collecting in Yunnan 云南植物

 

Caprifoliaceae or 忍 冬 科 rěn dōng kē (winter bearing)

The Caprifoliaceae includes many genera of horticultural and economic importance such as honey-suckles, snow berries and abelias. Some taxa are used medicinally, and Lonicera caerulea is cultivated for its edible fruits. In evolution and development research this family represents an important example in the evolution of the thyrsoid inflorescence.

Southwest China is a centre of diversity for the Caprifoliaceae sensu latu (including Dipsaceae, Linnaeaceae, Morinaceae and Valerianaceae). 15 of the 36 genera of Caprifoliaceae can be found in their natural habitat in Yunnan including many well-known, hardy, flowering shrubs such as Abelia, Dipelta, Leycesteria and Symphoricarpos. In addition, Yunnan is home to 30 species of Lonicera (honeysuckle) ranging from large-leaved, subtropical climbers such as Lonicera hildebrandiana to dense, shrubby alpines such as Lonicera litangensis, reflecting the great diversity of climate and habitat within the province.

Lonicera cyanocarpa Franch. Section Isika Subsection bracteatae Lonicera nigra L. Section Isika Subsection rhodanthae
Lonicera setifera Franchet Section Isika Subsection bracteatae Lonicera maackii (Ruprecht) Maximowicz, Section Isika Subsection rhodanthae
Lonicera maackii (Ruprecht) Maximowicz, Section Isika Subsection rhodanthae Lonicera tangutica Maximowicz Section Isika Subsection purpurascentes
flowers inflorescence
Leycesteria gracilis (Kurz) Airy-Shaw    Triosteum himalayanum Wall. 

Key to the genera of Caprifoliaceae s.s:

1a.    Flowers single and opposite in a spike.
2a.    Ovaries 5-, 7-, or 8-locular, all locules with many ovules ......................  Leycesteria
2b.    Ovaries 4-locular, 2 locules with 1 fertile ovule, other 2 locules with many sterile ovules     ........................................................................................................Symphoricarpos
1b.    Flowers paired (rarely single and then pedunculate; rarely in 3s and then bracteoles fused into a cupule) or in whorls of 6.
3a.    Flowers paired (rarely single and then pedunculate; rarely in 3s and then bracteoles fused into a cupule) ............................................................................................... Lonicera
3b.    Flowers in whorls of 6.
4a.    Flowers 6 in a single whorl, with 2 pairs of tight involucral bracts forming a capitulum; capitula paniculate; shrubs with triplinerved leaves ..................................... Heptacodium
4b.    Inflorescence different, if flowers in a capitulum then plant a climber with perfoliate involucral bracts and bilabiate flowers in 1–4 whorls.
5a.    Herbs; leaves pinnatifid, otherwise perfoliate .......................................... Triosteum
5b.    Shrubs or climbers; leaves entire, not perfoliate (involucral bracts sometimes perfoliate).
6a.    Shrubs; flowers infundibuliform ............................................................ Leycesteria
6b.    Climbers; flowers bilabiate ..................................................................... Lonicera

TRIOSTEUM

Triosteum is a small genus with only six species. It has a E Asia and North America disjunction distribution with very low level of morphological diversification. The genus is characterised by the terminal inflorescences of sessile 6-flowered whorls.It also shows some fruit adaptations like a drupe with 2-4 pyrenes with a thick stony endocarp

cyme

section of the nectary

corolla
Triosteum sinuatum cyme of three flowers Triosteum himalayanum section of the corolla tube with nectary bulge Triosteum sinuatum corolla

berry

section of the pyrene

section of the pyrene
Triosteum pinnatifidum drupe Triosteum himalayanum section of the pyrene Triosteum sinuatum section of the pyrene showing ribs

Some examples:

Triosteum himalayanum Wallich

Triosteum pinnatifidum Max.

Triosteum sinuatum Maximowicz

LEYCESTERIA & SYMPHORICARPOS

The two genera are also small genera and can be characterised by the spike of flowers. They also differ by their shrub like habit. Leycesteria is only found in Asia. Flowers can be single but they can also be in whorls of 6 flowers. The ovaries mostly have more than 3 locules.

fruits

Leycesteria formosa spike of berries (6 berries per whorl)

HEPTACODIUM

This genus is one of the most difficult group to place in the classification and it is though that it could have been the result of a past hybridization with tribe Linnaeeae. Though this shrub is easily distinguished by its triplinerved leaves the capitula of 6 to more flowers represent the key character. It also differ by the achene like fruit with a large persitent calyx.

Heptacodium miconioides capitulum of achenes

Some examples:

Heptacodium miconioides Rehder

LONICERA

The Genus Lonicera was treated in a monographic fashion by Rehder (Rehder, A. 'Synopsis of the genus Lonicera' Rep. Mo. Bot. Gard. xiv. : 27-232. 1903.). Below we have created a table with all the chinese species and their corresponding taxonomic groups according to Rehder classification. No monographic work has been carried out after Rehder but Nina Theis, Michael J.Donoghue and Jianhua Li published a Phylogentics paper on Lonicera in Systematic Botany (2008), 33(4): 776-783. As expected it raised many questions about Rehder's group delimitation but this work was limited in number of taxa as well as markers used. There is much work to be done to produce a satisfying infrageneric classification.

Some morphophological characters are constant and reliable like the paired ovaries versus a whorl of 6 flowers. Some others are more or less reliable like the brown pith or the acutely 4 angular winter buds.

Subgenus Section Subsection Species in China Distribution

Subgenus Chamaecerasus

 

 

 

 

 

Section Isoxylosteum

Subsection microstylae

L.angustifolia, L.rupicola, L.tomentella, L.tubuliflora

Himalayas

 

 

Subsection spinosae

L. spinosa

Himalayas

 

Section Isika

Subsection purpurascentes

L.microphylla, L.tangutica species complex,

Japan, Himalayas, China, Altai, North America, Mexico

 

 

Subsection coerulea

L.coerulea

Northern hemisphere

 

 

Subsection cerasinea

 

Japan

 

 

Subsection pileata

L.gynochlamydea, L.ligustrina

China, India

 

 

Subsection vesicariae

L.ferdinandi

China, Mongolia

 

 

Subsection chlamydocarpi

L.hypoleuca

Himalayas, W. Asia

 

 

Subsection fragrantissima

L.fragrantissima, L.mucronata

China

 

 

Subsection bracteatae

L.elisae, L.hispida species complex, L.humilis, L.praeflorens, L.setifera, L.subhispida

Central Asia, China, Himalayas, Japan

 

 

Subsection pyrenaicae

 

Pyrenees, Balearic islands

 

 

Subsection distegia

 

W North America

 

 

Subsection oblongifoliae

 

NE North America

 

 

Subsection alpigenae

L.alpigena species complex

Europe, Japan, China, Himalayas, Central Asia,

 

 

Subsection rhodanthae

L.maximowiczii species complex, L.nigra species complex, L.modesta, L.oblata

W North America, China, Japan, Asia minor, Himalayas, North Africa, Europe

 

Section Coeloxylosteum

Subsection tataricae

L.tatarica

Central Asia, Europe

 

 

Subsection ochranthae

L.chrysantha, L.maackii, L.ruprechtiana, L.trichosantha

China, Japan, Europe, Altai, Himalayas

 

Section Nintooa

Subsection calcaratae

L. calcarata

China

 

 

Subsection breviflorae

L.acuminata, L.crassifolia

China, Himalayas, India, Sumatra, Java

 

 

Subsection longiflorae

L.bournei, L.ferruginea, L.hildebrandiana, L.japonica, L.macrantha species complex, L.longiflora

India, Himalayas, China, Europe, North Africa, Java, Japan, Viet-Nam, Burma

Subgenus Periclymenum

 

Subsection Phenianthi

L. subaequalis

North America, Mexico, China

 

 

Subsection Cypheolae

L. yunnanensis

North America, China

 

 

Subsection Eucaprifolia

L. tragophylla

Europe, North Africa, China,

 

 

Subsection Thoracianthae

 

Afghanistan

Subgenus Chamaecerasus: Flowers paired (occasionally single), axillary.

Section Isoxylosteum: Shrubs; corolla actinomorphic, tube not gibbous and nectary of compact sessile glandular hairs in 5 regular lines; branches with solid pith; accessory buds absent.

Subsection microstylae and subsection spinosae seem not to be supported

corolla mouth

style and stigmas

style
Lonicera angustifolia var. angustifolia 4-merous flower Lonicera angustifolia var. angustifolia short style Lonicera rupicola var. syringantha short style

paired ovaries

berries pair

berry
Lonicera angustifolia paired ovaries and bracts Lonicera angustifolia var. myrtillus paired berries Lonicera angustifolia var. myrtillus paired berries

Some examples:

Lonicera angustifolia Wallich ex de Candolle

Lonicera angustifolia Wallich ex de Candolle

Lonicera myrtillus Hook.f. & Thomson

Lonicera rupicola J.D.Hooker & Thomson var. syringantha (Maxim.) Zabel

Section Isika: Shrubs; corolla zygomorphic (sometimes only at base), tube gibbous at base; branches solid.

Subsection Coeruleae; Bracteoles fused into a cupule enclosing ovaries at anthesis (note that this can give appearance of fused ovaries and absent bracteoles); bracteoles accrescent and enclosing paired berries in fruit. Winter bud with two conspicuous and large outer scales

paired flowers

cross section of the ovary

bracts and bracteoles
Lonicera coerulea paired ovaries enclosed in the cupular bracteoles Lonicera coerulea section of the paired free ovaries with the cupule around Lonicera coerulea paired berries enclosed in the fleshy pruinose cupular bracteoles

berries

petioles

axillary buds
Lonicera coerulea paired berries enclosed in the fleshy pruinose cupular bracteoles Lonicera coerulea, accessory buds Lonicera coerulea axillary winter bud with two large outer perulae

Some examples:

Lonicera caerulea L.

Subsection pileata; Bracteoles fused into a cupule enclosing ovaries at anthesis (note that this can give appearance of fused ovaries and absent bracteoles); bracteoles not accrescent and enclosing paired berries in fruit. Calyx forming a collarlike emergence at base.

paired ovaries

corolla mouth

calyx
Lonicera ligustrina paired ovaries enclosed in cupular bracteoles Lonicera ligustrina subregular corolla Lonicera ligustrina calyx forming a colar-like emmergence

longitudinal section of the corolla

berries

berries
Lonicera gynochlamydea corolla section showing bulging nectary Lonicera ligustrina paired berries without the cupular bracteoles Lonicera ligustrina paired berries without the cupular bracteoles

Some examples:

Lonicera gynochlamydea Hemsl. ex Forb. & Hemsl.

Lonicera ligustrina Wallich

Lonicera pileata Oliv.

Subsection vesicariae; Bracteoles fused into a cupule enclosing ovaries at anthesis (note that this can give appearance of fused ovaries and absent bracteoles); bracteoles sligthly accrescent, dry at maturity and enclosing paired berries in fruit. Calyx forming a collarlike emergence at base. inflorescence terminal, of 3-8 whorls

Subsection Bracteata; Ovaries free, outer bracteoles usually large and enveloping the ovaries.

calyx and bracts

corolla tube base

berries
Lonicera setifera paired ovaries enclosed by the large outer bracteoles Lonicera hispida nectary bulge at the base of the corolla tube Lonicera hispida paired berries with large accrescent outer bracteoles

Some examples:

Lonicera cf. hispida Roem. & Schult.

Lonicera setifera Franch.

Lonicera setifera Franch.

Subsection fragrantissima; Ovaries fused, bracteoles small

ovaries

Lonicera fragrantissima paired partly fused ovaries

Some examples:

Lonicera fragrantissima Lindley & Paxton subsp. standishii (Carriere) P.S.Hsu & H.J.Wang.

Subsection purpurascentes; Ovaries connate, corolla with regular erect lobes

bracts and bracteole

fruiting branch

abaxial leaf surface
Lonicera tangutica paired ovaries Lonicera litangensis paired berries and the striking resemblance with L.angustifolia Lonicera tangutica domatia on abaxial leaf surface

Some examples:

Lonicera obovata Royle

Lonicera obovata Royle

Lonicera tangutica Maxim.

Subsection rhodanthae; Winter bud acutely 4-angular

bracts and bracteoles

vegetative bud
Lonicera nigra paired berries and bracteoles Lonicera nigra 4-angular bud Lonicera nigra 4-angular bud

Some examples:

Lonicera lanceolata Wallich

Subsection alpignae; no distinctive characters found

Section Coeloxylosteum: Shrubs; corolla zygomorphic (sometimes only at base), tube gibbous at base; branches hollow

Subsection tataricae; no distinctive characters found

Some examples:

Lonicera cf. tatarica Linn.

Subsection Ochranthae; no distinctive characters found bud

longitudinal section of the stem

ovaries

paired berries
Lonicera trichosantha section of the stem showing brown pith Lonicera trichosantha var. deflexicalyx paired berries showing papery calyx Lonicera chrysantha paired berries
vegetative bud vegatative bud scales abaxial leaf surface
Lonicera chrysantha bud scales showing inner sericeous ones Lonicera trichosantha var. deflexicalyx bud scales Lonicera trichosantha domatia on abaxial leaf surface

Some examples:

Lonicera chrysantha Turcz. ex Ledebour subsp. chrysantha

Lonicera maackii (Rupr.) Maxim.

Lonicera quinquelocularis Hardw.

Lonicera trichosantha Bureau & Franchet var. deflexicalyx (Diels) P. S. Hsu & H. J. Wang

Section Nintooa: Climbers; corolla zygomorphic (sometimes only at base), tube gibbous at base.

Subsection calcaratae; corola with long basal spur

Subsection breviflorae; corolla not exceeding 3cm long. Lonicera crassifolia is a peculiar species that has trailing stems.

paired ovaries

longitudinal section of the corolla

longitudinal section of the stem
Lonicera acuminata paired ovaries Lonicera acuminata section of the corolla tube showing nectaries Lonicera crassifolia flowers
longitudinal section of the stem longitudinal section of the stem longitudinal section of the stem
Lonicera crassifolia paired ovaries Lonicera crassifolia bracteoles Lonicera crassifolia ovaries cross section

Some examples:

Lonicera acuminata Wall var. acuminata

Lonicera acuminata Wall var. acuminata

Lonicera acuminata Wall var. acuminata

Subsection longiflorae; corolla more than 3cm long

flowering branch

Lonicera similis flowers

Some examples:

Lonicera japonica Thunb. ex Murray

Subgenus Periclymenum: Flowers in 3-flowered cymes forming whorls at ends of branches

Subsection Phenianthi; Corolla with almost equal lobes or indistinctly 2-lipped

Subsection Cypheolae; Corolla 2-lipped, less than 4cm long

Subsection Eucaprifolia; Corolla 2-lipped, more than 4cm long

whorl

bracts and bracteoles

berries
Lonicera tragophylla whorl of 6 ovaries Lonicera tragophylla whorl of 6 ovaries and bracts Lonicera tragophylla whorl of berries

Some examples:

Lonicera tragophylla Hemsley

  Home page

Caprifoliaceae of Yunnan