JOURNAL OF THE ARNOLD ARBORETUM
THE GENERA OF VALERIANACEAE AND DIPSACACEAE
IN THE SOUTHEASTERN UNITED STATES 1
I. K. FERGUSON
VALERIANACEAE Batsch, Tab. Affin. Reg. Veg. 227. 1802, nom. cons.
(VALERIAN FAMILY)
Annual or perennial herbs, sometimes woody at base. Leaves in basal
rosettes or opposite, pinnately divided or entire, exstipulate, the bases
often sheathing. Inflorescence a monochasium, thyrse, or many-flowered
compound dichasial cyme, sometimes condensed and capitate, bracteate
and bracteolate [ebracteolate]. Flowers irregular or almost regular, bi-
sexual or unisexual. Calyx obsolete or developing late and becoming con-
spicuous only in fruit, annular [or toothed], adnate to ovary. Corolla
tubular, 5 [3 or 4]-lobed, imbricate, often basally spurred or saccate
[bilabiate]. Stamens epipetalous and alternate with the corolla lobes, vary-
ing in number, usually 3 in our genera [1, 2, 3, or 4]; anthers versatile,
2- or 4-lobed, 4-locular, introrse, dehiscing longitudinally; pollen tricolpate,
echinate. Gynoecium syncarpous, ovary inferior, 3-locular, with two
locules usually suppressed and one fertile, with a solitary, pendulous, an-
atropous ovule; style 1, stigma simple or lobed. Fruit dry, indehiscent, the
calyx often developing into a winged, awned, or plumose pappus. Seed 1;
endosperm absent; embryo large, straight, the cotyledons oblong, the
radicle superior. TYPE GENUS: Valeriana L.
A family of about ten genera and 370-400 species, widely distributed
but occurring mainly in the North Temperate regions and absent from
Australasia; three genera native and one introduced in North America;
two genera in our area.
Valerianaceae are a natural family closely related to Dipsacaceae but
1 Prepared for a generic flora of the southeastern United States, a joint project of
the Arnold Arboretum and the Gray Herbarium of Harvard University made possible
through the support of George R. Cooley and the National Science Foundation and
under the direction of Carroll E. Wood, Jr., and Reed C. Rollins. This treatment
follows the pattern established in the first paper in the series (Jour. Arnold Arb. 39:
296-346. 1958) and continued through those in volumes 40-46 (1959-1965). The
area covered is bounded by and includes North Carolina, Tennessee, Arkansas, and
Louisiana. The descriptions are based primarily on the plants of this area, with any
supplementary material in brackets. References which the author has not seen are
marked by an asterisk.
The author is indebted to Dr. Wood for his aid and valuable criticisms; to Dr.
George K. Brizicky, for his guidance and suggestions; and to Mrs. Gordon W. Dillon,
for her help in the preparation of the typescript.
[VOL. 46
218


1965] FERGUSON, VALERIANACEAE AND DIPSACACEAE 219
distinguished by the 3-locular ovary (one locule fertile), cymose inflores-
cences, and seeds without endosperm. The family also has affinities with
the Caprifoliaceae and Rubiaceae.
Members of the family often have a very characteristic unpleasant odor.
Volatile oils occurring mainly in the root and rhizome have been investi-
gated extensively, especially in Valeriana.
The cytogenetics of the family have not been very fully investigated,
although chromosome numbers for some species of Valeriana (21 species
of 300) and Valerianella (11 species of 60) are available. However, only
a few chromosome counts are reported for four additional genera: Centran-
thus, 2n = 14, 32 (three species); Fedia, 2n = 32 (one species); Patrina,
2n = 22 (one species); and Astrephia, 2n = 32 (one species).
The family is of little economic importance. A few members, among
them Centranthus ruber (red valerian) and Valeriana officinalis (garden
heliotrope or common valerian), are grown as ornamentals. The latter is
used as a source of the drug "valerian." Corn salad (Valerianella Locusta).
as its name implies, is used in salads. Spikenard (Nardostachys Jatamansi)
yields an oil which has been used in perfumery in the East.
REFERENCES:
ARNAL, C. Differenciation basipete des faisceaux liberoligneux stylaires de Cen-
tranthus angustifolius DC. Compt. Rend. Acad. Sci. Paris 222: 674-676.
1946.
ASPLUND, E. Studien fiber die Entwicklungsgeschichte der Bliiten einiger
Valerianaceen. Vet.-akad. Stockholm Handl. IV. 61(3): 1-66. 1920.
BAILLON, H. Valerianacees. Hist. P1. 7: 504-518. 1879.
BENTHAM, G., & J. D. HOOKER. Valerianeae. Gen. P1. 2: 151-156. 1874.
BORSINI. O. E. Valerianaceae. Inst. Paran. Catal. Gen. 12: 1, 2. 1956.
----. Revisi6n de las Valerianaceas de Brasil. Lilloa 31: 149-170. 1962.
BUCHENAU, F. G. P. Ueber die Bliithenentwickelung einiger Dipsaceen. Vale-
rianeen und Compositen. Abh. Senckenberg. Naturf. Ges. 55: 106-132.
pls. 5, 6. 1854.
CANDOLLE, A. P. DE. Valerianeae. Prodr. 4: 623-642. 1830.
----. Mmoire sur la famille des Valerianees. 24 pp. pls. 1-5. Paris. 1832.
CHATIN, J. Etudes botaniques, chimiques et medicales sur les Valerianees. 148
pp. pls. 1-14. Paris. 1872.
DEMPSTER, L. T. Dimorphism in the fruits of Plectritis. and its taxonomic
implications. Brittonia 10: 14-28. 1958.
DUFESNE. P. Histoire naturelle et midicale de la famille de Valerianees. 61 pp.
pls. 1-3. Montpellier. 1811.
DYAL, S. C. Studies in the family Valerianaceae. Abstr. Theses Cornell Univ.
1941: 333-336. 1942.*
EWAN, J. Centranthus: a new immigrant genus to California. Lead. West. Bot.
2: 195. 1939.
GRAEBNER, P. Die Gattungen der natiirlichen Familie der Valerianaceae. Bot.
Jahrb. 37: 464-480. 1906. [See also ibid. 436-451.]
HOCK, F. Beitraige zur Morphologie, Gruppirung und Geographischen Verbrei-
tung der Valerianaceen. Bot. Jahrb. 3: 1-73. 1882.
-. Valerianaceae. Nat. Pflanzenfam. IV. 4: 172-182. 1891.


JOURNAL OF THE ARNOLD ARBORETUM
Verwandtschaftsbeziehungen der Valerianaceen und Dipsacaceen. Bot.
Jahrb. 31: 405-411. 1902.
KILLIP, E. P. Valerianaceae. In: J. F. MACBRIDE, Flora of Peru. Publ. Field
Mus. Bot. 13(6): 287-321. 1937.
KROK, T. O. B. N. Valerianaceae. In: E. WARMING, Symbolae ad floram Brasiliae
27. Vid. Medd. Naturh. For. Kj0benh. 34: 139, 140. 1883.
LARSEN, K. Experimental and cytological studies in Centranthus. Bot. Not. 111:
301-305. 1958.
NIELSEN, S. D. Systematic studies in the Valerianaceae. Am. Midi. Nat. 42:
480-499. 1949. [Plectritis.]
PIZZOLONGo, P. Ricerche sulla cariologia del genere Centranthus e loro impor-
tanza tassonomica. Delpinoa II. 1: 149-164. pls. 1, 2. 1959.
POUCQUES, M. L. Recherches caryologiques sur les Rubiales. Revue Gen. Bot.
56: 97-138. 1949. [Le noyau des Valerianacees, 101-109; Fedia, Vale-
rianella, Valeriana, Centranthus.]
RENDLE, A. B. The classification of flowering plants, vol. 2. Dicotyledons. 636
pp. Cambridge, England. 1925. [Valerianaceae, 567-572.]
SCHERMERHORN, J. W., & M. W. QUIMBY, eds. Lynn Index. Monograph V.
67 pp. Boston. 1962. [Valerianaceae, 62-67.]
SoUEGES, R. Embryogenie des Valerianacees. Developpement de l'embryon
chez les Centranthus. Compt. Rend. Acad. Sci. Paris 212: 718-720. 1941.
VESQUE, J. Developpement du sac embryonnaire des phanerogames angio-
spermes. Ann. Sci. Nat. Bot. VI. 6: 237-285. pls. 1-8. 1878. [Valerianella
pumila, Centranthus ruber, 255, 256.]
VIDAL, L. Contribution a l'anatomie des Valerianacees. Ann. Univ. Grenoble
15: 561-605. 1903.*
WAGENITZ, G. Pollenmorphologie der mitteleuropaischen Valerianaceen. Flora
143: 473-485. pl. 5. 1956. [Valeriana, Valerianella, Centranthus.]
Die systematische Stellung der Rubiaceae. Bot. Jahrb. 79: 17-35.
1959. [Gentianales = Contortae + Rubiaceae; Dipsacales = Caprifolia-
ceae, Adoxaceae, Valerianaceae, Dipsacaceae.]
SValerianaceae. In: H. MELCHIOR, Engler's Syllabus der Pflanzen-
familien. ed. 12. 2: 475-477. 1964.
WEBERLING, F. Die Infloreszenzen der Valerianaceen und ihre systematische
Bedeutung. Akad. Wiss. Lit. Abh. Math.-Naturw. Mainz 1961: 151-281.
1961.
KEY TO THE GENERA OF VALERIANACEAE
Calyx inrolled in flower, developing in fruit to form a conspicuous pappus with
5-15 plumose awns; fruit 1-locular; plant perennial. ........ 1. Valeriana.
Calyx obsolete; fruit 3-locular; plant annual or biennial. .... 2. Valerianella.
1. Valeriana Linnaeus, Sp. P1. 1: 31. 1753; Gen. P1. ed. 5. 19. 1754.
Perennial herbs or vines with thickened, strong-scented, fleshy [woody
or tuberous] roots, sometimes stoloniferous; glabrous or sparsely hairy
with short, simple hairs. Leaves petiolate to nearly sessile, basal and
cauline, undivided, pinnate, pinnatifid, or ternate. Inflorescence usually
a dichasial cyme or thyrse, terminal or axillary, sometimes lax or sub-
capitate, bracteate and bracteolate. Flowers irregular, bisexual or uni-
[VOL. 46


1965] FERGUSON, VALERIANACEAE AND DIPSACACEAE 221
sexual. Calyx inrolled in flower, enlarging and developing in fruit to form
a pappus with short, sessile, patelliform, membranaceous limb and 5-15
plumose awns [or the limb cupuliform and irregularly dentate). Corolla
funnelform or campanulate [salverform or rotate], the tube slightly sac-
cate at the base [or straight], usually more or less hairy in the throat,
5 [3 or 4]-lobed. Stamens 3 [rarely 4], inserted toward the top of the
corolla tube, usually exserted (but included in V. scandens); anthers 2-
or 4-lobed. Stigma simple or 3-lobed; ovary 3-locular, with two locules
suppressed and one fertile with a solitary ovule. Fruit a unilocular, com-
pressed achene with 3 nerves on the outer and 1 on the inner surface.
LECTOTYPE SPECIES: Valeriana pyrenaica L.: see Britton & Brown, Illus.
Fl. No. U. S. ed. 2. 3: 284. 1913. (Name Medieval Latin, from valere,
to be strong; said also to be named for the Roman emperor Valerianus).
A genus of about 300 species of Europe, Asia, Africa. and America,
centered mainly in the temperate regions of the Northern Hemisphere and
the mountains of South America: eleven species in the United States. two
in our area.
Of the seven sections recognized by Hock, six (Pseudiastrephia H6ck,
Hybocarpus Hock, Valerianopsis Wedd., Phyllactis Pers., Porteria Hook..
and Aretiastrum DC.) are entirely South American and have been various-
ly treated by different authors as distinct genera or as sections within
Valeriana. The North American species belong to sect. VALERIANA
(� Euvaleriana Hick). Different authors have recognized a varying
number of series within this section (cf. Hock, Meyer).
The genus is well defined by the combination of perennial habit, three
stamens, and inrolled calyx usually enlarging and forming a pappus on
the one-locular fruit. It is probably most closely related to Valerianella
and to Centranthus, which is distinguished by one stamen and spurred
corolla.
Valeriana scandens L., readily recognized by its climbing habit, occurs
in Mexico, Central and South America, and the West Indies. northward in
peninsular Florida to a shell-mound in Duval County. It seems likely that
this species is a calciphile, but it does not appear to occur on the
Florida Keys. It is placed by Meyer in the Sorbifoliae, a series distin-
guished chiefly by included stamens and two-lobed anthers. Meyer recog-
nizes two varieties (which some authors have treated as distinct species):
var. scandens, with tripartite leaves, and var. Candolleana (Gard.) Muell..
with entire leaves. The widespread var. scandens occurs in Florida. but
var. Candolleana occurs northward only to Hispaniola and Cuba. Inter-
mediate specimens with both entire and tripartite leaves have been found.
however, both in our area and elsewhere. Valeriana pauciflora Michx..
distinguished by its long, funnelform corolla tube and stoloniferous habit.
occurs from Pennsylvania to Illinois, Tennessee, and Virginia. It is placed
in the series Officinales Hick, separated by the rhizomatous or stolonifer-
ous roots and four-lobed anthers.


JOURNAL OF THE ARNOLD ARBORETUM
Sexual polymorphism occurs in the genus in several ways, the plants
being completely hermaphrodite, polygamous, polygamodioecious, or di-
oecious, with dioecism least common. The presence of unisexual as well as
bisexual flowers is easily detected by the differential lengths of the corolla.
The mode of pollination appears to be unknown but is probably ento-
mophilous. Nectaries are reported to occur in the saccate base of the
corolla tube. The occurrence of proterandry assures cross-pollination in
bisexual flowers.
The cytogenetics of Valeriana oficinalis, 2n = 14, 28, 56, has been
extensively investigated, but the cytology of the genus as a whole is not
well known. Chromosome numbers of 2n = 14, 16, 18, 28, 32, 56, and
64 have been reported.
REFERENCES:
Under family references see POUCQUES, WAGENITZ (1956), and WEBERLING.
ARTIUSHENKO, Z. T., & I. N. KONOVALOV. Morphology of fruits belonging to
the nut and nutlet types. (In Russian.) Acta Inst. Bot. Acad. Sci. URSS.
7. Morphol. Anat. P1. 2: 170-192. 1951. [Valeriana fruit a dry syncarpous
drupe, 185, 186, 188.]
BORSINI, O. E. Addenda a las Valerianaceas argentinas I. Lilloa 12: 23-27.
1946. [Races of V. scandens.]
- Valerianaceas del estado de Santa Catarina (Brasil). Sellowia 15: 123-
136. 1963.
ELZENGA, G. Investigation of viability and germination rate of Atropa bella-
donna, Digitalis purpurea, Digitalis lanata, Valeriana officinalis and Viola
tricolor in connection with different seed treatments. (In Dutch.) Herba
11: 73-78. 1952.*
FAUCONNET, L. Variations saisonnieres chez Valeriana officinalis L. (German
summary.) Bull. Soc. Bot. Suisse 57: 122-131. 1947.
GSTIRNER, F. Evaluation of valerian root. Planta Med. 6: 81-85. 1958.*
HAUSCHILD, F. Die Problematik der sedativen Baldrianwirkung. Pharmazie
13: 420-422. 1958.*
HELWIG, B. Uber die Frage der Heterorhizie bei Radix Valerianae oficinalis.
Ber. Deutsch. Bot. Ges. 46: 595-609. 1928.
IRMIsCH, T. Beitrag zur Naturgeschichte der einheimischen Valeriana-Arten
insbesondere der Valeriana officinalis und dioica. Abh. Naturf. Ges. Halle
1: 19-44. pls. 1-4. 1854.
KATINA, Z. F. Anatomical data on the localization of essential oil in some
Valeriana species. (In Ukrainian; Russian summary.) Bot. Zhur. Kiev
10: 81-86. 1953.
KREPINSKY, J., V. HEROUT, & F. SORM. Plant substances. VI. The isolation of
neutral products from the root of valerian (Valeriana officinalis L.). Collect.
Czech. Chem. Commun. 24: 1884-1896. 1959.*
KUHNHOLTZ-LORDAT, G. Contribution a la biologie de Valeriana tripteris L.
Feuille Nat. II. 3: 101. 1948.*
LATTANZI, V. Valeriana officinalis L., substitutes and adulterations. (In Italian.)
Fitoterapia II. 26: 590-605. 1955.*
LAWALREE, A. Le groupe du Valeriana officinalis L. en Belgique. Bull. Jard.
Bot. Bruxelles 22: 193-200. 1952.
[VOL. 46
222


1965] FERGUSON, VALERIANACEAE AND DIPSACACEAE 223
MEIJERS, T. Een onderzoek van het linneon Valeriana officinalis L. in Nederland.
122 pp. 23 pls. 's Gravenhage. 1957.
. Methods to determine different cytotypes of Valeriana oficinalis L. s.
lato. (In Dutch; English summary.) Pharm. Weekbl. 95(9): 269-278.
1960.*
MEYER, F. G. Valeriana in North America and the West Indies (Valerianaceae).
Ann. Missouri Bot. Gard. 38: 377-503. 1951. [An extensive taxonomic
revision.]
. The genus Valeriana in East Tropical and South Africa. Jour. Linn.
Soc. Bot. 55: 761-771. 1958. [Distribution and taxonomy.]
PHILIPSON, W. R. Studies in the development of the inflorescence. III. The
thyrse of Valeriana officinalis L. Ann. Bot. II. 11: 409-416. pl. 3. 1947.
PITTIER, H., & E. P. KILLIP. Venezuelan species of Valeriana. section Porteria.
Jour. Wash. Acad. Sci. 16: 422-428. 1926.
PROTASSENJA. G. D. Zur Frage des karvologischen Unterschieds einiger von
Valeriana officinalis L. abgesonderter Baldrianarten. (In Russian.) Arbeit.
Bot. Kab. Centr. Moorversuchsstat. Minsk 1: 83-92. 1930.* [See German
summary in Bot. Centralbl. 162: 52. 1932.]
RUNQUIST. E. Zytologische und morphologische Ialeriana-Untersuchungen.
Hereditas 23: 279-286. 1937.
RUSSELL, N. H. Quantitative studies in angiosperm taxonomy. X. Valeriana.
XI. Geranium. XII. Mimulus. Castanea 29: 138-150. 1965.
SKALINSKA, M. Polyploidy in Valeriana officinalis Linn. in relation to its ecology
and distribution. Jour. Linn. Soc. Bot. 53: 159-186. 1947.
.Studies in cyto-ecology, geographic distribution and evolution of
Valeriana L. Bull. Acad. Polon. Sci. Lett. Sci. Math. Nat. B. I. 1950: 149-
175. pls. 3-5. 1951.
-- -. Meiosis in a polyhaploid twin plant and a hexaploid hybrid of I'aleri-
ana sambucifolia Mikan. Acta Soc. Bot. Polon. 23: 359-374. pl. 1. 1954.
SCytological studies in a chromosomal aberrant of laleriana and its
derivatives. Polsk. Akad. Nauk. Folia Biol. 10: 155-168. pls. 1, 2. 1962.
SPRAGUE. T. A. The British forms of Valeriana officinalis. Watsonia 2: 145-147.
1952.
STROH, G. Valeriana L. Provisorische Liste der alterweltlichen Arten. Repert.
Sp. Nov. 40: 225-233. 1936.
TODD, B. H. The cytology of the Valerianaceae with special reference to the
status of the British forms of Valeriana officinalis. Thesis, Univ. Durham.
1941.*
TROITSKY. N. Resultats des recherches morphologiques et biologiques sur le
Valeriana alliariaefolia Vahl du Caucase. (In Russian; French summary.)
Moniteur Jard. Bot. Tiflis 46/47: 77-100, (101). 1919.
VOROSKILOV, V. N. Medicinal Valeriana. 160 pp. Moscow. 1959.
WALTHER, E. Zur Morphologie und Systematik des Arzneibaldrians in Mittel-
europa. Mitt. Thiir. Bot. Ges. Beih. 1: 7-108. 1949. [V. oficinalis.]
2. Valerianella Miller, Gard. Dict. Abr. ed. 4. 1754.
Small annual or biennial, glabrous or pubescent herbs with dichasial
branching, appearing dichotomous, but the terminal bud aborting in the
lower branches. Basal leaves forming a rosette, petiolate to nearly sessile,
undivided or dentate; cauline leaves connate, undivided, dentate [or rarely


JOURNAL OF THE ARNOLD ARBORETUM
incised-pinnatifid]. Inflorescence capitate, terminal, usually a dichasial
cyme subtended by lanceolate to oblong, connate bracts. Flowers almost
regular, bisexual [or unisexual] in the axils of lanceolate to oblong, con-
nate bracteoles. Calyx obsolete [or forming a narrow, toothed rim]. Corol-
la funnelform, narrowly campanulate or tubular, the tube slightly saccate
at the base, 5-lobed. Stamens 3, inserted toward the top of the corolla
tube, exserted, anthers 4-lobed. Stigma simple or shortly 3-lobed; ovary
3-locular, one locule fertile with a solitary ovule. Fruit glabrous or pubes-
cent, 3-locular, two of the locules empty, the other 1-seeded. LECTOTYPE
SPECIES: Valeriana Locusta L. = Valerianella Locusta (L.) Betcke; see
Britton & Brown, Illus. Fl. No. U. S. ed. 2. 3: 286. 1913. (Name a diminu-
tive of Valeriana.)
A genus of temperate parts of the Northern Hemisphere, including about
60 species in two sections; 19 in North America, ten in our area.
The shape, relative size of the sterile locules, and indumentum of the
fruits are the most important characters used to distinguish the species.
The indumentum of the plant and the shape of the bracts (rounded,
pointed, or acuminate) are also important characters. Specimens with
both fruit and flowers are necessary for the determination of species.
Four species of sect. SIPHONELLA Krok, marked by salverform corollas
and strongly glandular fimbriate-serrulate bracts, occur in the southeastern
United States. Valerianella Bushii Dyal and V. ozarkana Dyal are strictly
calciphile and known only from Arkansas and Missouri, while V. Nuttallii
(Torr. & Gray) Walp. and V. longiflora (Torr. & Gray) Walp. are known
only from western Arkansas and eastern Oklahoma. The other six species
of our area belong to sect. VALERIANELLA, with funnelform corolla and
glabrous bracts with entire or ciliate margins. Valerianella Locusta (L.)
Betcke (V. olitoria (L.) Poll., Valeriana Locusta L. a olitoria L.), 2n =
14, an introduced European species widely scattered throughout our area,
is readily recognized by the corky mass on the fertile locule of the fruit.
Valerianella umbilicata (Sulliv.) Wood and V. patellaria (Sulliv.) Wood
occur from North Carolina and Tennessee, northward to New York, Ohio,
and Illinois. They are distinguished from the other species in the section
by the breadth of the sterile locules in comparison with the fertile locule
of the fruit. Valerianella intermedia Dyal, which ranges from Massachu-
setts to Kentucky and Illinois, occurs in scattered localities in North and
South Carolina, apparently as an introduction. It is closely related to
V. radiata (L.) Dufr., a widespread, polymorphic species occurring
throughout our area and extending to Pennsylvania, Kansas, and Texas.
Valerianella Palmeri Dyal, occurring only in Arkansas, appears to be
closely related to V. intermedia but is separated by its longer fruit with
abortive sterile locules.
Some authors have thrown doubt on the validity of some of the species
recognized in North America. Dempster has found dimorphism in the
fruits of species of the closely related Plectritis and has revised the taxono-
[VOL. 46


1965] FERGUSON, VALERIANACEAE AND DIPSACACEAE 225
my of that genus. In view of this, the species pairs Valerianella Bushii-
V. ozarkana and V. umbilicata-V. patellaria need further investigation,
for in each pair the two sometimes grow together, show a similar, limited
geographical distribution, and are distinguished by their fruits.
There is a lack of cytological information relating to North American
species of Valerianella. Chromosome numbers of 2n = 14, 16, 18, 32 have
been reported for European and Asiatic species. The method of pollination
appears to be unknown; insect visitors have been observed and self-pollina-
tion reported to occur.
Valerianella has affinities with Valeriana. It is very closely related to
Fedia and Plectritis, and early authors treated these variously as one or
more genera. More recent authors have treated Fedia as a monotypic
Mediterranean genus distinguished by its irregular corolla, two stamens,
and dimorphic fruits. Valerianella is separated from the predominantly
western North American Plectritis by the typically 3-locular ovary, 3-
lobed stigma, and dichotomously branched inflorescence vs. the 1-locular
ovary, 2-lobed stigma, and capitate or interruptedly spicate inflorescence.
REFERENCES:
Under family references see DEMPSTER, POUCQUES, VESQUE. \VAGENITZ
(1956), and WEBERLING.
BARTHOLEMEW, E. A. Three spermatophytes new to West Virginia. Castanea
5: 111. 1940. [V. olitoria (= V. Locusta), V. umbilicata.]
DYAL, S. C. Valerianella in North America. Rhodora 40: 185-212. 1938. [See
also ibid. 465-467.]
ELVERS, I. Chromosomenzahlen in der Gattung Valerianella nebst einigen sys-
tematischen Bemerkungen. Acta Horti Berg. 11: 81-87. 1932.
FREEMAN, O. M. Notes on the flora of Polk County, North Carolina. Castanea
20: 37-57. 1955. [V. intermedia as a field and roadside weed.]
GARJEANNE, A. J. M. Valerianella. (In Dutch.) Levende Nat. 53: 101-107.
1950.*
GOODMAN, G. J. Notes on Oklahoma plants. Proc. Okla. Acad. Sci. 32: 39,
40. 1952. [V. ozarkana recorded in Oklahoma.]
GRAVES, C. B. Valerianella in New England. Rhodora 4: 195, 196. 1902.
GRAY, A. Contributions to North American botany. Proc. Am. Acad. Arts Sci.
19: 1-96. 1883. [Valerianaceae-Valerianella, 81-83.]
KROK, T. O. B. N. Anteckningar till en Monografi ifver Viixlfamiljen Valeri-
aneae. 1. Valerianella. Vet.-akad. Stockholm Handl. IV. 5: 1-105. pls. 1-4.
1864.
PORTER, T. C. The Fedias of the northern United States. Am. Nat. 6: 385-388.
1872. [Valerianella.]
SHINNERS, L. H. Valerianella florifera Shinners, sp. nov. Field Lab. 21: 93.
1953. [Described from Texas.]
SOUEGES, R. Embryogenie des Valerianacees. Developpement de lembryon
chez le Valerianella olitoria Poll. Compt. Rend. Acad. Sci. Paris 176: 1081-
1083. 1923. [V. Locusta.]
SOYER-WILLEMET, H. F. Essai monographiques sur les Valerianella de France.
Precis Trav. Soc. Sci. Lett. Arts Nancy 1829-1832: 67-73. 288. 1833.


JOURNAL OF THE ARNOLD ARBORETUM
DIPSACACEAE A. L. de Jussieu, Gen. P1. 194. 1789, "Dipsaceae," nom. cons.
(TEASEL FAMILY)
Mostly perennial herbs, the flowers bisexual, irregular, with a calyx-
like involucel of fused bracteoles, and borne in the axils of imbricate recep-
tacular bracts, the calyx small [variable], the corolla gamopetalous, 4[5]-
lobed, the stamens 4 15], inserted toward the top of the corolla tube, and
the ovary inferior. TYPE GENUS: Dipsacus L.
An Old World family of about eight to twelve genera and 250-300
species, centered in the Mediterranean and Near East, extending to North
Europe, East Asia, and South Africa. About ten species in the genera
Dipsacus, Cephalaria, Scabiosa, Knautia, and Succisa are naturalized in
North America; one species of Dipsacus occurs in our area.
Dipsacaceae are placed in the Rubiales by most authors and appear
to be a very natural family, with the exception of the isolated genus
Morina, which some authors have placed in a separate family. Dipsacaceae
have close affinities with Valerianaceae. The family is distinguished from
Valerianaceae by the unilocular ovary and by the capitate inflorescence
surrounded by an involucre of bracts, characters which suggest affinities
with Compositae.
Chromosome numbers of 2n = 10, 14, 16, 18, 20, 34, 36, 38, 42. 43,
44, 46, 50, 54, and 64 have been reported. Ehrendorfer has suggested that
the basic chromosome number of the family is x = 9 and that euploidy,
aneuploidy on the diploid level, polyploidy followed by aneuploidy, gene-,
chromosome-, and genome-mutations have been important in the evolution
of Dipsacaceae.
The variation in the receptacular bracts and the diverse development of
the involucel and calyx are connected with fruit dispersal and give good
characters for taxonomic division of the family into genera.
The family is of little economic importance. A few genera are of horti-
cultural interest, and Scabiosa is widely cultivated for its ornamental
value.
REFERENCES:
ALVARADO. S. Constituci6n morfol6gica y filagenia del caliculo de las "dipsaci-
ceas." Trab. Mus. Ci. Nat. Bot. Madrid 21: 1-29. 1925. [Scabiosa,
Morina.]
---- . Der morphologische Aufbau des Hiillkelches der Dipsacaceen. Bot.
Jahrb. 61(Beibl. 138): 10-21. 1927.
BAILLON, H. Dipsacaces. Hist. P1. 7: 519-534. 1880.
BAKSAY, L. Monographie der Gattung Succisa. Ann. Hist.-Nat. Mus. Hungar.
II. 2: 237-260. 1952.
----. Anatomische und systematische Untersuchungen iiber die Gattung
Succisella. Ibid. 6: 167-176. 1955.
BENTHAM, G., & J. D. HOOKER. Dipsacaceae. Gen. P1. 2: 157-161. 1873.
CANDOLLE, A. P. DE. Dipsacaceae. Prodr. 4: 643-664. 1830.
CLAUSEN, R. T. Scabiosa columbaria in central New York. Rhodora 45: 220,
221. 1943.
[VOL. 46


1965] FERGUSON. VALERIANACEAE AND DIPSACACEAE 227
COULTER, T. Memoire sur les Dipsacees. Mim. Soc. Phys. Hist. Nat. Geneve
2(2): 13-60. pls. 1, 2. 1826.
DOLL, W. Beitrage zur Kenntnis der Dipsaceen und Dipsaceenihnlicher Pflan-
zen. Bot. Arch. 17: 107-146. 1927.
EHRENDORFER, F. Beitrage zur Phylogenie der Gattung Knautia (Dipsacaceae)
I. Cytologische Grundlagen und allgemeine Hinweise. Osterr. Bot. Zeitschr.
109: 276-343. 1962. [An extensive study of the genus in Europe.]
Cytologie, Taxonomie und Evolution bei Samenpflanzen. In: W. B.
TURRILL, ed., Vistas in botany 4: 99-186. New York. 1964. [Darstellung
einiger spezieller Probleme am Beispiel der Dipsacaceae. 121-124. Rela-
tionships and evolution of the genera of Dipsacaceae.]
FISCHER, J. Beitrige zur Systematik der Dipsaceen. Lotos Sitz-ber. Naturw.-
Med. Ver. Bohmen II. 26: 78-102. 1906.
FODOR, F. Beitriige zur Kenntnis der Histologie der Gattung Cephalaria. (In
Hungarian; German summary.) Bot. Kozlem. 9: 171-199. (45)-(46).
1910. [For leaf anatomy see F. VARGA, Bot. Kozlem. 22: 28-34. (17).
1925.]
FRANCOIS, L. Semences et jeunes plantes chez les Dipsacees et les Composees.
Ann. Agron. III. 17: 674-718. 1947.*
GUNTHART, A. Beitrage zur Blijtenbiologie der Dipsaceen. Flora 93: 199-250.
1904. [Scabiosa, Knautia, Succisa, Cephalaria, Dipsacus.]
HERR, C. Contribution a l'etude du pollen des Dipsacees. These Doct. Univ.
(Pharmacie) Strasbourg. 1934.*
HiCK, F. Dipsacaceae. Nat. Pflanzenfam. IV. 4: 182-189. 1891.
.Verwandtschaftsbeziehungen der Valerianaceen und Dipsacaceen. Bot.
Jahrb. 31: 405-411. 1902.
JAEGER, P. Nouveaux cas de gynodimorphisme chez les Dipsacees. Compt. Rend.
Acad. Sci. Paris 199: 1239-1241. 1934. [Knautia. Scabiosa. Succisa. Cepha-
laria, Dipsacus; for Succisa see P. MAGNUS. Bot. Jahrb. 3: 187. 1882.]
-------. ~tude de la sexualite chez les Dipsacacees. Imprimerie Arts graphiques
modernes. Nancy. 1937.*
KACHIDZE, N. Karyologische Studien uiber die Familie der Dipsacaceae. Planta
7: 482-502. 1929.
LAVIALLE, P. Sur le sac embryonnaire des Dipsacees. Compt. Rend. Acad. Sci.
Paris 180: 1127-1129. 1925. [See also ibid. 1606-1608.]
L'etamine chez Knautia arvensis Coult. Polymorphisme des fleurs et
des capitules. Ibid. 192: 176-178. 1931.
---- La g6neration haploide femelle chez les Dipsacees. Bull. Soc. Bot.
France 83: 274-277. 1936.
L'androcee et la generation haploide male chez les Dipsacees. Ibid.
562-570.
----- & P. JAEGER. L'origine du fruit dans ses rapports avec la pollinisation
chez Knautia arvensis Coult. Compt. Rend. Acad. Sci. Paris 192: 1474-
1476. 1931.
----- & . La fertilite et la sterilite de l'androcee: leurs rapports avec
le polymorphisme staminal chez Knautia arvensis Coult. Ibid. 198: 114-
116. 1934. [See also cytology of pollen, ibid. 199: 485-487. 1934.]
---- & G. KLEIN. Pistils bi-ovules et soudure des deux ovules chez Knautia
arvensis Coult. Bull. Soc. Bot. France 77: 593-597. 1930.
LUNT, J. R. Succisa pratensis in Massachusetts. Rhodora 14: 174. 1912.
PHILIPSON, W. R. Studies in the development of the inflorescence. II. The


JOURNAL OF THE ARNOLD ARBORETUM
capitula of Succisa pratensis Moench. and Dipsacus fullonum L. Ann. Bot.
II. 11: 285-297. 1947. [D. sativus.]
PLOUVIER, V. Sur la recherche du methylglucoside p chez quelques Dipsacacees
et Rhamnacees. Compt. Rend. Acad. Sci. Paris 256: 1397-1399. 1963.
PODDUBNAJA-ARNOLDI, V. Spermazellen in der Familie der Dipsacaceae. Planta
21: 381-386. 1933. [Scabiosa.]
RAZI, B. A., & K. SUBRAMANYAM. Embryology of the Dipsacaceae. Proc. Indian
Acad. Sci. B. 36: 249-257. 1952. [Cephalaria.]
RISSE, K. Beitrage zur Zytologie der Dipsacaceen. Bot. Arch. 23: 266-288.
1928.
SCHERMERHORN, J. W., & M. W. QUIMBY, eds. Lynn Index. Monograph V.
67 pp. Boston. 1962. [Dipsacaceae, 20-22.]
SCHNARF, K. Vergleichende Embryologie der Angiospermen. 350 pp. Berlin.
1931. [Dipsacaceae, 209, 210.]
SOUEGES, R. Embryogenie des Dipsacacees. Developpement de l'embryon chez
le Scabiosa columbaria L. Compt. Rend. Acad. Sci. Paris 245: 465-468.
1957.
---- Embryogenie des Dipsacacees. Developpement de l'embryon chez le
Cephalaria tatarica Schrad. Ibid. 256: 45-48. 1963. [Knautia, ibid. 1190-
1194.]
SZAB6, Z. Nouvelles observations concernant l'histologie et le developpement
des organes sur les especes du genre Knautia. (In Hungarian; French sum-
mary.) Bot. Kozlem. 9: 133-148, (25)-(41). 1910.
- A Knautia genusz MonografiAja. Mat. Termeszett. Kozlem. 31. 1911.*
----. The development of the flower of the Dipsacaceae. Ann. Bot. 37: 325-
334. 1923.
--- . Organografiai es genetikai vizsgalatok dipsacaceakon (Organographische
und genetische Untersuchungen an Dipsacaceen). Magyar Tudom. Akad.
Ertek. Termeszett. K6reb. 53: 571-636. 1935.*
SMonographie der Gattung Cephalaria. Mat. Termeszett. Kozlem. 38.
1940.*
TAMMES, T. Notiz ilber das Vorkommen von Dipsacan bei den Dipsaceae. Rec.
Tray. Bot. Neerl. 8: 369, 370. 1911.
TIEGHEM, P. VAN. Remarques sur les Dipsacacees. Ann. Sci. Nat. Bot. IX. 10:
148-200. 1909.
VARGA, F. Vergleichende anatomische Untersuchung der Gattungen Succisella
und Succisa mit Rucksicht auf die verwandten Gattungen. (In Hungarian;
German summary.) Bot. Kozlem. 21: 32-47, (4)-(8). 1923.
VIETH, J. Beitrag zur Kenntnis Dipsacaceenbliite. Ann. Univ. Sarav. Sci. 7:
215-274. 1959.
WAGENITZ, G. Die systematische Stellung der Rubiaceae. Bot. Jahrb. 79: 17-35.
1959. [Gentianales = Contortae + Rubiaceae; Dipsacales = Caprifolia-
ceae, Adoxaceae, Valerianaceae, Dipsacaceae.]
----. ipsacaceae. In: H. MELCHIOR, Engler's Syllabus der Pflanzenfamilien.
ed. 12. 2: 477, 478. 1964.
1. Dipsacus Linnaeus, Sp. P1. 1: 97. 1753; Gen. P1. ed. 5. 43. 1754.
Stout, erect, biennial herbs with spiny or prickly stems. Leaves simple,
entire [to pinnately divided], opposite, exstipulate; basal leaves petiolate,
stem leaves connate [petiolate]. Inflorescence a head with 1 or 2 rows of
[VOL. 46


1965] FERGUSON. VALERIANACEAE AND DIPSACACEAE 229
linear to lanceolate [narrowly triangular], subulate, erect or speading,
spine-tipped involucral bracts; each flower subtended by a chaffy recep-
tacular bract and surrounded by a 4-angled tubular involucel of fused
bracteoles, which is grooved with a median rib and truncate at the apex,
with an almost obsolete denticulate margin. Calyx cup-shaped, 4-angled.
with a ciliate margin. Corolla imbricate, lobes often unequal. Stamens 4.
alternate with the corolla lobes, exserted; filaments free; anthers 4-locular,
introrse, longitudinally dehiscent; pollen tricolpate, echinate. Stigma lat-
eral, entire; style filiform; ovary unilocular, ovule solitary, anatropous.
pendulous from the apex of the locule. Fruit indehiscent, dry, inclosed
within the calyx-like involucel and crowned with the persistent calyx. Seed
1; endosperm fleshy; embryo straight, the cotyledons oblong or ovate, the
radicle superior. Embryo sac development of the normal (Polygonum)
type. LECTOTYPE SPECIES: D. fullonum L.; see Britton & Brown. Illus.
Fl. No. U. S. ed. 2. 3: 289. 1913. (Name from Greek, dipsa, thirst, be-
cause the united cup-shaped bases of the leaves in some species hold
water.) - TEASEL.
About 12 species of Europe, western Asia, and Africa, three naturalized
in North America. Dipsacus fullonum L. (D. sylvestris Huds.),2 2n = 16,
18, a native of Europe, is probably widespread and scattered throughout
the United States, occurring sporadically in our area. Dipsacus fullonum
has close affinities with D. sativus (L.) Schkuhr (D. fullonum auct. non
L.), 2n = 16, 18, also a native of Europe, locally naturalized in the north-
eastern United States. The involucral bracts curve upward in D. fullonum
and the receptacular bracts end in a long straight spine, while in D.
sativus the involucral bracts spread more or less horizontally and the
receptacular bracts end in a stiff, recurved spine. Some authors have
treated D. sativus as a subspecies of D. fullonum. The very different
laciniate-leaved D. laciniatus L. is established in the northeastern United
States.
The genus is mainly a well-defined one recognized by the presence of
stem prickles and spine-tipped bracts. It is most closely related to Cepha-
laria, which is also distinguished from the rest of the family by chaffy
receptacular bracts and a four-angled involucel. Grenier and Godron
placed D. pilosus L. in Cephalaria, but few subsequent authors have
adopted this treatment.
The mode of flowering in the genus is characteristic, the flowers opening
first about halfway up the head and developing in sequence both upward
and downward.
The species are separated on the basis of the leaves, which may be con-
nate or stalked, entire or divided; the inflorescence, which may be oblong.
ovate, or globose; and the involucral bracts, which may be linear-lanceo-
late, subulate, keeled with spines on the margins and midrib, or narrowly
triangular, shorter or longer than the inflorescence.
2 The application of the name Dipsacus fullonum L. will be discussed in the next
issue of this Journal.


JOURNAL OF THE ARNOLD ARBORETUM
The ripened inflorescences of D. sativus, the fuller's teasel, are used in
fulling cloth (raising the nap), and the plants are grown to a limited ex-
tent for this purpose.
The flora and fauna of the rain water caught by the connate leaf bases
have been studied more or less extensively, but most biological aspects
of the genus have been neglected.
Chromosome numbers of 2n = 16, 18, 36 have been reported.
REFERENCES:
Under family references see GUNTHART, JAEGER (1934), and PHILIPSON.
BEAL, W. J., & C. E. ST. JOHN. A study of Silphium perfoliatum and Dipsacus
laciniatus in regard to insects. Bot. Gaz. 12: 268-270. 1887.
CHRISTY, M. The common teasel as a carnivorous plant. Jour. Bot. 61: 33-45.
1923. [Observations on whether the water held in the connate leaf bases
of D. fullonum operates as an insectivorous mechanism.1
GREENE, E. L. Nomenclature of the fuller's teasel. Pittonia 3: 1-9. 1896.
GRENIER, M., & M. GODRON. Dipsacaceae. Flore de France 2: 67-81. 1850.
[D. pilosus transferred to Cephalaria, 69. ]
GROSSIN, F. Virescence spontanee chez Dipsacus silvestris Mill. et virescence
induite experimentalement chez Zinnia elegans L. Compt. Rend. Acad. Sci.
Paris 242: 1349-1352. 1956. [Experiment with 2,4-D on D. fullonum.]
HOWELL, J. T. Dipsacus sylvestris Huds. Leafl. West. Bot. 2: 16. 1937. [Oc-
currence of D. fullonum in western U. S.]
JURICA, H. S. Development of head and flower of Dipsacus sylvestris. Bot. Gaz.
71: 138-145. 1921. [D. fullonum.]
MAGUIRE, B. Aquatic biotas of teasel (Dipsacus sylvestris) waters. Ecology
40: 506. 1959. [Algae, protozoa, nematodes, and other organisms found in
the water of the connate leaf bases of D. fullonum.]
MULLINS, D. Teasel growing, an ancient practice. World Crops 3: 146, 147.
1951.* [D. sativus as D. fullonum.]
PLANTEFOL, L. La soudure des feuilles dans la torsion de contrainte de la
cardere (Dipsacus silvestris Huds.). Compt. Rend. Acad. Sci. Paris 256:
3917-3921. 1963. [D. fullonum.]
La torsion de contrainte des tiges de cardere (Dipsacus silvestris
Huds.) et la theorie des helices foliaires multiples. Ibid. 3398-3403. [D.
fullonum.]
ROBINSON, B. B. Minor fiber industries. Econ. Bot. 1: 47-56. 1947. [Includes
Dipsacus.]
SCHWEITZER, I. Adatok a Dipsacus genus anatomiai es fejlodestani ismeretehez
[Beitrage zur Anatomie und Entwicklungsgeschichte der Gattung Dipsacus].
Kiilon. Egyet. Termeszett. Szovet. Evkonyv. 1910: 1-32. 1910.* [See
German summary in Bot. Centralbl. 116: 482. 1911.]
SNOW, R. Phyllotaxis of flowering teasels (Dipsacus laciniatus). New Phytol.
53: 99-107. 1954.
SOUiGES, R. Embryogenie des Dipsacacees. Developpement de l'embryon chez
le Dipsacus sylvestris Mill. Compt. Rend. Acad. Sci. Paris 256: 2268-
2273. 1963. [D. fullonum.]
SPOEL-WALVIUS, M. R. VAN DER, & R. J. DE VRIES. Description of Dipsacus
fullonum L. pollen. Acta Bot. Neerl. 13: 422-431. 1964.
STEYERMARK, J. A. An albino form of Dipsacus sylvestris. Rhodora 60: 174,
175. 1958. [D. fullonum.]
[VOL. 46


1965] FERGUSON, VALERIANACEAE AND DIPSACACEAE 231
TONGIORGI, E. Un caso di virescenza del Dipsacus silvestris Mill. Nuovo Gior.
Bot. Ital. II. 45: 221-223. 1938. [D. fullonum.]
VARGA, L. Recherches limnologiques sur la biocoenose des reservoirs de la
cardere (Dipsacus silvester Huds.). (In Hungarian; French summary.)
Erdesz. Kiser. 30: 353-369, 410. 1928. [D. fullonum.
VIETH, J. Persistance de morphoses induites chez Dipsacus Jullonum. Bull.
Sci. Bourgogne 20: 70-81. 1960. [D. sativus.]
SSur la feuille terminale presentee par Dipsacus silvestris. Ibid. 21:
39-44. 1962. [D. fullonum.]
VOROBJER, B. A. The inhabitation of larvae of Diptera in water bodies in axils
of the leaves of Dipsacus. (In Russian.) Entomol. Obozr. 39: 799-801.
1960.
WATTIEZ, N. Contribution a l'etude biochimique des dipsacees. Presence dans
Dipsacus arvensis de methylglucoside et de scabioside. Bull. Soc. Chim.
Biol. 8: 501-507. 1926.*
ARNOLD ARBORETUM
AND
GRAY HERBARIUM