivalent structures.
‘Perianth’: floral envelope(s), inclusive of calyx, corolla, tepals and all morphologically equivalent structures.
‘Tepal’: a segment or unit of a whorled perianth which is not resolvable into calyx and corolla.
In general, an effort has been made to encode this character straightforwardly, according to appearance when conscientiously examined by a trained botanist using a dissecting microscope. However, alternative interpretations of it (and of its dependents) have been encoded in the descriptions where this seemed appropriate. If in doubt about its use in identification with INTKEY, entering all the reasonable possibilities will usually still afford useful discrimination.
An attempt has been made on principle to encode family morphological descriptions so as to allow for likely alternative interpretations or ‘misinterpretations’. The same device has been used in families where the ‘true’ state of affairs seems sometimes to be deducible only from comparative morphology. Its implementation has necessarily been somewhat selective, however, in view of the complications arising from character dependencies.
#215. Perianth <regardless of its morphological interpretation, total number of parts: ‘many’ encoded as ‘to 50 or more’>/
‘Perianth’: floral envelope(s), inclusive of calyx, corolla, tepals and all morphologically equivalent structures.
The DELTA system currently has no device for encoding/interpreting the loose botanical convention ‘many’ (= ‘too many to bother counting’), application of which varies from character to character and from person to person. In compiling the descriptions, ‘many’ has been interpreted so as to encompass the number specified in the character list. The actual ranges entered usually represent guesswork, and are not reliable.
#216. Perianth <‘tepals’, whether free or joined: intended for cases where K and C not applicable>/
1. free/
2. joined <syntepalous>/
‘Perianth’: floral envelope(s), inclusive of calyx, corolla, tepals and all morphologically equivalent structures.
#217. Perianth <number of whorls>/
whorled/
‘Perianth’: floral envelope(s), inclusive of calyx, corolla, tepals and all morphologically equivalent structures.
The reliability of the Monocot data is very uncertain: ‘3+3’ in published descriptions has been assumed to imply two tangible whorls, rather than extrapolation from phylogenetic assumptions.
#218. Perianth <isomery of whorls>/
1. isomerous <the same number(s) in successive whorls>/
2. anisomerous <different numbers in the whorls>/
#219. Perianth <whether petaloid or sepaloid: mainly applied only when not resolvable into calyx and/or corolla>/
1. sepaloid/
2. petaloid/
#220. Perianth <concolorous/spotted: data from Dahlgren and Clifford 1982. Applied only to Monocots>/
1. without spots <implicit for Monocots, where applicable>/
2. <at least some members> spotted/
#221. Perianth <‘tepals’, similarity between the two whorls>/
1. similar in the two whorls/
2. <more or less markedly> different in the two whorls <heterochlamydeous>/
#222. Perianth <‘tepals’, colour - when ‘corolla’ inapplicable>/
1. green/
2. white/
3. cream/
4. yellow/
5. orange/
6. red/
7. pink/
8. purple/
9. violet/
10. blue/
11. brown/
12. black/
13. hyaline <non-exclusive state - to be removed>/
#223. Perianth <whether fleshy: for application when not resolved into ‘calyx’ and/or ‘corolla’>/
1. fleshy/
2. non-fleshy <<to become implicit>>/
#224. Perianth <whether persistent: for application when not resolved into ‘calyx’ and/or ‘corolla’>/
1. persistent/
2. deciduous/
#225. Perianth <whether accrescent: for application when not resolved into ‘calyx’ and/or ‘corolla’>/
1. accrescent <continuing to grow after flowering>/
2. non-accrescent <<to become implicit>>/
#226. Calyx <number of parts>/
This character conveys the ‘K’ numbers of floral formulae, and is usually unambiguously determinable. However, values recorded here sometimes involve morphological interpretation (relying on recognition of joined members, etc.), and inexperienced botanists are advised to avoid using it for identifications in connection with gamosepalous, zygomorphic calyces.
The DELTA system currently has no device for encoding/interpreting the loose botanical convention ‘many’ (= ‘too many to bother counting’), application of which varies from character to character and from person to person. In compiling the descriptions, ‘many’ has been interpreted so as to encompass the number specified in the character list. The actual ranges entered usually represent guesswork, and are not reliable.
#227. Calyx <representation by bristles>/
1. represented by bristles <a pappus, or ‘pappus-like’>/
2. not represented by bristles <<implicit>>/
#228. Calyx <number of whorls>/
whorled/
#229. Calyx <exclusive of hypanthium (q.v.), freedom (see Notes)>/
1. polysepalous <of free sepals>/
2. partially gamosepalous <some members joined, others free>/
3. gamosepalous <all members joined; monosepalous>/
‘Polysepalous’: consisting of free sepals.
‘Partially gamosepalous’: two or more calyx members joined (palpably, or readily interpreted as such with reference to lobes, venation and considerations of floral symmetry), the other(s) free.
‘Gamosepalous’ (monosepalous): reasonably interpretable as representing joining of all the calyx members into a single structure (as evidenced by free lobes, venation and considerations of floral symmetry). Nearly always associated with possession of a long or short calyx tube.
In the present descriptions, any tubular structure bearing corolla and/or stamens in addition to calyx ‘lobes’ is regarded as hypanthium (q.v.), not calyx tube. Calyx ‘lobes’ inserted on such a hypanthium at or below the level of insertion of the (uppermost cycle of) stamens are treated as free sepals, and the calyx is fairly unambiguously ‘polysepalous’. Likewise, calyx ‘lobes’ inserted directly on an inferior ovary (i.e. with no free tube) also represent polysepaly. Ambiguities arise when a free perianth tube is continued without textural change (etc.) above the level of insertion of stamens, so that it is hard to decide where ‘hypanthium’ ends and ‘calyx’ begins: an effort has been made in the descriptions to allow for alternative interpretations.
#230. Calyx <joined, whether entire>/
1. entire/
2. lobulate/
3. blunt-lobed/
4. toothed/
#231. Calyx lobes <(= teeth, lobes, segments) length relative to that of the tube>/
1. markedly shorter than the tube/
2. about the same length as the tube/
3. markedly longer than the tube/
#232. <Number of joined sepals, when only some joined:>/
of the members joined/
#233. Degree of gamosepaly (maximum length joined/total calyx length) <including any hypanthium>/
#234. Calyx <gamosepalous, shape: cf. Lindley’s Glossology>/
1. cupuliform/
2. cyathiform <cupular above, narrowed at the base>/
3. campanulate/
4. urceolate/
5. funnel-shaped/
6. tubular/
7. hypocrateriform/
#235. <Gamosepalous> calyx <symmetry>/
1. unequal but not bilabiate <e.g. unilateral, or with small laterals>/
2. bilabiate/
3. regular/
#236. Calyx <whether basally appendaged or spurred>/
1. basally appendaged/
2. spurred/
3. neither appendaged nor spurred <<implicit>>/
#237. Calyx <whether fleshy>/
1. fleshy/
2. non-fleshy <<to become implicit>>/
#238. Calyx <whether persistent>/
1. persistent/
2. not persistent/
#239. Calyx <accrescence>/
1. accrescent <continuing to grow after flowering>/
2. non-accrescent <<to become implicit>>/
#240. Calyx <joined, whether calyptrate>/
1. calyptrate/
2. not calyptrate <<implicit>>/
#241. Calyx <aestivation (see Notes)>/
1. imbricate/
2. contorted <convolute>/
3. valvate <including induplicate-valvate - see data comments>/
4. open in bud/
5. plicate in bud/
States 1 and 2 (together equivalent to ‘sepals or calyx lobes overlapping one another’) should be lumped for routine identificatory work. The available data imply that the ‘contorted’ state is much rarer than the corresponding state for the corolla. It is generally more easily observed in the corolla, however, and may be under-recorded for the calyx.
‘Imbricate’: where the edges of the sepals or calyx lobes overlap in the flower bud, with one or more members wholly external, one or more wholly internal and the other(s) with one side overlapped and the other overlapping.
‘Contorted’ (convolute, twisted): where the edges of the sepals or calyx lobes overlap in the flower bud in regular sequence, with each member having one edge overlapped and the other overlapping.
‘Valvate’: where the edges of the sepals or calyx lobes are contiguous in the flower bud without overlapping.
‘Induplicate-valvate’: where the edges of the sepals or calyx lobes in the flower bud are bent inwards along their length, and applied valvately to those of their neighbours.
‘Open’: where the edges of the sepals or calyx lobes are neither overlapping nor contiguous with one another in the flower bud.
‘Plicate’: where the sepals or calyx lobes are folded, fanlike, in the flower bud. (It remains to be established whether this condition is exclusive, or represents a qualification of one of the others.)
Precisely defined variants of ‘imbricate’ (quincuncial, cochlear), and left versus right handed orientation of these and of ‘contorted’, are potentially taxonomically informative, but are largely undocumented.
#242. Calyx <orientation>/
1. with the median member anterior <abaxial, usually implying a posterior petal or corolla lobe>/
2. with the median member posterior <adaxial>/
‘Anterior’: abaxial, = on the side away from the axis on which the flower (and its pedicel) are borne.
‘Posterior’: adaxial, = on the side against the axis on which the flower is borne.
Use with caution, since the ‘morphological’ orientation (encoded here) is sometimes different from the apparent orientation because of twisting (resupination) of the pedicel, e.g. in many orchids.
#243. Epicalyx <presence>/
1. present/
2. absent <<implicit>>/
‘Epicalyx’: an ‘accessory calyx’, more or less concrescent with or borne upon the calyx, more or less obviously or only theoretically interpretable as representing bracts, bracteoles or stipules.
#244. Corolla <number of petals or segments: ‘many’ encoded as ‘to 50 or more’>/
This character conveys the ‘C’ numbers of floral formulae, and is usually unambiguously determinable. However, values recorded here sometimes involve morphological interpretation (relying on recognition of joined members, etc.), and inexperienced botanists are advised to avoid using it for identifications in connection with gamopetalous, zygomorphic corollas.
The DELTA system currently has no device for encoding/interpreting the loose botanical convention ‘many’ (= ‘too many to bother counting’), application of which varies from character to character and from person to person. In compiling the descriptions, ‘many’ has been interpreted so as to encompass the number specified in the character list. The actual ranges entered usually represent guesswork, and are not reliable.
#245. Corolla <number of whorls>/
whorled/
#246. Corolla <members alternating with or opposite the calyx members>/
1. alternating with the calyx <<implicit>>/
2. opposite the calyx/
#247. Corolla <whether appendiculate>/
1. appendiculate/
2. not appendiculate <<implicit>>/
#248. Corolla <extent of gamopetaly>/
1. polypetalous <of free petals - includes insertion at the mouth of a hypanthium>/
2. partially gamopetalous <with both joined and free members>/
3. gamopetalous <all members joined; monopetalous>/
‘Polypetalous’: of free petals, inserted directly on the receptacle or hypanthium.
‘Partially gamopetalous’: two or more corolla members joined (palpably, or readily interpreted as such with reference to lobes, venation and considerations of floral symmetry), the other(s) free.
‘Gamopetalous’ (monopetalous): reasonably interpretable as representing joining of all the corolla members into a single structure (as evidenced by free lobes, venation and considerations of floral symmetry). Nearly always associated with possession of a long or short corolla tube.
#249. Corolla tube <Goodeniaceae: whether adaxially split>/
1. not noticeably adaxially split <<implicit>>/
2. adaxially deeply split/
#250. Corolla lobes <(= teeth, lobes, segments) length relative to that of the tube>/
1. markedly shorter than the tube/
2. about the same length as the tube/
3. markedly longer than the tube/
#251. <Number of joined petals, when only some joined:>/
of the petals joined/
#252. The joined petals <when not all joined>/
1. anterior/
2. posterior/
#253. Degree of gamopetaly <excluding any hypanthium: length joined/total corolla length>/
#254. Corolla <whether calyptrate>/
1. calyptrate/
2. not calyptrate <<implicit>>/
#255. Corolla <aestivation>/
1. imbricate/
2. contorted <convolute>/
3. valvate <including induplicate-valvate>/
4. with open aestivation/
5. plicate/
6. crumpled in bud/
‘Imbricate’: where the edges of the petals or corolla lobes overlap in the flower bud, with one or more members wholly external, one or more wholly internal and the other(s) with one side overlapped and the other overlapping.
‘Contorted’ (convolute, twisted): where the edges of the petals or corolla lobes overlap in the flower bud in regular sequence, with each member having one edge overlapped and the other overlapping.
‘Valvate’: where the edges of the petals or corolla lobes are contiguous in the flower bud without overlapping.
‘Induplicate-valvate’: where the edges of the petals or corolla lobes in the flower bud are bent inwards along their length, and applied valvately to those of their neighbours.
‘Open’: where the edges of the petals or corolla lobes are neither overlapping nor contiguous with one another in the flower bud.
‘Plicate’: where the petals or corolla lobes are folded, fanlike, in the flower bud.
Precisely defined variants of ‘imbricate’ (quincuncial, cochlear), and left versus right handed orientation of these and of ‘contorted’, are potentially taxonomically informative (e.g. Scotland et al. 1994 for Acanthaceae and relatives) but are undocumented for most families.
It remains to be established whether ‘plicate’ and ‘crumpled’ represent exclusive states, or qualifications of the others (as seems likely).
#256. Corolla <gamopetalous, shape: cf. Lindley’s Glossology>/
1. rotate/
2. cupuliform/
3. cyathiform/
4. campanulate/
5. urceolate/
6. funnel-shaped/
7. hypocrateriform/
8. tubular/
9. botuliform/
10. vase-shaped <vascularis>/
#257. Corolla <symmetry>/
1. unequal but not bilabiate <e.g. unilateral, or with reduced lateral members>/
2. bilabiate/
3. regular/
#258. Corolla <colours; data unreliable (see Notes)>/
1. green/
2. white/
3. yellow/
4. orange/
5. red/
6. pink/
7. purple/
8. blue/
9. of unusual colour <i.e. none of the above> <<specify>>/
This exemplifies a common situation, where a superficially attractive character has low reliability mainly through lack (or inaccessibility) of comprehensive data on large families. Many families remain uncoded, and some of the descriptions no doubt underestimate the extent of variability.
#259. Corolla <marking>/
1. plain/
2. with contrasting markings <streaks, spots, etc.>/
#260. Corolla <whether spurred>/
1. spurred/
2. not spurred <<implicit>>/
#261. Corolla <whether becoming fleshy>/
1. fleshy/
2. not fleshy <<implicit>>/
#262. Corolla <whether persistent>/
1. persistent <marcescent>/
2. deciduous/
#263. Corolla <accrescence>/
1. accrescent/
2. non-accrescent <<to become implicit>>/
#264. Petals <whether clawed>/
1. clawed <unguiculate, stalked>/
2. sessile <not clawed>/
#265. Petals <whether incised>/
1. deeply bifid/
2. bilobed/
3. fringed <deeply 3 or more -fid>/
4. entire <neither deeply bifid nor bilobed not fringed>/
#266. Androecium <presence - to cope with female-only flowers exhibiting staminodes etc., but poorly recorded>/
1. present/
2. absent/
‘Androecium’: the total male component of the flower, including all recognisable ‘derivatives’ of this. ‘Androecial members’ thus include the stamens (q.v.), plus any staminodial structures (q.v.).
#267. Fertile stamens <presence in individual flowers - determining applicability of stamen, anther, and pollen characters>/
1. present <<implicit>>/
2. absent/
#268. Androecium <of male-fertile flowers, number of members including any staminodes: ‘many’ encoded as ‘to 100 or more’>/
‘Androecium’: the total male component of the flower, including all recognisable ‘derivatives’ of this. ‘Androecial members’ thus include the stamens (q.v.), plus any staminodial structures (q.v.).
The DELTA system currently has no device for encoding/interpreting the loose botanical convention ‘many’ (= ‘too many to bother counting’), application of which varies from character to character and from person to person. In compiling the descriptions, ‘many’ has been interpreted so as to encompass the number specified in the character list. The actual ranges entered usually represent guesswork, and are not reliable.
#269. Androecial members <of male-fertile flowers, whether ‘branched’, proliferous, or ‘split’: as indicated by trunk bundles, ‘dedoublement’, pairs with half-anthers, etc.)>/
1. branched/
2. unbranched <<implicit>>/
#270. Androecial sequence <whether determinable: redundant character, temporarily required to cope with hierarchical dependence re maturation sequence>/
1. determinable <<implicit>>/
2. not <readily> determinable/
#271. Androecial members <of male-fertile flowers, developmental sequence. Note the lack of reliable data on this supposedly ‘important’ character: satisfactory representation would involve distinguishing between maturation and initiation, between whole androecia and cycles of stamens and clusters, and would also require states for ‘simultaneous’>/
1. maturing centripetally/
2. maturing centrifugally/
Note the lack of reliable data on this supposedly ‘important’ character. Satisfactory representation would involve implementing several characters, distinguishing between maturation and initiation and between whole androecia and cycles of stamens and clusters. Separate states would also be required for ‘simultaneous’.
‘Androecium’: the total male component of the flower, including all recognisable ‘derivatives’ of this. ‘Androecial members’ thus include the stamens (q.v.), plus any staminodial structures (q.v.).
#272. Androecial members <of male-fertile flowers, insertion>/
1. <all> free of the perianth <includes insertion at the mouth of a hypanthium>/
2. adnate <i.e. to the perianth, including perigone tube>/
#273. Androecial members <of male-fertile flowers, whether united with the gynoecium>/
1. united with the gynoecium/
2. free of the gynoecium <<implicit>>/
#274. Androecial members <of male-fertile flowers, relative in situ lengths>/
1. all equal/
2. markedly unequal/
#275. Androecial members <of male-fertile flowers, freedom>/
1. free of one another/
2. coherent <state how>/
#276. Androecial members <of male-fertile flowers, manner of coherence>/
adelphous/
#277. Androecial members <of male-fertile flowers, number of whorls: ‘many’ encoded as ‘to 15 or more’>/
whorled/
Refers to insertion only; thus, most legumes have only one whorl, although the ‘inner’ and ‘outer’ whorls may be detectable morphologically via differences in size, anther shape, etc.
The reliability of the data is very uncertain, especially for Monocots: e.g., ‘3+3’ in published descriptions has been assumed to mean two detectable whorls, rather than extrapolation from phylogenetic theory.
‘Androecium’: the total male component of the flower, including all recognisable ‘derivatives’ of this. ‘Androecial members’ thus include the stamens (q.v.), plus any staminodial structures (q.v.).
#278. The androecial bundles <position relative to the corolla members>/
1. alternating with the corolla members/
2. opposite the corolla members/
#279. Androecium <of male-fertile flowers, presence of staminodes>/
1. exclusively of fertile stamens <<implicit>>/
2. including staminodes/
‘Androecium’: the total male component of the flower, including all recognisable ‘derivatives’ of this. ‘Androecial members’ thus include the stamens (q.v.), plus any staminodial structures (q.v.).
‘Staminode’: a sterile stamen, or a modified structure identifiable as such, borne in the androecial region of the flower. May be merely ‘imperfect’, vestigial, or specialized (e.g. petaloid or nectariferous).
#280. Staminodes <of male-fertile flowers, number: ‘many’ encoded as ‘to 50 or more’>/
‘Staminode’: a sterile stamen, or a modified structure identifiable as such, borne in the androecial region of the flower. May be merely ‘imperfect’, vestigial, or specialized (e.g. petaloid or nectariferous).
The DELTA system currently has no device for encoding/interpreting the loose botanical convention ‘many’ (= ‘too many to bother counting’), application of which varies from character to character and from person to person. In compiling the descriptions, ‘many’ has been interpreted so as to encompass the number specified in the character list. The actual ranges entered usually represent guesswork, and are not reliable.
#281. Staminodes <position relative to the fertile stamens>/
1. external to the fertile stamens/
2. in the same series as the fertile stamens/
3. internal to the fertile stamens/
This character often demands recognition of morphological whorls, with reference to the rest of the ‘floral formula’. In particular, stamens inserted at different levels on a corolla tube may nevertheless be of the same ‘morphological series’.
‘Staminode’: a sterile stamen, or a modified structure identifiable as such, borne in the androecial region of the flower. May be merely ‘imperfect’, vestigial, or specialized (e.g. petaloid or nectariferous).
#282. Staminodes representing <Lamiiflorae etc.: position>/
1. the posterior <posticous, adaxial> median member/
2. the posterior-lateral <posticous-, adaxial-lateral> pair/
3. the anterior-lateral <anterior-, abaxial-lateral> pair/
4. the anterior <anticous, abaxial> median member/
#283. Staminodes <of male-fertile flowers, form>/
1. petaloid/
2. non-petaloid <<specify>> <<implicit>>/
‘Staminode’: a sterile stamen, or a modified structure identifiable as such, borne in the androecial region of the flower. May be merely ‘imperfect’, vestigial, or specialized (e.g. petaloid or nectariferous).
‘Didynamous’: stamens paired, with two long and two short (as in most Labiatae).
‘Tetradynamous’: stamens four long and two short, as in Cruciferae.
#284. Fertile stamens representing <Lamiiflorae etc.: position when some androecial members may be sterile or lacking>/
1. the posterior <posticous, adaxial> median member/
2. the posterior-lateral <posticous-, adaxial-lateral> pair/
3. the anterior-lateral <anterior-, abaxial-lateral> pair/
4. the anterior <anticous, abaxial> median member/
#285. Stamens <fertile, number: ‘many’ encoded as ‘to 100 or more’>/
The DELTA system currently has no device for encoding/interpreting the loose botanical convention ‘many’ (= ‘too many to bother counting’), application of which varies from character to character and from person to person. In compiling the descriptions, ‘many’ has been interpreted so as to encompass the number specified in the character list. The actual ranges entered usually represent guesswork, and are not reliable.
#286. Stamens inserted <fertile stamens of Lamiiflorae etc.: position of insertion (distal point of attachment) in the corolla tube>/
1. near the base of the corolla tube/
2. midway down the corolla tube/
3. in the throat of <near the top of> the corolla tube/
#287. <Unequal> stamens <fertile, special size relations>/
1. didynamous <two pairs, one pair longer>/
2. tetradynamous <six, the outer opposite pair small, the four inner members large>/
3. not didynamous, not tetradynamous <<implicit>>/
#288. Stamens <fertile only, number relative to the (inner) perianth - irrespective of relative position/symmetry>/
1. reduced in number relative to the adjacent <inner> perianth/
2. isomerous with the <adjacent> perianth/
3. diplostemonous <double the number of the adjacent perianth>/
4. triplostemonous <thrice the number of the adjacent perianth>/
5. polystemonous <more than thrice the number of the adjacent perianth>/
#289. Stamens <fertile only, insertion position relative to the perianth: applies only to flowers with identifiable calyx and/or corolla>/
1. <the outer or only whorl> alternisepalous <alternating with calyx and/or opposite the corolla: alternisepalous, antepetalous, obdiplostemonous etc.>/
2. <the outer or only whorl> oppositisepalous <opposite the calyx and/or alternating with the corolla: antesepalous, alternipetalous>/
#290. <Individual> stamens <fertile, position relative to the corolla members, regardless of the number of whorls>/
1. <all> alternating with the corolla members/
2. <all> opposite the corolla members/
3. both alternating with and opposite the corolla members/
#291. Stamens <fertile, insertion relative to the (inner) perianth whorl; currently applied only to Monocots>/
1. alterniperianthial/
2. oppositiperianthial/
#292. Stamens <orientation in bud>/
1. erect in bud/
2. inflexed in bud/
3. bent outwards in bud/
#293. Stamens <fertile, form>/
1. laminar/
2. petaloid/
3. filantherous <conventional, with anthers and filaments> <<implicit>>/
4. with sessile anthers/
#294. <Staminal> filaments <whether appendiculate>/
1. appendiculate/
2. not appendiculate <<implicit>>/
#295. Anthers <of coherent stamens, whether syngenesious>/
1. separate from one another <<implicit>>/
2. connivent <inclined toward one another, touching but not joined>/
3. cohering <joined, connate, syngenesious>/
#296. Anthers <fixing>/
1. dorsifixed <including peltate, epipeltate, hypopeltate>/
2. basifixed <impeltate>/
3. apicifixed/
4. adnate/
#297. Anthers <whether becoming inverted during development>/
1. becoming inverted during development, their morphological bases ostensibly apical in the mature stamens/
2. not becoming inverted during development <<implicit>>/
#298. Anthers <versatility>/
1. versatile/
2. non-versatile/
#299. Anthers <manner of dehiscence>/
1. dehiscing via pores/
2. dehiscing via short slits/
3. dehiscing <full length> via longitudinal slits/
4. dehiscing transversely/
5. dehiscing by longitudinal valves/
6. dehiscing irregularly/
‘Valve’: a flap (as opposed to a slit).
#300. Anthers <direction of dehiscence>/
1. extrorse/
2. latrorse/
3. introrse/
#301. Anthers <at anthesis, locule number>/
1. unilocular/
2. bilocular <<implicit>>/
3. four locular/
4. many locular/
#302. Anthers <number of anatomical sporangia>/
1. bisporangiate/
2. tetrasporangiate/
3. multisporangiate/
Data on anther development mainly from Davis 1966, with additions from more recent publications (see References).
#303. Anthers <appendages>/
1. appendaged/
2. unappendaged <<implicit>>/
#304. The anther appendages <position>/
1. apical/
2. dorsal/
3. lateral/
4. basal/
#305. Endothecium <whether developing fibrous thickenings>/
1. developing fibrous thickenings/
2. not developing fibrous thickenings/
#306. The endothecial thickenings <type: most data from Dahlgen, Clifford and Yeo 1985>/
1. spiral/
2. girdling/
#307. Anther epidermis <whether persistent>/
1. persistent/
2. degenerating/
#308. Microsporogenesis <type>/
1. successive/
2. simultaneous/
#309. The initial microspore tetrads <shape>/
1. tetrahedral/
2. isobilateral/
3. decussate/
4. T-shaped/
5. linear/
#310. Anther wall <number of middle layers>/
1. initially with one middle layer <‘dicot’ and ‘monocot’ types>/
2. initially with more than one middle layer <‘basic’ type>/
#311. Anther wall <formation type>/
1. of the ‘basic’ type/
2. of the ‘dicot’ type/
3. of the ‘monocot’ type/
4. of the ‘reduced’ type <no middle layer>/
#312. Tapetum <type>/
1. amoeboid/
2. glandular/
#313. Pollen <germination type>/
1. polysiphonous/
2. monosiphonous/
#314. Pollen <how shed>/
1. shed in aggregates/
2. shed as single grains <<implicit>>/
#315. Pollen grains <whether filamentous>/
1. lacking exine, and dispersed in the sea as long filaments/
2. not dispersed as long filaments <<implicit>>/
#316. Pollen <viscin>/
1. with viscin strands/
2. without viscin strands/
#317. Pollen <kinds of aggregates>/
1. in diads/
2. in triplets/
3. in tetrads/
4. in polyads/
5. in the form of pollinia/
#318. Pollen grains/
1. aperturate/
2. nonaperturate/
#319. Pollen grains <number of apertures>/
aperturate/
#320. Pollen grains <type, with regard to apertures>/
1. sulcate <distal-polar furrow (sulcus)>/
2. annulosulcate <with a continuous, meridional aperture>/
3. sulculate <2 (to 3) slitlike apertures, parallel to one another (or in a triangle), with the distal pole between>/
4. zoniaperturate <(-sulcate, -colpate or -porate) with one continuous aperture around the equator>/
5. ulcerate <distal-polar pore (ulcus)>/
6. colpate <more or less equatorial furrows (colpi)>/
7. porate <more or less equatorial pores. Including poroidate>/
8. colporate <more or less equatorial furrows with endopore. Including colporoidate>/
9. foraminate <(forate, pantoporate) scattered pores>/
10. rugate <(pantocolp(or)ate) scattered furrows>/
11. spiraperturate <one or more furrows, arranged spirally over the grain>/
#321. The ulcus <when pollen ulcerate, whether operculate>/
1. operculate/
2. without an operculum <<implicit>>/
‘Operculum’: a clearly defined thickening on the pollen aperture membrane, forming a (sometimes detatchable) lid.
Forms theoretically exhibiting vestigial opercula (in the form of coarse granules, exine fragments) have been encoded as inoperculate, with qualifying comment. The correct treatment would employ three ordered states (operculate/with vestigial operculum/inoperculate, without vestiges), but observations permitting reliable assignment may be unavailable for most of the 25 or so ‘ulcerate’ families.
#322. The ulcus <when pollen ulcerate, whether annulate>/
1. with an annulus/
2. without an annulus <<implicit>>/
#323. Pollen grains <surface>/
1. psilate/
2. scabrous/
3. muricate/
4. spinulose/
#324. Pollen grains <whether lophate>/
1. lophate/
2. not lophate/
#325. Interapertural exine < of the pollen, whether scrobiculate>/
1. scrobiculate/
2. not scrobiculate/
‘Scrobiculate’: exhibiting scrobiculi — small, open, more or less circular lumina, separated by sexinous areas or streaks several times as wide as the average diameter of a single scrobiculus (Erdtman 1969).
Recorded here only for families supposedly closest to Gramineae (see Linder and Ferguson 1985, Kellogg and Campbell 1987).
#326. Interapertural interstitium <of the pollen exine, whether columellate (= baculate)>/
1. columellate/
2. granulate/
‘Columellate’ (baculate): exhibiting more or less perpendicular (radial) rods (branched or unbranched, attached at both ends or free internally).
Recorded here only for families supposedly closest to Gramineae (see Linder and Ferguson 1985, Kellogg and Campbell 1987). A list of families with ‘at least one representative exhibiting columellae (bacules)’ could be drawn up from the usual compilations (notably from Erdtman 1966), but it would be useless in the absence of data on alternative intertitial types.
#327. Pollen grains <number of cells per grain: data mainly from Brewbaker 1967, supplemented from other sources>/
1. <at least some> 2-celled <when shed>/
2. <all> 3-celled <when shed: generally not readily germinated in vitro>/
#328. Fertile gynoecium <presence in individual flowers - determining applicability of ovary, ovule, fruit, and seed characters>/
1. present <<implicit>>/
2. absent/
#329. Gynoecium <of female-fertile flowers, number of carpels: ‘many’ encoded as ‘to 100 or more’. Based on sometimes dubious morphological theory - locule number is safer for identification, when applicable>/
carpelled/
‘Gynoecium’: a collective term for the total female component of the flower, comprising ovary(ies), style(s) and stigma(s), and including all recognisable derivatives of these.
‘Carpel’: the ovule-bearing unit of the gynoecium (including any associated style and stigma), representing a hypothetical ancestral ‘megasporophyll’ (a fertile, three-veined leaf with ovules near its edges associated with the lateral veins). Easily recognisable in practice when the gynoecium is ‘monomerous’