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Synopsis of the genus Lonicera



Alfred Rehder



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SCIENTIFIC PAPERS.



SYNOPSIS OF THE GENUS LONICEBA.

BY ALFRBD REHDER.
INTRODUCTION.

The last general review of the genus Lonicera was pub-
lished more than seventy years ago by A. P. De CandoUe
in the fourth volume of his Prodromus, where 53 species
are enumerated of which, however, one is inserted twice,
one does not belong to the genus and nine are now
referred to other species as varieties or synonyms. This
leaves only 42 recognized species. Since that time
this number has more than tripled, chiefly from discover-
ies made in central and eastern Asia, so that now there are
known more than 150 well distinguished species.

As the literature and references to these species are
widely scattered through numerous books and periodicals,
it is at present rather difficult to properly place and
determine unknown species, particularly those of Asiatic
origin, though the difficulty is somewhat relieved by more
or less comprehensive accounts of the species of certain
regions, as given by Maximowicz, Eegel, Clarke, Franchet,
Gray, Wolf, and by systematic enumerations of the culti-
vated species published by Koch, Dippel and Koehne,

In looking over the literature of the genus one cannot
help noticing the great diversity of opinion exhibited by
the different writers as regards the limitation of the divi-
sions and subdivisions of the genus. This is not because
the distinctive characters are obscure and slight ; on the
contrary, there is a great variety of obvious characters,

(27)

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28 MISSOURI BOTANICAL GARDEN.

but they are promiscuous and of very different systematic
value and the grouping will vary according to the charac-
ters to which preference is given. Unfortunately most
authors have seized upon characters of secondary value
and therefore their arrangement has not expressed the
natural affinities of the species to any great extent. Most
of the older herbalists, as Brunfels, Fuchs, Gresner,
Dodonaeus and Bauhin, distinguished two genera, Pericly^
menum or GaprifoUum and Xylosteum or Ohamaecerasus ;
only few, as Gerard and Clusius, recognized a single genus,
under the name Pericly menum ^ which is the oldest name
for the genus though it is not quite clear whether Dios-
corides, who used it first, intended it for a species of Loni^
cera. Tournefort in 1700 distinguished four genera, Cajp-
rifoUum^ Peinclymenum^ Xylosteum and Ohamaecerasus ^
which were united by Linn^ under CaprifoUum in his
Systema Plantarum (1735), but in his Genera Plant-
arum (1737) and Species Plantarum (1753) he substi-
tuted the name Lonicera for GaprifoUum and added
other genera and species afterwards again removed from
this genus. Few botanists, however, followed Linn^
and most divided the genus into at least two genera, one
containing the climbing, the other the upright species,
and it was not until the appearance of DeCandoUe's
Prodromus that Linux's conception of the genus was
generally accepted. Miller (1754) called the genera
Periclymenum and Xylosteum^ but changed the latter
name in 1759 to Lonicera. Delarbre (1800) named
them GaprifoUum and Ghamaecerasus ; Humboldt, Bon-
pland & Kunth (1818) GaprifoUum and Xylosteum;
Eoemer & Schultes (1819) GaprifoUum and Lonicera;
and Torrey (1825) Lonicera and Xylosteum. Another
division into two genera was essayed by Necker (1790),
who separated the species with almost regular limb of the
corolla from Lonicera as Gobaea. Three genera were
established by Adanson, who distinguished the species



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SYNOPSIS or THE GEIOJS LONIGEBA. 29

with connate ovaries as Isiha and called the other two
genera Xylosteum and Caprifolium^ while Borkhausen
and Boehling, who took up. his laika^ called them Lonicera
and Oaprifolium. A different division into three genera
was made by Sweet (1830), who separated the climbing
species with flowers in pairs from Lonicera and Caprifo- ; >^
Hum as Nintooa. Webb (1838) adopted NirUooa^ but ^'
substituted Xylosteum for Lonicera and Lonicera for -
Capri folium. The four genera of Tournefort were ac-
cepted by Medicus (1789), but for Xylosteum he substi-
tuted the name Isika. Bafinesque at first (1820) also
recognized all the Toumefortian genera, but changed
Periclymenum to Periclyma and Chamaecercums to
Chamerasia; and on Lonicera sempervirens as the type, he
founded a new genus, Phenianthus^ probably not realiz-
ing that Toumefort's Pe^nclymenum is based on the same
species. In his New Sylva (1836) he had changed his
view, using Lonicera for the climbing, and Xylosteum for
the upright honeysuckles, and he founds on L. involucrata
the genus Distegiaj a genus which has not been recognized
by any botanist except recently by Greene. The only
other botanist who took up the Toumefortian genera was
Spach (1839), who distinguished Xylosteum^ Lonicera
( CAamaecerewtw Toum.), Oaprifolium and Periclymenum^
referring Nintooa to Oaprifolium as a section.

Most botanists, however, especially after the first quar-
ter of the last century, accepted the genus Lonicera in the
Linnean sense with some emendations. The emended
genus is usually ascribed to Desfontaines, Flora Atlantica
(1798). There seems, however, no reason for this, since
he says nothing to indicate that he deviates from the Lin-
nean conception of the genus except that the two species
he describes happen to belong to Lonicera proper; but
for this same reason the emended genus could be cred-
ited as well to Hudson, Scopoli or Loureiro. The first
place where the genus appears in the now accepted concep-



II {



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30 MISSOURI BOTANICAL GABDEN.

tion is in Willdenow's Enumeratio Plantarum Horti Bero-
linensis (1809).

The subdivision of the genus as proposed by Linn^ into
Periclymena and Chamaecerasa (misspelled in the first edi-
tion of Species Plantarum ** Chamaecerosa ") has always
remained the principal subdivision, though usually with
the names changed according to De CandoUe (1805) into
Caprif olium and Xylosteum. In his Prodromus (1830),
De Candolle subdivided the subgenus Xylosteum further
into Isika, Cuphantha, Chamaecerasus and Nintooa. He
included Nintooa in Xylosteum, while other botanists, as
Eafinesque (1836), Maximowicz (1877) and Clarke (1882),
refer it less happily to Caprif olium to which it is similar
in habit and partly in the shape of the corolla, but not
closely allied. By Bafinesque, Nintooa is called Eunemium,
and Periclymenum (Tourn.^ Kantemon: he apparently had
forgotten that he had named the same group sixteen years
earlier Phenianthus. For the American species of Capri-
folium he proposes a new section Cjrpheola, which is iden-
tical with the later Caprif olium § Loniceroides of Spach
(1839). Another new section was based on L. Iberica by
Jaubert & Spach (1847) and named Chlamydocarpus.
Hooker f. & Thomson (1858) separated as Bracteatae
the Lonicera hiapida and allied species ; and Maximowicz
(1877) formed of the red-flowered species of the section
Chamaecerasus the group Bhodanthae. The latest addi-
tion to the sections is Vesicaria of Komarov (1900)
founded upon L. vesicaria and L. Ferdinandi. Quite
recently the unnamed sections proposed by Koehne (1893)
have been adopted with slight changes and named by Zabel
(1903). Of these only the sections Ochranthae, Subses-
fiiliflorae and Ebracteolatae can be considered new. The first
is substantially the same as Maximowicz's unnamed second
section of his Chamaecerasus, but the other two seem too
heterogeneous to be maintained.

There can be little difference of opinion about the



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SYNOPSIS or THE GENUS LONIGEBA. 31

division of the genus into the two subgenera, Periclyme-
num and Chamaecerasus, or as they are usually called,
Caprifolium and Xylosteum, which form two very well
defined and natural groups if based on the character of the
inflorescence and not on the habit.

The subgenus Periclymenum, which is comparatively
small and homogeneous, can be divided easily into closely
I related subsections ; but the much larger subgenus Chamae-
] cerasus has been subdivided in various and mostly not
very satisfactory ways. Most botanists take the shape of
the corolla, — whether the limb is two-lipped or almost
regular, — as the best character for the subdivision, but
this difference can scarcely be used to characterize even
subsections, since it would separate in several cases very
closely related species, while in other cases the shape of
the limb is intermediate. It is a character of but sec-
ondary value, as it has been probably acquired late in the
genesis of the genus by part of the species of several
already well differentiated groups as an adaptation to
certain insect visitors ; it therefore would furnish a very
artificial subdivision without any concomitant characters.
There is, however, a group of Loniceras with really actino-
morphous flowers, hitherto confounded with the pseudo-
actinomorphous species. Only Klotzsch seems to have
noticed this fact, as appears ^from a note by A. Garcke
in Reise des Prinzen Waldomar (1862), p. 86:
*^ Ausser diesen befindet sich in der Sammlung noch
eine, wie es scheint, von i. MyrtiUus nicht verschie-
dene Art, die Dr. Klotzsch neu benannt and worauf
er eine neue Gattung griindete, deren Namen ich zur Ver-
meidung unniitzer Synonyme mit StiUschweigen iibergehe."
Theactinomorphous character of the species of Isoxylosteum,
as Ipropose to call this section, is not only shown by the limb,
but much more markedly by the presence of five nectaries^
while all the other species have only one to three nectaries at
the base of the corolla below the lower lip, usually placed



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32 MISSOURI BOTANICAL GARDEN.

in a more or less pronounced depression forming outwards
a gibbosity of the tube. Besides this character there is a
difference in the vernation of the foliage, the leaves being
flat or carinate, while in all other species I have found them
convolute in bud or occasionally involute, and a further differ-
ence is the absence of the superposed accessory buds so com-
mon in most other species. The remaining species of the
subgenus Chamaecerasus can be easily divided into further
sections by the pith of the branches, which is either well
developed or evanescent : in the first case the branches are
solid, in the second they are hollow. This behavior of the
pith is a concomitant of a difference in the union of the
bractlets. In the solid-branched species there is a tendency
of the bractlets of one flower to unite more or less with the
bractlets of the second flower of the same pair or to form
altogether a complete cupula, and at the same time there is
a tendency of the ovaries to unite, while in the hollow-
branched species only the bractlets of the same flower unite,
or all are free, as are the ovaries. The hollow-branched
species form a remarkably homogenous section, for which I
propose the name Coeloxylosteum, in reference to the hollow
branches. The other section, however, for which the name
Isika of Adanson in a somewhat enlarged sense may be
used, presents a very great variety of different characters,
especially as regards the bractlets : sometimes these are en-
tirely wanting, sometimes perfectly distinct and sometimes
wholly connate, even so far as to form with the inclosed
ovaries a kind of pseudocarp. The limb of the corolla
varies from almost regular to distinctly two-lipped; the
ovary varies from two- to three-celled and the bud scales
and other vegetative characters also show great differ-
ences. Though these differences afford good characters
to subdivide the species of this section into a number of
well-defined subsections, it is rather difficult to group satis-
factorily the subsections according to their affinities, since
their relations cannot be expressed by linear sequence.



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SYNOPSIS or THE GENUS LONICERA. 33

On the geographical distribution and the morphological
and ecological aspects of the genus, which presents some
very interesting facts, I shall speak more fully in another
paper ; only a few notes on the geographical distribution
may follow here. The genus is distributed throughout the
temperate regions of the northern hemisphere, crossing the
equator only in the Malayan Archipelago, where it reaches
Java as its most southern point, and northwards extending
to the arctic circle. Its greatest segregation occurs in cen-
tral and eastern Asia, where all sections and most subsec-
tions are represented. In North and Central America only
twenty species occur, and the European together with the
Mediterranean flora contains but eighteen species. All the
rest are Asiatic. Of the two subgenera, Chamaecerasus is
found throughout the whole range of the genus, while Peri-
clymenum does not extend so far north or southward, and
in central Asia it is represented only by four species, each,
however, belonging to a different subsection. In Europe
and in North America, on the other hand, it is well repre-
sented by many more or less polymorphous species. Of
the four sections of Chaipaecerasus, the section Isika has the
widest range, but does not reach the Malayan Archipelago.
Isoxylosteum is restricted to the higher elevations of cen-
tral Asia and Coeloxylosteum extends from western cen-
tral Asia through the Mediterranean region to Spain and
Morocco. Nintooa is well represented in eastern and
southern Asia and appears again in the western part of the
Mediterranean region. In North America the subgenus
Chamaecerasus is represented only by the section Isika with
7 species, one of them circumpolar.

In accounting for all published specific and varietal
names, I have omitted purposely any reference to the 270
new species proposed by Gandoger,* since his names seem to
have their origin only in the wish to give to each specimen

• Flora Europaea 11 : 14-28 (1886).

8



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34 HI8SOURI BOTANICAL GARDEN.

of his private herbarium a separate name. As he has not
paid any attention to previously published names except
the Linnean ones, his nomenclature cannot be regarded as
resting on scientific principles and may therefore be passed
over, as has been done by the editors of the Index Kewen-
sis.

In the course of the study of the genus, the following
herbaria have been consulted: the Gray Herbarium and the
herbarium of the Arnold Arboretum, Harvard University,
the herbarium of the Missouri Botanical Garden, St. Louis,
the herbarium of Columbia University and of the New York
Botanic Garden, the herbarium of the Academy of Natural
Sciences, Philadelphia, the National Herbarium at Wash-
ington, the Biltmore Herbarium, Biltmore, N. C, the her-
barium of the Royal Botanic Museum at Berlin, the herba-
rium of the University of Goettingen (including the Grise-
bach Herbarium), the herbarium of the Eoyal Botanic
Garden at Kew, the herbarium of the Museum of Natural
History at Paris, and the herbarium of the Botanic
Garden at Brussels. From the herbarium of the Imperial
Botanic Garden at St. Petersburg and from the Herbier
Boissier at Chamb^zy near Geneva, a considerable number
of specimens, partly types, were kindly sent to me for ex-
amination. I beg to acknowledge my sincerest thanks to all
those in charge of the above named institutions. I am also
greatly indebted to Professor E. Koehne of Friedenau-
Berlin, who kindly loaned me the Loniceras of his herba-
rium and has furnished me moreover with much valuable in-
formation. I am likewise under obligation for material and
information to Mr. George Nicholson of Kew, Mr. M. L.
de Vilmorin of Paris, Mr. H. Zabel of Gotha, formerly of
Muenden, Mr. E. Wolf of St. Petersburg, Mr. J. Kessel-
ring of St. Petersburg, Mr. E. Bettig of Jena, Mr. L.
Beissner of Bonn, Mr. A. Purpus of Darmstadt, Professor
J. Battandier of Alger-Mustapha, Mr. W. Lipsky of St.
Petersburg and Mr. Joaquim de Mariz of Coimbra.



L.



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SYNOPSIS OF THE GENUS LONICERA. 35

A great aid in the study of the genus has been the
opportunity to observe more than one-third of all species
in a cultivated state , — most of them for years . The general
conception of the affinities of the different species and of
their most natural arrangement I gained first from the study
of the living plants, the close observation of which revealed
characters and combinations of characters which probably
would have remained unnoticed in herbarium specimens.
In addition to the living collections of the Arnold Arbore-
tum and of the Botanic Garden of the University of Goet-
tingen, where I began the study of the genus, I have made
valuable observations in such other living collections as those
of the botanic gardens of Kew, Berlin, Darmstadt, and
Muenden, the collection of Mr. M. L. de Vilmorin at Les
Barres, France, and other gardens.

With the exception of three species, L. Sumatrana^ L.
irichosantha and L, Kachkarovi^ I have seen specimens
of all species enumerated in the present paper. Where
I have seen type specimens the name of the collector is
marked with an exclamation point ( 1 ).

As the genus has not before been treated comprehen-
sively I have taken great pains to collect all bibliographical
references of taxonomic importance, to serve as a base for
further investigations and studies. All the references are
quoted from their original places of publication with a few
exceptions, when the source of information is always given.

LONICEEA.

Lonicera, Linn^ [Gen. PL 57 (1737); Hort. Cliff. 57
(1737); Syst. Veg. ed. 2. 19 (1740)]; Spec. PI. 173
(1753). — Muenchhausen, Hausvater 5:192 (1770). —
Scopoli, Fl. Cam. ed. 2. 1: 152 (1772). — Lamarck, Enc.
Meth. Bot. 1:727 (1783); Tabl. Enc. M^th. Bot. 2:149
(1793). — Loureiro, Fl. Cochinch. 1 : 149 (1793). — Will-
denow, Spec. PI. 12:982 (1797); Enum. PI. Hort. Berol.



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36 MISSOURI BOTANICAL GARDEN.

220 (1809). — DesfontaiDes, Fl. Atl. 1:183 (1798).-^
Persoon, Syn. PL 1:213 (1805).— A. P. De Candolle,
Fl. Fran§. 4 : 269 (1805) ; Prodr. 4: 330 (1830). — Poiret,
Enc. M^th. Bot. Suppl. 2:227 (1811). — Stokes, Bot.
Mat. Med. 1 : 370 (1812). — Link, Enum. PI. Hort. Berol.
1:221 (1821).— Wallich in Eoxburgh, Fl. Ind. ed. 2.
2:174 (1824). — Sprengel, Syst. Veg. 1:757 (1825).—
Ledebour, Fl. Alt. 1:247 (1829); Fl. Eoss. 2:387
(1844).— Loudon, Hort. Brit. 79 (1830); Arb. Frut.
Brit. 2:1042 (1838). — Hooker, Fl. Bor.-Am. 1:281
(1834)..— Bertoloni, Fl. Ital. 2:555 (1835). — Lindley,
Nat. Syst. Bot. ed. 2. 248 (1836). —Koch, Syn. Fl.
Germ. 324 (1837).— Endlicher, Gen. PI. 568 (1838)). —
Spenner in Nees von Esenbeck, Gen. PI. Fl. Germ. faso.
21 : 401. pi. 2 (1840). — Torrey & Gray, Fl. N. Am. 2 : 4
(1841). — Walpers, Eep. Bot. 2:447 (1843). — Grise-
bach, Fl. Eumel. 2:153 (1844).— Gray, Man. Bot. 171
(1848).— Kirillow, Lonic. Euss. Eeich. 9 ( 1849).— Grenier
i&Godron, Fl. France 2:8 (1850). — K. Koch, Linnaea
24:477 (1851); Dendr. 2^:7 (1872).— Miquel, Fl. Ned.
Lid. 2:125 (1856). — Hooker f. & Thomson, Jour. Linn.
Soc. 2:165 (1858).— Bentham & Hooker, Gen. PI. 2:5
(1873). — Franchet & Savatier, Enum. Plant. Jap. 1:203
(1875); 2:383 (1879).— Maximowicz, Bull. Acad. Sci.
St. P^tersb. 24:35; M^l. Biol. 10:55 (1877).— Baillon,
Hist. PI. 7:499 (1880).— Clarke in Hooker, Flor. Brit.
Ind.8:9(1882).— Gray, Syn. Fl. N.Am. 1^:14(1884). —
Nicholson, Diet. Gard. 2:296 (1887); Hand-list Arb.
Kew 2:11 (1896).— Dippel, Handb. Laubholzk. 1:200
(1889).— Laguna, Fl. For. Esp. 2:44 (1890).— Fritsch
in Engler & Prantl, Nat. Planzenf. IV. 4 : 166 (1891).—
Koehne, D. Dendr. 541 (1893).— Britton & Brown, 111.
Fl. N. U. S. Can. 3: 237(1898). — Wolf, Isv. St. Peterb.
Lesn. Inst. 3:3 (1898). — Bubani, Fl. Pyr. 2:331
(1900). — Rehder in Bailey, Cycl. Am. Hort. 2:939



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SYNOPSIS OF THE GENUS LONICEBA. 37

(1900). — Graebner, Bot. Jahrb. 29:594 (1901).—
Penkowsky, Derev. Kust. Eoss. 3:8 (1901).

Miller, Gard. Diet. ed. 7 (1759). See subgen.

Chamaecerasus.

Necker, Elem. Bot. 1:128 (1790), and his

Cohaea.*

Torrey, Fl. N. M. Sect. U. S. 1:242 (1824). See

subgen. Periclymenum.

Periclymenum^ Miller, Gard. Diet. ed. abridg. 3
(1754). See subgen. Periclymenum.

Xylosteum^ Miller, Gard. Diet. ed. abridg. 3 (1754).
See subgen. Chamaecerasus.

Caprifolium [Linn^, Syst. Veg. (1785)]. — Zinn,
Cat. PL Gotting. 210 (1757). — Haller, Hist. Stirp. Helv.
1:300(1768). — Lamarck, Fl. Fran§. 3:364(1778).—
Gaertner, Fruct. Sem. PI. 1 : 135 (1788). — Kuntze, Rev.
Gen. PI. 1:273 (1891). — Greene, Fl. Francisc. 345
(1892)..

Adanson, Fam. PI. 2:157 (1763). See subgen.

Periclymenum.

/«fA;a, Adanson, Fam. PI. 2:501 (1763). See sect.
Isika.

Chamaecerasus i Medicus, Phil. Bot. 1 : 126 (1789). See
subgen. Chamaecerasus.

Cobaea, Necker, Elem. Bot. 1:129 (1790). See
subgen. Chamaecerasus.

Chamerasia^ Eafinesque, Ann. Gen. Sci. Phys. 6 : 83
(1820). See subgen. Chamaecerasus.



* This method of qaotation is adopted to indicate that the two
genera together equal the conception (In this case Loniceray Linnd) to
which the first (in this case Lonicera^ Necker) is referred as a synonym;


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