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A monographaf
The genus BailloniaBy HAROLD N. MOLDENKE (t)
The following is the sixteenth (2) in my series of
monographicstudies of the genera of Verbeil'ta'ceae and
Avicenniaceae. In thelist of citations of herbarium specimens the
following abbreviationsof the names of herbaria are employed: B =
Botanisches Museum,Berlin; Bm = British 'Museum (Natural History),
London;Cb = Delessert Herbarium, Conservatoire et J ardin
Botaniques,Geneva; F= Field Museum of Natural History, Chicago; G =
GrayHerbarium of Harvard University, Cambridge, Mass.; K =
RoyalBotanic Gardens, Kew; Mu = Botanisches Museum, Munich;N =
Britton Herbarium, New York Botanical Garden, New YorkCity; P =
Museum National d 'Histoire Naturelle, Paris; S = Naturhistoriska
Riksmuseet, Stockholm; V = Naturhistorisches Museum, Vienna; W ,=
United States National Herbarium, W a s h i n g ~
(1) The New York Botanical Garden, Bronx Park, N. Y. City, U. S.
A.(.) For the benefit of readers wishing to consult previous
numbers in this series,they are briefly listed here: [1] Aegiphila,
Brittonia 1: 245-477 (1934), supplementsin Phytologia 1: 182-208,
222-240, 248-272 (1937), 289-304 (1938), and 364-368(1939). [2]
Tectona, Phytologia 1: 154-164 (1935). [3] Recordia, Phytologia 1:
171-174(1935), supplement in Rev. Sudam. Bot. 6: 24-25 (1939). [4]
Rehdera, I'edde's Re-pertorium 39: 47-55 (1935), supplement in Rev.
Sudam. Bot. 6:25-27 (1939). [5]Timotocia, Fedde's Repertorium 39:
129-153 (1936). [6] Callicarpa, Fedde's Reper-torium 39: 288-317
and 40: 38-131 (1936). [7] Cornutia, Fedde's Repertmum 40:153-205
(1936). [8] Priva, Fedde's Repertorium 41: 1-76 (1936). [9]
Svensonia,Fedde'8 Repertorium 41: 129-143 (1936). [10]
Rhaphithamnus, Fedde's Repertorium42: 62-82 (1937), supplement in
Rev. Sudam. Bot. 6: 27-29 (1939). [11] Petitia,Fedde's Repertorium
42: 229-251 (1937). [12] Petrea, Fedae's Reperwrium 43: 1-48and
161-221 (1938). [13] Chascanum, Fedde's Repl;rtorium 45: 113.156.
300-319(1938) and 46: 1-12 (1939), supplement in Rw. Sudam. Bot. 6:
15-24 (1939). [14]Amasoniu, Fedde's Repertorium, 1939 ined.? [15]
Bouchea, Fedde's Repertorium,1939 or 1940, ined.?
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H ~ N. MOLDENKE, A monograph of the genus Bailloni,a i 167ton; X
= Herbier Boissier, Universite de GenEwe, Geneva; andZ = H. N.
MOLDENKE herbarium, Watchung, New. Jersey. To thedirectors and
curators of the above-mentioned herbaria the writerextends his most
sincere thanks for their courtesy and kindness inallowing him to
study their material of this genus and for theircontinued and very
generous cooperation throughout the progressof this work. All
material thus far received from these 14 herbariais herein
accounted for and cited and all specimens so cited havebeen
annotated with uniform printed labels giving this
information.Eighty-eight other herbaria have been canvassed, but
did not containany material of this group, or, at least, did not
send any forreVISIOn. I t gives me great pleasure to dedicate this
paper to myvery esteemed and respected friend and colleague, Prof.
Dr. LORENzo R. PARlODI, who has been a pillar of South American
botany forfully 25 years and who we hope may still have before him
very manymore years in which to continue his valuable services to
the scienceof botany.
Baillonia BocQ. in BAILL., Adansonia 2: 251-252. 1862.Shrubs,
with slender tetragonal branchlets; leaves opposite,ter
nate, or scattered, borne on prominent spinescent sterigmata,
deciduous, exstipulate, net-veined, short-petiolate, the blades
usuallyvery narrow and elongate; inflorescence racemose and
indeterminate(centripetal), racemiform or spicate, axillary and
terminal, simple,erect or nutant, rather densely many-flowered,
pedunculate; flowerscomplete,perfect (hermaphroditic), hypogynous,
small; calyx persistent, gamosepalous, varying from tubular to
cyathiform or campanulate, usually irregular and more or less
2-lipped, the rim varyingfrom truncate and subentire or undulate to
5- or 6-apiculate or-toothed, the teeth unequal; corolla
gamopetalous, hypocrateriform,zygomorphic, its tube cylindric,
straight or incurved, ampliate atapex, its limb obliquely 5-parted,
the lobes unequal in size, alternatewith the calyx-teeth, the
anterior one largest, the posterior onessmallest, cochlear il'l
prefloration; fertile stamens 4" didynamous,inserted at the same or
slightly different levels in the corolla-tube,opposite the
calyx-teeth, included or slightly exserted, the fifth(posterior)
stamen represented by a rUdimentary staminode; filaments short,
filiform or flattened; anthers oblong, erect, dorsi fixednear the
base, introrse, 2-locular, the thecae parallel, dehiscing by
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168 Darwiniana, t. 5, 1941longitudinal slits; pistil one,
included, 2-carpellary, but one carpeloften aborted and the pistil
therefore appearing to be simple; stylecapillary, stoutish and
fleshy, ampliate at apex, included; stigmavery small and capitate,
flattened, or punctiform, sometimes obscurely 2-10bed with one
anterior and one posterior lobe; ovary elongateand conic or
subglobose, borne on a hypogynous annular disk, 1-celledor
incompletely 2-celled, with an anterior parietal 1- or
2-ovulateplacenta in each cell, a falBe and usually incomplete
posterior dissepiment separating the placentae; ovules ascending,
hemianatropous,with superior chalaza and inferior micropyle;
fruiting-calyx persistent, indurate, cupuliform; fruit drupaceous,
oblong or pyriform,succulent, the mesocarp fleshy, the endocarp
woody, incompletely2-pyrened, the pyrenes 1- or 2-seeded; seeds
oblong, ascending, theintegument very thin and membranous, without
elldosperm; embryoerect; cotyledons 2, elliptic, equal or unequal,
plane or contorted,the radicle inferior, very short, conic or
turbinate.
This monotypic genus was named by BOCQUILLON in honor ofHENRI
ERNEST BAILLON (Nov. 30, 1827-July 18, 1895), author ofthe classic
Hist'oire des Plantes in 13 volumes and a prodigiousworker on plant
systematics. The type and only known species isB. amabilis BocQ.,
distributed from Mattogrosso, Brazil, and Paraguay to adjacent
Bolivia and Argentina.
In spite of the full and excellent description of the genus
byBOCQUILLON in his paper entitled Description d'un genre nouveaude
la familie des VerbCnacees in Adansonia 2: 251-252 (1862) andin his
classic Revue du groupe des Verbenacees in Adansonia 2;81-165
(1861-1862) and 3: 177-264 (1863), this little genus has beenbadly
confused and misinterpreted by previous workers on thisfamily.
BENTHAM began this confusion when in BENTH. & HOOK.f., Gen. Plo
2; 1133 & 1143-1144 (1876) (1) he combined Diosteawith
Baillonia. The genus Diostea was published by MIERS inTrans. Linn.
Soc. Lond. 27: 102-103 (1870), as a group of some 6species of
plants from Argentina and Chile, segregated from thegenera Lippia
and Verbena. MIERS gave a splendid plate illustrating one species
and showing the generic characters very well. I t
. has nothing 'Yhatever to do with Baillonia and how BENTHAM
cameto combine it with Baillonia is almost inexplicable.
BENTIIAM'S
(1) See BENTHAM'S paper entitled On the joint and separate work
of the authorsof BENTHAM & HOOKER's Genera Plantarum in Journ.
Linn. Soc. Lond. 20:304-308. 1884.
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Hi. N. MOLDENKE, ..4. monograph of the genus B a i l l o n ~ a
169mistake, however, was perpetuated by even that ordinarily
verycareful and exact worker, J. BRIQUET, in his monograph of
theVerbena,ceae in ENGLER & PRANTL. N at. Pflanzenfam. 4, 3a;
152-1513(1895). S. LE M. MOORE Was the first to point out this
unfortunateerror in his paper entitled The phanerogamic botany of
the MattoGrosso Expedition, 1891.92 in Trans. Lrinn. Soc. Lond.,
ser. 2, 4:437 -439 (1895) . Here he compares Diostea with Baillonia
andproves that they differ generically both in floral anatomy
andmorphology and in vegetative characters. Among the
differenceswhich he points out are that Baillonia has branches
which arespinescent after the falling off of the leaves; leaves
which are long,narrow, thick-textured, frequently alternate or
subverticelIate;inflorescences which are usually racemiform j a
calyx which is relatively short and campanulate, with an undulate
or only obscurelytoothed rim; a corolla with a short, straight or
only slightly curvatetube and a large limb; a short fleshy style,
with a relatively largecapitate stigma; and a comparatively large
and fleshy fruit, surrounded only at its base by the persistent
cupuliform calyx, witha large woody endocarp. Diostea, on the other
hand, does not havespinescent branches; its leaves are small,
thin-textured or fleshy,and strictly opposite; the inflorescences
are strictly spicate; thecalyx is comparatively long and tubular,
with distinct lobes; thecorolla has a narrow, elongate, decidedly
curvate tube, and a smalllimb; the style is long and slender, with
a linear recurved stigma;and the fruit is small, 2-lobed, and dry,
completely enclosed by thepersistent calyx, with a very thin
endocarp. He also points outthat Baillonia inhabits tropical
regions in South America, whileDiostea inhabits temperate and
desert areas. He remarks Althoughplaced [by BENTHAM and by BRIQUET]
in different tribes of theVerbenaceae, one cannot fail to be struck
with the evident affinityof Baillonia to Citha1-exylum; the habit,
the racemose inflorescence,and the flowers of both are very
similar; in its 2-seeded fruitalone does Baillonia show any marked
difference . BRIQUET inAnn. Conserv. Jard. Bot. Genev. 7-8; 316
(1904) takes note ofMOORE'S remarks - although he mis-cites his
paper to . Trans.Linn. Soc. Lond. IV, 3, 437 ~ but still does not
plainly repudiatehis previous following Of BENTHAM in combining the
two genera.
In the very simple classification of DUMORTIER (3) Baillonia(1)
DUMORT., Anal. Fam. Plo 22, 1829.
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170 Darwiniana, t. 5, 1941would go into Tribe 1, Verbeneae
DUMORT. BARTLING (1) wouldhave placed it in his Section 2, Verbenea
BARTL., and LINDLEY e)in Tribe Ve,rbeneae BARTL. In the system
proposed by ENDLrCHER CS) it would fall into Tribe 2, Lantaneae
ENDL., which hecharacterized as comprising genera with indehiscent
drupaceousfruit, as contrasted with Tribe 1, Lippiea1e ENDL., whose
membershave dehiscent fruit. MEISNER (4) would also have classified
it illhis Tribe 2, Lantaneae ENDL. Coming now to more recent
andthorough classifications of the family, we find that in the
systemproposed by SCl:IAUER (5) B a ~ l l o n i a would come under
Tribe 1,Verbeneae DUMORT., Subtribe 6, Duranteae SCl:IAU. BENTHAM
(6),the first of the famous classifiers of the Verbenalceae to take
cognizance of Baillonia, made the unfortunate mistake referred to
aboveand placed it in the first section of this Tribe 4, Verbeneae,
alongwith such unrelated genera as Acharitea BENTH., Lantana L.,
LippiaHOUST., Neosparton GRISEB., Bouchea CHAM., and
StachytarphetaVAHL, characterized by Fructus pyrenae 2 v. abortu
solitariae,l-spermae. Fl,ores in spica, sessiles . Actually, he
should haveplaced Diostea in that position and have placed
Baillonia in hissixth section with the genera Citharexylum MILL.,
RhaphithamnusM:IERS, and Duranta L., characterized by Fructus
pyrenae 2-5,2-locellatae, 2-sperma1e. Flores in race,mo v. ad
axillas pedicellati.Calyx truncatus v. minute dentatus .
Following BENTHAM in his erroneous combination of Diostea
withBaillon,ia, BRIQUET (1) places Baillonia in his Subfamily 2,
Verbenoideae BRIQ., Tribe 2, LantaneaeENDIJ., with the genera
TateaF. MUELL., Ub,ochea BAILL., Lantana, Lippia, Bouchea,
Neosparton,and Stachytarpheta. Actually, he should have left
Diostea in thatposition and placed Baillonia in his Tribe 6,
Citharexyleae BRIQ.Needless to say, the generic diagnoses given by
BENTHAM and byBRIQUET are composite diagnoses, including characters
of both Diostea and Baillonia, and must therefore be discounted.
JUNELL, thelatest worker to investigate the taxonomy of the family
as a wnole
(1) BARTL., Ord. Nat. PI., 180, 1830.(2) LINDL., Nat. Syst.
Bot., ed. 2, 278, 1836.(3) ENDL., Ger . PI., 633-638, 1838.(4)
MEISN., PI. Vase. Gen., 290-292, 1840.(0) SCRAU. in A. DC., Prodr.
11: 524-525, 1847.(6) BENTH. in BENTH. & HOOK. f., Gen. PI. 2:
1132-1136 & 1143-1144,1876.(7) BRIQ. in ENGL. & PRANTL,
Nat. Pflanzenfam. 4, 3a: 144 & 152-153, 1895.
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H. N. MiOLDENKE, A monograph of the genu8 Baillonia 171and
related families -(1), proves that MOORE'S sugg8tion wascorrect and
that Baillonia actually is very closely related toCitharexylum and
is to be classified in the same subtribe with thatgenus, and not
with the genus Diostea, which goes in the same groupwith Lippia. He
has made careful morphologic and anatomic dissections of Baillonia
flowers and has found the ovary actually tobe morphologically
4-ovulate, but one ovule (or sometimes twoovules) usually aborted.
He found also (pp. 48-49) that the ovulesare not attached strictly
parietally on the margins of the carpels,as stated by BOCQUILLON,
but actually a slight distance away fromthe margins of the inner
face. He therefore classifies the genusin his Tribe 1, Verbenoideae
(BRIQ.) JUNELL, Su,btribe 6,.Citharexylea.e (BRIQ.) JUNEL.L (2),
along with the genera Citharexy.lum, Rhaphithamnus, and
Duranta.
I t is worthy of note here that the Baillonia spicata of BAILLON
inBull. Soc. Linn. Paris 2; 880 (1890) is actually Citharexylum
ligus-trinum VAN HouTTE, while the Citharexylum linearifolium
whichI published in PhytoZogia 1: 97-98 (1934) is actually
Bailloniaamabilis, indicating again the close relationship of these
two generaand the ease with which they may be confused.Some 38
persons have contributed to our knowledge of this genusand its
bibliography embraces some 22 titles to date. No fossilspecies are
as yet known. The only' economic use for the one knownspecies is
that it has been introduced into cultivation in Paraguayas an
ornamental plant.
1. Baillonia amabilis BocQ. ex BAILL., Adansonia 2: 252, pI. 7.
1862BaiUonia amabilis var. longi/olia KUNTZE, Rev. Gen. Plo 3(2):
250, 1898.Baillonia amabilis ~ r . heterophylla KUNTZE, Rev. Gen.
Plo 3(2): 250, 1898.Citharexylum linearifolium MOLDENKE Phytolorria
1: 97-98, 1934.
Slender shrub, to 4 m. tall; branches and branchlets often
ternate,slender, stramineous (drying white), terete, very glabrous
_ ndsmooth, not at all lenticellate, mostly conspicuously
longitudinallystriate; twigs very slender, stramineous or buff in
calor, opposite;nodes somew!hat incrassate, not annulate, usually
2- or 3-aculeate;
(1) JUNELL, Symb. Bot. Ups. 4: 37-38, 48-50, & 213-214,
1934.(2) See my note in FEDDE, Repert. 41: 135 (1936) in regard to
the invalid
transfers by JUNELL of 6 subfamily and 13 tribal names to lesser
rank withoutt h ~ required modification of the terminations.
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172 Darwiniana, t. 5, 1!)41principal internodes 1-5.5 cm. long;
leaf-scars borne on conspicuousspinescent sterigmata which are
spreading or ascending, slender,1-3.5 mm. long, glabrous, and bear
a long beak-like projection onthe abaxial side, which becomes a
definite spine after the leaf isshed; leaves decussate-opposite or
ternate (or sometimes suboppositeor subspirally scattered or even
alternate on young twigs) ; petiolesslender, 3-6 mm. long,
glabrous; blades chartaceous, dark green andvery llitid on both
surfaces, linear, linear-oblong, or very
narrowlylanceolate-elliptic, 4.5-20 cm. long, 2-11 mm. wide
(usually about15 cm. long and 6 mm. wide), subulate-acuminate at
apex, entireor bearing a few scattered minute (or widely spreading,
subulate,and hooked) teeth along the margins, long-attenuate at
base, bearinga pair of black discoid or elliptic glands at the very
base, perfectlyglabrous on both surfaces, sometimes much
abbreviated and spatulatein shape (0.8-4.5 cm. long), blunt at apex
and there 1-3-toothedwith rather l{mg sharp teeth, the leaf-size
often very variable onthe same twig, with the smallest leaves at
the base; midrib veryslender, prominulous on both surfaces;
secondaries numerous andvery slender, often not distinct, much
elongated and ascending; veinand veinlet reticulation sometimes
more or less prominulous on bothsurfaces, e8pecially above, often
very sparse and obscure on bothsurfaces, especially beneath;
inflorescence axillary and terminal andalso terminating very short
axillary twigs, racemiform Or spicate,erect or nutant, 4-17 cm.
long, 1-1.5 cm. wide in anthesis, simpleand unbranched, rather
densely many-flowered, bracteolate; peduncules very slender, 5"-9
mm. long, glabrous; rachis very. slender andglabrous, similar to
the peduncles and twigs; pedicels obsolete orto 2 mm. long, very
slender and glabrous; foliaceous bracts absent;bractlets
linear-subulate or setaceous, 1-2 mm. long, glabrate,
mostlyobscure; prophylla absent; flowers fragrant, each subtended
by asingle sessile bractlet; calyx membranous, tubular or
cyathiform,varying to cupuliform or campanulate, usually irregular
and moreor less 2-lipped, 2-2.8 mm. wide, the longer lip 4.1-5.5
mm. long,the shorter lip 3.6-5.2 mm. long, glabrous or subglabrate,
with 5(or 6) thicker ribs which often terminate in teeth or
apiculations,the 2 anterior teeth usually somewhat larger than the
3 posteriorteeth, the rim sometimes truncate and subentire or
undulate, sometimes 5 (or 6) -apiculate, mostly 2-lipped with 4
very shallow sinusesand one deep one (about 0.7 mm. deep), or the
longer lip 2-toothedwith ovate and sharply acute teeth (0.7-1 mm.
wide at base) or
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H. N. MOLDENKE, A monograph of ihe genus Baillo'nia 173merely
2-apiculate, and the shorter lip truncate or 3-apiculate oreven
3-toothed with teeth about 0.26 mm. long and 1 mm. wide atbase;
corolla hypocrateriform, zygomorphic, white, blackening indrying,
its tube cylindric, straight or slightly incurved, the abaxialside
6.3-8 mm. long, the adaxial side 5.9-7 mm. long, 1.8-2 mm. wideat
base, ampliate to 2.7-3.1 mm. at apex, glabrous outside,
glabrouswithin for the lowermost 0.5-0.7 mm., then gradually
becoming moreand more hirsutulous with hairs that become longer and
denser asthe limb is approached, its limb obliquely 5-parted,
usually about8 mm. in diameter, the lobes unequal in size,
alternate with thecalyx-teeth or -ribs, cochlear in prefloration,
venose, oblong- orelliptic-lingulate or subspatulate, rounded at
apex, pubescent onthe lower 1/4-1/3 of the inner surface
(especially along the midrib),glabrous outside, the anterior
(abaxial) lobe the largest, 3.8-5.7 mm.long, 2.6-3.3 mm. wide, the
4 remaining lobes smaller, 3.3-5.2 mm.long, 2-3.1 mm. wide; fertile
stamens 4, didynamous, opposite thecalyx-teeth or -ribs, inserted
at about the same level 2.8-3.8 mm.above the base of the
corolla-tube or the longer (posterior) pairinserted slightly higher
(2.9-4.8 mm. from the base of the corollatube), occasionally all 4
inserted near the apex of the corolla-tube,mostly included or
equaling the mouth of the tube, occasionally theupper pair (or all
4) slightly exserted; filaments filiform or flattened, glabrous,
the 2 lower ones 0.7-1.3 mm. long, the 2 upper ones1.5-1.8 mm.
long; anthers oblong or elliptic, 1-2.2 mm. long, about0.5 mm.
wide, 2-celled, dorsifixed very near the base, erect,
introrse,glabrous throughout, margined at apex, the connective not
dilated,the thecae p a r a l ~ e l , not spreading, opening by
longitudinal slits;staminode adaxial, flattened, usually inserted
at the same level as .the lower most pair of stamens, 0.5-0.6 mm.
long, glabrous, anantherous or with a very abortive sterile anther;
pistil single, included,compound and 2-carpellary, but one carpel
often aborted and thepistil then appearing to be 1-carpellary and
simple; style capillary,stoutish, 2-3.8 mm. long, lightly
puberulent or sparingly pilose withminute hairs, widened at apex;
stigma very small and capitate orflattened (about 0.7 mm. long and
0.5 mm. wide) or sometimespunctiform, mostly very shortly 2-10bed
with one anterior and oneposterior lobe, often only very obscurely
lobed; ovary superior,conic or subglobose, often slightly falcate,
1-1.5 mm. long, 1-1.8 mm.wide, glabrous or subglabrate,
incompletely 2-celled, with an anteriorparietal 1- or 2-ovulate
placenta in each cell; disk oblong, annular,
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174 DarwiMaM, t. 5, 1941hypogynous, usually not exceeding the
ovary in width, 1-1.5 mm.wide, flattened at both ends, glabrous,
glandulose; ovules ascending,hemianatropous, with a superior
chalaza and inferior micropyle;fruiting-calyx persistent,
indurated, cupuliform (sometimes onlyshallowly so), about 3 mm.
long, 6-6.5 wide, glabrous, its rim ratherdeeply 5-lobed with
rounded lobes or deeply 2-lipped with truncateor dentate lips,
sometimes cleft to. the very base; fruit drupaceous,oblong or
pyriform, 16-18 mm. long, 9-11 mm. wide, 2-lobed atapex, invested
at the base by the persistent fruiting-calyx, theexocarp glossy and
glabrous, the mesocarp fleshy, the endocarpwoody and very much
indurated, incompletely 2-pyrened; pyrenes1- or 2-seeded; seeds
oblong, ascending, glabrous, the integumentvery thin and.
membranous, without endosperm; embryo erect;cotyledons 2, elliptic,
equal or unequal, plane or contorted; radicleinferior, very short,
conic or turbinate.
This species, the type species of the genus, was based by
BocQUILLON on two collections of flUGR ALGERNON WEDDELL nQ 219'3
and32'08 from the banks of the Rio Paraguay and from dry plains
inParaguay, made in 1845 and deposited in the herbarium of
theMuseum National d'Histoire Naturelle at Paris. The types
ofKUNTZE'S so-called varieties heterophylla and longifolia were
collected by OTTO KUNTZE .at Puerto ,Esperanza, Paraguay, in
September,1892, and are deposited in the Britton Herbarium at the
New YorkBotanical Garden, the former described as 3 m. tall and the
latterat 1 m. tall. The type of Citharexylum linearifolium was
collectedby EDWARD PALMER in Paraguay during the exploration of the
Riode La Plata and adjacent countries between the years 1853
and1856, in charge of Capt. T. J. PAGE, and is deposited in the
UnitedStates National Herbarium at Washington.
BOCQUILDON regarded the ovary of B. amabilis as 1-celled and
thepartition which nearly divides it in half as a false one. Dr. H.
A.GLEASON, on examining some dissections made by the present
writer,maintains that the often falcate shape of the ovary bears
outBOCQUILWN in this view and is a sign of a 1-carpellary
condition,the incomplete cross-wall also indicating this.. I f the
ovary wereentirely symmetrical and the dissepiment were continuous,
it mightindicate a 2-carpellary condition. A definite statement,
however,as to the number of carpels actually making up the
gynoecium heremust await a careful anatomic study by a trained
morphologist.The present writer feels that JUNELL is probably
correct in assuminga 2-carpellary condition, with one of these two
carpels often aborted.
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H. N. MOLDENKE, A monograph of the genus Bailloni,a:l>
175BOCQUILlJON describes the leaves as sessile, but they are
probably
more properly regarded as being shortly and indistinctly
petiolate.He describes them as membranous, while MOORE calls them
coriaceous~ actually' they are firmly chartaceous. The
inflorescence ismostly racemiform, with distinct, although very
short, pedicelssubtending the flowers, as pointed out by MOORE in
Trans. Linn. Soc.Lond., ser. 2, 4: 437-439 (1895) and as
illustrated by BocQUILLONin his excellent figure (Adansonia 2: pI.
7. 1862), although BocQUILLON refers to it in some places in his
discussion as spicate. The4-toothed calyx and many other characters
mentioned by BENTHAMand by BRIQUET in their generic descriptions of
Baillonia, applyto the genus Diostea and not to Baillonia.MOORE, in
the reference cited above (page 438), has commentedon the strange
nature of the spines of Baillonia amabilis: Perhapsthe most
remarkable point about Baillonia is the curious nature ofits
spines. These seem to be branches which instead of separatingfrom
their mother branch, remain attached to it, except for a
splallprojecting flange at the top, upon which flange the leaf is
borne.After fall of the leaf, the flange remains permanently as a
strong,sharp, straight spine. Adhesions, possibly of branches to
theirparent branch, occur in some other Verbenaceae, e. g., in
DiosteaItself, in Symphorema, in some species of Avicennia, in
Premna,Petraea, and Citharexylum; while in V olkameria, and to some
extentin Citharexylum, spines are left after leaf-fall, as occurs
withBaillonia . The spines seen here remind one of those seen
inClerodet:tdrum aculeatum (L.) SCHLECHT., although they are notas
pronounced a.'? in that well-named plan.t.
Baillonia amabilis inhabits wet or dry sandy plains and
river-banks, ascending to 200 m. in Bolivia. I t has been collected
in
. anthesis from March to May and in July, September, October,
andDecember, and in fruit in September. The leaf-blades on
manyspecimens are badly chewed along t h ~ i r margins by some
insect pest,while the natural variation in calyx form and leaf size
is mostunique. BRIQUET in Ann. Conserv. Jard. Bot. Genev. 7-8:
316(1904) states that the genus Baillonia is not represented in
thelarge Paraguayan collection of HASSLER. Actually, however,
B.amabilis is represented in the HASSLER collection by nos. 2638
and2638 a, two sheets of the latter number being deposited in
theDielessert Herbarium at Geneva where BRIQUET worked!
BALANsAstates on the label of his no. 2089, that the species is
cultivated in
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176 Darwiniana, t. 5, 1941gardens at L'Assomption, Paraguay. In
all, 53 herbariumspecimens, including the original cotypes and also
the types of anthe synonyms, and 7 mounted photographs have been
examined bythe present writer in the preparation of this paper, the
best collections being at New York and Paris. Illustrations will be
found in(1) Adansonia 2: pI. 7.1862; (2) BocQ., Rev. Verbenac. pI.
6(1862); (3) ENGL. & PRANTL, Nat. Pflanzenfam, 4 3a: f. 58 E
& F.1895; and (4) Symb. Ant. Ups. 4: f. 89 & 90. -1934.
Citations: BRAZIL: Mattogrosso: SP. LE M. MOORE 1067 (B,Bm, P).
PARAGUAY: ANlSITS 2098 (S-2); FIEBRIG 1207 (B,Cb2,K, Mu), 1207 a
(B, Ob, K ); HASSLER 2638 (Bm, K, P), 2638 a(Bm, Cb-2, G, K, P, V)
; KUNTZE s. n. [Puerto Esperanza, IX. 92,3 m.] (B. N, N-photo, W,
Z-photo), s.n. [Po Esperanza, Sept. 1892,1 m.] (F. N, X); MALME s.
n. [3/29/1903] (S-2); EDW. PALMERS. n. [1856-6] (B-photo, K-photo,
N-fragment & 2 photos, S-photo,W, Z-photo) ; ROJAS 189'3 (B,
Mu, N) ; WEDDELL 2193 (N-2 cotypes,P-2 cotypes), 3208 (N-cotype,
P-3 cotypes). BOLIVIA: SantaCruz: KUNTZE s.n. [West-Velasco, 200
m., VII.92] (B, F, N).ARGENTINA: Formosa: Ji:iRGENSEN 1983 (N).
CULTIVATED:Paraguay: BALANSA 2089 (Cb-2, K, P).
GEOGRAPHIC D I S T R I D U T ~ O N OF Baillonia amabilisBrasil;
Mattogrosso; Paraguay: spontan. et cult.; Bolivia: San
ta Cruz; Argentina: Formosa.A LIST OF EXCLUDED SPECIES
1. Baillonia juncea (GILL. & HOOK.) BENTH. in BENTH &
HOOK. f., Gen. Plo 2:1144 (1876) apud HOOK. f. & .TA.CKS., loo.
Kew. 1: 264, 1895 (1) = DiQsteajuncea (GILL. & HOOK.) MIERS
.
2. Baillonia spicata (JACQUES) BAILL., Bull. Soc. Linn. Paris 2:
880, 1890 = Citha-rexylum ligustrinum VAN HOUTTE.
(1) This binomial is acredited by HOOKER f. and JACKSON in Ind.
Kew. 1: 264(1895) to BENTH. & HOOK. f. , but in Ind. Kew.
Suppl. 2: 23, in syn. (1904)it is corrected by Thiselton-DyeI to
BENTH. alone, in accordance with BENTRAM'S own account of the joint
and separate authorship of parts of BENTHAM& HOOKER'S <
Genera Plantarum in Journ. Linn. Soc. Lond. 20: 304-308(1884).
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H. N. MOLDENKE, A 'monograph of the genus Baillonia 177
RESUMENEl autor haee una minuciosa deseripci6n del genero de
Verbemieeas Baillonia
BOCQ. y de su unica especie B. amabilis BocQ.Reune los
antecedentes relacionados con el genero y su historia; habla de
laconfusi6n de ciertos autores al sinonimizarlo con Diostea MIERS y
de la comparaci6n que realiza MOORE de ambos generos, para
demostl'ar sus diferencias fundamentales. Indica la estrecha
relaci6n que tiene con Citharexylum. Establece que elovario es
bicarpelar, con un carpelo frecuentemente abortado y un falso
tabiqueque origina dos 16culos, segun la opini6n de JUNELL.La
distribuci6n geografica de la especie es dada con citaci6n del
material estudiado (Brasil: Matto Grosso, Paraguay, Bolivia
oriental, Argentina: Formo
sa). - N. S. T.
DARWINIANA. - T. V 12
