Takhtajan Armen. Flowering Plants

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514 Subclass VIII. LAMIIDAE

on the hilar side, mostly minutely papil-

late to areolate, with small straight

embryo and ﬂ eshy endosperm. Alkaloids

present or absent, iridoids detected,

n = 10. . . . . . . . . 10. geniostomaceae.

5 Fruits drupes or berries. Trees, shrubs, or

woody lianas, often with tendrils and/or

axillary spines or spreading prickles, gla-

brous or with simple hairs. Vessels with

simple perforations and with vestured

pits. Fibers with bordered pits. Leaves

opposite or on the main axis sometimes

ternate, entire, pinnately veined or

3–5-veined from or above the base, estip-

ulate or with persistent or caducous

stipules. Flowers in terminal or/and axil-

lary cymose inﬂ orescences, rarely soli-

tary, bisexual, actinomorphic or nearly

so, with scalelike to leafy bracts, perianth

and androecium 4- or 5-merous. Sepals

free or up to the middle connate, imbri-

cate, persistent. Corolla rotated to salver-

shaped, more or less ﬂ eshy, lobes valvate.

Stamens 4–5, inserted on the corolla tube

or in the throat of the corolla tube; ﬁ la-

ments shorter or longer than the anthers;

anthers introrse. Pollen grains 3-colpo-

rate or rarely 4- or 2-colporate, or syn-

colpate, usually psilate with widely

distributed perforations. Ovary superior,

2-locular, with an axile placenta with few

to many ovules or, if 1-locular, with a

basal placenta; ovules anatropous; style

usually short, with capitate to 2-lobed

stigma. Fruits berries or (Neuburgia)

drupes. Seeds variously shaped, with thin

or thick testa; embryo minute to very

large; endosperm thick, horny, with a

large central cavity. Colleters in the axils

of the leaves and bracts and at the base of

the sepals or not. Producing loganin and

related iridoid glycosides and indole

alkaloids n = 11. . . . . 3. strychnaceae.

4 Mostly herbs, rarely shrublets, shrubs, or

small trees.

8 Corolla lobes valvate. Herbs, rarely sub-

shrubs, sometimes swith-plants. Vessels

with simple perforations, but in Mitreola

sometimes occur scalariform perfora-

tions with a few bars. Wood diffuse

porous, not storied. Axial parenchyma

absent. Secondary thickening anomalous

from a single cambial ring. Leaves well

developed or much reduced, opposite, or

opposite and verticillate, simple, entire,

petiolate to subsessile, with stipular lines

or small, interpetiolar stipules. Colleters

in the axils of leaves and bracts and at the

base of inner side of sepals sometimes

present. Flowers in terminal or axillary

cymes or more or less unilateral spikes,

rarely solitary or fasciculate or subumbel-

late, 5-merous. Calyx persistent, some-

times glandular within the base. Corolla

valvate (the lobes pointed, erect), funnel-

shaped or tubular. Stamens ﬁ ve, attached

to the corolla tube or in the throat of the

corolla tube; anthers dorsiﬁ xed, introrse.

Pollen grains 2(3)4-colpate (Spigelia) or

3-colporate. Gynoecium of two carpels;

style biﬁ d or deeply 2-lobed or 2-partite,

with truncate, clavate, or capitate stig-

mas; ovary superior, sessile, 2-locular,

with numerous ovules densely arranged

on peltate placentas; ovules anatropous,

or campylotropous, or hemianatropous.

Fruits capsules, septicidal or (Spigelia) cir-

cumscissile at the base. Seeds wingless

and not particularly compressed, some-

times polyhedral; embryo straight;

endosperm ﬂ eshy, ruminate. Producing

iridoid compounds asperuloside, genipo-

side, and geniposidic acid. n = 8 (Spigelia),

10 (Mitreola). . . . . . . . . .5. spigeliaceae.

8 Corolla lobes contorted or imbricate.

Perennial or annual herbs, often mycor-

rhizal and sometimes strongly

mycotrophic and without chlorophyll,

rarely shrubs or trees. Glabrous or with

various types of trichomes, sometimes

glandular. Vessels with simple or some-

times (Saccifolium) scalariform perfora-

tions. Rays when present predominantly

uniseriate. Axial parenchyma scanty

vasicentric or (Saccifolium) wanting.

Leaves opposite or sometimes verticil-

late, very rarely alternate, usually entire,

simple, saccate (Saccifolium), in

Superorder LAMIANAE 515

mycotrophic genera reduced to scales.

Stomata anomocytic or sometimes aniso-

cytic. Flowers in cymose or rarely race-

mose inﬂ orescences, sometimes solitary,

bisexual or rarely (Veratrilla) unisexual,

actinomorphic or rarely slightly zygomor-

phic, 5–4(rarely 6–12)- merous. Calyx

mostly synsepalous, usually with a short

to well-developed tube and imbricate or

sometimes valvate or open lobes, some-

times the lobes much reduced or sup-

pressed, rarely sepals nearly free. Corolla

often with scales or nectary pits within the

tube. Stamens attached to the corolla tube

or throat, free or rarely connate by anthers;

anthers basiﬁ xed, introrse or rarely

(Saccifolium), latrorse, opening longitudi-

nally or rarely (as in Exacum and

Cotylanthera) by apical pores that extend

halfway to the base. Pollen grains 2-celled

or 3-celled, (2)3(4)-colporate or

2–3-porate, sometimes monoporate (as

in Voyriella and spp. of Leiphamos).

Intrastaminal nectary disc or nectary

glands usually present. Gynoecium of two

carpels; style with an entire or 2-lobed,

papillate stigma; ovary 1-locular, with two

parietal placentas that usually pro ject

more or less deeply into the cavity and

sometimes connate in its center, forming a

false partition as in Exacinae; in

Cotylanthera as a result of suppression of

the false partition the parietal placentation

transformed into free central. Ovules more

or less numerous, anatropous or some-

times hemitropous or orthotropous, rarely

(in some parasitic species) ategmic. Fruits

septicidal capsules or rarely baccate.

Seeds numerous, small, winged or not,

with small embryo and copious endosperm,

very simpliﬁ ed in mycotrophic genera.

Usually producing bitter substances con-

sisting of iridoid compounds, especially

gentiopicroside and related substances,

n = 5–13. . . . . . . . . . . 9. gentianaceae.

2 Intraxylary (internal) phloem absent.

9 Flowers bisexual or rarely unisexual.

10 Stamens (8)16–18(-25) in two cycles, free.

Trees up to 30 m tall, with dense white-

tomentose young branches. The wood is

light but extremely hard. Vessels with sim-

ple perforations; lateral pitting alternate.

Fibers septate. Rays heterogeneous. Axial

parenchyma scanty paratracheal strands.

Leaves opposite, entire, simple, with large,

intrapetiolar and interpetiolar stipules that

are laterally connate basally in pairs.

Stomata paracytic. Flowers in terminal

thyrses or cymose panicles, bisexual, acti-

nomorphic, subtended by three bracteoles.

Calyx of four free sepals decussate in two

cycles of two, imbricate, persistent. Petals

four, ﬂ eshy, white, free, in two cycles, imbri-

cate. Stamens 16–25 (usually 16–17), united

at their base into a short androecial ring atop

the ovary, free from the corolla; ﬁ laments

short, basally connate into a tube; anthers

elongate, 2-locular (the locules possess two

horn-like appendages at apex), basiﬁ xed,

introrse, opening by apical pores. Pollen

grains 3-colporate, reticulate. Nectary disc

forming a ﬁ mbriate ring at apex of ovary.

Gynoecium of two fused carpels, they are

ﬂ attened, grooved and densely covered with

uniseriate, white hairs; style elongate, with

very shortly 2-lobed stigma; ovary inferior,

2- locular, with numerous, ascending, anat-

ropous (Piesschaert et al. 1997) ovules, on

axile placentas. Fruits septifragal capsules

crowned by persistent calyx and sparsely

covered with hairs, after maturation fruits

glabrous. Seeds exotestal, numerous, elon-

gate, ﬂ at, winged, with straight, large

embryo and scanty, oily endosperm. Iridoids

not detected. . . 6. dialypetalanthaceae.

10 Stamens as many as corolla lobes, epipe-

talous. Trees and shrubs, often lianous or

epiphytic, or subshrubs and perennial or

annual herbs, mostly terrestrial, rarely epi-

phytic or aquatic (Limnosipanea and spe-

cies of Spermacoce). Secretory cells or

cavities of various types often occur in

parenchymatous tissues. Calcium oxalate

crystals are found as raphides, styloids,

druses, and crystal sand, and can be stores

in various plant parts (Delprete 2004).

Vessels almost always with simple perfo-

rations; lateral pitting alternate, pits

516 Subclass VIII. LAMIIDAE

vestured. Rays heterogeneous with long

ends to homogeneous. Axial parenchyma

apotracheal, paratracheal, or wanting.

Nodes unilacunar or less often trilacunar,

or more. Leaves opposite or rarely ver-

ticillate (Henriquezia), entire or serrate,

very rarely alternate by suppression of one

of a pair at each node, or dentate, very

rarely sinuate-lobate or pinnatiﬁ d, petio-

late to sessile, gland-dotted, or not, some-

times containing small galls with symbiotic

bacterial colonies; stipules present, entire,

divided, or ﬁ mbriate, connate, or sheath-

ing at base, often caduceus and only repre-

sented by scars, usually interpetiolar or

less often intrapetiolar, rarely with four

stipules in Condaminea, or leaﬂ ike in

Galium; often bearing colleters inside that

produce mucilage and protect the mer-

istematic tissues of the growing buds from

herbivory (Delprete 2004). The mesophyll

containing calcium oxalate crystals.

Stomata usually paracytic. Inﬂ orescences

terminal or axillary, cymes, panicles,

umbels, racemes, spikes, heads inﬂ ores-

cences, or rarely uniﬂ orous. Flowers

bisexual or rarely unisexual (then plants

usually dioecious), mostly actinomorphic,

often heterostylous, in many groups (e.g.,

the Gardenieae and related tribes)

protandrous with specialized stylar pollen

presentation mechanism similar to one in

Campanulaceae, Goodeniaceae, and

Asteraceae, mostly with bracts and bracte-

oles. Calyx 5–4-lobed, lobes open and often

small or sometimes obsolete (as in

Neogaillonia) or sometimes one or more of

them enlarged and brightly colored (e.g., in

Mussaenda), frequently with colleters inside

the base of the calyx. Corolla sympetalous

(very rarely with petals free to base, as in

Mastixiodendron and Hedstromia), (3)4–

5(8–10)-lobed, rarely 8–11-, or 12–15-

lobed, with valvate, imbricate or contorted

lobes, actinomorphic to bilabiate. Stamens

4–5, attached to corolla tube or at its throat,

sometimes at its base; anthers sometimes

slightly appendaged, basiﬁ xed or dorsiﬁ xed,

introrse or extrorse, opening by longitudinal

slits or by two apical pores (Rustia), or by

one common lateral pore (in Tresanthera).

Pollen grains 2-celled or 3-celled, of various

types, mostly 3-colporate or 3(-4)-zonocol-

porate, in monads or in tetrads (Dessein

et al. 2005). Nectary disc often present at

the top of ovary. Gynoecium of two or sel-

dom 3–5 or even more carpels; style slen-

der, with lobed or capitate stigma, rarely

stylodia free to base (as in Galium,

Hedstromia, and Coprosma); ovary inferior

or seldom semi-inferior (as in

Mastixiodendron) rarely almost completely

superior (Gaertnera and Pagamea), with as

many locules as carpels, rarely 1-locular

with parietal placentas (as in Gardenia,

etc.), each locule with one to many ovules

(commonly one ovule in Rubioideae).

Ovules pendulous, horizontal or ascending,

anatropous to hemitropous, very rarely cir-

cinotropous or hypertropous, often with

funicular obturator, nearly always without

endothelium. Endosperm nuclear or very

rarely cellular. Fruits of diverse types, but

mostly capsules (septicidal or loculicidal),

ﬂ eshy or leathery berries, or drupes. Seeds

wingless or rarely winged, exotesta alone

persisting, papillate hairy or not, mesotestal

cells thickened; embryo usually straight

to curved, embedded in usually copious,

oily, ruminate endosperm, or sometimes

endosperm scanty or wanting, in certain

groups without seed coat. Producing iri-

doids, various types of alkaloids, ﬂ avonols,

kaempferol and quercetin, ursolic acid, inu-

lin (Cinchona), n = 6–17 (mostly 11, less

often 9).. . . . . . . . . . . . . . . . . 7. rubiaceae.

9 Flowers unisexual. Perennial or annual herbs,

sometimes succulent with raphide sacs. Xylem

forming a continuous cylinder. Vessels very

numerous, small, with simple perforations.

Mechanical tissues lacking. Leaves opposite

below, alternate above by suppression of one

of each pair, simple, entire, ﬂ eshy, with large,

club-shaped glands at the apex; stipules inter-

petiolar, united, membranous, bearing col-

leters near the tip. Stomata paracytic. The

mesophyll containing calcium oxalate crystals

(raphides). Flowers monoecious, anemophil-

Superorder LAMIANAE 517

ous, and evidently apetalous. Male ﬂ owers

solitary or paired opposite the leaves of upper

nodes, female ﬂ owers mostly in simple axillary

dichasia at lower nodes. Calyx of male ﬂ owers

inconspicuous, closed in bud but soon valvately

splitting to the base into 2–5 broad, recurved

lobes. Calyx of female ﬂ owers membranous,

very oblique, tubular, shortly 2–4-dentate at the

apex. Corolla of female ﬂ owers 2–4-toothed,

valvate, unequal, but not bilabiate. Stamens

(2)7–12(-30), sometimes basally united into

groups of two, four, or six; ﬁ laments slender;

anthers erect in bud, but pendulous later, ver-

satile, 2-locular. Pollen grains 3-celled,

(3)4–6(7)-zonoporate. Gynoecium pseudomono-

merous; style gynobasic, slender, exerted from

mouth of calyx; ovary superior, 1-locular, with

one basal and more or less campylotropous or

amphitropous, unitegmic (the integument mas-

sive) ovule. Fruits subglobose, nutlike drupes,

pericarp with elaiosome. Seeds hippocrepiform,

with strongly curved, large embryo in copious,

oily end osperm. Producing iridoid compounds,

n = 11. . . . . . . . . . . . . . . . . 8. theligonaceae.

1 Plant with a well-developed latex-system (except in

Nerium), and commonly producing cardiotonic

glycosides. steroidal alkaloids, cardenolides, group

II decarboxylated iridoids present. Trees, shrubs,

herbs, very often lianas, sometimes succulents, gla-

brous or with various kinds of trichomes.). Stem

often of anomalous structure. Vessels with scalari-

form or simple perforations; Leaves opposite or

sometimes verticillate, rarely alternate, simple,

entire or rarely lobed or dentate; stipules wanting or

rarely small and interpetiolar or vestigial. Flowers

in cymose or racemose inﬂ orescences, or solitary,

bisexual or very rarely functionally unisexual, acti-

nomorphic or nearly so, commonly 5-merous.

Sepals (4)5, more or less connate, imbricate or val-

vate. Corolla sympetalous, lobes contorted or rarely

imbricate or valvate. Stamens as many as and alter-

nate with the corolla lobes, inserted on the corolla

tube, free or more often connate; ﬁ laments short;

anthers free or more or less closely coherent or con-

nate around the style head, basiﬁ xed or dorsiﬁ xed,

tetra sporangiate, bisporangiate or disporangiate,

introrse, opening longitudinally or apically. Pollen

grains 3-celled, in monads, tetrads or pollinia,

3-colporate or porate. Nectar secreted in alternis-

taminal troughs on staminal tube or staminal feet or

from disclike nectary around base of ovary, more

rarely from sides of ovary or absent. Gynoecium of

2(-8) carpels, these connate in varying degrees;

ovary superior to semiinferior, stigma mostly on

underside of style head, often restricted to ﬁ ve

chambers behind guide rails. Ovules two to many,

commonly pendulous, anatropous, amphitropous,

or hemitropous, very rarely orthotropous, with or

without endothelium. Fruits of diverse types. Seeds

small, exotestal, often with a terminal coma of long

hairs, naked, winged or arillate, with straight

embryo and oily, copious to scanty endosperm. n =

8–12. . . . . . . . . . . . . . . . . . . . . . 11. apocynaceae.

1. GELSEMIACEAE

L. Struwe et V.A. Albert 1995. 2/11. Tropical Africa,

Madagascar, Southeast Asia, southern North America,

and Central and northern South America.

Gelsemium, Mostuea

Probably the most archaic member of the order.

2. LOGANIACEAE

R. Brown ex C. Martius 1827. 1/15. New Caledonia

(1), New Zealand (1), and Australia (13).

Logania

3. STRYCHNACEAE

Perleb 1818 (including Gardneriaceae Perleb 1838).

4/250. Eastern and Southeast Asia, East Malesia,

Melanesia, Fiji.

Strychnos, Scyphostrychnos, Neuburgia, Gardneria

4. ANTONIACEAE

Hutchinson 1959. 4/8. West Malesia (Norrisia), tropi-

cal South America, and West and tropical Africa

(Usteria).

Norrisia, Bonyunia, Antonia, Usteria

Connected with the Loganiaceae and Gelsemiaceae,

but lacking both iridoids and alkaloids (Bisset 1975;

Jensen 1992).

518 Subclass VIII. LAMIIDAE

5. SPIGELIACEAE

C. Martius 1827. 3/96. Madagascar, southern, eastern,

and Southeast Asia, New Guinea, Australia, Tasmania,

New Zealand, New Caledonia, Caroline Islands, North

and South America.

Spigelia, Mitreola, Mitrasacme

Spigeliaceae are a rather heterogeneous and per-

haps not a natural family. Pollen grains of the

Spigeliaceae are of different types (Punt 1980), and

this together with diverse types of fruits and different

basic numbers of chromosomes does not indicate any

great coherence within the group (Bisset et al. 1980).

There are afﬁ nities with Loganiaceae but even more

with Rubiaceae, especially with the Hedyotideae in

habit, stipules, ﬂ owers, and wood anatomy (Bisset

et al. 1980). According to Mennega (1980: 119),

“ striking is the similarity in wood structure between

Spigelieae and Hedyotis” Here I am accepting the

family Spigeliaceae only tentatively.

6. DIALYPETALANTHACEAE

Rizzini et Occhioni 1948. 1/1. Amazonian Brazil and

Peru.

Dialypetalanthus

Very closed to the Rubiaceae (Dahlgren and Thorne

1984 [“Dialypetalanthus is probably an aberrant early

off-shoot of the Rubiaceae or a relict family closely

related to the Rubiaceae”, p. 690]). It is important to

mention that Dialypetalanthus has the same type of

biﬁ d stipules and many-seeded capsules as in the

Rondeletieae complex (Delprete 2004), but differs in

dimerous origin of calyx and corolla, free corolla

lobes, and numerous stamens.

7. RUBIACEAE

A.L. de Jussieu 1789 (including Aparinaceae

Hoffmannsegg et Link 1813–1829, Asperulaceae

Chamisso ex Spenner 1835, Catesbaeaceae Martynov

1820, Cephalanthaceae Raﬁ nesque 1820, Cinchonaceae

Batsch 1802, Coffeaceae Batsch 1802, Coutareaceae

Martynov 1820, Cynocrambaceae Endlicher 1841,

Galiaceae Lindley 1836, Gardeniaceae Dumortier

1829, Guettardaceae Batsch 1802, Hedyotidaceae

Dumortier 1829, Henriqueziaceae Bremekamp 1957,

Houstoniaceae Raﬁ nesque 1840, Hydrophylacaceae

Martynov 1820, Lippayaceae Meisner 1836–1843,

Lygodysodeaceae Bartling 1830, Naucleaceae

Wernham 1912, Nonateliaceae Martynov 1820,

Operculariaceae A.L. de Jussieu ex Perleb 1818,

Pagamaeaceae Martynov 1820, Psychotriaceae

F. Rudolphi 1830, Randicaceae Martynov 1820,

Sabiceaceae Martynov 1820). 650/13000. Cosmo-

politan, but mainly in tropical and subtropical regions

with some genera in temperate and cold regions (even

in the Arctic and Antarctic).

7.1 RUBIOIDEAE

Small trees, shrubs, lianas, epiphytes, or more often

herbs; hairs articulated. Raphides generally present.

Stipules entire, frequently biﬁ d or ﬁ mbriate

(Spermacoceea). Corolla mostly valvate, rarely imbri-

cate or contorted. Stamens inserted at middle or near

corolla mouth. Heterostyly very common. Placentas

pluriovulate or with a single (rarely two) erect ovules.

Fruits dry (capsules, dehiscent into mericarps, opening

with operculum, or indehiscent) or ﬂ eshy. Exotestal

cells often parenchymalike. – ophiorrhizeae:

Colletoecema, Ophiorrhiza, Neurocalyx, Xantho phy-

tum, Lerchea, Coptophyllum, Spiradiclis; pauridi-

antheae: Pauridiantha, Poecilocalyx, Rhipidantha,

Stelechantha; urophylleae: Urophyllum, Amphi-

dasya, Commitheca, Maschalocorymus, Pravinaria,

Praravinia; lasiantheae: Lasianthus, Saldinia,

Perama, Trichostachys, Metabolus; cruckshanks-

ieae: Cruckshanksia; coussareeae: Coussarea,

Faramea, Oreopolus, Heterophyllaea; cocco-

cypselea: Coccocypselum, Declieuxia, Hindsia; cra-

terispermeae: Craterispermum; psycho trieae:

Psychotria, Amaracarpus, Cephaelis, Chasallia,

Chazaliella, Geophila, Hydnophytum, Hymenocoleus,

Myrmecodia, Palicourea, Rudgea, Uragoga,

Anthorrhiza, Myrmephytum, Squamellaria, Readea,

Streblosa, Margaretopsis; morindeae: Morinda,

Appunia, Coelospermum, Damnacanthus, Gentingia,

Gynochthodes, Pogonolobus, Prismatometis, Renellia,

Motleyia, Mitchella,; schradereae: Schradera,

Lecananthus, Leucocodon; gaert nereae: Gaertnera,

Pagamea; danaideae:

Danais, Schismatoclada,

Payera; hedyotideae: Pentas, Batopedina, Carphalea,

Otiophora, Chaemepentas, Parapentas, Otomeria,

Knoxia, Calanda, Pentanisia, Neo pentanisia,

Chlorochorion, Pseudohedyotis, Tri ainolepis,

Agathisanthemum, Amphiasma, Arcyto phyllum,

Superorder LAMIANAE 519

Conostomium, Pentodon, Dentella, Scleromitrion,

Phylohydrax, Dibrachionostylus, Exallage,

Gomphocalyx, Gouldia, Hedyotis, Hedythyrsus,

Houstonia, Kohautia, Lelya, Manettia, Manostachya,

Mitrasacmopsis, Neohymenopogon, Neanotis, Olden-

landia; spermacoceae: Spermacoce, Bouvardia,

Diodella, Diodia, Emmeorhiza, Ernodea, Scandentia,

Galianthe, Hemidiodia, Mitracarpus, Paratriaina,

Pentanopsis, Placopoda, Psyl locarpus, Tobagoa,

Richardia, Crusea, Dolichometra, Hydrophylax,

Lucya, Nodocapaea, Schwendera, Synaptantha,

Staelia, Thecorchis, Thyridocalyx; antho spermeae:

Anthospermum, Co prosma, Carpacoce, Opercularia,

Galopina, Nenax, Nertera (including Peratanthe),

Phyllis, Durringtonia, Eleuthera nthus, Leptostigma,

Normandia, Pomax; argostemmateae: Argostemma,

Cyaneuron, Mouretia, Mycetia; paederieae: Paederia,

Gaillo nia, Saprosma, Neogaillonia, Plocama,

Aitchisonia, Putoria, Serissa, Spermadictyon,

Choulettia, Jaubertia, Kelloggia, Leptodermis,

Pseudogaillonia, Pseudopyxis, Pterogaillonia;

rubieae: Rubia, Asperula, Crucia nella, Cruciata,

Galium, Phuopsis, Sherardia, Valantia, Callipeltis,

Relbunium, Didymaea, Mericarpaea, Microphysa,

Warburgia.

7.2 IXOROIDEAE

Shrubs or less often trees or woody lianas. Raphides

nearly always absent. Stipules mostly entire, rarely

biﬁ d. Corolla contorted. Stylar pollen presentation

almost general. Stamens inserted at middle of tube or

near corolla mouth. Placentas mostly pluriovulate.

Fruits ﬂ eshy. Exotestal cells rarely parenchymalike,

generally with thickenings along the walls. – con-

damineae: Alseis, Bathysa, Calycophyllum, Capirona,

Chimarrhis, Condaminea, Dolichodelphys, Elaeagia,

Emmenopterys, Ferdinandusa, Hippotis,

Macrocnemum, Mastixiodendron, Parachimarrhis,

Pentagonia, Picardaea, Pinckneya, Pogonopus, Rustia,

Semaphyllanthe, Simira, Sommera, Tammsia, Tresan-

thera, Witt mackanthus; mussaendeae: Mussaenda,

Neo mussaenda, Pseudomussaenda, Schizomussaenda,

Aphaenandra, Heinsia, Landiopsis; sabiceeae:

Sabicea, Ecpoma, Pseudosabicea, Stipularia; virec-

tarieae: Virectaria, Hekisto carpa; sipa neeae:

Sipanea, Sipaneopsis, Limnosipanea; henriquezieae:

Gleasonia, Henriquezia, Platy carpum, ? Monopanthera,

Posoqueria; retini phylleae: Retiniphyllum; ixoreae:

Ixora, Captaincookia, Doricera, Myonima, Versteegia,

? Scyphiphora; vanguerieae: Canthium, Cuviera,

Fadogia, Fadogiella, Hutchinsonia, Keetia, Meyna,

Multidentia, Peponidium, Perakanthus, Psydrax,

Pygmaeothamnus, Pyrostria, Rytigynia, Scypho-

chlamya, Vangueria, Vangueriella, Vangueriopsis;

alberteae: Alberta, Nematostylis; coffeeae: Coffea,

Psilanthus (including Paracoffea), Diplospora,

Discospermum, Tricalysia, Bertiera, Sericanthe;

octotropideae: Jovetia, Lamprothamnus, Octotropis,

Paragenipa, Polysphaeria, Ramosmania; cre-

masporeae: Cremaspora; pavetteae: Pavetta,

Rutidea, Tarrena, Robbrechtia, Coptosperma,

Leptactina, Paracephaelis, Schizenterospermum,

Dictyandra, Cladoceras, Nichallea, Pachystylus,

Tennantia, Homollea, Homolliella, Coleactina. gar-

denieae: Aidia, Aidiopsis, Alibertia, Alleizettella,

Amaioua, Atractocarpus, Atractogyne, Aulacocalyx,

Benkara, Borojoa, Brachytome, Brenania, Burchellia,

Byrsophyllum, Calochone, Casasia, Catun aregam,

Ceriscoides, Coddia, Deccania, Didymo salpinx,

Dioecrescis, Duperrea, Duroia, Euclinia, Fagerlindia,

Ganguelia, Gardenia, Genipa, Adeno randia,

Glossostipula, Heinsenia, Himalrandia, Hyper-

acanthus, Kailarsenia, Kochummenia, Kutchubaea,

Larsenaikia, Macrosphyra, Mantalania, Massularia,

Melanopsidium, Mitriostigma, Monosalpinx, Morelia,

Oligocodon, Oxyanthus, Oxyceros, Pelagodendron,

Phellocalyx, Pleiocoryne, Porterandia, Preussiodora,

Pseudomantalania, Randia, Riodocea, Rosenber-

giodendron, Rothmannia, Rubovietnamia, Schumannio-

phyton, Sherbournia, Sphinctanthus, Stachyarrhena,

Sukunia, Sulitia, Tamilnadia, Tarennoidea, Tocoyena,

Trukia, Vidalasia, ? Argocoffeopsis.

7.3 CINCHONOIDEAE

Trees, shrubs, woody lianas, or sometimes epiphytic

shrubs, very rarely herbaceous. Raphides mostly

absent (present in Hillia and Hamelia). Stipules

mostly entire, rarely biﬁ d. Corolla imbricate or valvate,

rarely contorted. Stamens inserted at base of tube, or

near corolla mouth. Heterostyly frequent. Stylar pollen

presentation rare. Placentas with numerous ovules.

Fruits mostly capsular, sometimes ﬂ eshy, or drupa-

ceous. Seeds numerous, often winged. Exotestal cells

always with thickenings along tangential walls. –

cinchoneae: Cinchona, Dolicholobium, Joosia,

Ladenbergia, Remijia, Stilpnophyllum; isertieae:

Isertia, Kerianthera, Raritebe, Schizostigma,

Temnopteryx; catesbaeae: Asemnantha, Badusa,

520 Subclass VIII. LAMIIDAE

Bikkia, Catesbaea, Ceratopyxis, Chiococca, Couta-

portla, Coutarea, Erithalis, Exostema, Hintonia,

Isidorea, Morierina, Nernstia, Osa, Portlandia,

Salzmannia, Schmidtottia, Scolosanthus, Syringantha;

hymenodictyeae: Hymenodictyon, Paraco rynanthe;

naucleeae: Adina, Adinauclea, Breo nadia, Breonia,

Burttdavya, Cephalanthus, Corynanthe, Cubanola,

Gyrostipula, Haldina, Hallea, Janotia, Ludekia,

Metadina, Mitragyna, Myrmeconauclea, Nauclea,

Neolamarckia, Neonauclea, Ochreinauclea,

Pausinystalia, Pertusadina, Sarcocephalus, Sinoadina,

Uncaria; hamelieae: Hillia, Hamelia, Cosmibuena,

Hoffmannia, ? Balmea, Chione; rondeletieae:

Rondeletia, Blepharidium, Roigella, Suberanthus, ?

Acunaeanthus, Glionnetia, Habroneuron, Rogiera (pro

parte), Stevensia; guettardeae: Arachnothryx,

Chomelia, Gonzalagunia, Guettarda, Javorkaea,

Timonius, ? Dichilanthe.

Uncertain Position: Acranthera, Airosperma,

Aleisanthia, Aleisanthiopsis, Aoranthe, Aphanocarpus,

Ariadne, Augusta, Calycosiphonia, Coryphothamnus,

Crossopteryx, Dunnia, Greeniopsis, Jackiopsis,

Kajewskiella, Lathraeocarpa, Leucolophus, Lindenia,

Mitchella, Mussaendopsis, Myrioneuron, Pagameopsis,

Petitio coden, Placocarpa, Sarcosperma, Stephanococcus,

Trailliaedoxa, Wendlandia.

Very closely related to the Loganiaceae and even

more to the Spigeliaceae. It is also interesting that

Gaertnera and Pagamea, which are characterized by

superior ovary, are sometimes transferred to the

Loganiaceae. All the available evidence leads to the

conclusion that the Rubiaceae share a common origin

with the lower Rubiales from the hydrangealean

ancestor.

8. THELIGONACEAE

Dumortier 1829. 1/3. Canary Islands, Mediterranean,

Crimea, western Asia, southwestern China, Japan.

Theligonum

Very closely related to the Rubiaceae, especially to

the Rubioideae (see Wunderlich 1971; Kooiman 1971;

Cronquist 1981). Robbrecht (1993) and Thorne (1992b,

2006) even include Theligonum in Rubiaceae-

Rubioideae. However, Theligonum is remarkable in

having a large number of stamens arranged in bundles

and pseudomonomerous gynoecium and 1-locular

ovary without a trace of second locule (a unique fea-

ture of Theligonum according to Robbrecht [1993: 90])

as well as highly specialized and reduced anemophil-

ous ﬂ owers, the absence of mechanical tissues and

campylotropous ovules).

9. GENTIANACEAE

A.L. de Jussieu 1789 (including Chironiaceae

Horaninow 1847, Coutoubeaceae Martynov 1820,

Obolariaceae Martynov 1829, Potaliaceae C. Martius

1827, Saccifoliaceae Maguire and Pires 1978).

88/1600–1690. Cosmopolitan, but mainly in tempe-

rate and subtropical regions and also in tropical

mountains.

saccifolieae: Curtia, Hockinia, Saccifolium,

Tapeinostemon, Voyriella; exaceae: Cotylanthera,

Exacum, Gentianothamnus, Ornichia, Sebaea,

Tachiadenus; chironieae: Bisgoeppertia, Black-

stonia, Centaurium, Chironia, Cicendia, Eustoma,

Exaculum,Geniostemon, Ixanthus, Orphium, Sabatia,

Zygostigma; Canscora, Cracosna, Hoppea, Micro-

phium, Phyllocyclus, Schinziella; Coutoubea,

Deianira, Schultesia, Symphyllophyton, Xestaea;

helieae: Adeno lisianthus, Aripuana, Calolisianthus,

Celiantha, Chelo nanthus, Chorisepalum, Helia,

Irlbachia, Lagenanthus, Lehmanniella, Macrocarpaea,

Neblin antha, Prepusa, Purdieanthus, Rogersonanthus,

Senaea, Sipapoantha, Symbolanthus, Tachia,

Tetrapollinia, Wurdackanthus, Zonanthus; potalieae:

Congolanthus, Djaloniella, Enicostema, Faroa,

Karina, Neurotheca, Oreonesion, Pycnosphaera,

Urogentias, Lisianthus, Anthocleista, Fagraea, Potalia,

gentianeae: Crawfurdia, Gentiana, Tripterospermum,

Anagallidium, Bartonia, Comastoma, Frasera,

Gentianella, Gentianopsis, Halenia, Jaeschkea,

Lotouchea, Lomatogonium, Megacodon, Obolaria,

Pterygocalyx, Swertia, Veratrilla; voyrieae: Voyria.

Gentianaceae are closely connected with the

Gelsemiaceae and evidently derived from the gelsemi-

aceous stock.

10. GENIOSTOMACEAE

L. Struwe et V.A. Albert 1995. 2/75. Mascarene

Islands, Malesia (except for the Malay Peninsula),

Kyushu, Bonin Islands, Micronesia (Mariana Islands

and Caroline Islands), New Guinea, Solomon Islands,

Superorder LAMIANAE 521

Australia (eastern Queensland), New Caledonia, New

Hebrides, Fiji, Lord Howe Island, northern New

Zealand, Polynesia (including Hawaii).

Geniostoma, Labordia

A rather isolated family that is probably related to

the Gentianaceae.

11. APOCYNACEAE

A.L. de Jussieu 1789 (including Asclepiadaceae

Borkhausen 1797, Cerberaceae Martynov 1820,

Cryptostegiaceae Schlechter 1905, Cynanchaceae

G. Meyer 1836, Ophioxylaceae Perleb 1838, Pacouri-

aceae Martynov 1820, Periplocaceae Schlechter 1905,

Plumeriaceae Horaninow 1834, Stapeliaceae Horani-

now 1834, Vincaceae Vest 1818, Willughbeiaceae

J.G. Agardh 1858). 432/5100. Tropical and subtropical

regions with relatively few members in temperate areas.

Classiﬁ cation after M.E. Endress and P.V. Bruyns

(2000) and M.E. Endress and W.D. Stevens (2001).

11.1 RAUVOLFIOIDEAE

Anthers not adherent to style head by a reticulum of

viscid exudates, full of pollen, rarely with spines.

Fruits dehiscent or indehiscent, berries, drupes, folli-

cles, or capsules. Seeds naked, with wings or arils, but

almost never with coma at one end; holmindole

alkaloids present or absent, n = 9 (Alyxieae), 10, 11,

(23). – alstonieae: Alstonia, Aspidosperma,

Geissospermum, Haplophyton, Laxoplumeria,

Microplumeria, Strempeliopsis, Tonduzia, Vallesia;

vinceae: Amsonia, Catharanthus, Kopsia,

Neisosperma, Ochrosia, Petchia, Rauvolﬁ a, Vinca;

wullughbeieae: Ancylobotrys, Bousigonia,

Chamaeclitandra, Clitandra, Couma, Cyclocotyla,

Cylindropsis, Dictyophleba, Hancornia, Lacmellea,

Landolphia, Willughbeia, Leuconotis, Orthopichonia,

Paucouria, Parahancornia, Saba, Vahadenia; taber-

naemontaneae: Ambelania, Bonafousia, Callichilia,

Calocrater, Carvalhoa, Crioceras, Macoubea, Mucoa,

Molongum, Neocouma, Rhigospira, Schizozygia,

Spongiosperma, Stenosolen, Tabernaemontana,

Stemmadenia, Tabernanthe, Voacanga, Woytkowskia;

melodineae: Craspidospermum, Diplorhynchus,

Dyera, Gonioma, Kamettia, Melodinus, Pycnobotrya,

Stephanostegia; hunterieae: Hunteria, Picralima,

Pleiocarpa; plumerieae: Allamanda, Anechites,

Cerbera, Cerberiopsis, Cameraria, Thevetia,

Himatanthus, Mortoniella, Plumeria, Skytanthus;

Carisseae: Carissa, Acokanthera; alyxieae:

Chilocarpus, Alyxia, Pteralyxia, Condylocarpon,

Lepinia, Lepiniopsis, Plectaneia.

11.2 APOCYNOIDEAE

Corolla left-contorted, valvate. Anthers tetrasporan-

giate, adherent to style head by a retinaculum of

viscoid exudates, empty at base and with spines.

Nectaries, if present, in ring around base of ovary.

Seeds ecomose or comose, n = (6-) 10, 11 (12). –

wrightieae: Nerium, Isonema, Pleioceras,

Stephanostema, Strophanthus, Wrightia, Adenium;

malouetieae: Mascarenhasia, Funtumia Kibatalia,

Holarrhena, Malouetia, Malouetiella, Alaﬁ a,

Allowoodsonia, Carruthersia, Farquharia,

Pachypodium, Spirolobium; apocyneae: Beaumontia,

Aganonerion, Anodendron, Urceola, Ecdysanthera,

Odontadenia, Baissea, Apocynum, Trachomitum,

Aganosma, Baharuia, Chonemorpha, Epigynum,

Forsteronia, Ichnocarpus, Vallaris, Vallariopsis,

Trachelospermum, Cleghornia, Dewevrella, Elytropus,

Epigynum, Eucorymbia, Ixodonerium, Montandra,

Oncinotis, Papuechites, Parameria, Parepigynum,

Sindechites, Baissea; mesechiteae: Allomark graﬁ a,

Galactophora, Mandevilla, Secondatia, Macro-

siphonia, Mesechites, Tintinnabularia, Quiotania,

Telosiphonia; echiteae: Prestonia, Echites,

Laubertia, Cycladenia, Parsonsia, Pottsia, Pentalinon,

Thenardia, Angadenia, Temnadenia, Rhabdadenia,

Stipecoma, Amalocalyx, Neobracea, Macropharynx,

Salpinctes, Asketanthera, Fernaldia, Peltastes, Artia,

Ecua, Hylaea.

11.3 PERIPLOCOIDEAE

Corolla valvate, tube formation intermediate. Filaments

free from each other. Anthers tetrasporangiate. Two

pollinia (consisting of loosely coherent tetrads) in each

locule, transferred by spoon-shaped translator ending

in a sticky disc, n = 11 (mostly). – Cryptostegia,

Telectadium, Pentopetia, Stomatostemma, Ectadium,

Maclaudia, Omphalogonus, Cryptolepis (including

Mangenotia), Schlechterella, Gymnanthera, Strepto-

caulon, Hemidesmus, Decalepis (including Utleria),

Epistemma, Finlaysonia (including Atherolepis,

Stelmacrypton, Gongylosperma, Hanghomia, Mela-

derma), Sarcorhiza, Phyllanthera (including

Streptomanes, Pentanura), Baroniella, Sacleuxia,

522 Subclass VIII. LAMIIDAE

Batesanthus, Baseonema, Rhaphionacme, Campto-

carpus, Mondia, Myriopteron, Periploca, Tacazze

(including Zacateza), Petopentia, Atherandra,

Ischnolepis (Peptopentia), Buckollia, Zygostelma.

11.4 SECAMONOIDEAE

Corolla left-contorted. Filaments connate. Anthers

tetrasporangiate. Two very small pollinia in apical part

of each locule, transferred by translator with arm that

has two clefts. – Secamone, Secamonopsis, Toxocarpus,

Goniostemma, Pervillea, Trichosandra, Rhynchostigma,

Calyptranthera, Genianthus.

11.5 ASCLEPIADOIDEAE

Filaments connate. Anthers bisporangiate, with one

solid pollinium in each locule (pollen in tetrads –

Fockeae), transferred by translator, orbicules present

or (Riocreuxia) absent, granular layer of exine thin;

n = 9–14. – fockeeae: Fockea: marsdenieae:

Gymnema, Dischidia, Hoya, Marsdenia, Sarcolobus,

Cosmostigma, Asterostemma, Rhyssolobium, Gongro-

nema, Cathetostemma, Pseusmagennetus, Pycnorha-

chis, Lygisma, Treutlera, Absolmsia, Anisopus,

Telosma, Anatropanthus, Micholitzia, Campestigma,

Spirella, Stigmatorhynchus, Clemensiella, Oreosparte,

Heynella, Cibirhiza, Gunnessia, Madangia; CERO-

PEGIEAE: Leptadenia, Brachystelma, Ceropegia,

Macropetalum, Hoodia, Piaranthus, Caralluma,

Stapella, Huernia, Duvalia, Orbea, Anisotoma,

Conomitra, Orthanthera, Pectinaria, Pentasachme,

Riocreuxia, Sisyranthus, Heterostemma, Desmidorchis,

Frerea, Tridentea, Tromotriche, Tenaris, Tavaresia,

Quaqua, Echidnopsis, Edithcolea, Emplectanthus,

Dittoceras, Neoschumannia, Stapeli opsis, Stape-

lianthus, Whitesloanea, Rhytidocaulon, Pseudolithos,

Duvaliandra, Lavrania, Ophionella, Notechidnopsis;

asclepiadeae: Tylophora, Ade lostemma, Matelea,

Oxystelma, Sarcostemma, Cynanchum, Vincetoxicum,

Vincetoxicopsis, Calotropis, Lachnostoma, Macro-

scepis, Metastelma, Microloma, Gonolobus, Ditassa,

Macroditassa, Pergularia, Schubertia, Funastrum,

Asclepias, Gomphocarpus, Kanahia, Morrenia, Penta-

stelma, Merrillanthus, Biondia, Pentatropis, Seutera,

Ampelamus, Calostigma, Cyathella, Cordylogyne,

Glossonema, Pachyglossum, Schizostemma, Tweedia,

Urostelma, Odontanthera, Pentarrhinum, Pachycarpus,

Metaplexis, Pherotrichis, Solenostemma, Tassadia,

Turrigera, Nematostemma, Raphistemma, Philibertia,

Schizoglossum, Schisto gyne, Trichosacme, Aspido-

glossum, Holostemma, Graphistemma, Xysmalobium,

Oxypetalum, Diplolepis, Oncinema, Stenomeria,

Eustegia, Astephanus, Acrocoryne, Blyttia, Dictyanthus,

Diploglossum, Fischeria, Araujia, Parapodium,

Nautonia, Pycnostelma, Rhyssostelma, Orthosia, Mito-

stigma, Peplonia, Melinia, Hemipogon, Ibatia,

Barjonia, Blepharodon, Polystemma, Nematuris, Rhyn-

charrhena, Nephradenia, Amblistigma, Sphaerocodon,

Fanninia, Lugonia, Margaretta, Metalepis, Calatho-

stelma, Lagoa, Glossostelma, Husnotia, Telminostelma,

Cyathostelma, Lorostelma, Stelmation, Sattadia,

Bustelma, Kerbera, Lagenia, Anomotassa, Dactylo-

stelma, Hypolobus, Jobinia, Odontostelma, Petalo-

stelma, Diplostigma, Grisebachiella, Pentabothra,

Podandra, Pleurostelma, Stelmagonum, Platykeleba,

Scyphostelma, Steleostemma, Stenostelma, Prosthe-

cidiscus, Stathmostelma, Woodia, Vailia, Widgrenia,

Tetraphysa, Trachycalymma, Meresaldia, Karimbolea,

Schistonema, Stelmatocodon, Labidostelma, Mahawoa,

Rajasia, Stuckertia, Rojasia, Miraglossum, Seshagiria,

Emicocarpus, Folotsia, Gonioanthela, Amblyopetalum,

Dicarpophora, Corollonema, Hickenia, Goydera,

Aidomene, Aspidonepsis, Stephanotis.

Here I follow Hallier (1905, 1912), Stebbins (1974),

Thorne (1983, 1992a, b, 2000, 2006), Endress and

Bruyns (2000), and Endress and Stevins (2001) in

combining Apocynaceae and Asclepiadaceae into one

family. They are connected by a series of transitional

forms, and as Cronquist (1981: 861) states, “There is a

fairly straight-line evolutionary series in ﬂ oral mor-

phology within the Apocynaceae and Asclepiadaceae

collectively, from the Plumerioideae to the

Apocynoideae to the Periplocoideae to the Secamoneae

and thence to the other tribes of the Asclepiadoideae.”

Closely related to the Strychnaceae, Gentianaceae

and Geniostomaceae clade (see Struwe et al. 1995).

Some genera, such as Tabernaemontana, are quite

similar in their wood anatomy to the genus Neuburgia

(Mennega 1980). It is interesting that the fruit mor-

phology of Neuburgia also resembles that of some

apocynaceous genera, such as Cerbera, Ochrosia, and

Rauvolﬁ a (Bisset et al. 1980). Formerly Neuburgia

was even included in Apocynaceae.

Bibliography

Achille F, TJ Motley, PP Lowry II, and J Jérémie. 2006.

Polyphyly in Guettarda L. (Rubiaceae, Guettardeae) based

on nrDNA ITS sequence data. Ann. Missouri Bot. Gard. 93:

103–121.

Superorder LAMIANAE 523

Agababian VS and KT Tumanian. 1977. Contributions to the

palynomorphological study of the family Gentianaceae, part

4. Biol. Zhurn. Armenia 30(8): 43–53 (in Russian).

Albers F and U Meve. 2001. A karyological survey of

Asclepiadoideae, Periplocoideae, and Secamonoideae, and

evolutionary considerations within Apocynaceae s.l. Ann.

Missouri Bot. Gard. 88: 624–656.

Albert VA and L Struwe. 1996. Morphological cladistics of

Gentianaceae. III. Phylogeny and biogeography of the sapro-

phytic tribe Voyrieae. Am. J. Bot. 83(6): 135–136

(abstract).

Albert VA and L Struwe. 2002. Gentianaceae in context. In:

L Struwe and VA Albert, eds. Gentianaceae: systematics and

natural history, pp. 1–20. Cambridge University Press,

Cambridge.

Anderson CL, JHE Rova, and L Andersson. 2001. Molecular

phylogeny of the tribe Anthospermeae (Rubiaceae): system-

atic and biogeographic implications. Aust. Syst. Bot. 14:

231–244.

Andersson L. 1995. Tribes and genera of the Cinchoneae com-

ples (Rubiaceae). Ann. Missouri Bot. Gard. 82: 409–427.

Andersson L. 1996. Circumscription of the tribe Isertieae

(Rubiaceae). Opera Bot. Belg. 7: 139–164.

Andersson L and A Antonelli. 2005. Phylogeny of the tribe

Cinchoneae (Rubiaceae), its position in Cinchonoideae, and

description of a new genus, Ciliosema. Taxon 54: 17–28.

Andersson L and C Persson. 1991. Circumscription of the tribe

Cinchoneae (Rubiaceae) – a cladistic approach. Plant Syst.

Evol. 178: 65–94.

Andersson L and JHE Rova. 1999. The rps16 intron and the

phylogeny of the Rubioideae. Plant Syst. Evol. 214:

161–186.

Andreasen K and B Bremer. 1996. Phylogeny of the subfamily

Ixoroideae (Rubiaceae). Opera Bot. Belgica 7: 119–138.

Andreasen K and B Bremer. 1999. Phylogenetic utility of the

nuclear rDNA ITS region in the subfamily Ixoroideae

(Rubiaceae): comparisons with cpDNA rbcL sequence data.

Plant Syst. Evol. 217: 119–135.

Andreasen K and B Bremer. 2000. Combined phylogenetic anal-

ysis in the Rubiaceae-Ixoroideae: morphology, nuclear and

chloroplast DNA data. Am. J. Bot. 87: 1731–1748.

Andronova NN. 1977. Structure of the ovule in Rubiaceae Juss.

Bot. Zhurn. 62: 1461–1469 (in Russian).

Andronova NN. 1988. Comparative embryology of Loganiaceae

and Rubiaceae. Bot. Zhurn. 73: 937–951 (in Russian with

English summary).

Backlund M, B Oxelman, and B Bremer. 2000. Phylogenetic

relationships within the Gentianales based on ndhF and rbcL

sequences, with particular reference to the Loganiaceae. Am.

J. Bot. 87: 1029–1043.

Behnke H-D. 1975. Elektronenmikroskopische Untersuchungen

zur Frage der verwandtschaftlichen Beziehungen zwischen

Theligonum und Rubiaceae: Feinbau der Siebelement-

Plastiden und Anmerkungenzur Strucktur der Pollenexine.

Plant Syst. Evol. 123: 317–326.

Bendre AM. 1973. Studies in the family Loganiaceae. I.

Trichomes. J. Indian Bot. Soc. 52: 225–234.

Bendre AM. 1975. Studies in the family Loganiaceae: II.

Embryology of Buddleia and Strychnos. J. Indian Bot. Soc.

54: 272–279.

Bisset NG. 1958, 1961. The occurrence of alkaloids in the

Apocynaceae. Ann. Bogor. 3: 105–236, 1958; 4: 65–144, 1961.

Bisset NG. 1975. Chemical structures and biosynthesis of

Loganiaceae alkaloids. Pharm. Weekbl. 110: 425–441.

Bisset NG. 1980a. Alkaloids of the Loganiaceae. In: JD

Phillipson and MH Zenk, eds. Indole and biogenetically

related alkaloids, pp. 27–61. Academic, London.

Bisset NG. 1980b. Phytochemistry of the Loganiaceae. In:

A Engler and K Prantl, eds. Die natürlichen Pﬂ anzenfamilien,

28b: 211–237.

Bisset NG and AK Choudhury. 1974. Alkaloids and iridoids

from Strychnos nux-vomica fruits. Phytochemistry 13:

265–269.

Bisset NG, Th WJ Gadella, AJM Leeuwenberg, AMW Mennega,

and W Punt. 1980. General discussion of relationships

between taxa inside and with taxa outside the family

Loganiaceae. In: A. Engler and K. Prantl, eds. Die natürli-

chen Pﬂ anzenfamilien, 28b: 3–7.

Boppré M. 2005. Pyrrolizidine alkaloids in Apocynaceae. In:

XVII International Botanical Congress, Vienna, Abstracts,

p. 398.

Bouman F and S Schier. 1979. Ovule ontogeny and seed coat

development in Gentiana, with a discussion on the evolu-

tionary origin of the single integument. Acta Bot. Neerl. 28:

467–478.

Bouman F, L Cobb, N Devente, V Goethals, PJM Maas, and E

Smets. 2002. The seeds of Gentianaceae. In: L Struwe and VA

Albert, eds. Gentianaceae: systematics and natural history, pp.

498–572. Cambridge University Press, Cambridge.

Bremer B. 1992. Phylogeny of the Rubiaceae (Chiococceae)

based on molecular and morphological data: useful

approaches for classiﬁ cation and comparative ecology. Ann.

Missouri Bot. Gard. 79: 380–387.

Bremer B. 1996a. Combined and separate analyses of morpho-

logical and molecular data in the plant family Rubiaceae.

Cladistics 12: 21–40.

Bremer B. 1996b. Phylogenetic studies within Rubiaceae and

relationships to other families based on molecular data. In:

E Robbrecht, C Puff and E Smets, eds. Second International

Rubiaceae Conference Proceedings, pp. 33–50. Opera Bot.

Belgica 7.

Bremer B. 1997. Combined and separate analyses of morpho-

logical and molecular data in the plant family Rubiaceae.

Cladistics 12: 21–40.

Bremer B and O Eriksson. 1992. Evolution of fruit characters

and dispersal modes in the tropical family Rubiaceae. Biol.

J. Linn. Soc. 47: 79–95.

Bremer B and RK Jansen. 1991. Comparative restriction site

mapping of chloroplast DNA implies new phylogenetic rela-

tionships within Rubiaceae. Am. J. Bot. 78: 198–213.

Bremer B and J-F Manen. 2000. Phylogeny and classiﬁ cation of

the subfamily Rubioideae (Rubiaceae). Plant Syst. Evol.

225: 43–72.

Bremer B and L Struwe. 1992. Phylogeny of the Rubiaceae and

the Loganiaceae: congruence or conﬂ ict between morpho-

logical and molecular data? Am. J. Bot. 79: 1171–1184.

Bremer B and M Thulin. 1998. Collapse of Isertieae, re-

establishment of Mussaendeae, and a new genus of Sabiceeae

(Rubiaceae): phylogenetic relationships based on rbcL data.

Plant Syst. Evol. 211: 71–92.

Bremer B, RG Olmstead, L Struwe, and JA Sweere. 1994. rbcL

sequences support exclusion of Retzia, Desfontainia, and

Nicodemia from the Gentianales. Plant Syst. Evol. 190:

213–230.