JOURNAL
OF THE
ARNOLD ARBORETUM
VOLUME 69 JANUARY 1988 NUMBER 1
THE JUNCAGINACEAE IN THE SOUTHEASTERN
UNITED STATES'
JOHN W. THIERET2
JUNCAGINACEAE L. C. Richard, Demonstr. Bot. ix. 1808, "Juncagines,"
nom. cons.
(ARROWGRASS FAMILY)
Perennial [or annual], glabrous, typically perfect-flowered [dioecious], sca-
pose herbs, the axis sympodially branched, bulbous or rhizomatous, sometimes
with runners, often covered with old leaf bases, the roots fibrous [sometimes
tuberiferous]; starch grains of pteridophyte type. Leaves basal, alternate, dis-
tichous or spirodistichous, sessile, linear to almost filiform [filiform], terete to
semiterete [dorsiventrally or laterally flattened (i.e., ensiform)], ligulate [elig-
ulate]; axillary scales (squamules) present; stomata mostly paracytic. Inflores-
'Prepared for the Generic Flora of the Southeastern United States, a long-term project currently
made possible through the support of National Science Foundation grants BSR-8415769 (Carroll E.
Wood, Jr., principal investigator) and BSR-8415637 (Norton G. Miller, principal investigator). The
119th in the series, this paper follows the format established in the first one (Jour. Arnold Arb. 39:
296-346. 1958) and continued to the present.The area covered by the Generic Flora includes North
and South Carolina. Georgia. Florida. Tennessee, Alabama, Mississippi. Arkansas. and Louisiana.
The descriptions are based primarily on plants of this area, with information about extraregional
members of a family or genus in brackets [ ]. References that I have not verified are marked with an
asterisk.
1 am indebted to Carroll Wood and Norton Miller for their editorial and bibliographic help: to the
staff of the Lloyd Library. Cincinnati: to David M. Brandenburg. Brooklyn Botanic Garden, and
Barney L. Lipscomb. Southern Methodist University. for bibliographic help; and to Northern Ken-
tucky University, which provided a research grant that made possible the acquisition of reference
materials and travel to the University of North Carolina, Chapel Hill. where J. R. Massey. of the
Department of Biology. and William Burk and Jeff Beam, of the Botany Library, were of invaluable
help. The illustration was prepared in 1987 by lhsan A. Al-Shehbaz. of the Arnold Arboretum, from
specimens collected by James R. Burkhalter near Pensacola, Florida.
'Department of Biological Sciences. Northern Kentucky U1niversity, Highland Heights. Kentucky
41076.
c President and Fellows of Harvard College. 1988.
Journal of the Arnold Arboretum 69: 1-23. January, 1988.


2 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 69
cence of terminal, bractless, pedunculate racemes [spikes] [rarely reduced to a
single terminal flower]. Flowers actinomorphic to zygomorphic, hypogynous,
perfect to imperfect, anemophilous, proterogynous. Perianth of 6 [4] (or fewer
by abortion) distinct and mostly similar tepals in 2 whorls. Androecium usually
of 6 [4] subsessile stamens (or fewer by abortion) in 2 whorls, each filament
adnate at its base to the base of the subtending tepal; anthers tetrasporangiate,
bilocular at anthesis. extrorse, opening by lengthwise slits, the tapetum ame-
boid, microsporogenesis of the successive type: pollen grains globose to ellip-
soid. monocolpate. inaperturate, binucleate (sometimes trinucleate?) when shed.
Gynoecium of 6 [4] carpels in 2 whorls (occasional flowers with 3 to 12 carpels),
these more or less adnate to the central axis (carpophore) [or central axis
lacking], some commonly sterile, the fertile carpels eventually separating from
the axis [the carpels connate at least basally, forming a compound ovary in
Maundia and Tetroncium]; styles short to lacking [well developed, subulate,
and persistent as beaks on the fruit in 'Tetroncium]; stigmas distinct, plumose
to papillose: ovary 1-locular [2-4-locular in Maundia, incompletely 4-locular
in 7itroncium], each locule with a single ovule, this bitegmic, crassinucellar,
anatropous [orthotropous], and erect [pendulous in Maundia]; megagameto-
phyte (embryo sac) development of the Polygonum (normal) type; endosperm
development nuclear; embryogeny of the caryophyllad type. Fruit schizocarpic,
the mericarps indehiscent, achenelike [fruit essentially capsular(?) in Maundia
and Tctroncium]: seed linear to ovoid, without endosperm [this present in
letroncium], the embryo straight. (Excluding Lilaeaceae Dumortier, 1829, and
Scheuchzeriaceae Rudolphi. 1830, nomina conservanda.) TYPE GENUS: Juncago
Seguier, nom. illeg. = Triglochin L.
A small family of three genera and about 22 species: the monotypic Maundia
F. Mueller (M. triglochinoides F. Mueller), of coastal eastern Australia (New
South Wales and Queensland); the likewise monotypic Tetroncium Willd. (T.
magellanicum Willd.). of the Falkland Islands and southern South America
(from Tierra del Fuego. northward in the Andes to at least 400S in Argentina
and Chile), unique in the family because it is dioecious; and the widely dis-
tributed Triglochin L. (including the Australian Cy(cnogeton Endl.). about 20
species, one of which occurs in the southeastern United States.
The circumscription of the Juncaginaceae here accepted (with three genera)
has ample precedent in the literature (e.g., by Chant; Cook: Engler & Diels;
Hutchinson: Kimura; Mason: Tomlinson, 1982), but the relationships of two
additional genera, Sccheuchzeria L. and Lilaea Humb. & Bonpl., remain un-
settled. Both (e.g., by Agrawal: Britton: Buchenau, 1903; Buchenau & Hieron-
ymus; Hegi; Johri; Lotsy; Rendle; Suessenguth; Thorne, 1976, 1983; and Wett-
stein) or either one of these (Scheuchzeria only, by authors including Abrams;
or Lilaea only, by those including Cronquist, 1968, 1981; Dahlgren & Clifford;
Dahlgren, Clifford, & Yeo; Markgraf, 1981; Eckardt; Stebbins; and Takhtajan)
may be included in an amplified version of the Juncaginaceae.
That Scheuchzeria represents a family by itself is accepted by the majority
of workers after 1940 (see summary of the various monocot systems in Dahlgren
& Clifford). Among the "large" number of important features (Tomlinson,
_ �______~__
__


THIERET, JUNCAGINACEAE
1982) in which this genus differs from Triglochin and Lilaea are three- (vs.
two-)celled pollen; biovulate (vs. uniovulate) carpels; helobial (vs. free-nuclear)
endosperm; the presence (vs. absence) of hypodermal crystals, a leaf apical
pore, and floral bracts; and the absence (vs. presence) of squamules (minute
scale- or gland-like outgrowths occurring in the axils of leaves; unique in the
Helobiae). Floral developmental evidence further confirms the separate familial
status of Scheuchzeria (Posluszny). A character common to all three genera,
however, is the occurrence of the cyanogenic glucoside triglochinin.
The placement of Lilaea appears to be less certain. With Triglochin it shares
two-celled pollen, uniovulate carpels, free-nuclear endosperm, and squamules.
Both genera have an unusual type of embryo development in which the "first
embryonic root does not originate from the point where the suspensor is at-
tached, but from the lateral position . . . directly from the epidermis of the
embryo, therefore not in the manner of adventitious roots, which develop from
the deep inner tissue" (Yamashita, p. 205). Lilaea differs from Triglochin in
the number of carpels (one vs. six) and perianth segments (one vs. six) and in
the presence of laticifers (also lacking in Scheuchzeria). Tomlinson (1982),
whose discussion of the three genera is outstanding, concluded that it is better
to retain the separate familial status of Lilaea until a more extensive study has
been made. (See also Posluszny et al. and Singh.) It should be noted that
knowledge ofJuncaginaceae (sensu stricto) is based mostly on Northern Hemi-
sphere representatives of the family: more data are needed on species of Tri-
glochin of the Southern Hemisphere, especially those of Australia, most of
which are diminutive annuals, as well as on Tetroncium and Maundia.
Many authors accept a close relationship between the Juncaginaceae and the
Potamogetonaceae. Takhtajan suggested that the latter may well be derived
from the former (and that the Posidoniaceae probably represent an extreme
marine stage of the Juncaginaceae). (Engler & Diels and Skottsberg even in-
cluded the Juncaginaceae in the Potamogetonaceae.) Such a relationship was
postulated over a century ago by Bentham (p. 165): "Triglochin ... appears
to me to be much more nearly connected with Potamogeton [than with Alis-
maceae], in the peculiar structure of the flowers, differing chiefly in habit, and
even in that respect, the half-floating leaves of T. procera show a near approach
to that genus." Hutchinson (p. 678) expressed the opinion that "the primitive
tribe Helonieae of the Liliaceac has been developed from the same stock as
the Juncaginaceae: the general facies of its genera is the same and in both groups
the flowers are quite ebracteate." Joseph D. Hooker pointed out that the habit
of Tetroncium is precisely that of iarthecium of the Liliaceae.
Fruit types in the family have been variously, and often questionably, de-
nominated and described. I find. for example. no convincing evidence that the
fruits of any of the three genera are indeed "follicles" or that they are "de-
hiscent." The fruits of Triglochin appear to me to be schizocarpic, the mericarps
indehiscent (see under Triglochin below). Those of the other genera are essen-
tially capsular according to some authors: in Maundia, "carpels 2-4, united
along the whole length of their inner faces. Fruiting carpels ... remaining united
and together forming a 2- to 4-angled squat cylinder truncate at the summit
with 2-4 short spreading beaks" (Aston, p. 241); and in Tetroncium, "carpels
1988]


JOURNAL OF THE ARNOLD ARBORETUM
4, united at base into incompletely 4-locular ovary.... Fruit indehiscent,
4-celled, seed usually single, with three aborted ovules" (Moore. 1968, p. 137).
Both genera were included among those with capsular fruits by Dahlgren &
Clifford, but three years later Dahlgren, Clifford, & Yeo (p. 312) wrote of them:
"carpels ... more or less fused centrally . . . but becoming separate in the fruit
stage." Bentham (p. 169) described the carpels of Maundia as being "almost
drupaceous, each with a thinly cartilaginous endocarp." According to Thomp-
son (p. 80), the "fruiting carpels [are] joined at the adaxial angle [and have] a
spongy covering on the ventral surface." Developmental study of the fruits of
both llaundia and Tetroncium is called for.
Although seeds of the Juncaginaceae are usually described as lacking en-
dosperm, Tetroncium is apparently an exception. Of this genus J. D. Hooker
(p. 359) wrote "albumen farinaceum." Buchenau (1882, p. 472) acknowledged
this exception in a footnote to his assertion that "Der Samen der Juncaginaceen
enthalt bekanntlich kein Albumen." Later, however, he and Hieronymus (p.
223) were skeptical ("Ter'ronciumn mit kleinem Nahrgewebe?"). Finally, how-
ever, Buchenau (1903, pp. 2, 6). apparently on the basis of his own observation.
agreed with J. D. Hooker: "Embryo ... in TeIroncio albuminosus" and "Nur
bei 7Teroncium liegt der ... Embryo ... in einem stirkemehlreichen Nahr-
gewebe."
Noteworthy anatomical features of the Juncaginaceae include a single ring
of collateral vascular bundles in the inflorescence axis, imparting an "exceed-
ingly dicotyledonous" look (Chrysler. p. 180); the reported occurrence of an
"incipient cambium" in the rhizome bundles (Andersson, Hill, Salisbury) ("...
perhaps ... a case of incipient secondary thickening which is found developed
to a much greater extent in Monocotyledons like Aristea. Dracaena, Yucca,
etc."; Hill, p. 88); the presence of so-called axillary squamules, which are of
wide occurrence in the Alismatidae (and one species ofAraceae) (Arber [1923]
considered them as appendages originating from the abaxial epidermis of the
leaf just distal to the leaf in the axil of which they are found): and inflorescence
formation by apical bifurcation of the axis (the original terminal meristem
forming the inflorescence, the axillary bud of the distalmost leaf continuing
further vegetative growth: reported in other Alismatidae).
Adoption of the name Scheuchzeriaceae (e.g., by Kartesz & Kartesz) when
Scheuchzeria and Triglochin are included in the same family is incorrect. The
name Juncaginaceae predates Scheuchzeriaceae by 22 years. If the Juncagi-
naceae are merged with the Potamogetonaceac Dumort. (1829), the latter name
must be used (see International Code of Botanical Nomenclature, 1983, Ap-
pendix II).
REFEREN( ES:
ABRAMS, L. An illustrated flora of the Pacific States Washington. Oregon, and California.
Vol. 1. 557 pp. Stanford. (alifornia. 1923. [Scheuchzeriaceae. including T'rilochin,
96, 97.]
AGRAWAL.. J. S. The embryology of Lilaea suiulata H.B.K. with a discussion on its
systematic position. Phytomorphology 2: 15-29. 1952. IilIaea closely related to
Trilochin. ]
[VOL. 69


1988] THIERET, JUNCAGINACEAE 5
ANDERSSON, S. Om de primira karlstringarnes utveckling hos monokotyledonerna.
Bihang Sv. Vet.-Akad. Handl. 13. 1(12). 23 pp. 2 pis. 1888. [T. maritimum, 8, 9,
21, pl. 2. fig. 1: vestigial cambium. German summary in Bot. Centralbl. 38: 586,
587, 618, 619. 1889.]
ARBER, A. On the 'squamulae intravaginales' of the Helobieae. Ann. Bot. 37: 31-41.
figs. 1-5. 1923. [Triglochin, passim.]
SMonocotyledons. A morphological study. 258 pp. Cambridge, England. 1925.
[Tetroncium, 111, fig. 84: Triglochin, passim.]
AsTON, H. I. Aquatic plants of Australia. 368 pp. Melbourne. 1973. [Juncaginaceae,
241-248, 346; 11aundia. Triglochin.]
BAILLON, H. Triglochineae. Hist. Pl. 12: 99-101, 119, 120. figs. 146-150. 1894. [Te-
tronciumn, Scheuchzeria, and Triglochin (as a "serie" of Najadacees), 101, 102, 120,
121: Lilaea (in serie Lilaeeae). 101, 102, 120, 121.]
BENTHAM, G. Flora Australiensis. Vol. 7. Roxburghiaceae to Filices. 806 pp. London.
1878. [Triglochin, 165-169: author "assisted by Baron Ferdinand von Mueller."]
S& J. D. HOOKER. Naiadaccae. Gen. PI. 3(2): 1009-1019. 1883. [Tribus I. Jun-
cagineae, including Lilaea, Triglochin, and Scheuchzeria, 1010, 1012, 1013.]
BREWBAKER, J. L. The distribution and phylogenetic significance of binucleate and
trinucleate pollen grains in the angiosperms. Am. Jour. Bot. 54: 1069-1083. 1967.
[Lilaea trinucleate: discounts earlier report of binucleate pollen in Triglochin but
does not document.]
BRITTON, N. L. Scheuchzeriaceae. N. Am. Fl. 17:41,42. 1909. [Scheuchzeria, Triglochin:
Lilaea in Lilaeaceae, 37.]
BUCHENAU, F. Index criticus Juncaginacearum hucusque descriptarum. Abh. Naturwiss.
Ver. Bremen 1: 213-224. 1867.
Index criticus Butomacearum. Alismacearum, Juncaginacearumque hucusque
descriptarum. Jour. Bot. London 7: 219-232. 1869. [Juncaginaceae. 230-232.]
- Nachtrage zu den im crsten und zweiten Bande dieser Abhandlungen verfflent-
lichen kritischen Zusammenstellungen der bis jetzt beschriebenen Butomaceen, Al-
ismaceen und Juncaginaceen. Abh. Naturwiss. Ver. Bremen 2: 481-503. 1871. [Jun-
caginaceae, 491-498.]
Beitrage zur Kenntniss der Butomaceen. Alismaceen und Juncaginaceen. Bot.
Jahrb. 2: 465-510. 1882.
Scheuchzeriaceae. Pflanzenr. IV. 14(Heft 16): 1-20. 1903. [Lilaea, tlaundia.
Scheuchzeria, Tetroncium. Triglochin.]
-- & G. HIERONYMUS. Juncaginaceae. Nat. Pflanzenfam. II. 1: 222-227. 1889.
[Lilaea, Scheuchzeria, Triglochin (including Aaundia), lTTroncium.]
BURGER, W. C. The Piperales and the monocots. Alternate hypotheses for the origin of
monocotyledonous flowers. Bot. Rev. 43: 345-393. 1977. [Juncaginaceae, 351, 352,
and passim.]
CAMP. W. H. Distribution patterns in modern plants and the problem of ancient dis-
persals. Ecol. Monogr. 17: 159-183. 1947. [Juncaginaceae, 168, fig. 13 (map of
world distribution of family).]
CAMPBELL, D. H. The development of the flower in Lilaea suhulata. H.B.K. Ann. Bot.
12: 1-28. p/s. 1-3. 1898.
CHANT, S. R. Juncaginaceae, Scheuchzeriaceae. P. 273 in V. H. HEYWOOD, consultant
ed., Flowering plants of the world. New York. 1978. [Inaccurate maps of world
distribution of these families.]
CHARLTON, W. A. Flower and inflorescence in Lilaca, Triglochin and Zostera. Helobiae
Newslett. 1: 10. 1976. [Trivial.]
CHATIN, G.-A. Anatomic comparec des vcgetaux. Plantes aquatiques. Monocotyledones.
96 pp. Paris. 1856, 1862. [Ordre des Juncaginees, 75-96. pls. 20-22.]
CHRYSLER, M. A. The structure and relationships of the Potamogetonaceae and allied


JOURNAL OF THE ARNOLD ARBORETUM
families. Bot. Gaz. 44: 161-188. p/s. 14-18. 1907. [1'. maritimum. 179. 180, p1. 17,
/ig. 27, p1. 18, fig. 33.]
COOK, C. D. Lilaeaceae. Pp. 273, 274 in V. H. HEYWooD, consultant ed., Flowering
plants of the world. New York. 1978. [Lilacaceae can be "considered to represent
a bridge between the Zannichelliaceae and the Najadaceae."]
(CORREA, M. N. Juncaginaceac. In: M. N. CORREA, ed., Fl. Patagonica 2: 33-37. 1969.
[ 'etroncium, Triglochin.
CRONQUIST. A. The evolution and classilication of flowering plants. 396 pp. Boston.
1968. [Najadales. 327-330. including Juncaginaceae.]
An integrated system of classification of flowering plants. 1262 pp. New York.
1981. [Juncaginaceae, 1062-1064, fig. 7.6.]
CzAJA. A. T. Structure of starch grains and the classification of vascular plant families.
Taxon 27: 463-470. 1978. [Juncaginaceae, including Lilaea, 467.]
)DA(HLIAN, C. P. A review of the fossil record of monocotyledons. Bot. Rev. 47: 517-
555. 1981. [Juncaginaceae: no reliable fossils (but cf. MARKGRAF, 1981).]
1)AHI.REN, R. M. T., & H. T. CLIFFORD. The monocotyledons: a comparative study.
378 pp. New York. 1982. [Juncaginaceae. passim.]
S& P. F. YEO. The families of monocotyledons. Structure, evolution, and
taxonomy. 520 pp. Berlin. 1985. [Juncaginaceae, 310-312, passim: (Cycnogeton.
l.iaea. \laundia. T troncium. Trigilochin: errors of fact.]
DAVIS, (. L. Systematic embryology of the angiosperms. x + 528 pp. New York. 1966.
[Juncaginaceae. 150.]
D)MIN, K. Morphologische und phylogenetische Studien iiber die Stipularbildung. Ann.
Jard. Bot. Buiten/org, II. 9: 117-326. p/s. 23-33. 1911. [Juncaginaceae. 188-191:
I.ilaca, Scheu'hzeria. Tri'/ochin, 7i'trncium.]
ECKARDT, T. Reihe Helobieae. Pp. 499-512 in H. MELCHIOR, A. Engler's Syllabus der
Pflanzenfamilien. ed. 12. Vol. 2. Berlin. 1964. [Juncaginaceae, 506, 507. fig. 211.
(G-R: including Lilaea.]
EI( HLER. A. W. Bliithendiagramme. Erster Theil. 348 pp. Leipzig. 1875. [Juncaginaceae,
101, 102: Scheuchrzria, Tc~ronciumn. Trigl,.chin.]
ENGLER, A., & L. DIEls. Syllabus der Pflanzenfamilien. ed. 11. xliii - 419 pp. Berlin.
1936. [Juncaginaceae (sen'.v stricto) included in Potamogetonaceae as tribe Triglo-
chineae, 133; Lilaea in tribe Zostereae, 133; Scheuchzeria in Scheuchzeriaceae, 135.]
ERDTMAN. G. Pollen morphology and plant taxonomy. Angiosperms. xii ' 539 pp.
Stockholm. 1952. [Juncaginaceac, 21 7: 'Tcroncium. Trig/ochin.]
GIBus. R. D. Chemotaxonomy of flowering plants. Vols. 1-4 (paged continuously). 2372
pp. Montreal. 1974. [Juncaginaceae. 2: 1131. 1156: 3: 1858.]
(iOLDBLATT, E., ed. Index to plant chromosome numbers 1975-1978. Monogr. Syst.
Bot. 5. vii + 553 pp. 1981. [Juncaginaceac, 287.]
Slii-c, G. Illustrierte Flora von Mittel-Europa. Band I. clviii 4 411 pp. Miinchen. 1906.
[Juncaginaceae, 146-149. pI. IS, if. 4: 1)'rilochin. See SUESSENGUT H and MARKGRAF
(1981) for eds. 2 and 3.]
HEG;NAUER, R. Chemotaxonomie der Pflanzen. Band 2. Monocotyledoneae. 540 pp.
Basel and Stuttgart. 1963. [Juncaginaceae. 432-434.]
--- & H. W. L. RUIJOROK. Lil.aca scilloimdc und .Juncus bulhosuts wei neue cyanogene
Pllanzen. Phytochemistry 10: 2121-2124. 1971. [L. scilloides contains a cyanogenic
glycoside that is probably identical with triglochinin.]
HiL.. T. G. The structure and development of I'rig/ochin maritimum, L. Ann. Bot. 14:
83-107. p/s. 6. 7 1900.
HITCHCOCK, C. L. Juncaginaceae. Pp. 153-157 in C. L. HIT(CHCOCK. A. (RONQUIST, M.
OWNBEY, &J. W. THOMPSON, Vase. P. Pacific Northwest. Vol. 1. Seattle and London.
1969. [Lilaea. Tr'ig/ochin.]
HOOKER, J. D. The botany of the Antarctic voyage of H. M. discovery ships Erecus
and T'error in the years 1839-1843 under the command of Sir James Clarke Ross.
3 parts (Flora Antarctica, Flora Novae-Zelandiae. Flora Tasmaniae) in 6 vols. 528
[VOL. 69


THIERET, JUNCAGINACEAE
pls. London. 1844-1859. (Facsimile ed. 3 vols. + atlas vol. with all pls. J. Cramer.
1963.) [Tetroncium, Part I (Fl. Antarctica), 359, pl. 128. Triglochin, Part I (Fl.
Antarctica), 360; Part II (Fl. Novae-Zelandiae), 235, 236; Part III (Fl. Tasmaniae),
Vol. 2, 39-41.]
HOOKER, W. J. Tetroncium magellanicum. Ic. P1. II. 2: t. 534. 1843.
HUTCHINSON, J. The families of flowering plants. ed. 3. xx + 968 pp. Oxford. 1973.
[Juncaginaceae, 678, 679, fig. 349.]
JOHANSEN, D. A. Plant embryology. Embryogeny of the Spermatophyta. 305 pp. Wal-
tham, Massachusetts. 1950. [Juncaginaceae, 237, 238.]
JOHRI, B. M. Embryology and taxonomy. Pp. 395-444 in P. MAHESHWARI, ed., Recent
advances in the embryology of angiosperms. Delhi. 1963. [Comparison of Lilaea,
Scheuchzeria, and Triglochin. 396.]
KARTESZ. J. T.. & R. KARTESZ. A synonymized checklist of the vascular flora of the
United States, Canada, and Greenland. 498 pp. Chapel Hill, North Carolina. 1980.
[Scheuchzeriaceae, 438; Lilaea, Scheuchzeria, Triglochin.]
KIMURA, Y. Systeme der phylogenie des monocotyledones. Not. Syst. Paris 15: 137-
159. 1956. [Scheuchzeriaceae and Triglochinaceae (Juncaginaceae) in Scheuchzeri-
ales, Lilaeaceae in Najadales, 138.]
KUPRIANOVA, L. A. Pollen morphology and phylogeny of the monocotyledons. (In
Russian.) Acta Inst. Bot. Acad. Sci. URSS 1. Syst. 7: 163-262. 1948. [Juncaginaceae,
254, pl. 14, fig. 6 (T. palustris), fig. 7 (Tetroncium magellanicum); descriptions and
illustrations of pollen.]
LARSEN, K. Cytotaxonomical note on Lilaea. Bot. Not. 119: 496, 497. 1966. [2n = 12;
"cytological support" for uniting Lilaeaccae with Juncaginaceae.]
LOTSY, J. P. Vortrage liber botanische Stammesgeschichte. Dritter Band. Cormophyta
Siphonogamia. Erster Theil. 1055 pp. Jena. 1911. [Juncaginaceae, 649-653.]
MARKGRAF. F. Bliitenbau und Verwandtschaft bei den cinfachsten Helobiae. Ber. Deutsch.
Bot. Ges. 54: 191-229. pls. 1-8. 1936. [Mlaundia, Tetroncium, Triglochin, passim.]
- Juncaginaceae. In: G. HEGI, Illus. Fl. Mitteleuropa. ed. 3. 1(2): 198-203. pl.
18, jig. 4. 1981. [Useful account.]
MASON, H. L. A flora of the marshes of California. viii + 878 pp. Berkeley and Los
Angeles. 1957. [Scheuchzeriaceae, Juncaginaceae, Lilaeaceae, 95-103, figs. 39 (Tri-
glochin maritimum), 40 (T. concinnum), 41 (T. striatum), 42 (Lilaea scilloides).
MICHELI, M. Juncagineae L. C. Richard. In: A. & C. DE CANDOLLE, Monogr. Phanerog.
3: 94-112. 1881. [Systematic account with discussion of affinities.]
MOORE, D. M. The vascular flora of the Falkland Islands. British Antarctic Surv. Sci.
Rep. 60. 202 pp. 6 pls., map. London. 1968. [Tetroncium. 137, pl. 6k.]
Flora of Tierra del Fuego. 396 pp. St. Louis. 1983. [Juncaginaceae, 267, 268;
Tetroncium, Triglochin.]
MUELLER, F. Fragmenta phytographiac Australiae. Vol. 1. 252 pp. Melbourne. 1858-
1859. [Maundia, 22, 23: original description.]
SFragmenta phytographiac Australiae. Vol. 6. 276 pp. Melbourne. 1867-1868.
[Juncaginaceae, 81-83; M.aundia, Triglochin.]
MUENSCHER, W. C. Aquatic plants of the United States. x - 374 pp. Ithaca. New York.
1944. [Juncaginaceae, 72-78, figs. 30-32, maps 62-65: Lilaea, Scheuchzeria, Tri-
glochin.]
MuNoz PIZARRO, C. Sinopsis de la flora chilena. ed. 2. 500 pp. 243 p/s. Santiago. 1959.
[Juncaginaceae, 175, pl. 232 (Tetroncium magellanicum).]
PECK, M. E. A manual of the higher plants of Oregon. ed. 2. 936 pp. [Portland.] 1961.
[Juncaginaceae, 76. 77; Triglochin (including T. striatum), Scheuchzeria.]
PICHI SERMOLLI, R. E. G., & M. P. BIZZARRI. The botanical collections (Pteridophyta
and Spermatophyta) of the AMF .Mares-G.R.S.T.S. expedition to Patagonia, Tierra
del Fuego and Antarctica. Webbia 32: 455-534. 1978. [Tetroncium magellanicum,
519.]
POSLUSZNY, U. Re-evaluation of certain key relationships in the Alismatidae: floral
1988]


JOURNAL OF THE ARNOLD ARBORETUM
organogenesis of .'heuch:eria pui.stris (Scheuchzeriaceae). Am. Jour. Bot. 70: 925-
933. 1983. [Reconfirms retention of Schcuchzeria in a unigeneric family.]
-- & W. A. CHARLTON. Inflorescence in the Helobiae (Alismatidae)-convergences
and divergences. (Abstract.) Am. Jour. Bot. 73: 617. 1986. [Includes data on Lilaea
and Triglochin.]
. , & D. K. JAIN. Morphology and development of the reproductive shoots
in Lilaca scilloidcs (Poir.) Hauman (Alismatidae). Bot. Jour. Linn. Soc. 92: 323-
342. fig. 1-43. 1986. [Li/aea close to Triglochin; "if one views the eccentricities of
Lilaea as specializations then it is reasonable to keep the assignation to the separate
family Lilacaceae."]
RAVOLOLOMANIRAKA. D. Contribution l'etude de quelques feuilles de monocotyle-
dones. Bull. Mus. Hist. Nat. Paris, 3e ser., no. 46, Bot. 2: 29-69. 1972. [Tetroncium.
48, fig. 40: leaf ontogeny.]
RENDLE. A. B. The classification of flowering plants. ed. 2. Vol. 1. Gymnosperms and
monocotyledons. 412 pp. Cambridge. England. 1956. [Juncaginaceae, 208. 209.]
SALISBURY, E. J. Floral construction in the Helobiales. Ann. Bot. 40: 419-445. 1926.
[Triglochin, passim; comments on various degrees of syncarpy, on vestigial cam-
bium, and on the "strikingly dicotyledonous" ring of bundles in the inflorescence
axis.]
SAINDERS, E. R. On carpel polymorphism. III. Ann. Bot. 43: 459-481. 1929. [Juncag-
iraceac, 468, figs. 38-44; Scheuch-eria, 7Triglochin.]
S(OGGAN, H. J. Juncaginaceae. Fl. Canada 2: 209. 1978. [Schcucheria. rriglochin:
Lilaea in Lilaeaceae, 210.]
SINGH, V. Morphological and anatomical studies in Helobiae. V. Vascular anatomy of
the flower of Liltaa scilloicds (Poir.) Hamm. Proc. Indian Acad. Sci. B. 61: 316-
325. 1965. [Lilaca probably derived from Triglochin; now sufficiently far removed
from present-day Triglochin to justify recognition of Lilaeaceae.]
-- & R. SATTLER. Floral development of A.ponogeton nalans and 1. undullaus.
Canad. Jour. Bot. 55: 1106-1120. 1977. [Among the Helobiae Aponogetonaceae
appear most closely related to Scheuzeriaceae and Juncaginaceae.]
SKOTTSBERG. C., ed. Vixternas liv. Vol. 5. 735 pp. Stockholm. 1940. [Juncaginaceae
(Mlaundia, Tetronciun,. Triglochin) included in Potamogetonaceae as subfamily
Triglochinoideae. 617.]
SOI EREDER, H.. & F. MEYER. Scheuchzeriaceae. In: Svstematische Anatomie der Mo-
nokotyledonen 1(1): 147-155. Berlin. 1933. [Lilaca, Alaundia, Schcuchzeria, Te-
tronciumt. 7Trig/oc hin.]
STEBBINS, G. L. Flowering plants. Evolution above the species level. 399 pp. Cambridge,
Massachusetts. 1974. [Scheuchzeriaceae and Juncaginaceae (including Lilaca). 354.]
- & G. H. KHUSH. Variation in the organization of the stomatal complex in the
leaf epidermis of monocotyledons and its bearing on their phylogeny. Am. Jour.
Bot. 48: 51-59. 1961. [Juncaginaceae, 54, t. 2: among families with two subsidiary
cells as the predominant condition.]
SUESSENGUTH, K. G. Hegi. Illustrierte Flora von Mittel-Europa. ed. 2. Band 1. 528 pp.
Munich. 1935. [Juncaginaceae, 212-215, t. IS, fig. 4; 7r-igrochin.]
TAKHTAJAN, A. L. Outline of the classification of flowering plants (Magnoliophyta). Bot.
Rev. 46: 225-359. 1980. [Juncaginaceae, 303.]
THOMPSON, J. Juncaginaceae. Fl. New S. Wales No. 16. Contr. New S. Wales Natl.
Herb. Fl. Ser. 1-18: 77-80. 1961. [Maundia, Triglochin.]
THORNE. R. F. A phylogenetic classification of the Angiospermae. Evol. Biol. 9: 35-
106. 1976. [Juncaginaceae, 65.]
Proposed new realignments in the angiosperms. Nordic Jour. Bot. 3: 85-117.
1983. [Juncaginaceae placed in order Zosterales, suborder Potamogetonineae. along
with Potamogetonaceae, Posidoniaceae, Zannichelliaceae, Cymodoccaceae; subfam-
ilies Scheuchzerioideae and Juncaginoideae (including Lilaca) recognized.]
[vol.. 69


THIERET, JUNCAGINACEAE
TOMLINSON, P. B. Development of the stomatal complex as a taxonomic character in
the monocotyledons. Taxon 23: 109-128. 1974. [Juncaginaceae, 127.]
Helobiac (Alismatidae). Vol. 7 in C. R. METCALFE, ed., Anatomy of the mono-
cotyledons. 522 pp. 16 pls. Oxford. 1982. [Juncaginaceae, 13, 242-257, fig. 7.1-7.4;
Lilaeaceae, 13, 258-269, fig. 8.1-8.3: Scheuchzeriaceae, 13, 226-241, fig. 6.1-6.4.]
UHL, N. W. Studies in the floral morphology and anatomy of certain members of the
Helobiae. Unpubl. Ph.D. thesis. Cornell University. 1947. [Triglochin, 16-30, pas-
sim, figs. 1-19; also includes data on Li/aea and Scheuchzeria; often cited, but
unfortunately never published.
WETTSTEIN, R. Handbuch der systematischcn Botanik. ed 4. x + 1152 pp. Leipzig and
Vienna. 1935. [Scheuchzeriaceae, including Lilaea. Scheuchzeria, and Triglochin,
974, 975, p/. 651, figs. 1-4.]
YAMASHITA, T. Eigenartige Wurzelanlage des Embryos bei Lilaea subulata Humb. et
Bonpl. und Triglochin maritimum L. Jour. Fac. Sci. Univ. Tokyo, III. 10: 181-205.
1970.
1. Triglochin Linnaeus, Sp. PI. 1: 338. 1753; Gen. P1. ed 5. 157. 1754.'
Perennial [annual], perfect-flowered, herbaceous plants of wetlands, axis
sympodially branched, somewhat [to strongly] bulbous [rhizomatous], with
[without] runners, the roots fibrous [sometimes tuberiferous]; vessels confined
to the roots. Leaves basal, distichous, sessile, linear, terete or semiterete [flat],
ligulate [eligulate], sheathing, the sheath open. Inflorescence of terminal, bract-
less, pedunculate racemes [spikes] [rarely reduced to a single terminal flower].
Flowers actinomorphic, perfect to imperfect. Perianth of 6 tepals (or fewer by
abortion) in 2 whorls, the upper whorl seemingly distal to the lower whorl of
stamens. Androecium of 6 subsessile stamens (or fewer by abortion) in 2 whorls,
each filament adnate at its base to the base of the subtending tepal, the stamen
and tepal typically deciduous as a unit; pollen inaperturate, spheroidal, retic-
ulate, binucleate (sometimes trinucleate?) when shed. Gynoecium of 6 carpels
(occasional flowers with 3 to 12), these more or less adnate to the prolonged
central axis (carpophore) [or central axis not prolonged, the carpels free], the
lower 3 usually sterile, remaining attached to the axis as wings or ridges, the
upper 3 [or all 6] eventually separating from the axis; styles short or lacking;
stigmas distinct, plumose to papillose. Fruit schizocarpic, the mericarps in-
dehiscent, achenelike [gynoecium apocarpous, the carpels achenelike in T. pro-
cerum]. TYPE SPECIES: 7. palustre L.; see N. L. Britton, N. Am. Fl. 17: 41. 1909.
(Name from Greek treis, three, and glochis, point, from the pointed carpel
bases of 7T palustre, which spread from the axis at fruit maturity, the gynoecium
then slightly recalling a three-barbed spear-point.)- ARROWGRASS.
A genus of about 20 species in two subgenera, widely distributed but best
represented in the Southern Hemisphere, especially in Australia. Subgenus
CYCNOGETON (Endl.) Buch. (characterized by perennial habit, free carpels, and
lack of a carpophore) embraces a single species, Triglochin procerum R. Br.,
which occurs in southern New Guinea and Australia (including Tasmania).
'The name Triglochin has been-and continues to be-treated as feminine by many authors, either
because glochin is a feminine noun or. more likely. because others do it that way. Linnaeus, however,
used the name as neuter, and his choice should be followed (ICBN, 1983, Rec. 75A. 3).
1988]


JOURNAL OF THE ARNOLD ARBORETUM
o
1 2
W�i  , i \ 3
r ^ tIa
.' -i !A t-iil
FIC;ItiE 1. TIriglochin. a-h, T'. ,slrialnnr: a, habit, � /2; b. portion of leaf sheath to
show ligule. � 4: c. tfower. the fertile stamen and its subtending tcpal to the right, x 12;
d, floral diagram (after Lieu). showing 6 tepals (5 with sterile stamens). 3 sterile and 3
tfertile carpels, the latter each with a single 2-integumented ovule; e, portion of infruc-
tescence. x 6; f, mature fruit with 3 fertile and 3 sterile carpels, from above, x 12; g.
persistent sterile carpels after removal of fertile mericarps, x 12; h, fertile mericarp.
S12.
[VOL. 69


THIERET, JUNCAGINACEAE
Triglochin procerum var. dubium (R. Br.) Bentham is recognized by some (e.g.,
Cunningham et al.) as a distinct species. Triglochin procerum, water-ribbons,
is the most aquatic of its congeners, with its leaves up to 2(-3.5) m long (Aston)
when the plant grows in deep, flowing water, the distal portion floating or the
entire leaf submersed and trailing downstream (in the manner of Sparganium
L. or I allisneria L.).
Subgenus TRIGLOCHIN (characterized by annual or perennial habit and-
except in one annual-the adnation of carpels to a carpophore) is found on all
continents except Antarctica. The single species of the southeastern United
States, Triglochin striatum Ruiz & Pavon (7T. loridanum Gand., T. sessile
Gand., T. triandrum Michaux) is one of several species (number uncertain: see
below) of its genus in North America. In our area it occurs along the coast
from Maryland to Louisiana, typically in brackish and saline habitats (marshes,
meadows, ditches, swamps), often in shallow water. An impressively wide-
ranging plant, it is also distributed in the western United States (Oregon and
California), the Valley of Mexico, the Bahamas, Cuba, South America (Peru
and southeastern Brazil south to Tierra del Fuego), southern Africa (Cape
Peninsula to Angola and Mozambique), and other extratropical regions, mostly
coastal Australia (Western Australia to Tasmania and southeastern Queens-
land), New Zealand, and the Auckland and Chatham islands.
The taxonomy of Triglochin, at least in North America, is unsettled; estimates
of the number of species here range from three (e.g., Britton) to six (e.g., Kartesz
& Kartesz). The circumboreal T. maritinmum complex, source of most of the
uncertainty, is in need of critical study worldwide (see Love & Love. 1958a).
The North American species of Triglochin, all perennials, typically grow in
brackish, saline, and alkaline hydric habitats (less frequently in fresh water),
especially in the western United States and western Canada, and to the north
of our area.
The flowers of Triglochin have one or two whorls of three "bractiform peri-
anth-like appendages" (Mason. p. 95). the nature of which has been much
discussed. Each appendage subtends and is adnate at its base to a stamen (these
reduced in some species), the two typically falling as a unit. The upper whorl
of appendages appears to diverge from the axis above the lower whorl of sta-
mens: its vascular supply, too, arises above that of the lower stamens (Uhl).
This morphology led Uhl to interpret the flower of Triglochin as a reduced
inflorescence of six staminate flowers and one or more naked carpellate flowers,
each carpel probably representing a single flower (see also Burger). Such an
interpretation has been questioned by Lieu, who found (p. 1418) that both
whorls of tepals are initiated in acropetal succession before initiation of the
stamens, that the anomalous position of the upper tepals results from "differ-
ential timing and rates of growth of the inner tepals and outer stamens," and
that "the evidence does not support the interpretation of the flower of Triglochin
as a reduced lateral branch of [an] inflorescence." Lieu did note that emphasis
on vasculature would support interpretation of the flower as an inflorescence,
but that developmental studies refute such an interpretation and lend support
to regarding the flower of Triglochin as comparable to the ordinary trimerous
monocotyledonous flower.
1988]


JOURNAL OF THE ARNOLD ARBORETUM
The flowers of those perennial species for which the floral biology has been
studied ( Triglochin maritinmum . T palustre, and T. striatum) are proterogynous,
the penicillate stigmas protruding from the perianth two to three days before
stamen maturation. Anthers of the lower whorl of stamens dehisce after the
stigmas have turned brown, their pollen falling not onto the stigmas of that
flower but into the deeply concave tepal below each stamen. The wind finally
carries the pollen away or shakes it onto more proximal flowers. The stamens,
each often attached to its tepal, eventually drop off. This sequence of events
is then repeated for the upper whorl of stamens. Pollen deposition on stigmas
of the same flower was reported for the Australian annuals studied by Keighery.
These species and at least T. maritimum, T. palustre, and T. striatum are
autogamous and show no significant difference in seed set between open pol-
lination and enforced selfing. In contrast, T. gaspense Lieth & Love, when grown
in a greenhouse, did not set seed except when pollinated by hand (Love &
Lieth).
Carpel number in Triglochin, although variable, is usually six (but as few as
three and as many as 12 have been observed in occasional flowers). In some
species (e.g., T. maritiimum) all six carpels are fertile, eventually falling as
achenelike mericarps from the persistent and terete or merely angled carpo-
phore. In others (e.g., T. palustre, T. striatum) the three carpels of the outer
whorl are sterile (even vestigial) and remain adnate to the thereby winged or
ridged carpophore after the inner, fertile carpels have fallen: they often then
assume "the appearance ofdissepiments of a capsule" (Bailey, 1902. p. 1705).
Of the species in subg. TRIGLOCHIN, only T. turrjnfra Ewart, one of the Aus-
tralian annuals, is said to lack a carpophore.
Chromosome counts, some undocumented, have been published for about
ten species of Triglochin. Triglochin procerum (subg. CYCNOGETON) has diploid,
tetraploid, and octoploid "morphological forms" (2n = 16, 32. 64; Robb &
Ladiges).
In subg. TRIGLOCHIN the most counted taxon is Triglochin maritimum agg.,
for which seven out of 12 possible numbers in a polyploid series from diploid
(12) to 24-ploid (144) have been noted, some from the Old World, some from
the New. (Not fitting into this series are counts of 2n = 30 [Bolkhovskikh et
al.] and 56 [Looman] in T. maritinmun, or = 75-77 [Gervais & Cayouette]
in T. clatum Nutt., a member of the T. maritimum aggregate.) According to
Love & Love (1958a. p. 20). 7. maritimun is "one of the most collective
species known and so heterogeneous that its inclusion in the so-called circum-
polar element ofdistribution is highly misleading." They ( 1958b, p. 162) further
stated that this "supposedly circumpolar [taxon] is in fact an aggregate of at
least more than a dozen good species."
Counts for 'riglochin palustrc, which shares with 1T. maritinmin a distri-
bution in both the Old and New worlds, are 2n = 12, 18, 24, 26. 28, 36. 48:
perhaps this taxon, too, is an aggregate (Hess et al.).
A strictly North American taxon for which a count is available is Triglochin
gaspense, 2n = 96. Counts for Old World taxa include '. Barrelieri Loisel.,
2n 30; 7. bulbosum L.. 2n = 18; and '. la.villorum Guss.. 2n = 18. 1 know
of no counts for either T. strialtu or any of the Australian annuals.
[VOL. 69


THIERET, JUNCAGINACEAE
Dispersal of species of Triglochin needs further study. Some are certainly
hydrochorous. Trig/ochin procerun has been seen to drop its disseminules into
the water of a stream, where they were carried away (Keighery). Pojar observed
floating seedlings of 7. maritirum, and mericarps of this species have been
collected from drift-line debris (Boorman). However, "seeds" of T. maritimum
(and of 7T palustre) float for less than a week in fresh water (perhaps a bit longer
in salt water), according to Praeger. Epizoochory in mud and anemochory have
been suggested for T. maritimum (Pojar).
Disseminules of species of Triglochin with mucronate, spurred, or even hooked
apices or bases (e.g., T. palustre and some of the Australian annuals) have been
assumed to be epizoochorous by attachment, an assumption that requires ver-
ification. Animal dispersal of the Australian annuals has been questioned by
Keighery, who found that in these plants, which occupy shallow, sandy, winter-
wet depressions, seed dispersal "has been reduced to a minimum ... ensuring
that the species will occupy the same favourable area next year" (p. 83). It
seems possible that in these annuals the structural devices that seem to fit them
for animal dispersal serve rather as "anchors" in favorable habitats.
Triglochin maritimumn and T. palustre, considered in some regions to be
"fairly" good forage, may be relished by livestock because of the salt content
of the leaves. In the western United States and western Canada, however, both
species (but especially 7. marilimum)n are among the major plants poisonous
to cattle and sheep, the toxic principle being hydrocyanic acid. Death results
from asphyxia or, in severe cases, respiratory failure. Triglochin striatum seems
not to have been implicated in poisoning; it is grazed by sheep in Chile (Al-
duante et al.) and is considered a good salt-marsh fodder in Australia (Bailey,
1883).
Species of Triglochin may be of importance as food for wildlife in some
regions. At James Bay, Canada, for example, the perennating bulbs of T. pa-
lustre were found to be the most strongly selected food item of snow geese
(Prevett et al.).
The young leaves of Triglochin maritimum--their dubious qualities not-
withstanding-can be cooked and eaten as a vegetable. They emit an unpleasant
odor when cooking ("an Chlor erinnerende"; Suessenguth, 1935, p. 214) and
are reputed to be a "blood-purifier." Ashes from this species are rich in sodium
carbonate and can be used in making soap (Fedchenko). The fruits were parched
and eaten by Oregon's Klamath Indians, who also roasted them as a substitute
for coffee (Coville). Triglochin procerum produces globose root tubers 2-3 cm
across (Van Steenis), of which Mueller (1867-1868, p. 83) said, "tubera edulia
nativis avide quaesita."
REFERENCES:
Also see family references, especially ARIBER 1923, 1925: ASTON: BAIILON: BENTHAM
& HOOKER: BRITTON: BUCHENAU. 1867. 1869, 1871, 1882. 1903, BUCHENAU &
HIERONYMUS: BURGER; (CHANT; (CHATIN; ( RONQUIST, 1981: DAHLGREN & CLIFFORD:
DAHLGREN, CLIFFORD, & YEO: ECKARDIT EI(CHLER; ERDTMAN: HEGI; HEGNAUER; HII.;
HITCHCOCK: HUTCHINSON: JOHRI: KART1SZ & KARTESZ: MARKGRAF. 1981: MASON: MI-
1988]


JOURNAL OF THE ARNOLD ARBORETUM
CHELu: MUELLER. 1867-1868: MtUENSCHFR: RENDLE; SUESSENGUTH: TOMLINSON, 1982:
and UHI..
ADAM, P. The ecological significance of'halophytes' in the Devensian flora. New Phytol.
78: 237-244. 1977. [Notes on present-day ecology of 7. maritimum.]
ADAMSON, R. S. The Cape species of T7rigiochin. Jour. S. Afr. Bot. 5: 29-31. 1939.
AHMAD, I., F. LARHER. A. F. MANN. S. F. M(NALLY, & (. R. STEWART. Nitrogen
metabolism of halophytes. IV. Charactcristics of glutamine synthetase from Tri-
glochin maritima 1 . New Phytol. 91: 585-595. 1982.
ADi .IANTE, C., J. J. ARMESTO, V. CASTRO, & C. VILLA(RAN. Ethnobotany of Pre-
Altiplanic community in the Andes of northern Chile. Econ. Bot. 37: 120-135. 1983.
[T. striatum, 131: forage for sheep.]
ANGELY, J. Flora analitica do Parana. 728 pp. (uritiba, Brazil. 1965. [Juncaginaceae,
134: 7. strialtm only.]
ARBER, A. The phyllode theory of the monocotyledonous leaf, with special reference to
anatomical evidence. Ann. Bot. 32: 465-501. 1918. [/'. maritimum, 478, 482, fig.
10: included among genera in which "phyllodic leaf anatomy has been observed
among monocotyledons."]
. Leaves of the lelobicae. Bot. Gaz. 72:31-38. pl 1. 1921. [Juncaginaceae. 34.]
.Leaves of Trilochin. Mid. 77: 50-62. p/s. 8-10. 1924. [7. striatum, 54. pl. 9,
figs. 11-13: anatomy and morphology.]
BAI.EY, F. M. A synopsis of the Queensland flora. 890 pp. Brisbane. 1883. [Triglochin.
572, 573; 7. striatum "a good salt-marsh fodder."]
The Queensland flora. Part VI. Alismaccae to Filicales. Pp. 1701-2015. Bris-
bane. 1902. [Triglochin. 1705-1707.]
The weeds and suspected poisonous plants of Queensland. 245 pp. Brisbane.
1906. [T. striatum, 206, /ig. 355: neither a weed nor a poisonous plant.]
BATE-SMITH, E. C. The phenolic constituents of plants and their taxonomic significance.
II. Monocotyledons. Jour. Linn. Soc. Bot. 60: 325-356. 1968. [7. palustre, 328:
chromatography of leaf hydrolysates.]
BEAI.. E. O. A manual of marsh and aquatic vascular plants of North Carolina. N.
Carolina Agr. Exper. Sta. Tech. Bull. 247. iv + 298 pp. 1977. [T. striatum, 55. 58.]
tBEAH, O. A., J. H. DRAIZE, & H. F. EPPSON. Arrow grass-chemical and physiological
considerations. Wyoming Agr. Exper. Sta. Bull. 193. 36 pp. 1933. [7. maritimum.]
BECHiET, M.-A., & P. BINFT. Croissance de Triglochin maritimum L. apres refrigeration
des semences imbihbes. Physiol. Veg. 2: 25-34. 1964.
IBELi, F. G. The occurrence of southern, steppe and halophyte elements in Weichselian
(last-glacial) floras from southern Britain. New Phytol. 68: 913-922. 1969. ([. mari-
imummn in deposits 41.900 * 800 to 45.000 - 1890 years old.]
BINET, P. Dormances primaire et secondaire des semences de Triglochin maritimum
L.: action du froid et de la lumiere. Bull. Soc. Linn. Normandie. IX. 10: 131-142.
1959.
. L'eau de mer et Ie froid, facteurs de levee de la dormance pour les semences
de Trig/lchin mariiunum L. (Compt. Rend. Acad. Sci. Paris 251: 422-424. 1960.
Rapports entre l'eau de mer et la germination des semences de Triglochin
imaritimum L. Bull. Soc. Linn. Normandie, X. 1: 117-132. 1961a.
Acquisition de I'aptitude a germer en milieu sale par les semences de Triglochitn
iaritimuo L. Ibid. 2: 124-128. 1961 b.
Les semences de 7rig/ochin pailstri L. et de Triglochin manritilmu L.: etude
comparce de leur germination. Ibid. 148-160. 1962.
Bi A(CK. J. M. Flora of South Australia. Part 1. Cyatheaceae-Orchidaceae. 253 pp. Ade-
laide. 1960. 1Triglochin, 49-51.]
li ANiKSMA, . J.. Blauw/suur in /outgras (l'rig/lochlin). Pharm. Weekbl. 50: 1295-1302.
1913. [HCN.]
BOi KHOVSKIKH, Z., V. (RiF, T. MATVEIJEVA, & O. ZAKHiARYEVA. Chromosome numbers
[vot.. 69


THIERET, JUNCAGINACEAE
of flowering plants. A. A. FEDEROV, ed. (Russian and English prefaces.) 926 pp.
Leningrad. 1969. [Juncaginaceae, 361. 362: includes most chromosome counts through
1964.]
BOORMAN, L. A. Some aspects of the reproductive biology of Limonium vulgare Mill.
and Limonium humile Mill. Ann. Bot. II. 32: 803-824. 1968. IT. maritimum "seeds"
in drift-line debris, 821, 822.]
BOUTARD, B., M.-L. BOUILLANT, J. CHOPIN, & P. LEBRETON. Chimiotaxinomie flavo-
nique des Fluviales [Helobiae]. Biochem. Syst. 1: 133-140. 1973. [Triglochin fla-
vonoid patterns, tables II, I .]
BOYD, L. Monocotylous seedlings. Trans. Proc. Bot. Soc. Edinburgh 31: 1-224. 1932.
[7. maritimurn, 13-15, fig. 3, ix-xi.]
BRUNKENER, L. Beitrige zur Kenntnis der friihen Mikrosporangienentwicklung der An-
giospermen. Svensk Bot. Tidskr. 69: 1-27. 1975. [Triglochin. 20.]
BUCHANAN, J. On the flowering plants and ferns of the Chatham Islands. Trans. Proc.
New Zealand Inst. 7: 333-341. pls. 12-15. 1875. [7. triandrum (i.e., T. striatum),
340.]
BUCHENAU, F. [Ueber die Sprossverhiltnisse in der Gattung Triglochin.] Amtl. Ber.
Versamml. Deutsch. Naturf. Arzte 40: 178-180. 1886.
SUebersicht der in den Jahren 1855-57 in Hochasien von den Briidern Schlagint-
weit gesammelten Butomaceen, Alismaceen, Juncaginaceen und Juncaceen. Nachr.
Kanigl. Ges. Wiss. Georg-Augusts-Univ. 1869(13): 237-258. 1869. [T. maritimum
in Tibet, 241, 242.]
Ein Fall von Saison-Dimorphismus in der Gattung Trig/ochin. Abh. Naturw.
Ver. Bremen 13: 408-412. 1896.
BUCHET. S., & C.-L. GATIN. Un cas de polyembryonie chez le 7riglochin palustre L. et
une germination anormale de l'. risarum rulgare Targ.-Tozz. Bull. Soc. Bot. France
55: 164-169. 1908. [T. palustre, 164, 165.]
BURKE, F. An inland record of Triglochin maritimum L. Nature 150: 405. 1942. [From
a salt spring in Cheshire: nonlittoral occurrence exceptional.]
CABRERA, A. L. Juncaginaceae. In: A. L. (ABRERA. ed.. Fl. Prov. Buenos Aires 1: 291-
295. 1968. [Including Triglochin palustre, T. striatum, Lilaea scilloides.]
CASTRO, D., & F. C. FONTES. Primeiro contacto citologico com a flora hal6fila dos
salgados de Sacavem. Broteria Ci. Nat. 15: 38-46. 1946. [T. Barrelieri. 40; n = 15.]
CHAPMAN, V. J. Salt marshes and salt deserts of the world. 392 pp. London. 1960.
[Triglochin, passim: many mentions.]
CHARLTON. W. A. Studies in the Alismataceae. I. Developmental morphology of Echi-
nodorus tenellus. Canad. Jour. Bot. 46: 1345-1360. 1968. [T. maritimum, 1358.]
Features of the inflorescence of Triglochin maritima. Ibid. 59: 2108-2115. 1981.
[The inflorescence has a number of characters "more usually associated with flowers."]
CHAUVEAUD, G. Recherches sur le mode de formation des tubes cribles dans la racine
des monocotyledones. Ann. Sci. Nat. Bot. VIII. 4: 307-381. pls. 4-9. 1897. [T.
palustre, 362, pl. 8, fig. 29.]
Sur le passage de la disposition alterne des elements liberiens et ligneaux a leur
disposition superpos~e dans le trocart (Triglochin). Bull. Mus. Hist. Nat. Paris 7:
124-130. 1901. [Root and stem anatomy.]
CHEADLE, V. I. The occurrence and types of vessels in the various organs of the plant
in the Monocotyledoneae. Am. Jour. Bot. 29: 441-450. 1942. [Including T mari-
timum.]
CHEESEMAN, T. F. Manual of the New Zealand flora. ed. 2. 1163 pp. Wellington. 1925.
[Triglochin, 125, 126.]
CLARKE, L. D., & N. J. HANNON. The mangrove swamp and salt marsh communities
of the Sydney [Australia] district. II. Plant growth in relation to salinity and wa-
terlogging. Jour. Ecol. 58: 351-369. 1970. [T. striatum. passim; autecology.]
1988]


JOURNAL OF THE ARNOLD ARBORETUM
CLAUSEN, P. Ueber das Verhalten des Antheren-Tapetums bei einiger Monocotylen und
Ranales. Bot. Arch. 18: 1-27. 1927. ['1. maritimum, 9. figs. 18, 19.]
CLAWSON, A. B.. & E. A. MORAN. Toxicity of arrowgrass for sheep and remedial treat-
ment. U.S. Dep. Agr. Tech. Bull. 580. 16 pp. 1937. [T. maritimum.]
CLEWELL, A. F. Guide to the vascular plants of the Florida Panhandle. 605 pp. Talla-
hassee. 1985. ['. striatum, 175.]
COOPER, A. The effects of salinity and waterlogging on the growth and cation uptake of
salt marsh plants. New Phytol. 90: 263-275. 1982. [Includes data on Triglochin.]
(ORDEMOY, J. DE. OrganogCnie des Triglochin. Adansonia 3: 12-14. 1862-1863. [Floral
morphology.]
CORRELL, D. S., & H. B. CORRELL. Aquatic and wetland plants of southwestern United
States. Frontisp. + xvi t 177 pp. Environmental Protection Agency, Washington,
D. C. 1972. (Reissued in 2 vols. by Stanford Univ. Press, 1975.) [Juncaginaceae,
129-133, fig. 54; Triglochin.]
- & - . Flora of the Bahama archipelago. 1692 pp. Vaduz. 1982. [Juncagi-
naceae (T. striatum), 80, fig. 34.]
COVILLE, F. V. Notes on the plants used by the Klamath Indians of Oregon. Contr.
U. S. Natl. Herb. 5: 87-108. 1897. [1. maritimum. 90.]
CRANWELL, L. M. New Zealand pollen studies. I. Key to the pollen grains of families
and genera in the native flora. Rec. Auckland Inst. Mus. 2: 280-308. 1942. [Tri-
glochin. 293.]
(ROW, (. E., & C. B. HELLQUIST. Aquatic vascular plants of New England: part 4.
Juncaginaceae, Scheuchzeriaceae, Butomaceae, Hydrocharitaccac. New Hampshire
Agr. Exper. Bull. 520. 20 pp. 1982. [Juncaginaceae. 2, 3; T'riglochin.]
CUNNINGHAM, G. M.. W. E. MULHAM, P. L. MILTHORPE, & J. H. LEIGH. Plants of
western New South Wales. 761 pp. [Sydney?] 1981. [Juncaginaceae, 42, 43: pho-
tographs of Triglochin calcitrapa Hooker. T. hexagonum J. M. Black, 7. dubium
R. Br., and 7. procerum R. Br.]
CUTLER, A. J.. W. HiSEL, M. STERNIBERG, & E. E. CONN. The channeled biosynthesis
of taxiphyllin in Triglochin maritima. (Abstract.) Pl. Physiol. (Suppl.) 65(6): 27.
1980.
DANDY. J. E. Juncaginaceae. In: K. H. RECHINGER, ed., Fl. Iranica, no. 83.3 pp. 1971.
[Triglochin.]
SJuncaginaceae. Fl. Europaea 5: 6, 7. 1980. [Triglochin, 7: includes 7. striatum,
"naturalized in W. Portugal."]
DELAY, J. Triglochin maritlmum L.: etude ultrastructurale des caryocineses. Inform.
Ann. Caryosyst. Cytogendt. 6: 25-32.*
DEWIT, H. C. D. Aquarium plants. 255 pp. London. 1964. [Triglochin, 93-95.]
DYER, R. A. The genera of southern African flowering plants. Vol. 2. Gymnosperms
and monocotyledons. Pp. 757-1040. Pretoria. 1976. [Juncaginacac. 776. 777: Tri-
glochin.]
EAMES. A. J. Morphology of the angiosperms. 518 pp. New York. 1961. [Triglochin,
passim.]
EBER, E. 1934. Karpellbau und Plazentationsverhiltnisse in der Reihe der Helobiac.
Flora 127: 273-330. 1934. [Triglochin, 301-305, figs. 15, 16.]
ELEIrrERIUS, I. N. Tidal marsh plants. 131 pp. Place of publication not indicated. 1980.
[T. striatum, 32, 33.]
EMBERGER, L. Les plantes fossiles dans leurs rapports avec les vegetaux vivants. 758
pp. Paris. 1968. [Triglochin, 606, fig. 716.]
EYJOi FSSON, R. Investigations on cyanogenic plants. (In Danish: English abstract.) Dansk
Tidsskr. Farm. 42: 301-307. 1968. [Including '. maritimum.]
Isolation and structure determination of triglochinin, a new cyanogenic glucoside
from Triglochin maritimunm. Phytochemistry 9: 845-851. 1970.
FEDCHENKO, B. A. Juncaginaceae Lindl. In: V. L. KOMAROV. ed.. Fl. USSR 1: 216-218.
1968. [Triglochin; English translation by N. LANDAU of Fl. URSS 1: 275-279. 1934.]
[VOL. 69


THIERET, JUNCAGINACEAE
FELDMANN, G. Une nouvelle espece de plasmodiophorale parasite du Triglochin mari-
timum L.: Plasmodiophora maritima nov. sp. Rev. Gen. Bot. 65: 634-650. 1958.
FERNALD, M. L. Some variations of Triglochin maritima. Rhodora 5: 174, 175. 1903.
SPreliminary lists of New England plants.-XXII. Ibid. 10: 168-172. 1908.
[Includes notes on Triglochin.]
SA century of additions to the flora of Virginia. Ibid. 42: 355-416, 419-498,
503-521.pls. 626-649. 1940. [T striatum, 387, 388, 407, 506-508. map 19(p. 388;
world distribution of T. striatum).]
-- & A. C. KINSEY. Edible wild plants of eastern North America. (Revised by
R. C. ROLLINS.) 452 pp. New York. 1958. [T. maritima, 60, 86; "The fresh seeds
have a strong taste or odor suggesting watchmaker's oil or turpentine."]
FLEMING, C. E., & N. F. PETERSON. Arrow-grass. A new stock-poisoning plant (Triglochin
maritima). Nevada Agr. Exper. Sta. Bull. 98. 21 pp. 1920.
FLOWERS, T. J., M. A. HAJIBAGHERI, & N. J. W. CLIPSON. Halophytes. Quart. Rev. Biol.
61: 313-337. 1986. [Triglochin, passim; physiology.]
FREIDENFELT, T. Der anatomische Bau der Wurzel in seinem Zusammenhange mit dem
Wassergehalt des Bodens. Bibliot. Bot. 12(61). 118 pp. 5 pls. 1904. [T. palustre, 36,
37, p1. 2, fig. 29, p1. 3, fig. 31.]
FRENCH, J. C. Stem elongation in the Helobiae. Helobiae Newsletter 1: 11. 1976. [Data
on Triglochin.]
GANDOGER, M. Sertum plantarum novarum. Pars secunda (suite). Bull. Soc. Bot. France
66: 286-307. 1919. [T. floridanum, sp. nov., 293, type from Florida; T. sessile, sp.
nov., 293, type from Mississippi; both T. striatum.]
GARDNER, R. O. Binucleate pollen in Triglochin L. New Zealand Jour. Bot. 14: 115,
116. 1976. [T. striatum.]
GASCOIGNE, R. M., E. RITCHIE, & D. E. WHITE. A survey of anthocyanins in the Aus-
tralian flora. Jour. Proc. Roy. Soc. New S. Wales 82: 44-70. 1949. [Anthocyanin
present in very small amount in flowers of T. procerum, 55.]
GERARD, R. Recherches sur le passage de la racine a la tige. Ann. Sci. Nat. Bot. VI. 11:
279-430. pls. 16-19. 1881. [T. palustre, 404, 405, pl. 19, figs. 67, 68.]
GERVAIS, C., & J. CAYOUETTE. Liste annotee des nombres chromosomiques de la flore
vasculaire du nord-est de l'Amirique. IV. Nat. Canad. 112: 319-331. figs. 62-87.
1985. [7: elatum, 328, 329, fig. 85: n = 75-77.]
GIBSON, R. J. H. The axillary scales of aquatic monocotyledons. Jour. Linn. Soc. Bot.
37: 228-236. pls. 5. 6. 1905. [Tniglochin, 230.]
GLOCK, H. Die Stipulargebilde der Monokotyledonen. Verh. Naturhist.-Med. Vereins
Heidelberg, II. 7: 1-96. pls. 1-5. 1902. [Triglochin, 29.]
GOAS, G., M. GOAS, & F. LARHER. Formation de l'acide pipecolique chez Triglochin
maritima. Canad. Jour. Bot. 54: 1221-1227. 1976.
GODFREY, R. K., & J. W. WOOTEN. Aquatic and wetland plants of southeastern United
States. Monocotyledons. 712 pp. Athens, Georgia. 1979. [Juncaginaceae, 43; T.
striatum.]
GRESHOFF, M. Een nieuwe natuurlijke groep van blauwzuurplanten: de Juncaginaceae.
Pharm. Weekbl. 45: 1165-1171. 1908. [Cyanogen in Trigiochin and Scheuchzeria.]
GROENEDIJK, A. M. Primary production of four dominant salt-marsh angiosperms in
the SW Netherlands. Vegetatio 57: 143-152. 1984. [One of the four is T. maritimum.]
GUILLAUD, A. Recherches sur I'anatomie comparee et le developpement des tissus de
la tige dans les Monocotyledones. Ann. Sci. Nat. Bot. VI. 5: 5-176. p/s. 1-6. 1878.
[T. maritimum, 77-85, pl. 4, figs. 3, 4.]
Guppy, H. B. Observations of a naturalist in the Pacific between 1896 and 1899. 627
pp. London. 1906. [Triglochin, 34, 36, 537, 541, 545; flotation experiments.]
GUTTENBERG, H. VON. Embryologische Studien an Monokotyledonen. III. Die Embryo-
genese von Triglochin maritimum L.. Arum maculatum L. und Tvpha latifoiia L.
Flora 149: 243-281. figs. 1-37. 1960. [7T maritimum, 243-255, figs. 1-13.]
1988]


JOURNAL OF THE ARNOLD ARBORETUM
HARA. H. Notes on Triglochin maritinmui of Japan. (In Japanese: English summary.)
Jour. Jap. Bot. 35: 190-192. 1960. [2n = 120.]
Racial differences in widespread species, with special reference to those common
to Japan and North America. Am. Jour. Bot. 49: 647-652. 1962. [7. maritimum,
649.]
HAYNES, R. R. Aquatic and marsh plants of Alabama. I. Alismatidae. Castanca 45: 31-
51. 1980. [7. striatuim, 41. fg. 2a.]
HEGNAUER, R. Die cyanogenen Verbindungen der Liliatac und Magnoliatae-Magnoli-
idae: zur systcmatischen Bedeutung des Mcrkmals der Cyanogenese. Biochem. Syst.
1: 191-197. 1973. [Triglochin, passim.]
HEMSLEY, W. B. The flora of Tibet or high Asia. Jour. Linn. Soc. Bot. 35: 124-265.
1902. [T. maritinum, T. palustre, passim; growing at 15,000-16,000 ft.]
HE SLOP-HARRISON, Y.. & J. HESLOP-HARRISON. Ring formation by Triglochin maritima
in eastern Irish salt marsh. Irish Nat. Jour. 12: 223-229. 1958.
--- & K. R. SHIVANNA. The receptive surface of the angiosperm stigma. Ann. Bot.
41: 1233-1258. 1977. [Trilochin. 1237: stigmatic surface dry, with unicellular pa-
pillae.]
HEss. H. E., E. LANDOilT, & R. HIRZEL. Flora der Schweiz und angrenzender Gebiete.
Band I. Pteridophyten bis Caryophyllaceae. 858 pp. Basel and Stuttgart. 1967. [Jun-
caginaceae. 209-211.]
HFISSER, C. J. Palynology of cushion bogs of the Cordillera Pelada, Province of
Valdivia, Chile. Quaternary Res. 17: 71-92. 1982. [Fossil pollen of Tetronciunm, 10,
424 B.P.]
HORN, P. Beitrage zur Kenntnis der Triglochin-Biithe. Arch. Ver. Freunde Naturgesch.
Mecklenburg 30: 120-136. pi. 2. 1876. [Morphology.]
HORN AF RANTZIEN. H. Notes on the African species of Triglochin. Svensk Bot. Tidskr.
55:81-117. 1961.
HOTCHKIss, N. Common marsh plants of the United States and Canada. 1. S. Dep.
Interior. Fish Wildl. Serv. Bur. Sport Fish Wildlife. Resource Publ. 93. 99 pp. 1970.
[Triglochin, 1 1.]
HOWEI.L, J. T. Remarks on Triglochin cocinnaa. Leafl. West. Bot. 5: 13-19. 1947.
HItLTEN, E. The circumpolar plants. I. Vascular cryptogams, conifers, monocotyledons.
Sv. Vet.-Akad. Handl. IV. 8(5): 1-275 [+ 5 unnumbered pp.]. 1964. [7T palustre,
112, map 104: '. maritimunm, 120, 248, map 112.]
HuiiTN. E.. & K. TORSELI.. Alkaloid-screening of Swedish plants. Phytochemistry 4:
425-433. 1965. [T. maritirunm, 430: alkaloid content too low to warrant further
study.]
IRMISC H, T. Ueber das Vorkommen von schuppen- oder haarformigen Gebilden in-
nerhalb der Blattscheiden bei monokotylischen Giewachsen. Bot. Zeit. 16: 177-179.
I858. [Data on Trig/ochin.]
JAvRi. S. M. H. Juncaginaceae. No. 48 in E. NASIR & S. I. ALI, eds., Fl. W. Pakistan. 5
pp. 1973. ['Trnilochin.]
Juncaginaceae. No. 11 in S. 1. ALI & S. M. H. JAFRI, eds., Fl. Libya. 4 pp. 1977.
[ Triglochin.]
JAROSZEWSKI, J. W.. & M. G. ETTLINGER. Ring cleavage of phenols in higher plants:
biosynthesis of triglochinin. Phytochemistry 20: 819-821. 1981. [In Triglochin; com-
pound also in Thalictrum aquil/gifolium.]
JEFFRIES, R. L., A. JENSEN. & K. F. ABRAHAM. Vegetational development and the effect
of geese on vegetation at La Perouse Bay, Manitoba. Canad. Jour. Bot. 57: 1439-
1450. 1979. [7'. maritinnor, passim: ecology.]
JENNIN;S. O. E. Note on the occurrence of Triglochin palustris Linnaeus in Pennsylvania.
Ann. Carnegie Mus. 3: 482. 1906.
JOHNSON. P. N., & D. J. CAMPBELL. Vascular plants of the Auckland Islands. New
Zealand Jour. Bot. 13: 665-720. 1975. ['. striatum at 50�S.]
[voL. 69


1988] THIERET, JUNCAGINACEAE 19
JONES, S. B., JR. Mississippi flora. I. Monocotyledon families with aquatic or wetland
species. Gulf Res. Rep. 4: 357-379. 1974. [T. striatum. 367: map of distribution in
Mississippi, 365.]
KEIGHERY, G. J. Breeding systems of the Western Australian flora, I. Triglochin L.
(Juncaginaceae). West. Austral. Nat. 13(4): 81-84. 1975.
KERNER VON MARILAUN, A., & F. W. OLIVER. The natural history of plants. 983 pp.
New York. 1895. [Triglochin, 149, 150, ig. 479 (p. 874).]
KINGSBURY, J. M. Poisonous plants of the United States and Canada. 626 pp. Englewood
Cliffs, New Jersey. 1964. [Triglochin, 501-503,, ig. 130.]
KNUTH, P. Handbook of flower pollination. (English translation by J. R. AINSWORTH
DAVIS.) Vol. 3. 644 pp. Oxford. 1909. [Triglochin, 506.]
KOLKWITZ, R. Ueber die Standorte der Salzpflanzen. Ber. Deutsch. Bot. Ges. 35: 518-
526. 1917. [Triglochin, passim.] III. Triglochin maritima. Ibid. 37: 343-347. 1919.
LAGERBERG. T. Vilda vaxter i norden. ed. 3. Band I. Polypodiaceae-Myricaceae. 455
pp. Stockholm. 1956. [Juncaginaceae, 124-128, figs. 64-67: Triglochin.]
LEON, HERMANO. Juncaginaceae. Fl. Cuba 1: 78, 79. 1946. [T. striatum.]
LIEU, S. M. Organogenesis in Triglochin striata. Canad. Jour. Bot. 57: 1418-1438. 1979.
LoGx;E. R. W., A. McLEAN. & A. JOHNSTON. Stock-poisoning plants of western Canada.
Canada Dep. Agr. Publ. 1361. 34 pp. 1968. [Triglochin, 5-7: a "major trouble-
maker."]
LOEw, E. Juncaginaceac. Pp. 556-584 in O. VON KIRCHNER. E. LorE, & C. SCHROTER.
Lebensgeschichte der Blutenptlanzen Mitteleuropas. Band I, Abteilung 1. Stuttgart.
1908. [Triglochin. 571-584, figs. 310-323.]
LONG, R. W., & O. LAKELA. A flora of subtropical Florida. 962 pp. Coral Gables, Florida.
1971. [T. striatum, 123, 124 (rather poor illustration).]
LOOMAN. J. Biological flora of the Canadian prairie provinces. IV. Triglochin L.. the
genus. Canad. Jour. P1. Sci. 56: 725-732. 1976.
LOVE, A., & D. LOVE. An unusual polyploid series in Triglochin maritimum agg. Genet.
Soc. Canada Proc. 3(2): 19-21. 1958a.
S& - . Biosystematics of Triglochin maritimum agg. Nat. Canad. 85: 156-
165. 1958b.
LOVE, D., & H. LIETH. Triglochin gaspcnsc, a new species of arrow grass. Canad. Jour.
Bot. 39: 1261-1272. 1961.
MABBOTT, D. C. Food habits of seven species of American shoal-water ducks. U. S.
Dep. Agr. Bull. 862. 67 pp. 1920. ["Seeds" of 7. maritimum eaten by baldpate and
pintail ducks, 14, 33.]
MACBRIDE, J. F. Flora of Peru. Part I. Field Mus. Nat. Hist. Bot. Ser. 8. 320 + ii pp.
1936. [Scheuchzeriaceae. 90: Lilaca and Triglochin. including T. striatum.]
MACLEOD, J. Bestuiving en insectenbezoek der blomen van het kempisch gedeelte van
Vlaanderen. Bot. Jaarb. 5: 282-452. 1893. [T. palustre, 291: species is proterogynous
and anemophilous.]
MARKGRAF, V., & H. L. D'ANTONI. Pollen flora of Argentina. 208 pp. 43 pls. Tucson.
1978. [T. palustre and T. striatum, 43, pl. 6-64 (T. palustre).]
MARSH, C. D.. A. B. CLAWSON, & G. C. Roi. Arrow grass (Triglochin mnaritima) as a
stock-poisoning plant. U. S. Dep. Agr. Tech. Bull. 113. 14 pp. 1929.
MARTIN. A. C. The comparative internal morphology of seeds. Am. Midi. Nat. 36: 513-
660. 1946. [Juncaginaceae (Scheuchzcria, Triglochin), 550: no endosperm.]
MASON. E. Notes on the presence of mycorrhiza in the roots of salt marsh plants. New
Phytol. 27: 193-195. 1928. [None seen in 7. maritimum, 195.]
MCATEE. W. L. Food habits of the mallard ducks of the United States. U. S. Dep. Agr.
Bull. 720. 36 pp. 1918. [7. maritimnm (part not specified) a "minor item" found
in stomachs of the common mallard.]
McDONNELL. S. M. The taxonomy and life history of Triglochin procera R. Br. B.Sc.
(Honors) thesis, University of Sydney. Australia. 1969.*


JOURNAL OF THE ARNOLD ARBORETUM
MESSERI. E. Richerche sullo sviluppo del sislema vascolare in alcuni monocotiledoni.
Nuovo Giorn. Bot. Ital. 11. 32: 317-362. p/s. 15-20. 1925. [T. laviflorum, 339. 340,
pl. 19, /i,. 55.1
MICHAUX, A. Flora Boreali-Americana. Vol. 1. 330 pp. Paris. 1803. [Description of T.
striatum (as . triaunruwm). 208.]
MIKKEISEN, V. M. The distribution within Denmark of the Juncaginaceae, Alismataceae.
and Hydrocharitaceac. Bot. Tidsskr. 47: 65-93. figs. [maps] 1-14. 1943. [T. mari-
limum, 70. 71. 87. fig. 3: '. palustre. 71, 72, 87. f/i. 4: mericarps of T. imaritimnum
have been found in lateglacial Drwas clay.]
MIRHEL. M. Examen de la division des vegetaux en endorhizes el exorhizes. Ann. Mus.
Hist. Nat. Paris 16: 419-458. p/s. 16-21. 1810. [1'. maritimum, 435, p/. 16 (ger-
minating "seed").]
MOHR, C. Plant life of Alabama. ('ontr. I'. S. Natl. Herb. 6.921 pp. 1901. [T1 striatumi,
128.]
Motl IARD, M. I.ne nouvelle Plasmodiophorce, parasite du Triglochin pulustre L. Bull.
Soc. Bot. France 56: 23-25. 1909. [7Ttramyxa triglochinis.]
MONTFORT, C. Die Xeromorphic der Hochmoorpflanzen als Voraussetzung der "phy-
siologischen Trockenheit" der Hochmoore. Zeitschr. Bot. 10: 257-352. 1918. [7.
pu/lustre. 316. 317.]
MOORE, L. B.. & E. EDGAR. Flora of New Zealand. Vol. 2. Indigenous Tracheophyta.
Monocotyledones except Gramineae. 354 pp. Wellington. 1970. [Juncaginaceae. 7,
8: including 7. striatum.]
MOORE. R. J.. ed. Index to plant chromosome numbers 1967-1971. Reg. Veg. 90. 539
pp. 1973. [Tri'ilochin, 49.]
- ed. Index to plant chromosome numbers for 1972. Ibid. 91. 108 pp. 1974.
[Juncaginaceae. 8.]
, ed. Index to plant chromosome numbers for 197374. Ibid. 96. 257 pp. 1977.
[Juncaginaceae. 128.]
MoRON(. T. The Naiadaceae of North America. Mem. Torrex Bot. Club 3(2): 1-65.
p/s. 20-74. 1893. [Tri,/ochin, 5-8. pls. 20-22.]
M;EINSCHER, W. (. Poisonous plants of the United States. 277 pp. New York. 1962.
[Juncaginaceae, 32-34. fi,. 4: Tri'g/lchin.]
MuNz, P. A. Supplement to a (alifornia flora. 224 pp. Berkeley. 1968. [Triglochin. 172.]
S& D. D. KECK. A California flora. 1681 pp. Berkeley. 1959. [Juncaginaceae.
1320. 1321.]
NAHRSTEDT, A., W. HOSEL, & A. WAALHER. (haracterization ofcyanogenic glucosides
and i-glucosidases in Triglochin maritiia seedlings. Phytochemistry 18:1 137-1141.
1979. [In addition to triglochinin. taxiphyllin has been detected as a cyanogenic
glucoside.]
--- . J. D. KAN . & W. HlSEt. Aspects of the biosynthesis of the cyanogenic glucoside
triglochinin in Triglo/chin maritima. PI. Med. 50: 394-398. 1984.
NAPPER, D. M. Juncaginaceae. In: W. B. Tt RRII. & E. MILNE-REDHEAD, eds., FI. Trop.
E. Africa. 3 pp. 1971. [Trig/ochin.]
NORTON, A. H. Notes on Trig/lochin pa/ulrins and Montia lamprosperma in Maine.
Rhodora 35: 291, 292. 1933.
)BERMEYER, A. A. Juncaginaceae. In: L. E. CoinD & B. DEWINTER. eds.. Fl. Southern
Africa 1: 92-95. 1966. [7Trigi,,hin.]
)STENFELD, C. H. A revision of the West Australian species of Triglochin. Crassula
(Tillaea) and Irankenia. Dansk Bot. Ark. 2(8): 30-55. pl. 4. 1918. [Triglochin. 30-
39, annual species only: p/. 4 ("Australian Annual Triglochins").]
PACKER, J. G. Chromosome numbers and taxonomic notes on western Canadian and
arctic plants. (anad. Jour. Bot. 42: 473-494. 1964. [lrig/ochin, 477, 478, fg. 1; n =
24.]
PARK. .. Notes on salt marsh plants III. T'riglochin maritimum Linn. Trans. Proc. Bot.
[VOL. 69


THIERET, JUNCAGINACEAE
Soc. Edinburgh 30: 320-325. 1931. [Leaf anatomy, germination, seedling growth
and anatomy.]
PERRIER DE LA BATHIE, H. Scheuchzeriacees (Scheuchzeriaceae). Famille 24 in H.
HUMBERT, Fl. Madagascar Comores (Pl. Vasc.). 3 pp. I fig. 1946. [7. striatum.]
POJAR, J. Reproductive dynamics of four plant communities of southwestern British
Columbia. Canad. Jour. Bot. 52: 1819-1834. 1974. [T. maritimum, passim: wind-
or self-pollinated, markedly proterogynous, hydrochorous (floating seedlings), ane-
mochorous. epizoochorous in mud.]
PoPP, M., & R. ALBERT. Freie Aminosauren und Stickstoff-Gehalt in Halophyten des
Neusiedlersee-Gebietes. Flora 170: 229-239. 1980. [Proline in T. mnaritimum.]
PRAEGER, R. L. On the buoyancy of the seeds of some Britannic plants. Sci. Proc. Roy.
Dublin Soc. 14: 13-62. 1913. [Triglochin, 41, 54.]
PREVETT, J. P., I. F. MARSHALL, & V. G. THOMAS. Spring foods of snow and Canada
geese at James Bay. Jour. Wildlife Managem. 49: 558-563. 1985. ["Bulbs" of T.
palustre the "most strongly selected food item."]
RAINHA, B. V. A Triglochin laxillora Guss. em Portugal. Bol. Soc. Brot. II. 19: 289-
294. 1944. [Four species of Trig/ochin, including T. striatum, in Portugal.]
REIMOLD, R. J., & W. H. QUEEN. Ecology of halophytes. 605 pp. New York. 1974. [T.
maritiimum, 274-277.]
REUTER, E. Ueber Vorblattbildung bei Monokotylen. Flora 110: 193-261. 1918. [T.
maritimum, 197, figs. 4-7.]
REVEAL, J. Juncaginaceae. In: A. CRONQUIST et al., Intermountain Fl. 6: 18-21. 1977.
[Triglochin.]
RICHARD, M. Analyse botanique des embryons endorhizes ou monocotyledones, et
particulierement de celui des Graminees. Ann. Mus. Hist. Nat. Paris 17: 223-251.
pls. 5-10. 1811. [T. maritimum, 231. p/. 5, fig. 24 (longisection of "seed," germi-
nating "seed").]
RIDLEY. H. N. The dispersal of plants throughout the world. xx + 744 pp. Ashford,
Kent. 1930. [Triglochin. 190, 231. 490, 492.]
RILEY. H. P. Families of flowering plants of southern Africa. 269 pp. Lexington, Ken-
tucky. 1963. [Juncaginaceae, 193, 194: Triglochin.]
RINGIUS, G. S. Vegetation survey of a James Bay coastal marsh. Canad. Field-Nat. 94:
110-120. 1980. [T. maritimum, passim: phytosociology.]
ROBB, L., & P. Y. LADIGES. Morphological forms and polyploidy in Triglochin procera
agg. in Victoria. Austral. Jour. Bot. 29: 639-651. 1981. [2n = 16, 32, 64.]
ROSS-CRAIG, S. Drawings of British plants. Part 31. 31 pls. - index of 2 unnumbered
pp. London. 1973. [Juncaginaceae. p/s. 13 (T. maritimum). 14 (7. palustre).]
RZEDOWSKI. J. Extinci6n de cspccics vegetales. Pp. 43-45 in J. RZEDOWSKI & G. C. DE
RZEDOWSKI. Flora fanerogamica del Valle de Mexico. Mexico, D. F. 1981. [T. stria-
lum listed among species that are endangered.]
SANCHEZ SANCHEZ. O. La flora del Valle de Mexico. 519 pp. Mexico, D. F. 1968. [T.
striatum, 28.]
SCHNARF, K. Kleine Beitrage zur Entwicklungsgeschichte der Angiospermen. V. Ueber
zwei kritische Falle der Endospermentwicklung (V1erbena und Triglochin). Oesterr.
Bot. Zeitschr. 74: 40-50. 1925. [7. maritinum, 45-50. pi. 2, figs. 1-8: embryo-sac
development.]
Embryologic der Angiospermen. Band X/2 in K. LINSBAUER. ed., Handbuch
der Pflanzenanatomie, II. Abteilung. 2. Teil: Archegoniaten. 689 pp. Berlin. 1929.
[Triglochin. passim.]
Variation im Bau des Pollenkornes der Angiospermen. Tabul. Biol. 17: 72-89.
1939. [T. palustre, 81: 2-nucleate pollen.]
SCHUMANN, K. Neue Untersuchungen iber den Blutenanschluss. 519 pp. 10 p/s. Leipzig.
1890. [Triglochin. 141, 160. p/. 5. figs. 3-5.]
1988]


JOURNAL OF THE ARNOLD ARBORETUM
SCIITHORPE, C. D. The biology of aquatic vascular plants. xviii 4 610 pp. London.
1967. [Juncaginaccac. mainly Trigilohin. passim.]
SERBANESCII-JITARII. G. Betrachtungen uber Gynaizeum. Frucht und Keimung bei Tri-
,l'chin maritinium IL. und T'nglochin palustre L. Revue Roum. Biol. Ser. Bot. 18:
9-20. 1973.
SEiUERT, M. Alismaccae. In: C. F. P. VON MARTIuS. Fl. Brasil. 3(1): 101-112. 1847.
[Tribus I. Juncagineae Rich., 103, pl. 12 (7. montevidense).]
SHARMA, A. Chromosome census of the plant kingdom. I, Monocotyledons. Part 1.
Butomales to Zingiberales. Nucleus 15(ii): 1-20. 1972. [Juncaginaceae, 3.]
SiNnH, V. Development of gynoecium of 'riglochin in three dimensions. (urr. Sci.
Bangalore 42: 813-815. 1973.
SMALL, J. K. Manual of the southeastern flora. 1554 pp. New York. 1933. (Reprinted
by Univ. N. Carolina Press. [1961].) [Scheuchzeriaceae, 20: 7. striatum.]
SoIF'GES, R. Embryogenie des Scheuch/eriacees. Developpement de l'embryon che/ le
Trigochin imaritima L. Compt. Rend. Acad. Sci. Paris 216: 746-748. 1943.
SPEuHT, R. L. The vegetation of South Australia. ed. 2. 328 pp. Adelaide. 1972. [l'i,.
'2, photograph of 7. procera growing with 7I phu.]
STAUDERMANN. W. Die Haare der Monocotyledonen. Bot. Arch. 8: 105-184. 1924.
[ T'ri/)c/hin, 106; review of squamules.]
STEENIS. C. (i. C. J. VAN. JuncaginaceaC . In C. (. G. J. VAN STEENIS, general ed.. Fl.
Malesiana. I. 4: 57. fig. I. (p. 56). 1949. [1. procera.]
STEWSART, G. R.. & J. A. LEE. The role of proline accumulation in halophytes. Planta
120: 279-289. 1974. [Including I. mariimuni.]
----. -. & T. O. OREBAMJO. Nitrogen metabolism of halophytes. II. Nitrate
availability and utilization. New Phytol. 72: 539-546. 1973. [I'. laritnotn, passim;
autecology.]
T:i CKHOL M, G., & E. S(Dni IE RO;G. Neue Beispiele der simultanen und sukzessiven Wand-
hildung in den Pollenmutterzellen. Svensk Hot. Tidskr. 12: 189-201. 1918. [1.
pulastre. 199: successive.]
TIEGHEM, P. VAN. Recherches sur la symetrie de structure des plantes vasculaires. Ann.
Sci. Nat. Bot. V. 13: 5-314. pls. 3-8. 1870-1871. [7. laritimumn, 160, 161, pl. 6,
i 3:. '3: root anatomy.]
S& H. DI)ouLOT. Recherches comparatives sur l'origine des membres endogenes
dans les plantes vasculaires. Ann. Sci. Nat. Bot. VII. 8: 1-660. pl/. 1-40. 1888. [1.
,naritin iun, 305. 509. 510, pi 33. fi/s. 503, 504: origin of lateral roots.]
U N(AR, I. A. Halophyte communities of Park County, Colorado. Bull. Torrev Bot. Club
101: 145-152. 1974. [T. uairiliiun. 147, 148, passim: ecology.]
I . S. DEPARTMENT OF AGR .I i .TRE FOREST SERVI(E. Range plant handbook. Unpaged.
Washington. 1937. [Tri,g/' chin, (iL17 (3 text pp., I pl).]
VAczv. C. The correct grammatical gender of some generic names. Phytocoenologia 2:
442-450. 1975. [Triulochin, 450: neuter.]
VIEQSUE, .1. Nouvelles recherches sur le developpcment du sac embryonnaire des pha-
nerogames angiospermes. Ann. Sci. Nat. Bot. VI. 8: 261-390. pls. 12-21. 1879. [1
tiarrclicri. 310, pl. 15I figv. 11/-6: description of embryo sac.]
WAGNER, P. Vessel types of the monocotyledons: a survey. Bot. Not. 130: 383-402.
1977. [Trigl/ochii, 385: vessels occur in the roots.]
WAISEL. Y. Biology of halophytes. 395 pp. New York. 1972. [17. Taritinlun, passim.]
WAI.i, M. E., J. W. (ARVIN, J.. J. WILLAMAN, Q. JONES. & B. G. SCHUiBERT. Steroidal
sapogenins LX. Survey of plants for steroidal sapogenins and other constituents.
Jour. Pharm. Sci. 50: 1001-1034. 1961. [71. vtriatuo, 1030: flavonoids, alkaloids.
and tannins not found in leaves. inflorescences, or stems.]
WARD, D. B. Checklist of the vascular flora of Florida. Part 1. Psilopsida, Lycopsida,
Sphenopsida. Filicinae. (Gmnospermae, Monocotyledoneae. Univ. Florida Agr.
Exper. Sta. Tech. Bull. 726. 72 pp. 1968. ['. striatum, 20.]
[voL. 69


1988] THIERET, JUNCAGINACEAE 23
WATT, J. M., & M. G. BREYER-BRANDWIJK. The medicinal and poisonous plants of
southern and eastern Africa. 1457 pp. Edinburgh and London. 1962. [Juncaginaceae,
513, 514; South African species of Triglochin have not "so far been implicated in
the production of stock poisoning."]
WILDER, G. J. Phylogenetic trends in the Alismatidae (Monocotyledoneae). Bot. Gaz.
136: 159-170. 1975. [Triglochin, t. 1 ("a summary of some putative primitive
characters within the Alismatidae").]
WIILIAMSON, H. B. The aquatic plants of Victoria. Part III. Victoria Nat. 44: 308-315.
figs. 1-7. 1928. [Triglochin, 312-315, figs. 1-7; includes T. striatum.]
WILLIS, J. H. A handbook to plants in Victoria. 448 pp. Melbourne. 1962. [Juncagi-
naceae. 71-74; 'riglochin.]
WVUFF, H. D. Die Entwicklung der Pollenkorner von Triglochin palustris L. und die
verschiedenen Typen der Pollenkornentwicklung der Angiospermen. Jahrb. Wiss.
Bot. 88: 141-168. 1939.
WUNDERLICH. R. Ueber das Antherentapetum mit besonderer Beriicksichtigung seiner
Kernzahl. Ocsterr. Bot. Zeitschr. 101: 1-63. 1954. [Triglochin, 37.]
WUNDERLIN, R. P. Guide to the vascular plants of central Florida. 472 pp. Tampa,
Florida. 1982. [Juncaginaceae, 48; 7. striatum.]
ZINDEREN BAKKER. E. M. VAN. South African pollen grains and spores. Part 1. 88 pp.
16 p/A. [Juncaginaceae, 18. 19. pl. I1, g. i. 21: Triglochin.]