| # The families of flowering plants |

L. Watson and M.J. Dallwitz

### Character list

#### Nomenclature

#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 included in or reduced to’; cf. Airy Shaw (1973). 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>/

#### Habit and leaf form

#5. <Habit - trees, shrubs, herbs>/

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, without coloured juice /

#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> /

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.

The cactaceous areole is a highly specialized, non-photosynthesizing short-shoot, usually borne at the tip of, or adaxially on, or in the axil of, a more or less conspicuous tubercle. The tubercles seem to represent morphologically the expanded bases of main stem leaves, which except in Pereskia and Maihuenia have no green lamina. The laterally bi-symmetrical areole (which lacks an axillant leaf, except in the Opuntioideae where this is ephemeral) is represented by a pair of buds which may be closely apposed or more or less clearly separated. The 'leaves' from the lower (abaxial) bud are represented by spines, while a flower or flowers may develop from the upper, adaxial one.

#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 /

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 /

2. rootless/

#14. Plants <succulence>/

1. succulent/

2. non-succulent /

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 <autotrophic, parasitic, carnivorous>/

1. autotrophic <implicit>/

2. parasitic/

3. carnivorous/

4. utilizing the nutritious exudates of trapped or harboured animals/

#16. Plants <parasitic, type>/

1. mycoheterotrophic <"saprophytic">/

2. haustorially parasitic/

3. endoparasitic/

#17. Plants <whether photosynthesizing>/

1. green and photosynthesizing <implicit>/

2. not green <not photosynthesizing, totally parasitic>/

#18. Parasitic <host parts>/

1. on roots of the host/

2. on aerial parts of the host/

#19. Trapping mechanism <whether active or passive>/

1. active/

2. passive/

#20. 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/

6. consisting of the sticky-glandular, non-irritable (flypaper-like) inflorescence branches or pedicels/

#21. <Herbaceous plants, life form:>/

1. annual/

2. biennial/

3. perennial <multiplying vegetatively>/

‘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.

#22. <Plants, presence of leaf rosettes:>/

1. with a basal <rosette or> aggregation of leaves <‘rosulate’, not necessarily vegetatively ‘acaulescent’>/

2. without conspicuous <basal or terminal> aggregations <rosettes> of leaves /

3. with terminal aggregations <rosettes> of leaves/

#23. Young stems <whether breaking easily at nodes>/

1. breaking easily at the nodes/

2. not breaking easily at the nodes /

#24. <Height of plants:>/

m high/

#25. <Perennial herbs, mode of perennation:>/

1. bulbaceous/

2. cormous/

3. rhizomatous/

4. tuberous/

5. pseudobulbaceous/

‘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).

‘Pseudobulbaceous’: rhizomes with conspicuous bulb-like structures derived from thickening of one or more internodes.

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.

#26. Rhizome and root tissues <red pigment, presence>/

1. brightly red-pigmented/

2. not red-pigmented /

#27. <Whether epiphytic or climbing:>/

1. self supporting /

2. epiphytic/

3. climbing/

#28. <Types of climbers:>/

1. stem twiners/

2. petiole twiners/

3. tendril climbers/

4. sucker climbers/

5. root climbers/

6. scrambling/

#29. <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.

#30. Stem growth <whether conspicuously sympodial>/

1. conspicuously sympodial/

2. not conspicuously sympodial /

#31. <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.

#32. <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.

#33. <Hydrophytic habitat:>/

1. marine/

2. non-marine /

#34. <Hydrophytes, plant habit:>/

1. free floating/

2. rooted <to the substrate> /

#35. Leaves <of rooted hydrophytes, habit>/

1. submerged/

2. emergent/

3. floating/

#36. <Whether conspicuously heterophyllous:>/

1. conspicuously heterophyllous/

2. not conspicuously heterophyllous /

‘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.

#37. Leaves <longevity>/

1. persistent <evergreen>/

2. <seasonally> 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.

#38. 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.

#39. 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.

#40. 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.

Published taxonomic descriptions and illustrations are essentially non-comparative and very inadequately informative over conspicuous aspects of monocotyledonous leaf morphology, notably phyllotaxy and orientation of blades relative to sheaths. Consequently, the data compiled here are unreliable, and there is abundant scope for original observations.

#41. Leaves <whorled, number per whorl>/

per whorl/

#42. Leaves <mature form of lamina>/

1. flat/

2. folded/

3. rolled/

4. terete/

#43. Leaves <texture>/

1. ‘herbaceous’ /

2. leathery/

3. fleshy/

4. membranous/

5. modified into spines/

#44. Leaves <insertion, distance>/

1. imbricate/

2. not imbricate/

#45. 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.

#46. Leaves <bases>/

1. connate/

2. not connate /

‘Connate’: united, congenitally or subsequently. Here, referring to opposite or whorled leaves having their bases joined around the supporting axis.

#47. 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/

#48. Leaf sheaths <form>/

1. tubular/

2. not tubular/

#49. Leaf sheaths <whether margins joined>/

1. with free margins /

2. <at least partially tubular> with <at least partially> joined margins/

#50. Leaves <whether gland-dotted (pellucid-punctate)>/

1. gland-dotted/

2. not gland-dotted /

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.).

#51. Leaves <odour>/

1. aromatic/

2. without marked odour /

3. foetid/

#52. Leaves <primary orientation, plane of flattening>/

1. with blades borne edgewise to the stem <equitant, flattened at right angles to the ‘normal’ plane - e.g. Iris, Eucalyptus (exclusive of reorientation by twisting)>/

2. with blades ‘normally orientated’ <not equitant: flattened in the ‘normal’ plane>/

Published taxonomic descriptions and illustrations are essentially non-comparative and very inadequately informative over conspicuous aspects of monocotyledonous leaf morphology, notably phyllotaxy and orientation of blades relative to sheaths. Consequently, the data compiled here are unreliable, and there is abundant scope for original observations.

The combination of distichous and ensiform can result in Monocots being described as "fan-shaped".

#53. Leaves <simple or compound>/

1. simple <without subdivision into leaflets; excluding ‘unifoliolate’>/

2. compound <comprising leaflets, or exhibiting evidence of representing a single leaflet>/

#54. Leaves <whether peltate>/

1. peltate/

2. not peltate /

‘Peltate’: having the petiole attached to the lower surface of the blade, instead of to its end or margin.

#55. 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.

#56. 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).

#57. Leaves <pinnate, pari- or imparipinnate>/

1. imparipinnate <unevenly pinnate, with a 'normal' terminal leaflet>/

2. paripinnate <lacking a 'normal' terminal leaflet>/

#58. 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.

#59. <Leaf> lamina <inversion>/

1. <consistently> inverted/

2. <consistently> twisted through 90 degrees/

3. neither <consistently> inverted nor twisted through 90 degrees /

Published taxonomic descriptions and illustrations are essentially non-comparative and very inadequately informative over conspicuous aspects of monocotyledonous leaf morphology, notably phyllotaxy and orientation of blades relative to sheaths. Consequently, the data compiled here are unreliable, and there is abundant scope for original observations.

#60. <Leaf> lamina <simple, dissection>/

1. dissected <-fid, -sect, lobed etc.>/

2. entire <i.e. not dissected - exclusive of the margin>/

#61. <Leaf> lamina <symmetry>/

1. conspicuously asymmetric/

2. not <generally> conspicuously asymmetric /

#62. <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.

#63. <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>/

#64. <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.

#65. <Leaf> lamina <venation system, closed or open>/

1. <conspicuously> cross-venulate <currently very inclusive - including ‘reticulate’>/

2. without <conspicuous> cross-venules/

#66. <Leaf> lamina <form, base>/

1. auriculate at the base/

2. cordate/

3. hastate/

4. sagittate/

5. attenuate to the base/

6. cuneate at the base/

7. oblique at the base/

8. rounded at the base/

#67. Leaves <whether ligulate>/

1. ligulate/

2. eligulate /

‘Ligule’: a collarlike or tonguelike projection near the junction of sheath or petiole and blade.

#68. 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.

#69. 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>/

#70. Stipules <intrapetiolar, freedom>/

1. free of one another/

2. concrescent/

#71. Stipules <concrescent, whether sheathing>/

1. ochreate <joined to form a cup-shaped structure round the stem>/

2. not ochreate /

#72. Stipules <with or without colleters>/

1. with colleters <secreting mucilage>/

2. without colleters /

‘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.

#73. Stipules <form>/

1. scaly/

2. leafy/

3. spiny/

4. represented by glands/

#74. Stipules <persistence>/

1. caducous/

2. persistent/

‘Stipules’: appendages of a leaf, borne one on each side of its insertion.

‘Caducous’: falling off early.

#75. Axillary scales <intravaginal squamules, presence>/

1. present/

2. absent /

#76. Lamina margins/

1. entire/

2. crenate/

3. serrate/

4. dentate/

#77. 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.

#78. Vegetative buds <investment>/

1. scaly/

2. not scaly/

#79. <Vegetative> prophylls <number - no data entered>/

#80. <Vegetative> prophylls <orientation - no data entered>/

1. adaxial/

2. lateral/

#81. Leaf development <intercalary meristems: see Notes>/

1. ‘graminaceous’ <with growth and tissue maturation of the lamina basipetal, involving a proximal intercalary meristem>/

2. not ‘graminaceous’ <development of the lamina not involving a proximal intercalary meristem>/

Associated with persistent ‘intercalary meristems’ involving stem nodes and leaf bases. These have been much studied in herbaceous grasses, where they are of universal occurrence, but there seems to be little hard information on their occurrence elsewhere. Judging from external appearances, however, their presence may represent an important ontogenetic difference between typical Dicots and many Monocots, and perhaps among Monocots.

#82. 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. coiled inwards from the tip <circinnate>/

#83. 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 /

‘Circinnate’: coiled basipetally, with the apex near the centre of the coil, as in fern fronds.

#84. Leaves <compound, developmental type>/

1. becoming compound by necrosis/

2. becoming compound by ontogenetically predetermined splitting/

3. becoming compound from primordial lobes /

#85. 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 /

‘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).

#86. Domatia manifested as <types>/

1. pits/

2. pockets/

3. hair tufts/

#### General anatomy

#87. 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 been treated as implicitly exhibiting the negative state. Families undescribed or poorly known anatomically have been tagged as ‘unknown’.

#88. The laticifers <location: very incompletely recorded>/

1. in leaves/

2. in stems/

3. in roots/

4. in flowers/

5. in the fruits/

#89. 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 been treated as implicitly exhibiting the negative state. Families undescribed or poorly known anatomically have been tagged as ‘unknown’.

#90. Plants <whether exhibiting silica bodies: recorded only for Monocots>/

1. with silica bodies/

2. without silica bodies <implicit for Monocots, where anatomy known>/

#91. Chlorenchyma <of leaves and/or stems, presence of ‘peg cells’>/

1. including ‘peg cells’/

2. without ‘peg cells’ /

#### Vegetative anatomy

Leaf

#92. The leaf lamina <general layout in transverse section>/

1. dorsiventral/

2. bifacial <including 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.

‘Bifacial, 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 tissue distinction.

‘Centric’: with more or less radial anatomical organization.

The equitant leaves characteristic of many Monocots, sometimes described as ‘unifacial’, here fall under ‘isobilateral’ or ‘centric’, according to the extent of flattening.

Published taxonomic descriptions and illustrations are essentially non-comparative and very inadequately informative over conspicuous aspects of monocotyledonous leaf morphology which are closely linked to anatomical layouts, notably phyllotaxy and orientation of blades relative to sheaths. Consequently, the data compiled here are unreliable, and there is abundant scope for original observations.

#93. The leaf lamina <presence of epidermal salt glands>/

1. exhibiting epidermal salt glands/

2. without epidermal salt glands <implicit>/

Reliable comparative observations on presence/absence of secretory hairs and epidermal glands, and identifications of substances secreted, do not exist for most families, and the data compiled here are inevitably unreliable. In particular, a study of ‘microhairs’ in grasses (Amarasinge and Watson, 1989) suggests that secretory hair types will often have gone unrecognised as such.

#94. Leaves <presence of ‘pearl glands’>/

1. with ‘pearl glands’/

2. without ‘pearl glands’ <implicit when anatomy ‘known’>/

#95. Extra-floral nectaries <presence on leaves>/

1. present /

2. absent/

Data mainly from Metcalfe and Chalk (1950, 1979).

#96. Hydathodes <presence on leaves>/

1. present/

2. absent/

#97. <Lamina> epidermis <differentiation>/

1. conspicuously differentiated into ‘long’ and ‘short’ cells/

2. without differentiation into ‘long’ and ‘short’ cells /

#98. <Lamina> epidermis <of plants exhibiting silica bodies, whether containing silica bodies>/

1. containing silica bodies/

2. without silica bodies /

#99. <Lamina> epidermis <presence of crystal idioblasts>/

1. with crystal idioblasts/

2. without crystal idioblasts/

Data mainly extracted from Metcalfe and Chalk (1950, 1979).

The data on hairs encoded for the present descriptions are inevitably unreliable. While the diversity of hair forms affords infinite scope for taxonomic classificatory applications, and has been extensively exploited for identification (notably in pharmacology), across-the-board comparative observations of the standard required in the present context do not exist.

#100. Abaxial epidermis <of the lamina, whether papillose>/

1. papillose/

2. not papillose/

#101. Mucilaginous epidermis <of the lamina, presence>/

1. present/

2. absent/

#102. <Foliar> stomata <presence>/

1. present /

2. absent/

#103. <Foliar> stomata <distribution on the lamina>/

1. mainly confined to one surface/

2. on both surfaces/

#104. <Foliar> stomata <subsidiary cell patterns>/

1. anomocytic <ranunculaceous: subsidiaries lacking or indstinguishable from other epidermal cells>/

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).

#105. <Stomatal> guard-cells <of leaves, shape>/

1. ‘grass type’ <dumb-bell shaped>/

2. not ‘grass type’ /

#106. Hairs <foliar, presence>/

1. present <recorded in the family>/

2. absent/

#107. <Leaf> hairs <type>/

1. eglandular/

2. glandular/

#108. <Leaf> hairs <form>/

1. unicellular/

2. multicellular/

#109. Unicellular <leaf> hairs <whether branched>/

1. branched/

2. simple <unbranched>/

#110. Multicellular <leaf> hairs <uni- or multiseriate>/

1. uniseriate/

2. multiseriate/

#111. Multicellular <leaf> hairs <whether branched>/

1. branched/

2. simple <unbranched>/

#112. Complex <leaf> hairs <presence>/

1. present/

2. absent/

#113. Complex <leaf> hairs <specialised forms>/

1. peltate/

2. stellate/

3. clavate/

4. capitate/

#114. Urticating hairs <presence on leaves>/

1. present/

2. absent /

#115. Adaxial hypodermis <presence in lamina>/

1. present <or the epidermis ‘horizontally divided’>/

2. absent/

#116. <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’>/

#117. Secretory cavities <of leaves>/

1. containing oil/

2. containing mucilage/

3. containing resin/

4. containing latex/

Data mainly from Metcalfe and Chalk (1950, 1979).

#118. Secretory cavities <of leaves, development>/

1. schizogenous/

2. lysigenous/

#119. Cystoliths <presence in leaves>/

1. present/

2. absent /

Cystoliths: conspicuous calcium carbonate or calcium carbonate+silica concretions more or less occluding the lumina of specialised cells (lithocysts), found in ground parenchyma or representing modified hairs. Seemingly characteristic of a few families, notably Acanthaceae, Cannabinaceae, Moraceae, Urticaceae.

Information mainly from Metcalfe and Chalk (1950) and (1979). Data may be unreliable through lack of direct information on families observed to exhibit ‘negatives’. When unmentioned in anatomical family descriptions, it has been treated as implicitly exhibiting the negative state. Families undescribed or poorly known anatomically have been tagged as ‘unknown’.

#120. The mesophyll <whether with etherial oil cells>/

1. with spherical etherial oil cells/

2. without etherial oil cells <implicit when anatomy ‘known’>/

#121. 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’>/

#122. The mesophyll <sclerenchymatous idioblasts: few negatives coded>/

1. with sclerenchymatous idioblasts/

2. without sclerenchymatous idioblasts <implicit when anatomy ‘known’>/

#123. The mesophyll <presence of crystals>/

1. containing crystals <nearly always calcium oxalate>/

2. without 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.

#124. The <mesophyll> 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.

#125. Midrib <of lamina, whether conspicuous>/

1. conspicuous/

2. not conspicuous/

#126. Midrib <of lamina, comments on vascularisation>/

#127. Main veins <of leaf lamina, excluding midribs, whether transcurrent>/

1. vertically transcurrent/

2. embedded/

#128. 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.

#129. Foliar vessels <end-wall perforation>/

1. with reticulately perforated end-walls/

2. with scalariform end-walls/

3. with simple end-wall perforations/

#130. 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.

#131. Young stems <shape in transverse section>/

1. <more or less> cylindrical/

2. oval in section/

3. tetragonal in section/

4. flattened/

5. triangular in section/

#### Axis (stem)

#132. Young stems <internodes, solidity>/

1. with solid internodes/

2. with spongy internodes/

3. with hollow internodes/

#133. The cortex <of young stems, presence of cristarque cells, with U-shaped thickenings and containing crystals>/

1. containing cristarque cells/

2. without cristarque cells <implicit when anatomy ‘known’>/

#134. Pith <of stems, whether with diaphragms: few negatives coded>/

1. with diaphragms/

2. without diaphragms <implicit when anatomy ‘known’>/

#135. Pith <whether homogeneous or heterogeneous>/

1. homogeneous/

2. heterogeneous/

#136. Pith <crystalline inclusions>/

1. with crystalline inclusions/

2. without crystalline inclusions/

#137. Secretory cavities <in stems, presence (as distinct from laticifers)>/

1. present/

2. absent/

#138. Secretory cavities <of stems, contents>/

1. with oil/

2. with resin/

3. with mucilage/

4. with latex/

#139. Cork cambium <phellogen, presence in stems>/

1. present/

2. absent/

Several ostensibly unambiguous and potentially useful anatomical characters associated with secondary thickening of stems, including this one, are unsatisfactorily represented in this data compilation, owing to the scarcity of precise, unambiguous published data.

Data mainly from Metcalfe and Chalk (1950 and 1979).

#140. Cork cambium <position of origin in young stems>/

1. initially deep-seated <in the cortex or pericycle>/

2. initially superficial <originating subepidermally, or in the epidermis>/

Several ostensibly unambiguous and potentially useful anatomical characters associated with secondary thickening of stems, including this one, are unsatisfactorily represented in this data compilation, owing to the scarcity of precise, unambiguous published data.

Data mainly from Metcalfe and Chalk (1950 and 1979).

#141. Nodes <bearing foliage leaves, vascularization, lacunar type: see Notes>/

1. unilacunar/

2. bilacunar/

3. tri-lacunar/

4. penta-lacunar/

5. multilacunar/

The number of gaps (‘leaf gaps’) in the vascular cylinder of young stems associated with (immediately distal to) the leaf traces ‘entering’ each leaf base. For the present purpose, treated as inapplicable to Dicot herbs with mature vascular systems comprising isolated bundles with no interfascicular cambium, as well as to Monocots.

Subsequently to Sinnott (1914), taxonomists have devoted more attention to evolutionary speculation than to publishing comparative data. This is unfortunate, because the requisite information is usually as easy to obtain via serial hand sections as are the routinely sought details of ovary placentation and ovule orientation, with the advantage of being available from vegetative material.

While there is usually only one trace per gap, there may be two or more; e.g., a unilacunar node may exhibit two (‘two-trace unilacunar’), and the median leaf gap of a trilacunar node may exhibit three. When the leaves are opposite or whorled, in the present context the nodes are classified according to the number of gaps associated with each leaf (unilacunar opposite, unilacunar whorled, etc,). Nodal vascular anatomy distal to the origins of the leaf traces is complicated by the vascularization of stipules and axillary buds. Most stipules derive their vascular supply as branches from the lateral traces of trilacunar nodes, and the vascular supply to the axillary bud is usually derived via two strands departing from the margins of the median (or only) leaf gap.

#142. Nodes <with opposite leaves, presence of shared split-lateral leaf traces>/

1. exhibiting on either side a trace which divides, contributing the outermost lateral traces to each of the opposite leaves/

2. without split-lateral traces /

#143. Primary vascular tissues <of stems, whether in discrete bundles>/

1. in a cylinder, without separate bundles/

2. comprising a ring of bundles/

3. comprising two or more rings of bundles/

4. consisting of scattered bundles/

Re states 1 and 2: ostensibly unambiguous and potentially useful anatomical characters associated with secondary thickening of stems (open versus closed vascular bundles, presence/absence of interfascicular cambium) and characters depending on them are unsatisfactorily represented in this data compilation, owing to the scarcity of precise, unambiguous published data.

#144. Primary vascular tissues <of stems, layout within bundles or cylinders>/

1. collateral <centrifugal, with xylem inside and phloem confined to the outside>/

2. bicollateral <with both external and internal phloem>/

#145. Internal <‘intraxylary’> phloem <presence to the inside of the stem xylem: not to be confused with ‘included’ (‘interxylary’) phloem, q.v.>/

1. present/

2. absent <implicit when anatomy ‘known’>/

#146. Cortical bundles <presence in stems: see Notes>/

1. present/

2. absent <implicit when anatomy ‘known’>/

For a relevant, concise discussion of comparative morphological interpretations of ‘cortical bindles’, see De Barry (1884).

In the inevitable absence of complete three-dimensional reconstructions of vascular systems, acquisition of properly comparative data requires precisely specifying the location at which transverse sections are to be made; viz., at mid-internode, or immediately beneath a node. The data on 'cortical vascular bundles' accumulated here from compilations in standard works on systematic plant anatomy (Metcalfe and Chalk (1950) and (1979), cross referenced with Solereder, 1908) are of questionable value, because they provide no information on samping in relation to vascular morphology. No attention has been paid to the question of ‘cortical bundles’ versus ‘leaf traces’ and leaf trace complexes, which they presumably sometimes (or always?) represent; hence, no consideration is given to the extent to which observed presence of ‘cortical bundles’ reflects variations in angles at which ‘leaf traces’ depart nodes, and the distances they traverse longitudinally through the internode(s) above.

Data on this character are further unreliable through lack of direct information on families observed to exhibit ‘negatives’. When unmentioned in anatomical family descriptions, it has been treated as implicitly exhibiting the negative state. Families undescribed or poorly known anatomically have been tagged as ‘unknown’.

#147. Medullary bundles <presence in stems>/

1. present/

2. absent <implicit when anatomy ‘known’>/

#148. Secondary thickening <of stems, presence and type>/

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.>/

Dicot forms with an interfascicular cambium producing little or no woody tissue have been awarded state 1 (secondary thickening absent). Available descriptions have not permitted more satisfactory character state definitions with reference to presence/absence of interfascicular cambium.

#149. The anomalous secondary thickening <of stems, pattern: states incomplete>/

1. via concentric cambia/

2. from a single cambial ring/

#150. 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).

#151. The axial <stem or wood> 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’.

#### Wood

#152. The wood <porosity>/

1. ring porous/

2. semi-ring porous/

3. diffuse porous/

#153. The <wood> vessels <mean diameter>/

1. small <less than 100 microns>/

2. medium <100–200 microns>/

3. large <more than 200 microns>/

#154. The <wood> vessels <arrangement>/

1. solitary/

2. radially paired/

3. in radial multiples/

4. clustered/

5. in tangential arcs <comment on dendritic>/

#155. The <stem or wood> vessel end-walls <orientation>/

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).

#156. The <stem or wood> 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.

#157. The <wood> vessels <presence of vestured pits: best data (including negatives!) from Brazier and Franklin 1961>/

1. with vestured pits/

2. without vestured pits /

#158. The <wood> vessels <presence of spiral thickening>/

1. with spiral thickening/

2. without spiral thickening/

#159. The axial <stem or wood> 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.

#160. The axial <stem or wood> xylem <vasicentric tracheids, presence>/

1. with vasicentric tracheids/

2. without vasicentric tracheids <implicit>/

#161. The axial <stem or wood> 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’.

#162. The axial <stem or wood> 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’.

#163. The axial <stem or wood> xylem <presence of septate xylem fibres>/

1. including septate fibres/

2. without septate fibres /

#164. The <wood> fibres <presence of spiral thickening>/

1. with spiral thickening/

2. without spiral thickening/

#165. The <wood> parenchyma <distribution>/

1. apotracheal <including subtypes diffuse, banded, marginal-terminal, marginal-initial, confluent>/

2. paratracheal <including subtypes scanty, vasicentric, aliform, confluent>/

Distribution of wood parenchyma as viewed in transverse sections, especially with reference to association with vessels. Data mainly from Metcalfe and Chalk (1950).

#166. 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> /

#167. ‘Included’ <‘interxylary’> phloem <presence within the secondary xylem, sensu lato (see Notes)>/

1. present/

2. absent /

‘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 been treated as implicitly exhibiting the negative state. Families undescribed or poorly known anatomically have been tagged as ‘unknown’.

#168. Tile <ray> cells <presence in stems - data from Carlquist 1988>/

1. present /

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’.

#169. The 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’.

#170. Tyloses <extending from parenchyma into tracheary elements, presence>/

1. present/

2. absent/

#### Roots

#171. Roots <velamen>/

1. with velamen/

2. without velamen /

#172. Lateral roots <endodermis>/

1. with a conspicuous endodermis/

2. without a conspicuous endodermis/

#173. 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?

#174. 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.

#### Reproductive type, pollination

#175. 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.

#176. 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.

#177. 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.

#178. Female flowers <presence of staminodes>/

1. with staminodes/

2. without staminodes/

#179. 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.

#180. Plants <whether viviparous>/

1. viviparous/

2. not viviparous/

#181. Plants <whether heterostylous>/

1. homostylous/

2. heterostylous/

'Heterostyly' is indicative of a breeding system which involves individuals of the same species having relatively short styles enclosed by the anthers, or relatively long ones exceeding them.

#182. 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.

#183. <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.).

#184. Pollination <type>/

1. anemophilous/

2. entomophilous/

3. ornithophilous/

4. chiropterophilous/

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.

#185. Pollination <entomophilous>/

1. via beetles/

2. via hymenoptera/

3. via lepidoptera/

4. via diptera/

#186. Pollination mechanism/

1. conspicuously specialized /

2. <relatively> unspecialized /

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.

#### Inflorescence, floral, fruit and seed morphology

#187. Flowers <aggregation>/

1. solitary/

2. aggregated in ‘inflorescences’/

#188. Flowers <when solitary, whether terminal or axillary>/

1. terminal/

2. axillary/

‘Axillary’ implies from a leaf axil, rather than from a bract axil.

#189. Flowers <inflorescence, general description: data in process of development>/

1. in cymes/

2. in racemes/

3. in spikes/

4. in heads <or in capitula>/

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 are an inevitable evolutionary consequence of continuing precocious maturation (neoteny), and it is easier to locate erudite morphological discussions and elegant terminologies than to obtain taxonomically satisfactory 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

#190. The fruiting inflorescences <whether conelike (strobiloid)>/

1. conelike <compact, woody, persistent>/

2. not conelike /

#191. Flowers <whether in ‘spikelets’>/

1. in ‘spikelets’/

2. not in ‘spikelets’ /

‘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.

#192. The ultimate inflorescence units <whether racemose or cymose>/

1. cymose <determinate>/

2. racemose <indeterminate>/

. Unsatisfactory character delimitation, inadequate character state definitions and unreliable data on ‘inflorescence’ features are an inevitable evolutionary consequence of continuing precocious maturation (neoteny), and it is easier to locate erudite morphological discussions and elegant terminologies than to obtain taxonomically satisfactory comparative data.

#193. Inflorescences <whether a scape>/

1. scapiflorous <main axis elongate, leafless>/

2. not scapiflorous /

#194. Inflorescences <position>/

1. terminal/

2. axillary/

3. leaf-opposed/

4. intercalary/

5. epiphyllous/

6. cauliflorous/

Axillary’ implies from a leaf axil, rather than from a bract axil.

‘Intercalary’ (of inflorescences): where the flower-bearing axes are not terminal but continue to grow, reverting to the vegetative state after giving rise to the aggregates of flowers.

‘Cauliflorous’ (of trees and shrubs): flowering from old wood, i.e. from trunk or branches.

#195. Inflorescences <type>/

Unsatisfactory character delimitation, inadequate character state definitions and unreliable data on ‘inflorescence’ features are an inevitable evolutionary consequence of continuing precocious maturation (neoteny), and it is easier to locate erudite morphological discussions and elegant terminologies than to obtain taxonomically satisfactory 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).

#196. Inflorescences <whether involucrate>/

1. with involucral bracts/

2. without involucral bracts /

‘Involucre’: one or more whorls of modified leaves or bracts subtending a cluster of flowers (or florets) or a cluster of inflorescence branches.

#197. Inflorescences <whether comprising or incorporating pseudanthia>/

1. pseudanthial <consisting of one or more flower-like units>/

2. not pseudanthial <implicit>/

‘Pseudanthium’: an aggregation of flowers superficially resembling (and perhaps fuctionally, biologically equivalent to) a single flower Spectacularly exemplifying the evolutionary process of precocious maturation ('neoteny').

#198. The involucres <accrescence>/

1. accrescent/

2. non-accrescent /

‘Accrescent’: increasing in size after flowering.

#199. Inflorescences <whether spatheate>/

1. spatheate/

2. espatheate /

‘Spathe’: a conspicuous bract or specialized leaf, enclosing an inflorescence.

#200. 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.

#201. Bracts <whether calyptrate and covering the flower bud: not to be confused with calyptrate K or C>/

1. calyptrate/

2. not calyptrate /

‘Calyptrate’: forming a hood or lid over the inner parts of the flower.

#202. 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.

#203. 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>/

#204. Flowers <whether calyptrate/operculate>/

1. calyptrate <operculate>/

2. not calyptrate /

#205. Flowers <odour>/

1. fragrant/

2. malodorous/

3. odourless/

#206. 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.>/

#207. 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>/

#208. Flowers <whether resupinate>/

1. resupinate <consistently inverted, with the pedicel twisted through (50-)180 degrees>/

2. not resupinate /

#209. The floral irregularity <components most conspicuously involved>/

1. involving <all or part of> the perianth/

2. involving the androecium/

#210. Flowers <organization: whether the androecium is external or internal to the gynoecium>/

1. not exhibiting the androecium internal to the gynoecium /

2. uniquely organized, with the stamens inserted internally to the gynoecium <Lacondoniaceae>/

#211. Flowers <whether papilionaceous>/

1. papilionaceous <the complete syndrome: see Notes>/

2. ‘pseudo-papilionaceous’/

3. neither papilionaceous nor pseudo-papilionaceous /

‘Papilionaceous’: exhibiting a C5, imbricate-descending corolla as in typical Leguminosae-Papilionoideae, the butterfly shape deriving from an outer, adaxial (posterior) ‘standard’ petal (the vexillum), two lateral ‘wings’, and a ‘keel’ constituted by the anterior (lowermost, abaxial) pair; the latter enclosing the declinate androecium and gynoecium.

‘Pseudo-papilionaceous’: denotes a floral structure exhibiting some but not all of the features of, or superficially resembling, that typical of Leguminosae-Papilionoideae (q.v.).

#212. Flowers <-merous: K, C and A - somewhat interpretive, not for identification>/

merous/

#213. Flowers <whether cyclic>/

1. <strictly> cyclic /

2. partially acyclic <i.e. in part more or less spiral>/

3. acyclic <i.e. more or less spiral>/

#214. <Distribution of acyclic (i.e. more or less spiral) components:>/

1. the perianth acyclic/

2. the androecium acyclic/

3. the gynoecium acyclic/

#215. Flowers <number of cycles (whorls): applicable only to hermaphrodite flowers>/

1. tricyclic/

2. tetracyclic/

3. pentacyclic/

4. polycyclic/

#216. 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 /

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.

#217. Floral receptacle <shape>/

1. markedly hollowed/

2. not markedly hollowed <elongated, domed, flattish to slightly depressed> /

#218. 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.

#219. Perigone tube <presence - applied only to Monocotyledons>/

1. present/

2. absent/

See comments under ‘hypanthium’.

#220. 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.

#221. Hypogynous disk <position>/

1. extrastaminal/

2. intrastaminal/

#222. Hypogynous disk <coherence>/

1. of separate members <e.g. ‘scales’>/

2. <more or less> annular/

#### Perianth

#223. 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.

#224. 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.

#225. 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.

#226. 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.

#227. Perianth <isomery of whorls>/

1. isomerous <the same number(s) in successive whorls>/

2. anisomerous <different numbers in the whorls>/

#228. Perianth <whether petaloid or sepaloid: mainly applied only when not resolvable into calyx and/or corolla>/

1. sepaloid/

2. petaloid/

#229. 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/

#230. Perianth <‘tepals’, similarity between the two whorls>/

1. similar in the two whorls/

2. <more or less markedly> different in the two whorls <heterochlamydeous>/

#231. 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>/

#232. Perianth <whether fleshy: for application when not resolved into ‘calyx’ and/or ‘corolla’>/

1. fleshy/

2. non-fleshy /

#233. Perianth <whether persistent: for application when not resolved into ‘calyx’ and/or ‘corolla’>/

1. persistent/

2. deciduous/

#234. Perianth <whether accrescent: for application when not resolved into ‘calyx’ and/or ‘corolla’>/

1. accrescent <continuing to grow after flowering>/

2. non-accrescent /

#235. Tepal apex trichomes (TAT) <presence: Macfarlane and Conran, 2016>/

1. present/

2. absent/

#236. 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.

#237. Calyx <representation by bristles>/

1. represented by bristles <a pappus, or ‘pappus-like’>/

2. not represented by bristles /

#238. Calyx <number of whorls>/

whorled/

#239. 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.

#240. Calyx <joined, whether entire>/

1. entire/

2. lobulate/

3. blunt-lobed/

4. toothed/

#241. 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/

#242. <Number of joined sepals, when only some joined:>/

of the members joined/

#243. Degree of gamosepaly (maximum length joined/total calyx length) <including any hypanthium>/

#244. 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/

#245. <Gamosepalous> calyx <symmetry>/

1. unequal but not bilabiate <e.g. unilateral, or with small laterals>/

2. bilabiate/

3. regular/

#246. Calyx <whether basally appendaged or spurred>/

1. basally appendaged/

2. basally spurred/

3. neither appendaged nor spurred /

#247. Calyx <whether fleshy>/

1. fleshy/

2. non-fleshy /

#248. Calyx <whether persistent>/

1. persistent/

2. not persistent/

#249. Calyx <accrescence>/

1. accrescent <continuing to grow after flowering>/

2. non-accrescent /

#250. Calyx <joined, whether calyptrate>/

1. calyptrate/

2. not calyptrate /

#251. Calyx <aestivation in flower buds (see Notes)>/

1. imbricate <including quincuncial>/

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.

#252. 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.

#253. Epicalyx <presence>/

1. present/

2. absent /

‘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.

#254. 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.

#255. Corolla <number of whorls>/

whorled/

#256. Corolla <members alternating with or opposite the calyx members>/

1. alternating with the calyx /

2. opposite the calyx/

#257. Corolla <whether appendiculate>/

1. appendiculate/

2. not appendiculate /

#258. 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.

#259. Corolla tube <e.g., Goodeniaceae: whether adaxially split>/

1. not noticeably adaxially split /

2. adaxially deeply split/

#260. 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/

#261. <Number of joined petals, when only some joined:>/

of the petals joined/

#262. The joined petals <when not all joined>/

1. anterior/

2. posterior/

#263. Degree of gamopetaly <excluding any hypanthium: length joined/total corolla length>/

#264. Corolla <whether calyptrate>/

1. calyptrate/

2. not calyptrate /

#265. Corolla <aestivation in flower buds>/

1. imbricate <including quincuncial>/

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).

#266. 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>/

#267. Corolla <symmetry>/

1. unequal but not bilabiate <e.g. unilateral, or with reduced lateral members>/

2. bilabiate/

3. regular/

#268. 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> /

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.

#269. Corolla <marking>/

1. plain/

2. with contrasting markings <streaks, spots, etc.>/

#270. Corolla <whether spurred>/

1. spurred/

2. not spurred /

#271. Corolla <whether becoming fleshy>/

1. fleshy/

2. not fleshy /

#272. Corolla <whether persistent>/

1. persistent <marcescent>/

2. deciduous/

#273. Corolla <accrescence>/

1. accrescent/

2. non-accrescent /

#274. Petals <whether clawed>/

1. clawed <unguiculate, stalked>/

2. sessile <not clawed>/

#275. Petals <whether incised>/

1. deeply bifid/

2. bilobed/

3. fringed <deeply 3 or more -fid>/

4. entire <neither deeply bifid nor bilobed not fringed>/

#### Androecium

#276. 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.).

#277. Fertile stamens <presence in individual flowers - determining applicability of stamen, anther, and pollen characters>/

1. present /

2. absent/

#278. 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.

#279. 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 /

#280. Androecial sequence <whether determinable: redundant character, temporarily required to cope with hierarchical dependence re maturation sequence>/

1. determinable /

2. not <readily> determinable/

#281. 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.).

#282. 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>/

#283. Androecial members <of male-fertile flowers, whether united with the gynoecium>/

1. united with the gynoecium/

2. free of the gynoecium /

#284. Androecial members <of male-fertile flowers, relative in situ lengths>/

1. all equal/

2. markedly unequal/

#285. Androecial members <of male-fertile flowers, freedom>/

1. free of one another/

2. coherent <state how>/

#286. Androecial members <of male-fertile flowers when cohering, manner of coherence>/

adelphous/

#287. 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.).

#288. The androecial bundles <position relative to the corolla members>/

1. alternating with the corolla members/

2. opposite the corolla members/

#289. Androecium <of male-fertile flowers, presence of staminodes>/

1. exclusively of fertile stamens /

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).

#290. Staminodes <of male-fertile flowers, number when present: ‘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.

#291. 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).

#292. 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/

#293. Staminodes <of male-fertile flowers, form>/

1. petaloid/

2. non-petaloid /

‘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.

#294. 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/

#295. 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.

#296. 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/

#297. <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 /

#298. 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>/

#299. 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>/

#300. <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/

#301. Stamens <fertile, insertion relative to the (inner) perianth whorl; currently applied only to Monocots>/

1. alterniperianthial/

2. oppositiperianthial/

#302. Stamens <orientation in bud>/

1. erect in bud/

2. inflexed in bud/

3. bent outwards in bud/

#303. Stamens <fertile, form>/

1. laminar/

2. petaloid/

3. filantherous <conventional, with anthers and filaments> /

4. with sessile anthers/

#304. <Staminal> filaments <whether appendiculate>/

1. appendiculate/

2. not appendiculate /

#305. Anthers <of coherent stamens, whether syngenesious>/

1. separate from one another /

2. connivent <inclined toward one another, touching but not joined>/

3. cohering <joined, connate, syngenesious>/

#306. Anthers <fixing>/

1. dorsifixed <including peltate, epipeltate, hypopeltate>/

2. basifixed <impeltate>/

3. apicifixed/

4. adnate/

#307. 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 /

#308. Anthers <versatility>/

1. versatile/

2. non-versatile/

#309. 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).

#310. Anthers <direction of dehiscence>/

1. extrorse/

2. latrorse/

3. introrse/

#311. Anthers <at anthesis, locule number>/

1. unilocular/

2. bilocular /

3. four locular/

4. many locular/

#312. 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).

#313. Anthers <appendages>/

1. appendaged/

2. unappendaged /

#314. The anther appendages <position>/

1. apical/

2. dorsal/

3. lateral/

4. basal/

#315. Endothecium <whether developing fibrous thickenings>/

1. developing fibrous thickenings/

2. not developing fibrous thickenings/

#316. The endothecial thickenings <type: most data from Dahlgen, Clifford and Yeo 1985>/

1. spiral/

2. girdling/

#317. Anther epidermis <whether persistent>/

1. persistent/

2. degenerating/

#318. Microsporogenesis <type>/

1. successive/

2. simultaneous/

#319. The initial microspore tetrads <shape>/

1. tetrahedral/

2. isobilateral/

3. decussate/

4. T-shaped/

5. linear/

#320. 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>/

#321. 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>/

#322. Tapetum <type>/

1. amoeboid/

2. glandular/

#323. Pollen <germination type>/

1. polysiphonous/

2. monosiphonous/

#324. Pollen <how shed>/

1. shed in aggregates/

2. shed as single grains /

#325. Pollen grains <whether filamentous>/

1. lacking exine, and dispersed in the sea as long filaments/

2. not dispersed as long filaments /

#326. Pollen <viscin>/

1. with viscin strands/

2. without viscin strands/

#327. Pollen <kinds of aggregates>/

1. in diads/

2. in triplets/

3. in tetrads/

4. in polyads/

5. in the form of waxy pollinia/

#328. Pollen grains/

1. aperturate/

2. nonaperturate/

#329. Pollen grains <number of apertures>/

aperturate/

#330. 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>/

#331. The ulcus <when pollen ulcerate, whether operculate>/

1. operculate/

2. without an operculum /

‘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.

#332. The ulcus <when pollen ulcerate, whether annulate>/

1. with an annulus/

2. without an annulus /

#333. Pollen grains <surface>/

1. psilate/

2. scabrous/

3. muricate/

4. spinulose/

#334. Pollen grains <whether lophate>/

1. lophate/

2. not lophate/

#335. 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).

#336. 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.

#337. 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>/

#### Gynoecium

#338. Fertile gynoecium <presence in individual flowers - determining applicability of ovary, ovule, and fruit characters>/

1. present /

2. absent/

#339. 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’ or ‘apocarpous’ (i.e. when the carpels are solitary, or multiple but separate), but becoming increasingly theoretical with increasing degrees of (hypothetical) congenital fusion in ‘syncarpous’ (compound) gynoecia. Thus, where the ovary is unilocular the ‘number of carpels’ represents phylogenetic speculation based on the lobing of the ovary, vestigial features, the number of styles or stigmas, the placentation pattern, the vasculature, the symmetry of the flower, etc.

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.

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.

#340. Gynoecium <whether (partly) petaloid - e.g. Iridaceae, Taccaceae>/

1. partly petaloid/

2. non-petaloid /

#341. Carpels <isomery with the (inner) P: data fairly incomplete>/

1. reduced in number relative to the <inner> perianth <whorl>/

2. isomerous with the <inner> perianth/

3. increased in number relative to the <inner> perianth <whorl>/

‘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’ or ‘apocarpous’ (i.e. when the carpels are solitary, or multiple but separate), but becoming increasingly theoretical with increasing degrees of (hypothetical) congenital fusion in ‘syncarpous’ (compound) gynoecia. Thus, where the ovary is unilocular the ‘number of carpels’ represents phylogenetic speculation based on the lobing of the ovary, vestigial features, the number of styles or stigmas, the placentation pattern, the vasculature, the symmetry of the flower, etc.

#342. The <single> pistil <monomerous or syncarpous, total number of cells (whether loculi or locelli) visible in transverse sections>/

celled/

‘Locellate’: of ovaries having secondary compartments (locelli) derived via ‘false septa’, the latter representing ingrowths from the ovary walls, or complete (fused) intrusions or extrusions of the placentas. In mature ovaries, locelli may be hard to distinguish from primary locules without examining the vascularization of the ovary wall and noting the number and location of styles (i.e. without establishing the number of carpels involved).

#343. Gynoecium <constitution: less precise but more accessible than the alternative, preferred version>/

1. monomerous/

2. apocarpous <of more than one free carpel. Excludes forms joined only at the stylehead or stigmas>/

3. syncarpous <carpels at least partially joined>/

‘Gynoecium’: a collective term for the total female component of the flower, including all recognisable derivatives of this.

‘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’ or ‘apocarpous’ (i.e. when the carpels are solitary, or multiple but separate), but becoming increasingly theoretical with increasing degrees of (hypothetical) congenital fusion in ‘syncarpous’ (compound) gynoecia. Thus, where the ovary is unilocular the ‘number of carpels’ represents phylogenetic speculation based on the lobing of the ovary, vestigial features, the number of styles or stigmas, the placentation pattern, the vasculature, the symmetry of the flower, etc.

With few exceptions (e.g. Hernandiaceae), inferior unilocular ovaries seem reasonably interpretable as pseudomonomerous and syncarpous (q.v.), and have been treated as such in the descriptions.

‘Apocarpous’: of a gynoecium composed of two or more separate carpels.

‘Monomerous’: of a gynoecium consisting of a solitary carpel.

‘Semicarpous’: of a gynoecium of two or more carpels, whose ovaries are only partially (basally) joined.

‘Syncarpous’: of a gynoecium composed of two or more (theoretically) congenitally united carpels.

‘Pseudomonomerous’ (a term often appearing as data comment in the present descriptions): of an ostensibly monomerous gynoecium, which is supposed on comparative morphological grounds (etc.) to represent a reduced syncarpous one. An effort has been made to encode the monomeric alternative and its dependent characters, at least as an alternative, for all families exhibiting ‘pseudomonomery’ deducible only from comparative morphology. All inferior unilocular ovaries seem reasonably interpretable as pseudomonomerous and syncarpous (q.v.), and have been treated as such in the descriptions. (See ‘carpel’ for further information.)

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.

#344. Gynoecium <monocarpy, apocarpy, syncarpy: states 4, 5 and 6 form a continuum, with ‘5 to 6’ often reflecting difficulty in distinguishing ‘style’ from ‘stigma’>/

1. of one carpel/

2. eu-apocarpous <of two or more free carpels>/

3. semicarpous <of two or more carpels, only partially fused>/

4. synovarious <ovary fused, styles and stigmas free>/

5. synstylovarious <ovaries and styles fused, stigmas free>/

6. eu-syncarpous <ovary, styles and stigmas fused>/

7. synstylous <carpels free below, but joined by their styles>/

#345. Gynoecium <superior or inferior>/

1. superior <free of P, K, C>/

2. partly inferior <adnate below to P, K, C>/

3. inferior <adnate to epigynous P, K, C and/or receptacle>/

#### Apocarpous, monomeric and synstylous gynoecia

#346. Carpel <in monomeric and apocarpous gynoecia, whether closed>/

1. fully closed /

2. incompletely closed/

#347. Carpel <of apocarpous/semicarpous or monomeric G, whether stylate>/

1. non-stylate/

2. stylate/

#348. Carpel <of apocarpous or monomeric G, stigma>/

1. with a longitudinal stigmatic surface/

2. apically stigmatic <i.e. with an apical style and/or stigma>/

3. with a lateral style/

4. with a gynobasic style/

#349. Styles <of apocarpous/semicarpous ovaries, whether free or united>/

1. united/

2. free /

#350. Carpel <monomeric or apocarpous, number of ovules: ‘many’ encoded as ‘to 100 or more’>/

ovuled/

#351. Placentation <of monomeric and apocarpous/semicarpous gynoecia>/

1. apical/

2. marginal <longitudinal>/

3. basal/

4. dispersed <over the inner carpel surface: ‘laminar’>/

#### Syncarpous gynoecia

#352. Ovary <syncarpous, uni/plurilocular>/

1. unilocular/

2. plurilocular/

‘Ovary’: the ovule-bearing part of a syncarpous, semicarpous or pseudomonomeric gynoecium, as distinct from its style and stigma.

#353. Ovary <locule number of syncarpous gynoecia: ‘many’ encoded as ‘to 50’ or ‘to 100’>/

locular/

‘Ovary’: the ovule-bearing part of a syncarpous, semicarpous or pseudomonomeric gynoecium, as distinct from its style and stigma.

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.

#354. Locules <of syncarpous ovary, whether exhibiting ‘false septa’>/

1. secondarily divided by ‘false septa’/

2. without ‘false septa’ /

‘False septa’: secondary septa, representing ingrowths from the ovary walls or complete (fused) extrusions or intrusions from the placentas, which subdivide the primary locules into locelli (q.v.), and result in cell counts in excess of (often double) the number of carpels. Rarely, they occur as horizontal partitions, detectable only via vertical sections of the ovary. As distinct from the true septa, which divide the ovary into primary locules each supposedly representing one carpel (q.v.).

#355. Locules <horizontal septa>/

1. each divided horizontally into one-ovulate locelli <Siphonodontaceae, Malvaceae-Malopeae>/

2. not horizontally divided /

#356. Gynoecium <when G2 or G1, orientation relative to the axis/median plane of flower>/

1. median <the carpels aligned anterior-posterior>/

2. oblique/

3. transverse/

‘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.

#357. The ‘odd’ carpel <when G3, orientation>/

1. posterior <adaxial, against the axis, on top in conventional diagrams>/

2. anterior <abaxial, away from the axis, at the bottom in conventional diagrams>/

3. obliquely positioned/

#358. Ovary <insertion>/

1. sessile/

2. subsessile/

3. stipitate/

#359. Epigynous disk <presence>/

1. present/

2. absent/

#360. Gynoecium <of syncarpous G: possession of style(s)>/

1. non-stylate/

2. stylate/

#361. Styles <number: intended for syncarpous gynoecia>/

#362. Styles <whether bearing an ‘indusium’>/

1. bearing an ‘indusium’ beneath the stigma/

2. without an indusium /

#363. Styles <of syncarpous G: when more than one>/

1. free/

2. partially joined/

#364. Styles <of syncarpous G: ovary junction>/

1. attenuate from the ovary/

2. from a depression at the top of the ovary/

#365. Styles <of syncarpous G: insertion>/

1. apical/

2. lateral/

3. ‘gynobasic’/

#366. Styles <of syncarpous G: length relative to that of the ovary at anthesis>/

1. shorter than the ovary/

2. about as long as the ovary/

3. much longer than the ovary/

#367. Stylar canal/

1. present/

2. absent/

#368. Stigmas <of syncarpous G: number>/

#369. Stigmas <whether commissural>/

1. dorsal to the carpels <not commissural> /

2. commissural <lateral to the carpels>/

#370. Stigmas <of syncarpous G: single, number of lobes; ‘1-lobed’ = ‘entire’>/

lobed/

#371. Stigmas <of syncarpous G: single, form>/

1. truncate/

2. clavate/

3. capitate/

4. subpeltate/

5. peltate/

#### Gynoecium, general

#372. Stigmas <type: data mainly from Heslop-Harrison and Shivanna 1977>/

1. wet type <exhibited by most gametophytic self-incompatibility systems>/

2. dry type <associated with sporophytic self-incompatibility systems>/

#373. Stigmas <whether surface papillate: Heslop-Harrison and Shivanna 1977>/

1. papillate/

2. non-papillate/

#374. Stigmas <type: Heslop-Harrison and Shivanna 1977>/

1. Group I type/

2. Group II type/

3. Group III type/

4. Group IV type/

#375. Placentation <when syncarpous and unilocular>/

1. basal/

2. parietal/

3. apical/

4. laminar-dispersed/

5. free central/

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.

#376. Placentation <when syncarpous and bi- or plurilocular>/

1. basal/

2. axile/

3. apical/

4. parietal <including where each of the two placentae is split by a replum>/

#377. Ovules <whether differentiated prior to fertilization>/

1. differentiated /

2. not differentiated <prior to fertilization>/

#378. Ovules in the single cavity <number in the unilocular ovary: ‘many’ indicated by 100>/

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.

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.

#379. Ovules <number per locule, when more than one-locular: ‘many’ indicated by 50>/

per locule/

#380. Ovules <carriage>/

1. funicled /

2. sessile/

3. <more or less> sunken in the placenta/

#381. Ovules <orientation>/

1. pendulous <descending>/

2. horizontal/

3. ascending <including ‘erect’>/

Orientation with regard to the relative positions of the top of the ovule (i.e. its distal end, regardless of the position of the micropyle), its point of attachment to the placenta, and the proximal/distal axis of the ovary. Not to be confused with ‘morphological orientation’.

‘Pendulous’ (descending): (ovule) fixed to an axile, free central, apical or parietal placenta, with the its top hanging or turned towards the base of the ovary.

‘Horizontal’: (ovule) fixed to an axile, free central or parietal placenta, with its axis more or less at right angles to that of the ovary.

‘Ascending’: (ovule) fixed to a basal, axile, free central or parietal placenta, with its top facing or turned towards the distal end of the ovary.

See Geesink et al. (1981) for diagrams.

#382. Ovules <non-orthotropous, orientation: data very incomplete>/

1. apotropous <the raphe ventral when an ovule is ascending, dorsal when it is descending>/

2. epitropous <the raphe dorsal when an ovule is ascending, ventral when it is descending>/

‘Apotropous’: of ovules with the raphe ventral when the ovule is ascending, dorsal when it is descending.

‘Epitropous’: the raphe dorsal when the ovule is ascending, ventral when it is descending. The latter case is equivalent to ‘micropyle upwards and outwards’ in older literature; but ‘micropyle superior’ does not distinguish apotropous from epitropous. (See Geesink et al. (1981) for excellent diagrams.)

This nineteenth century character is largely ignored in modern accounts. It implies the existence of a taxonomically important distinction among non-orthotropous ovules, between those seeming to represent a phylogenetic rotation and adhesion to its funicle of the ancestral orthotropous ovule ‘upwards and inwards’, i.e. with the micropylar end rotating towards the distal end of the ovary (ovule involute; epitropous), and those where it appears to have rotated and adhered ‘downwards and inwards’, i.e towards the base of the ovary (ovule revolute; apotropous). The taxonomic/phylogenetic significance of the distinction was recognised in the Engler and Prantl classification, where it was emphasized as the major difference between the important orders Geraniales and Sapindales, together involving some 50 families. For the present purpose, the available data seem applicable only to pendulous and ascending ovules, and then only in the context of axile placentation, as expressed by the character states defined above. Where not stated explicitly in published family descriptions, the condition has been tentatively deduced by combining compiled (sparse) information on ovule and raphe orientation. However, the principle seems independent of whether ovules ascend or descend, and ought sometimes also to be applicable where the placentation is apical, basal or parietal, provided there are more than two ovules on the placenta. It is difficult or impossible to glean much reliable data on this interesting aspect from existing taxonomic descriptions, and there is evidently a fine opportunity for making original observations.

‘Raphe’: of (more or less) anatropous ovules (q.v.), being a cord or ridge of fibrovascular tissue connecting the base of the nucellus with the placenta which morphologically represents the adherent funicle. It is ‘ventral’ if located on the side of the ovule against the axis, ‘dorsal’ if away from it, or ‘lateral’ if somewhere intermediate.

‘Orthotropous’: (ovule) straight, with the micropyle at one end and the basal funicular attachment at the other. Embryo straight.

#383. Ovules <non-orthotropous, position of raphe>/

1. with ventral raphe <against the placenta>/

2. with lateral raphe/

3. with dorsal raphe <away from the placenta>/

‘Raphe’: of (more or less) anatropous ovules (q.v.), being a cord or ridge of fibrovascular tissue connecting the base of the nucellus with the placenta, morphologically representing the adherent part of the funicle. It is ‘ventral’ if located on the side of the ovule against the placenta, ‘dorsal’ if away from it, or ‘lateral’ if somewhere intermediate.

#384. Ovules <alignment on the placenta>/

1. collateral/

2. superposed/

3. over the carpel surface/

4. biseriate <alternating in two close longitudinal ranks>/

#385. Ovules <whether arillate: currently includes funicular, micropylar and ‘strophiolar’ forms>/

1. arillate/

2. non-arillate/

Data on ovule development and embryology mainly from Davis 1966, with more recent additions (see References).

#386. Ovules <morphological orientation: data on this and ovule developmental features extensively from Davis 1966>/

1. orthotropous/

2. hemianatropous <semianatropous, hemitropous>/

3. anatropous/

4. campylotropous/

5. amphitropous/

#387. Ovules <number of integuments>/

1. without integuments/

2. unitegmic/

3. bitegmic/

#388. Ovules <type: definitions and most data from Davis 1966>/

1. tenuinucellate <no parietal cell cut off from the primary archesporial cell>/

2. crassinucellate <parietal cell (usually leading to tissue) cut off from the primary archesporial cell>/

3. pseudocrassinucellate <no parietal cell or parietal tissue, but the epidermis forming a nucellar cap>/

#389. Outer integument <of bitegmic ovules: data mainly from Davis 1966>/

1. contributing to the micropyle/

2. not contributing to the micropyle/

#390. Endothelium <of ovule>/

1. differentiated <from inner layer of integument>/

2. not differentiated/

#391. Embryo-sac development <type>/

1. Polygonum-type/

2. Allium-type/

3. Adoxa-type/

4. Penaea-type/

5. Scilla-type/

6. Drusa-type/

7. Fritillaria-type/

8. Oenothera-type/

9. Peperomia-type/

10. Plumbago-type/

11. Chrysanthemum-type/

12. Clintonia-type/

#392. Polar nuclei <timing of fusion: data from Davis 1966>/

1. fusing prior to fertilization/

2. fusing only after one has been fertilized/

3. fusing simultaneously with the male gamete <‘triple fusion’>/

4. not fusing/

Data from Davis (1966). The ambiguous ‘fusing in association with a male gamete’ and ‘fusing at fertilization’ have been encoded ‘2/3’.

#393. Antipodal cells <with walls, whether formed: data from Davis 1966>/

1. formed/

2. not formed/

#394. Antipodal cells <number, when not proliferating: data from Davis 1966>/

#395. Antipodal cells <whether proliferating: data from Davis 1966>/

1. proliferating/

2. not proliferating/

#396. Antipodal cells <whether persistent beyond fertilization: data from Davis 1966>/

1. ephemeral/

2. persistent/

#397. Synergids <shape. ‘Filiform apparatus’ recorded as text comment: data from Davis 1966>/

1. pear-shaped/

2. hooked/

#398. Synergids <whether haustorial: data from Davis 1966>/

1. haustorial/

2. non-haustorial/

#399. Hypostase <presence: data from Davis 1966 (no negatives)>/

1. present/

2. absent/

#400. Endosperm formation <data mainly from Davis 1966>/

1. cellular/

2. nuclear/

3. helobial/

#401. Endosperm haustoria <presence: data from Davis 1966 (no negatives)>/

1. present/

2. absent/

#402. Endosperm haustoria <type, with ‘terminal’ encoded as 1&2: data from Davis 1966>/

1. chalazal/

2. micropylar/

3. lateral/

#403. Embryogeny <type>/

1. onagrad/

2. asterad/

3. caryophyllad/

4. solanad/

5. chenopodiad/

6. piperad/

#### Fruit

#404. Fruit <texture at maturity>/

1. <at least partially> fleshy/

2. non-fleshy <when mature>/

#405. Fruit <whether an aggregate>/

1. an aggregate <of more or less discrete units on a common receptacle: the product of an apocarpous, semicarpous or synstylous gynoecium>/

2. not an aggregate <i.e. a single one-carpelled or syncarpous entity> /

‘Aggregate fruit’: one representing aggregation (clustering or coherence) of the discrete units (carpels) from a single synstylous, semicarpous or apocarpous gynoecium (e.g. blackberry, buttercup). Not to be confused with ‘multiple fruits’ (q.v.) derived from more than one flower.

#406. The fruiting carpels <of the aggregate fruit, whether coalescing>/

1. coalescing into a secondary syncarp/

2. not coalescing /

#### From apocarpous, semicarpous and monomeric gynoecia

#407. The fruiting carpel <of monomeric G, or individual carpels of aggregate fruits, whether dehiscent>/

1. dehiscent/

2. indehiscent/

#408. The fruiting carpel <of monomeric G, or individual carpels of aggregate fruits, type>/

1. a legume/

2. a loment/

3. a follicle/

4. an achene <small, hard, dry, from one free carpel>/

5. samaroid/

6. nucular/

7. drupaceous/

8. baccate <a berrylet>/

‘Legume’: a fruit derived from a single carpel, which dehisces both dorsally (along the placenta) and ventrally (the commonest type in Leguminosae).

‘Loment’: a fruit derived from a single carpel, which at maturity breaks transversely into (usually one-seeded) segments.

‘Follicle’: a fruit derived from a single carpel, which dehisces only along the ventral suture.

‘Achene’: a fruit derived from a single carpel which is small, hard, dry, one-seeded, and with a tight, thin pericarp.

‘Samaroid’: a winged fruit derived from a single carpel.

‘Nucular’: a fruit which is nutlike, but derived from a single carpel.

‘Drupaceous’: a fruit which is drupelike, but derived from a single carpel.

‘Baccate’: a fruit which is berrylike, but derived from a single carpel (a ‘berrylet’).

‘Aggregate fruit’: one representing aggregation (clustering or coherence) of the discrete units (carpels) from a single synstylous, semicarpous or apocarpous gynoecium (e.g. blackberry, buttercup). Not to be confused with ‘multiple fruits’ (q.v.) derived from more than one flower.

#### Syncarpous fruits

#409. Fruit <syncarpous, dehiscence>/

1. dehiscent <opening to release the seeds>/

2. indehiscent <neither opening nor breaking up in any precise fashion at maturity>/

3. a schizocarp <splitting longitudinally into non- or tardily dehiscent, fruitlike mericarps>/

4. lomentaceous <breaking transversely into segments>/

‘Dehiscent’: a fruit opening in anatomically organized, predetermined fashion to release the seeds.

‘Indehiscent’: a fruit with no precise, anatomical mechanism for opening to release the seeds (rotting, irregularly rupturing, ruptured by seedling germination, opened by external agencies, etc.).

‘Schizocarp’: a syncarpous fruit which splits longitudinally into non- or tardily dehiscent, fruitlike parts (mericarps, these generally corresponding with locules or locelli). Useage restricting the term to dry, bipartite fruits (Umbelliferae, Aceraceae, e.g. Lawrence 1951) is idiosyncratic and inconvenient.

‘Lomentaceous’: a syncarpous fruit which breaks transversely into segments, resembling a loment (q.v.).

#410. Mericarps <number>/

‘Mericarps’: the separate, fruitlike derivatives of a schizocarp (q.v.), regardless of their form and texture (cf. Jackson 1928 etc.). Useage restricting the term to dry, bipartite fruits (e.g. Lawrence 1951) is idiosyncratic and inconvenient.

#411. Mericarps <of schizocarpic fruits, type>/

1. comprising achenes/

2. comprising berrylets/

3. comprising follicles/

4. comprising nutlets/

5. samaroid/

6. comprising ‘legumes’/

7. comprising drupelets/

‘Achene’: a fruit derived from a single carpel which is small, hard, dry, one-seeded, and with a tight, thin pericarp.

‘Berrylet’: a berrylike mericarp.

‘Follicle’: a fruit derived from a single carpel, which dehisces only along the ventral suture.

‘Nutlet’: a nutlike mericarp.

‘Samaroid’: winged.

‘Legume’: a fruit derived from a single carpel, which dehisces both dorsally (along the placenta) and ventrally: the commonest type in Leguminosae.

‘Drupelet’: a drupelike mericarp.

#412. Fruit <syncarpous type, representing two or more carpels>/

1. a capsule <dry, dehiscent, from two or more carpels>/

2. capsular-indehiscent/

3. achene-like/

4. a silicula <dry, bilocular, of two carpels, longitudinally two-valved, length less than three times breadth>/

5. a siliqua <dry, bilocular, of two carpels, longitudinally two-valved, length at least three times breadth>/

6. a berry <fleshy, without a stony layer, usually several to many seeded>/

7. a drupe <fleshy, with each of the one or more seeds surrounded by a stony layer>/

8. a nut <dry, hard, indehiscent, one-seeded>/

9. a cypsella <‘achene’ from two carpels, invested by the adnate calyx>/

10. a samara <indehiscent, winged by extension of the fruit wall>/

11. a caryopsis/

‘Capsule’: a dry, dehiscent fruit, from two or more carpels (but exclusive of ‘silicula’ and ‘siliqua’, see below).

‘Capsular-indehiscent’: cf. ‘capsule’ — relatively thin walled and containing free seeds, but indehiscent.

‘Achene-like’: small, hard, dry, one-seeded, the pericarp tight and thin — cf. an achene (q.v.), but from a syncarpous gynoecium.

‘Silicula’: dry, bilocular, of two carpels, longitudinally two-valved (q.v. ‘valved’), length less than three times breadth.

‘Siliqua’: dry, bilocular, of two carpels, longitudinally two-valved (q.v. ‘valved’), length at least three times breadth.

‘Berry’: a fleshy syncarp, without a stony layer, usually several to many seeded.

‘Drupe’: a fleshy syncarp, with the one or more seeds surrounded by a stony layer.

‘Nut’: an indehiscent, one-seeded syncarp, with a hard, dry, woody pericarp from which the seed is free.

‘Cypsella’: an ‘achene’ from two carpels, invested by the adnate calyx.

‘Samara’: an indehiscent syncarp, winged by extension of the fruit wall.

‘Caryopsis’: syncarpous but one celled, one seeded, superior, and with the thin pericarp united to the seed (the ‘grain’ typical of grasses.

‘Syncarp’: a fruit representing two or more fused carpels.

‘Valved’: opening by a flap (as opposed to a slit).

#413. Fruit <superior or perigynous, fleshy investment external to the original superior or half-inferior ovary>/

1. enclosed in the fleshy receptacle/

2. enclosed in the fleshy hypanthium/

3. enclosed in the fleshy perianth/

4. without fleshy investment external to the original ovary /

#414. Capsules <syncarpous: dehiscence type - important, but currently unsatisfactory in that states 2, 4, 5, 6 are not mutually exclusive, and published descriptions are often imprecise (and unreliable?)>/

1. splitting irregularly/

2. denticidal/

3. poricidal/

4. septicidal/

5. loculicidal/

6. valvular <including ‘septifragal’>/

7. circumscissile <operculate, a pyxis>/

#415. The drupes <whether exhibiting separable pyrenes>/

1. with separable pyrenes/

2. with one stone <not separable pyrenes>/

#### Fruits general

#416. Fruit <dehiscent, whether dehiscence elastic>/

1. elastically dehiscent/

2. passively dehiscent/

#417. Gynoecia of adjoining flowers <whether forming a multiple fruit>/

1. combining to form a multiple fruit/

2. not forming a multiple fruit /

‘Multiple fruit’: one derived from the gynoecia of more than one flower (e.g. mulberry, pineapple), or representing an inflorescence. Not to be confused with ‘aggregate fruit’, q.v.

#418. The multiple fruits <from separate flowers, whether coalescing>/

1. coalescing/

2. not coalescing /

#419. Dispersal unit <type; states inadequately defined (see Notes)>/

1. the seed/

2. the mericarp/

3. the fruit/

4. the flower/

5. the inflorescence/

This character was included for its potential biological interest. However, the current states ‘fruit’ and ‘flower’ are inadequate, for example to deal with the various combinations of inferior ovaries and persistent perianth parts. Consequently, few families have been encoded.

#420. Dispersal <manner> /

#421. Fruit <number of seeds maturing: ‘many’ indicated by 100>/

seeded/

#422. Seeds <when mature, whether endospermic>/

1. endospermic/

2. non-endospermic/

#423. Endosperm <whether ruminate>/

1. ruminate/

2. not ruminate /

#424. <Mature> endosperm <whether oily>/

1. oily/

2. not oily/

#425. Perisperm <presence>/

1. present/

2. absent /

#426. Seeds <size: a ‘casual’ character, reflecting their greatest dimension (see Notes)>/

1. minute <1mm or less>/

2. small <1mm to 1cm>/

3. medium sized <5mm to 4cm>/

4. large <3 to 12cm>/

5. very large <greater than 10cm>/

#427. Seeds <whether conspicuously hairy>/

1. conspicuously hairy/

2. not conspicuously hairy /

#428. Seeds <presence of testa>/

1. with a testa /

2. without a testa/

#429. Seeds <whether winged>/

1. winged/

2. wingless /

#430. <Mature> seeds <whether storing starch>/

1. with starch/

2. without starch/

#431. <Mature> seeds <occurrence of amyloid protein: see Kooiman 1960 - no negatives available>/

1. with amyloid/

2. without amyloid <implicit when chemistry ‘known’>/

#432. Embryo <development>/

1. rudimentary at the time of seed release <not differentiated into organs>/

2. weakly differentiated/

3. well differentiated /

#433. Cotyledons <number>/

#434. Cotyledons <in the seed, how borne>/

1. flat/

2. folded /

3. rolled/

4. twisted/

5. crumpled/

#435. Embryo <chlorophyllous or achlorophyllous: data from Yakovlev and Zhukova 1980 - species/genera sampled in parenthesis>/

1. chlorophyllous/

2. achlorophyllous/

#436. Embryo <shape>/

1. straight/

2. curved/

3. bent/

4. coiled/

5. other than straight, curved, bent or coiled /

#437. The radicle <position when turned up against the cotyledons>/

1. lateral <aligned with the commissure of the cotyledons; cotyledons accumbent>/

2. dorsal <on the back of a cotyledon; cotyledons incumbent>/

#438. Embryo <length/total seed length>/

the length of the seed/

#439. Testa <whether operculate>/

1. operculate/

2. non-operculate/

#440. Testa <surface>/

1. smooth/

2. with spines/

3. with tubercles/

4. hairy/

#441. Testa <colour>/

1. homogeneous in colour <ignore raphe>/

2. conspicuously coloured - patterned/

#442. Testa <of the seed, whether with phytomelan>/

1. encrusted with phytomelan/

2. without phytomelan /

Data mainly from Dahlgren et al. (1985), supplemented from Stevenson and Loconte (1995).

#443. Testa <colour>/

1. grey/

2. black/

3. green/

4. brown/

5. orange/

6. yellow/

7. red/

#444. Micropyle <whether zigzag>/

1. zigzag/

2. not zigzag /

#445. Polyembryony <whether observed>/

1. recorded/

2. not recorded/

#446. Seed germination <whether 'orchidaceous'>/

1. resulting first in development of an endomycorrhizal 'protocorm' <orchidceous>/

2. not 'orchidaceous' <not involving an endomycorrhizal 'protocorm': implicit>/

#### Seedling

#447. Germination <type, phanerocotylar or cryptocotylar: Dicots only>/

1. phanerocotylar <epigeal, epicotylar>/

2. cryptocotylar <hypogeal>/

Data mainly communicated by H.T. Clifford (1987).

This character is widely inapplicable to Monocots, where the single cotyledon is commonly partly below ground and partly raised above it, and where structures of obscure homologies may be involved (e.g. a coleoptile, q.v.).

#448. <More or less elongated> hypocotyl internode <presence>/

1. present/

2. absent/

Data mainly for Monocots, from Tillich (1995).

#449. <More or less elongated> mesocotyl <presence (see Notes)>/

1. present/

2. absent/

‘Mesocotyl’: a structure representing congenital fusion of the epicotyl (q.v.) with the basal part of the coleoptile (q.v.), which may be conspicuously elongated (e.g. in many grass seedlings).

Data from Tillich (1995).

#450. Seedling collar <between shoot and primary root, conspicuous or not>/

1. conspicuous <massively developed, or appendaged>/

2. not conspicuous/

‘Collar’: the transitional region between the hypocotyl and the primary root of a seedling, or the lowest part of the hypocotyl when the latter is present. Usually detectable as a narrow line, but sometimes appendaged or otherwise conspicuously developed (‘conspicuous’).

Data for Monocots only, from Tillich (1995).

#451. Cotyledon hyperphyll <elongated or compact>/

1. <more or less> elongated <proximally to the haustorium, if this present>/

2. compact <= mainly haustorial>/

Data for Monocots only, from Tillich (1995).

#452. Cotyledon hyperphyll <whether assimilatory (see Notes)>/

1. assimilatory <at least partially green, implying epigeal>/

2. non-assimilatory <not green, implying hypogeal>/

Data for Monocots only, from Tillich (1995).

#453. Cotyledon hyperphyll <whether dorsiventrally flattened>/

1. more or less circular in t.s./

2. dorsiventrally flattened/

Data for Monocots only, from Tillich (1995).

Unfortunately, ‘circular in t.s.’ and ‘dorsiventrally flattened’ are not reliably indicative of anatomically unifacial versus bifacial.

#454. Coleoptile <in seedling, presence>/

1. present/

2. absent/

‘Coleoptile’: a tubular elongation of the (single), closed cotyledon sheath, peculiar to a few families of Monocotyledons.

Data from Tillich (1995).

#455. <Monocot> seedling <whether macropodous>/

1. macropodous <with nutrients stored in both hypocotyl and cotyledon>/

2. non-macropodous <with stored nutrients confined to the cotyledon>/

Data from Tillich (1995).

#456. Seedling cataphylls <presence>/

1. present/

2. absent/

Data mainly for Monocots, from Tillich (1995).

#457. First <seedling> leaf <excluding cataphylls, form>/

1. ensiform <bilateral>/

2. centric <unifacial, oval to circular in t.s.>/

3. dorsiventral <bifacial, ‘normally’ orientated>/

Data for Monocots only, from Tillich (1995).

#458. Primary root <persistence in seedlings>/

1. persistent <developing laterals>/

2. ephemeral/

Typically persistent and robust in Dicots, and treated here as unreliably implicit for most of them. Data for Monocots from Tillich (1995).

#459. Primary root <number of poles>/

polar/

#460. Nitrogen-fixing root nodules <presence>/

1. present/

2. absent /

#### Physiology, phytochemistry

#461. <Photosynthetic pathway (see Notes):>/

1. C3 <as evidenced by physiological experiment, or reliably indicated by ‘non-C4’ anatomy or high negative delta13C values - omitting cases with grounds for suspecting CAM>/

2. C4 <as evidenced by physiology, or predicted by unambiguous anatomical data or cautious interpretation of delta13C values - omitting forms suspected capable of CAM>/

3. <shown to be capable of> CAM <see Teeri 1982 for definition: dubious records of CAM omitted>/

4. <reliably demonstrated> C3-C4 intermediate/

The data represent an attempt at comprehensive compilation from reliable photosynthetic pathway-related data, except that no large scale, organized attempt has been made to pursue the leaf anatomical literature for illustrations adequate for anatomical predictions.

For sources of data, see references tagged PP in the References. Re problems in recording CAM/C3, see Martin et al. 1982/

#462. C3 physiology recorded directly in <generic list>/

For sources of data, see references tagged PP.

#463. C4 physiology recorded directly in <generic list>/

For sources of data, see references tagged PP.

#464. <A capacity for> CAM recorded directly in <generic list>/

For sources of data, see references tagged PP. The records originally entered here have been updated with reference to the list of CAM positive genera provided by Smith and Winter (1996). Unfortunately, as with so many features of potential taxonomic and phylogenetic interest, the necessity for recording hard data on negatives continues to be overlooked.

#465. C3-C4 intermediacy <convincingly demonstrated> in <generic list>/

For sources of data, see references tagged PP.

#466. Anatomy <photosynthetic pathway-linked pattern>/

1. C4 type <indicated by ‘Kranz’, or maximum cells-distant count = 1: used here as reliable indication of C4>/

2. non-C4 type <non-Kranz, maximum cells-distant count more than 1: taken to indicate C3 only in the absence of evidence of (or reason to suspect) CAM>/

No organized, large scale attempt has been made here to pursue the literature of plant leaf anatomy for illustrations of transverse sections adequate for reliable C3/C4 assignment.

For an exposition on applying the 'cells-distant' criterion to predicting photosynthetic pathways, see Hattersly and Watson (1975 and 1976). For further information, including pursuit of 'unconventional' C3/C4 anatomical configurations via immunofluorscent labelling of Rubisco in the PCR cells, see Hattersley, Watson and Osmond (1977).

#467. <RuBisCO Km(CO2) and Km(RUBP) data. To view tables with comparative data, display via Intkey the illustrations associated with this character>/

Comparative data consist of tables from Yeoh, Badger and Watson (1980, 1981), and can be displayed as illustrations of this character.

#468. 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’/

#469. Sugars transported <substances involved in phloem transport, cf. Ziegler 1974>/

1. <almost entirely> as sucrose <only>/

2. <almost entirely> as oligosaccharides <raffinose, stachyose, verbascose, ajugose> + sucrose/

3. as <significant proportions of> sugar alcohols <D-mannitol, sorbitol, dulcitol, myoinositol> + oligosaccharides + sucrose/

Data mainly from Zimmermann and Ziegler (1975). The difficulty of interpreting quantitative chromatographic data has rendered the encoding fairly subjective. For example, tr (trace) records have been ignored. Any taxonomic patterns emerging from Intkey interrogations of the descriptions as encoded here should certainly not be accepted as meaningful without first comparing them in detail with the original publications.

#470. Inulin <polysaccharide, whether recorded for the family>/

1. recorded/

2. not found /

Data mainly from Gibbs (1974).

#471. Mustard-oils <myrosin cells: glucosinolate–myrosinase complex, presence>/

1. present/

2. absent <implicit when chemistry ‘known’>/

See especially Kjaer (1973).

#472. <Whether ‘cyanogenic’ (data mainly from Gibbs 1974):>/

1. cyanogenic/

2. not cyanogenic/

Gibbs (1974) provides extensive data on cyanogenesis, including negative records, derived via consistent application of his published method. ‘Not mentioned’ in the present data cannot be interpreted as indicating the negative state.

Some published compilations and taxonomists’ generalizations (e.g. Cronquist 1981) are probably bedevilled by spurious positives.

#473. Cynogenic constituents <derivation: Hegnauer 1977>/

1. tyrosine-derived /

2. phenylalanine-derived/

3. of the gynocardin group/

4. of Hegnauer’s ‘Group C’/

5. leucine-derived/

#474. Polyacetylenes <presence in the family>/

1. recorded <for the family>/

2. not found <implicit when chemistry ‘known’>/

#475. Alkaloids <presence>/

1. present/

2. absent/

#476. Alkaloids <type>/

1. C6C2-N-C2C6/

2. C6C1-N-C2 C6/

3. C6C3-N-C2C6/

#477. Berberine <yellow isoquinoline alkaloid, presence>/

1. present <i.e., recorded in the family>/

2. absent /

#478. <Amino acid profiles of leaves. To view tables of comparative data, display via Intkey the illustrations associated with this character>/

Comparative data consist of tables from Yeoh, Wee and Watson (1986), and can be displayed as illustrations of this character.

#479. Anthraquinones <presence in the family>/

1. detected/

2. <sought, but> not detected <in any of the family representatives screened>/

Data mainly from Jensen, 1992 and pers. comm. As he points out, negative results are usually not reported, so it is inappropriate to treat ‘absent’ as the implicit state (cf. comments under ‘iridoids’).

#480. Anthraquinones <type>/

1. derived from shikimic acid/

2. polyacetate derived/

Data mainly from Jensen, 1992 and pers. comm.

#481. Verbascosides <sensu Jensen 1992 - caffeic acid derivatives, orobanchoside, verbascoside>/

1. detected/

2. <sought, but> not detected <in any of the family representatives screened>/

Data mainly from Jensen, 1992 and pers. comm. As he points out, negative results are usually not reported, so it is inappropriate to treat ‘absent’ as the implicit state (cf. comments under ‘iridoids’).

#482. Cornoside <presence in the family>/

1. detected/

2. <sought, but> not detected <in any of the family representatives screened>/

Data mainly from Jensen, 1992 and pers. comm.

#483. Arbutin <phenolic heterosidic glycoside, presence>/

1. present/

2. absent /

Data from Gibbs (1974).

#484. Iridoids <glycosides, presence in the family>/

1. detected/

2. <sought, but> not detected <in any of the family representatives screened>/

Data mainly from S.R. Jensen and collaborators 1975–1996, including compilations, original observations and unpublished records; and Gibbs 1974 (see the accompanying References).

Encoding has been organized here on the assumption that the most generally useful expression for taxonomic purposes will be ‘iridoids sought and found in at least one representative of the family’, contrasted with ‘iridoids sought in the family, but not found’.

Though of much taxonomic interest, as with most data on ‘esoteric’ features, these require careful interpretation for use in taxonomic analyses. There is often variation re presence and constitution of iridoids within families, but for most relatively few representatives have been screened. Most of the information in the source references seems to represent analyses of leaf material, but a positive record for Menyanthes (used in published cladistic studies) represents loganin reported for the rhizome, the leaves having been found Ehrlich-negative.

Explicit information on ‘absence’ is scarce (in the present data, much of it derives from the negative Ehrlich tests recorded by Gibbs). In employing the present compilation, therefore, ‘iridoids absent’ cannot be routinely regarded as implicit.

#485. Iridoids <type (see Jensen 1991)>/

1. ‘Route I’ type <seco-iridoids and CIAs, derived from iridodial>/

2. ‘Route II’ type <derived from epi-irododial>/

Data mainly from Jensen 1991, 1992 and pers. comm.: see References.

#486. Betalains <(betacyanin and betaxanthin glycosides) presence>/

1. present/

2. absent <implicit when chemistry ‘known’>/

#487. Saponins/sapogenins <(triterpenoid glycosides) presence>/

1. present/

2. absent/

#488. Proanthocyanidins <(= leucoanthocyanidins, condensed tannins) presence in leaves or equivalent structures of leucodelphinidin and/or leucocyanidin>/

1. present/

2. absent/

Data from Bate-Smith (1962, 1968).

#489. Proanthocyanidins <type present in leaves or equivalent structures>/

1. cyanidin <2-hydroxy>/

2. delphinidin <3-hydroxy>/

Data from Bate-Smith (1962, 1968).

#490. Flavonols <presence in leaves or equivalent structures>/

1. present/

2. absent/

Data from Bate-Smith (1962, 1968).

#491. Flavonols <type present in leaves or equivalent structures>/

1. kaempferol <1-hydroxy>/

2. quercetin <2-hydroxy>/

3. myricetin <3-hydroxy>/

Data from Bate-Smith (1962, 1968).

#492. Ellagic acid <(3-hydroxy) presence in leaves or equivalent structures (indicative of ellagitannins)>/

1. present/

2. absent/

Judging from Bate-Smith’s (1962, 1968) data, the mono- and di-hydroxy compounds p-coumaric acid and caffeic acid, along with sinapic and ferulic acid, seemed unlikely to justify taxonomically the effort of encoding them. But see ‘verbascosides’.

#493. Andromedotoxin <grayonotoxin, toxic diterpene: presence in the family>/

1. recorded/

2. not recorded <implicit when chemistry ‘known’>/

#494. Ursolic acid <(triterpene) presence>/

1. present/

2. absent/

Data from Gibbs (1974).

#495. Aluminium accumulation <whether demonstrated>/

1. demonstrated <in at least some members of the family: the leaves becoming conspicuously yellow-green on drying>/

2. not found <i.e. in any of the species tested>/

As indicated by the leaves becoming conspicuously yellow-green on drying. Encoding here of ‘aluminium accumulation not found’ implies the existence of at least one explicit negative record for the family. Data mainly from Chenery (1948), Webb (1954), Gibbs (1974), Chenery and Sporne (1976).

#496. Plants <whether accumulating free oxalates>/

1. accumulating free oxalates/

2. not accumulating free oxalates/

#497. 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.

#498. 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>/

#### Peculiar features

#499. <Seeds, whether borne on elongated, indurated, hook-shaped funicles (‘retinacula’) - Acanthaceae:>/

1. the seeds on elongated, indurated, hook-shaped funicles (‘retinacula’)/

2. the funicles not as in <the majority of> Acanthaceae /

#500. <Whether mangrove, with pneumatophores etc.:>/

1. mangroves <with pneumatophores, vivipary etc.>/

2. non-mangrove species /

#501. <Whether the young, syncarpous unilocular gynoecium and later the capsule are open - Resedaceae:>/

1. the young, syncarpous unilocular gynoecium and later the capsule open/

2. gynoecium and fruit not open as in typical Resedaceae /

#502. Lamina <whether abaxially pouched at the tip - Saccifoliaceae>/

1. exhibiting revolute margins continuous with a characteristic, abaxially pouched tip <Saccifoliaceae>/

2. tip not abaxially pouched (i.e., not as in Saccifoliaceae) /

#503. Epicalyx <whether double - Triplostegiaceae type>/

1. double, comprising an outer one of four conspicuous, basally connate, capitate-glandular, apically uncinate cuspid, persistent bracts, and an urceolate, 8-ribbed, persistent inner one <Triplostegiaceae>/

2. not double and complex as in Triplostegiaceae /

#### Geography, cytology, taxonomy

#504. <Floristic Kingdoms (see Notes):>/

1. Holarctic/

2. Paleotropical/

3. Neotropical/

4. Cape/

5. Australian/

6. Antarctic/

Intended to reflect ‘natural’ distributions, insofar as these are determinable. The phytogeographical Kingdoms, Subkingdoms, Regions and Subregions are those of Takhtajan (1969). Data for Liliiflorae provided by J.G. Conran (pers. comm., 1993); for the rest, the data are very incomplete.

#505. <Holarctic Subkingdoms (see Notes):>/

1. Boreal/

2. Tethyan <ancient Mediterranean>/

3. Madrean <Sonoran: SW U.S.A. and Mexico>/

#506. <Paleotropical Subkingdoms (see Notes):>/

1. African/

2. Madagascan/

3. Indomalesian/

4. Polynesian/

5. Neocaledonian/

#507. <Boreal Subkingdom regions (see Notes):>/

1. Arctic and Subarctic/

2. Euro-Siberian/

3. Eastern Asian/

4. Atlantic North American/

5. Rocky Mountains/

#508. <Tethyan Subkingdom regions (see Notes):>/

1. Macaronesian/

2. Mediterranean/

3. Irano-Turanian/

#509. <African Subkingdom regions (see Notes):>/

1. Saharo-Sindian/

2. Sudano-Angolan/

3. West African Rainforest/

4. Namib-Karroo/

5. Ascension and St. Helena/

#510. <Indomalesian Subkingdom regions (see Notes):>/

1. Indian/

2. Indo-Chinese/

3. Malesian <Malayan>/

4. Papuan/

#511. <Polynesian Subkingdom regions (see Notes):>/

1. Hawaiian/

2. Polynesian/

3. Fijian/

#512. <Neotropical regions (see Notes):>/

1. Caribbean/

2. Venezuela and Surinam/

3. Amazon/

4. Central Brazilian/

5. Pampas/

6. Andean/

7. Fernandezian/

#513. <Australian regions (see Notes):>/

1. North and East Australian/

2. South-West Australian/

3. Central Australian/

#514. <Antarctic regions (see Notes):>/

1. New Zealand/

2. Patagonian/

3. Antarctic and Subantarctic/

#515. <Euro-Siberian Subregions:>/

1. European Subregion/

2. Siberian Subregion/

#516. <Atlantic North American Subregions:>/

1. Canadian-Appalachian Subregion/

2. Southern Atlantic North American Subregion/

3. Central Grasslands Subregion/

#517. <Sudano-Angolan Subregions:>/

1. Sahelo-Sudanian Subregion/

2. Somalo-Ethiopian Subregion/

3. South Tropical African Subregion/

4. Kalaharian Subregion/

#518. <North and East Australian Subregions:>/

1. Tropical North and East Australian Subregion/

2. Temperate and South-Eastern Australian Subregion/

#519. <Antarctic and Subantarctic Subregions:>/

1. South Temperate Oceanic Islands Subregion/

2. Antarctic Subregion/

#520. <Climatic distribution:>/

1. frigid zone/

2. temperate/

3. sub-tropical/

4. tropical/

#521. <World distribution:>/

This text character has been comprehensively recorded, and incorporates ‘colloquial’ summaries of natural distribution, cf. Willis’s ‘Dictionary’.

#522. <Geographical distribution (encoding mandatory, but intended for keys only):>/

1. Western and Northern Eurasia, U.S.S.R./

2. Middle East <Turkey to North Africa, and eastwards to Iran>/

3. Mediterranean/

4. Eastern and Southern Asia <including Japan, China, India>/

5. North Africa/

6. Central, East tropical and West tropical Africa/

7. South Africa/

8. Madagascar/

9. Malay Archipelago <including New Guinea>/

10. Australasia <Australia, New Zealand>/

11. Pacific Islands/

12. North America/

13. Central America/

14. West Indies/

15. South America/

16. Arctic/

17. Antarctic/

Comprehensively encoded, but intended only for use in identification and for generating useful geographic subsets of the data. Assignments to these pragmatically defined world regions are intended to reflect likelihood of the family being encountered ‘in the field’, regardless of floristic status (i.e., including introduced weeds). Requires continuing updating ......

#523. Chromosomes <presence of diffuse centromeres: Dahlgren and Clifford 1982>/

1. with diffuse <non-localized> centromeres/

2. with normal <concentrated> centromeres <implicit when cytology ‘known’>/

#524. <Generic chromosome numbers (preferably X (basic); n or 2n when X not specified):>/

#525. Supposed basic chromosome number of family:/

#526. Ploidy levels recorded:/

#527. Subclass <Monocot/Dicot>/

1. Dicotyledonae <Magnoliopsida, Magnoliatae>/

2. Monocotyledonae <Liliopsida>/

This package incorporates the complete classifications to ordinal and supra-ordinal levels of Dahlgren (1980, for Dicots) and of Dahlgren, Clifford and Yeo (1985, for monocots), that of Cronquist (1981) for Dicots only, and the formal and informal suprafamilial groupings of the APG (APG III and IV, 2009 and 2016). The later of these aims to account for the plethora of recent cladistic and (more significantly) molecular-cladistic molecular phylogenetic studies. It replaces the earlier version of their treatment, and relatively few families remain unassigned at various levels. Note that the lists of family synonyms provided in APG papers (many of which are represented by separate descriptions in the present package) do not indicate which of them have been sampled in DNA sequencing studies: some have, but many probably have not. Rather than presenting them all as unclassified under the APG scheme, they are referred here to the (presumably) appropriate groups with the comment ‘ . . . as a synonym of . . . ’ against the ordinal assignment. While the Ordinal assignments here are intended to be consistent with theirs in terms of generic content, numerous APG re-alignments at family level, some of which are no doubt justified, simply cannot be implemented here in the absence of fully reorganized, detailed descriptions. The latter would have to be laboriously extacted from a mountain of publications which mostly purvey non-comparative and often hopelessly incomplete data.

It has long been apparent (cf. Young and Watson 1970) that Cronquist’s largest Subclasses of the Dicotyledons, Rosidae and Dilleniidae, had little or no taxonomic merit, and that his Asteridae constituted a less informative grouping than the Gamopetalae or Sympetalae of Nineteenth Century systems. Dahlgren’s scheme (viewed in conjunction with his diagrammatic representation) was obviously better, but his classification could be much improved, especially in connection with experimentalists’ requirements, by superimposing our ‘Crassinucelli’/‘Tenuinucelli’ subdivision on his 24 Superorders. For the present purpose, ‘Crassinucelli’/‘Tenuinucelli’ status has been assigned on correlated attributes, the nucellus being the best predictor (though not absolutely reliable) when data are available. Classification of Dicots into meaningful Orders is easier, and the Dahlgren and Cronquist schemes extensively coincided. Comparisons between the contents of their orders, of theirs with the APG classification, and assessments of the character state distributions they all supposedly reflect, are easily conducted by applying INTKEY.

Various details of the Dahlgren scheme for Dicots were adjusted in the course of compiling these descriptions, mainly to account for new information (e.g. families more recently described, referal of glucosinolate families to Capparales, and re-assignments in relation to the Araliiflorae/Corniflorae conundrum). For Monocots, the Dahlgren et al (1985) classification alone was incorporated, being obviously better than Cronquist’s, although it was evidently far from the last word. Earlier schemes (including Cronquist’s) are rendered extensively useless by gross sensu lato treatment of the family Liliaceae.

The exciting classificatory ideas now emerging from comparative nucleic acid studies exeplified by Chase et al. (1993 et seq.) will result in completely revised system within the next few years; but will demand congruent results from DNA comparisons (nuclear, as well as chloroplast) which can reasonably be expected to represent whole genomes, as well as thorough coverage of genera and species. Definitive phylogenetic conclusions will not be achieved by comparing one or two genes, and phylogenies based entirely on plasid DNA in particular do not inspire confidence unless the results are consistent with other comparative data. Furthermore, it seems that no attention has yet been paid to likely past occurrences of lateral transfer of genetic information at high hierarchical levels, for example through the agency of parasitic bacteria, fungi and viruses. A commonsense approach to assessing the phylogenetic credentials of purported phylogenies and revised classifications will require properly organised scrutiny of conventional character state distributions at all levels, as will preparation of comparative group descriptions: operations which are rendered relatively easy by the present package, and which were specifically evisaged in its design (Watson and Dallwitz, 1992).

#528. <Main Dicot grouping, Crassinucelli or Tenuinucelli (see Notes):>/

1. Crassinucelli <cf. Archichlamydeae>/

2. Tenuinucelli <cf. Gamopetalae, Sympetalae>/

#### Classification after Dahlgren et al.

#529. Dahlgren’s Superorder <of Dicotyledonae, modified from Dahlgren 1980>/

1. Magnoliiflorae/

2. Nymphaeiflorae/

3. Ranunculiflorae/

4. Caryophylliflorae/

5. Polygoniflorae/

6. Malviflorae/

7. Violiflorae/

8. Theiflorae/

9. Primuliflorae/

10. Rosiflorae/

11. Podostemiflorae/

12. Fabiflorae/

13. Proteiflorae/

14. Myrtiflorae/

15. Rutiflorae/

16. Santaliflorae/

17. Balanophoriflorae/

18. Araliiflorae/

19. Asteriflorae/

20. Solaniflorae/

21. Corniflorae/

22. Loasiflorae/

23. Gentianiflorae/

24. Lamiiflorae/

#530. Dahlgren et al. Superorder <of Monocots: Dahlgren, Clifford Order and Yeo 1985>/

1. Alismatiflorae/

2. Triuridiflorae/

3. Ariflorae/

4. Liliiflorae/

5. Zingiberiflorae/

6. Commeliniflorae/

7. Areciflorae/

8. Cyclanthiflorae/

9. Pandaniflorae/

10. Bromeliiflorae/

#531. <Dahlgren’s Order of Magnoliiflorae:>/

1. Annonales/

2. Aristolochiales/

3. Magnoliales/

4. Illiciales/

5. Laurales/

6. Nelumbonales/

#532. <Dahlgren’s Order of Nymphaeiflorae:>/

1. Piperales/

2. Nymphaeales/

#533. <Dahlgren’s Order of Ranunculiflorae:>/

1. Ranunculales/

2. Papaverales/

#534. <Dahlgren’s Order of Caryophylliflorae:>/

1. Caryophyllales/

#535. <Dahlgren’s Order of Polygoniflorae:>/

1. Polygonales/

#536. <Dahlgren’s Order of Malviflorae:>/

1. Paeoniales/

2. Dilleniales/

3. Malvales/

4. Urticales/

5. Rhamnales/

6. Elaeagnales/

7. Plumbaginales/

8. Thymelaeales/

9. Euphorbiales/

#537. <Dahlgren’s Order of Violiflorae:>/

1. Violales/

2. Salicales/

3. Tamaricales/

4. Capparales/

5. Salvadorales/

#538. <Dahlgren’s Order of Theiflorae:>/

1. Theales/

2. Droserales/

#539. <Dahlgren’s Order of Primuliflorae:>/

1. Ebenales/

2. Primulales/

#540. <Dahlgren’s Order of Rosiflorae:>/

1. Trochodendrales/

2. Hamamelidales/

3. Fagales/

4. Balanopales/

5. Juglandales/

6. Myricales/

7. Casuarinales/

8. Buxales/

9. Cunoniales/

10. Saxifragales/

11. Gunnerales/

12. Rosales/

#541. <Dahlgren’s Order of Podostemiflorae:>/

1. Podostemales/

#542. <Dahlgren’s Order of Fabiflorae:>/

1. Fabales/

#543. <Dahlgren’s Order of Proteiflorae:>/

1. Proteales/

#544. <Dahlgren’s Order of Myrtiflorae:>/

1. Myrtales/

2. Haloragales/

3. Rhizophorales/

#545. <Dahlgren’s Order of Rutiflorae:>/

1. Rutales/

2. Sapindales/

3. Balsaminales/

4. Polygalales/

5. Geraniales/

6. Tropaeolales/

#546. <Dahlgren’s Order of Santaliflorae:>/

1. Celastrales/

2. Vitidales/

3. Santalales/

#547. <Dahlgren’s Order of Balanophoriflorae:>/

1. Balanophorales/

#548. <Dahlgren’s Order of Araliiflorae:>/

1. Pittosporales/

2. Araliales/

#549. <Dahlgren’s Order of Asteriflorae:>/

1. Campanulales/

2. Asterales/

#550. <Dahlgren’s Order of Solaniflorae:>/

1. Solanales/

2. Boraginales/

#551. <Dahlgren’s Order of Corniflorae:>/

1. Fouquieriales/

2. Ericales/

3. Eucommiales/

4. Sarraceniales/

5. Cornales/

6. Dipsacales/

#552. <Dahlgren’s Order of Loasiflorae:>/

1. Loasales/

#553. <Dahlgren’s Order of Gentianiflorae:>/

1. Goodeniales/

2. Oleales/

3. Gentianales/

#554. <Dahlgren’s Order of Lamiiflorae:>/

1. Scrophulariales/

2. Hippuridales/

3. Hydrostachyales/

4. Lamiales/

#555. <Order of Alismatiflorae (Dahlgren, Clifford and Yeo 1985):>/

1. Hydrocharitales/

2. Alismatales/

3. Zosterales/

#556. <Order of Triuridiflorae (Dahlgren, Clifford and Yeo 1985):>/

1. Triuridales/

#557. <Order of Ariflorae (Dahlgren, Clifford and Yeo 1985):>/

1. Arales/

#558. <Order of Liliiflorae (Dahlgren, Clifford and Yeo 1985):>/

1. Dioscoreales/

2. Asparagales/

3. Liliales/

4. Burmanniales/

5. Orchidales/

6. Melanthiales/

#559. <Order of Zingiberiflorae (Dahlgren, Clifford and Yeo 1985):>/

1. Zingiberales/

#560. <Order of Commeliniflorae (Dahlgren, Clifford and Yeo 1985):>/

1. Commelinales/

2. Hydatellales/

3. Cyperales/

4. Poales/

#561. <Order of Areciflorae (Dahlgren, Clifford and Yeo 1985):>/

1. Arecales/

#562. <Order of Cyclanthiflorae (Dahlgren, Clifford and Yeo 1985):>/

1. Cyclanthales/

#563. <Order of Pandaniflorae (Dahlgren, Clifford and Yeo 1985):>/

1. Pandanales/

#564. <Order of Bromeliiflorae (Dahlgren, Clifford and Yeo 1985):>/

1. Velloziales/

2. Bromeliales/

3. Philydrales/

4. Haemodorales/

5. Pontederiales/

6. Typhales/

#### Cronquist's classification

#565. Cronquist’s Subclass <of Magnoliopsida (1981):>/

1. Magnoliidae/

2. Hamamelidae/

3. Caryophyllidae/

4. Dilleniidae/

5. Rosidae/

6. Asteridae/

#566. <Cronquist’s Order of Magnoliidae:>/

1. Magnoliales/

2. Laurales/

3. Piperales/

4. Aristolochiales/

5. Illiciales/

6. Nymphaeales/

7. Ranunculales/

8. Papaverales/

#567. <Cronquist’s Order of Hamamelidae:>/

1. Trochodendrales/

2. Hamamelidales/

3. Daphniphyllales/

4. Didymales/

5. Eucommiales/

6. Urticales/

7. Leitneriales/

8. Juglandales/

9. Myricales/

10. Fagales/

11. Casuarinales/

#568. <Cronquist’s Order of Caryophyllidae:>/

1. Caryophyllales/

2. Polygonales/

3. Plumbaginales/

#569. <Cronquist’s Order of Dilleniidae:>/

1. Dilleniales/

2. Theales/

3. Malvales/

4. Lecythidales/

5. Nepenthales/

6. Violales/

7. Salicales/

8. Capparales/

9. Batales/

10. Ericales/

11. Diapensiales/

12. Ebenales/

13. Primulales/

#570. <Cronquist’s Order of Rosidae:>/

1. Rosales/

2. Fabales/

3. Proteales/

4. Podostemales/

5. Haloragales/

6. Myrtales/

7. Rhizophorales/

8. Cornales/

9. Santalales/

10. Rafflesiales/

11. Celastrales/

12. Euphorbiales/

13. Rhamnales/

14. Linales/

15. Polygalales/

16. Sapindales/

17. Geraniales/

18. Apiales/

#571. <Cronquist’s Order of Asteridae:>/

1. Gentianales/

2. Solanales/

3. Lamiales/

4. Callitrichales/

5. Plantaginales/

6. Scrophulariales/

7. Campanulales/

8. Rubiales/

9. Dipsacales/

10. Calycerales/

11. Asterales/

#572. APG III <informal major grouping>/

1. core angiosperms/

2. peripheral angiosperms <no informal name>/

#573. APG III <informal eudicot grouping above Superorder>/

1. peripheral eudicot <no informal APG name>/

2. core eudicot/

#574. APG III <Superorder>/

1. Superorder Amborellanae/

2. Superorder Nymphaeanae/

3. Superorder Austrobaileyanae/

4. Superorder Magnolianae/

5. Superorder Lilianae <Monocots>/

6. Superorder Ceratophyllanae/

7. Superorder Buxanae/

8. Superorder Proteanae/

9. Superorder Ranunculanae/

10. Superorder Myrothamnanae/

11. Superorder Rosanae/

12. Superorder Berberidopsanae/

13. Superorder Caryophyllanae <Centrospermae>/

14. Superorder Santalanae/

15. Superorder Asteranae <= Tenuicelli (1970), cf. Gamopetalae (1876)!>/

16. unplaced at Superordinal level/

#575. APG III <informal Monocot grouping of Superorder Lilianae>/

1. commelinid Monocot/

2. non-commelinid Monocot <no informal APG name>/

#576. APG III <informal core eudicot grouping below superorder>/

1. fabid <eurosid I>/

2. malvid <eurosid II>/

3. lamiid <euasterid I>/

4. campanulid <euasterid II>/

#577. APG IV Order/

1. Acorales/

2. Amborellales/

3. Nymphaeales/

4. Austrobaileyales/

5. Chloranthales/

6. Canellales/

7. Piperales/

8. Magnoliales/

9. Laurales/

10. Alismatales/

11. Petrosaviales/

12. Petrosaviales/

13. Dioscoreales/

14. Pandanales/

15. Liliales/

16. Asparagales/

17. Arecales/

18. Commelinales/

19. Zingiberales/

20. Dasypogonales/

21. Poales/

22. Ceratophyllales/

23. Ranunculales/

24. Sabiales/

25. Proteales/

26. Trochodendrales/

27. Buxales/

28. Gunnerales/

29. Dilleniales/

30. Saxifragales/

31. Vitales/

32. Zygophyllales/

33. Fabales/

34. Rosales/

35. Fagales/

36. Cucurbitales/

37. Celastrales/

38. Oxalidales/

39. Malpighiales/

40. Geraniales/

41. Myrtales/

42. Crossosomatales/

43. Picramniales/

44. Sapindales/

45. Huertiales/

46. Malvales/

47. Brassicales/

48. Berberidopsidales/

49. Santalales/

50. Caryophyllales/

51. Cornales/

52. Ericales/

53. Icacinales/

54. Metteniusales/

55. Boraginales/

56. Vahliales/

57. Garryales/

58. Gentianales/

59. Solanales/

60. Lamiales/

61. Aquifoliales/

62. Asterales/

63. Escalloniales/

64. Bruniales/

65. Paracryphiales/

66. Dipsacales/

67. Apiales/

68. unassigned <to Order>/

#### Numbers of genera and species, examples

#578. <Number of extant> species/

#579. <Number of extant> genera/

#580. <Genera included (see Notes):>/

The generic lists were originally intended merely to exemplify family contents, with particular reference to the circumscriptions employed in compiling the descriptions. However, in terms of authoritatively accepted generic names in current use, i.e. excluding synonyms, they are now virtually complete, and the present Intkey package permits most Angiosperm generic names likely to be encountered by Intkey users to be referred to a taxonomically defensible family. In this connection, we extensively consulted Gunn et al. (1992 et seq.: Families and Genera of Spermatophytes recognized by the Agricultural Research Service), subsequently The Kew Plant List (http://www.theplantlist.org/), and more recently The World Flora Online (WFO) (http://www.worldfloraonline.org); but with numerous adjustments necessitated by our preference for sensu stricto family circumscriptions as explained in our Introduction. These mostly correspond with the treatments of Airy Shaw (1973), who also provided near-comprehensive cross referencing of family and generic synonyms. We list the genera of the traditional, hopelessly inadequate sensu lato ‘Liliaceae’ under the segregate families circumscribed by Dahlgren et al (1985). The latter represent a marked improvement, but family limits in this vicinity are evidently far from settled.

#### General remarks

#581. <Literature, comments:>/

#### Economic uses, etc.

#582. <Economic uses, etc. (see Notes):>/

There is much scope for extending the data, which are at present largely restricted to ‘general-interest’ remarks of the kind purveyed by Airy Shaw (1973) and Lawrence (1951). However, many supermarket ‘exotic fruits’ will be referable to families by searching for their English commercial names.

#### Quotations

#583. <Quotations:>/

From Shakespeare unless otherwise indicated. Many of the delightful and botanically astute quotations labelled ‘Ann Pratt 1857, unattributed’ no doubt derive from John Clare (1793–1864), especially from ‘The Shepherd’s Calender’, which has not yet been directly consulted in the present context.

Tropical and southern hemisphere families are poorly represented — contributions for these are especially welcome!

#### History of the encoded description

#584. <History of the encoded description (see Notes)>/

Descriptions were originally encoded and maintained by Watson between 1988 and 2008, and little of the subsequent, continuing editing of the data is recorded here.

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 8th June 2026. delta-intkey.com’.

Contents

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