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BOTANY
OF
THE NORTHERN UNITED STATES.
MANUAL
OF
THE BOTANY
OF THE
NORTHERN UNITED STATES,
INCLUDING THE DISTRICT EAST OF THE MISSISSIPPI AND
NORTH OF NORTH CAROLINA AND TENNESSEE.
BY ASA GRAY,
LATE FISHER PROFESSOR OF NATURAL HISTORY IN HARVARD UNIVERSITY.
Sixth Edition.
REVISED AND EXTENDED WESTWARD TO THE 100th MERIDIAN,
BY
SERENO WATSON,
CURATOR OF THE GRAY HERBARIUM, HARVARD UNIVERSITY,
AND
JOHN M. COULTER,
PROFESSOR OF BOTANY IN WABASH COLLEGE,
_ASSISTED BY SPECIALISTS IN CERTAIN GROUPS_.
WITH TWENTY-FIVE PLATES,
ILLUSTRATING THE SEDGES, GRASSES, FERNS, ETC.
IVISON, BLAKEMAN, AND COMPANY,
_PUBLISHERS_,
NEW YORK AND CHICAGO.
1890.
_Copyright, 1889_,
BY THE PRESIDENT AND FELLOWS OF HARVARD COLLEGE.
CONTENTS.
Page
PREFACE 1
SYNOPSIS OF THE ORDERS 5
ANALYTICAL KEY TO THE ORDERS 19
EXPLANATION OF ABBREVIATIONS OF AUTHORS' NAMES 30
EXPLANATION OF SIGNS 32
FLORA.--PHAENOGAMOUS OR FLOWERING PLANTS 33
Dicotyledonous or Exogenous Plants 33
Angiospermous, Polypetalous 33
Gamopetalous 216
Apetalous 425
Gymnospermous Plants 489
Monocotyledonous or Endogenous Plants 495
CRYPTOGAMOUS OR FLOWERLESS PLANTS 675
Vascular Acrogens, or Pteridophytes 675
Cellular Acrogens, or Bryophytes (Hepaticae) 702
ADDITIONS AND CORRECTIONS 733
TABLE OF ORDERS 736
GLOSSARY 738
INDEX 749
PLATES, WITH EXPLANATIONS 761
PREFACE.
The first edition of Gray's Manual was published in 1848. It was to a
great extent rewritten and its range extended in 1856, and it was again
largely rewritten in 1867. The great advances that have since been made
in systematic botany and in the knowledge of our flora have for several
years past made another revision desirable, which Dr. Gray before his
death was purposing to undertake.
The present editors, acting to the best of their ability in his stead,
have endeavored throughout to follow his methods and views. The original
plan, so long retained by Dr. Gray and so generally approved, has been
closely adhered to, the characters and descriptions of the last edition
have been left essentially unchanged so far as possible, and in the
numerous alterations and additions that have been considered necessary
or advisable, his conclusions and principles have governed in every
matter of importance, so far as they could be known. The effort
especially has been to maintain that high standard of excellence which
has always made the Manual an authority among botanists.
In the treatment of the genera and species, Gray's Synoptical Flora has
been made the basis in the revision of the Gamopetalous Orders, and the
manuscript in continuation of that work, so far as prepared, for the
Polypetalous Orders which precede _Leguminosae_ (excepting _Nuphar_, the
_Cruciferae, Caryophyllaceae, Vitis_, and the small Orders numbered 18,
22, 23, 25-27, and 29). The genus _Salix_ has been rewritten for this
edition by M. S. BEBB, Esq., the genus _Carex_ by Prof. L. H. BAILEY,
and the Ferns and allied orders by Prof. D. C. EATON. For the rest, all
known available sources of information have been made use of, and much
willing help has been received from botanists in all parts of our
territory.
The increasing interest that is taken in the study of the Cellular
Cryptogams, and the desire to encourage it, have led to the inclusion
again of the Hepaticae, which were omitted in the last edition. These
have been prepared through the kindness of Prof. L. M. UNDERWOOD, though
the limits of the volume have necessitated somewhat briefer descriptions
than he considered desirable. The three fine plates illustrating the
genera of these Orders, which were used in the early editions, are also
added, with a supplementary one, as well as an additional one in
illustration of the Grasses, thus increasing the number of plates to
twenty-five. A Glossary of botanical terms is appended, to meet an
expressed need of those who use the Manual alone, and a Synopsis of the
Orders in their sequence is given, to contrast more clearly their
characters, and to show the general principles which have determined
their present arrangement. This should be a useful adjunct to the more
artificially arranged Analytical Key.
GEOGRAPHICAL LIMITS, AND DISTRIBUTION.--The southern limit of the
territory covered by the present work is the same as in the later
previous editions, viz. the southern boundary of Virginia and Kentucky.
This coincides better than any other geographical line with the natural
division between the cooler-temperate and the warm-temperate vegetation
of the Atlantic States. The rapid increase of population west of the
Mississippi River, and the growing need of a Manual covering the flora
of that section, have seemed a sufficient reason for the extension of
the limits of the work westward to the 100th meridian, thus connecting
with the _Manual of the Flora of the Rocky Mountain Region_ by Prof.
Coulter. These limits, as well as that upon the north, have been in
general strictly observed, very few species being admitted that are not
known with some degree of certainty to occur within them. The extreme
western flora is no doubt imperfectly represented.
The distribution of the individual species is indicated somewhat more
definitely than heretofore in many cases, so far as it could be
satisfactorily ascertained. The extralimital range is also sometimes
given, but the terms "northward," "southward," and "westward" are more
frequently employed, signifying an indefinite range in those directions
beyond the limits of the Manual. Where no definite habitat is specified,
the species may be understood as found more or less generally throughout
the whole area, or at least to near the western limits.
NOMENCLATURE, ACCENTUATION OF NAMES, etc.--In case of question
respecting the proper name to be adopted for any species, Dr. Gray's
known and expressed views have been followed, it is believed, throughout
the work. While reasonable regard has been paid to the claims of
priority, the purpose has been to avoid unnecessary changes, in the
belief that such changes are in most cases an unmitigated evil. Synonyms
are rarely given except where changes have been made. As a guide to
correct pronunciation, the long sound of the accented vowel (modified
often in personal names) is indicated, as heretofore, by the grave
accent (`), and the short sound by the acute ('). In regard to the
derivations of generic names, many valuable suggestions have been due to
W. R. Gerard, Esq., of New York.
PROMINENT CHARACTERS are indicated by the use of _Italic type_ for the
leading distinctions of the Orders, and generally in the specific
descriptions for those points by which two or more nearly allied species
may be most readily distinguished. The ready discrimination of the
genera is provided for by a Synopsis of their leading characters under
each order. Whenever a genus comprises several species, pains have been
taken to render important differences conspicuous by proper grouping,
and when needed by a series of subordinate divisions and subdivisions.
The headings of these various groups are to be considered as belonging
to and forming a part of the specific characters of the several species
under them,--a fact which the student should always bear in mind.
ARRANGEMENT OF THE ORDERS.--The Natural Orders are disposed in very
close accordance with the method followed by Bentham and Hooker in the
_Genera Plantarum_, the principles of which are concisely shown in the
Synopsis of Orders which precedes the Analytical Key. The _Gymnospermae_
are retained as a Subclass following the Angiospermous Dicotyledons,
with which they have an obvious relationship, in preference to placing
them, as some authorities would do, next before the Pteridophytes, to
which their affinity, if no less certain, is nevertheless obscure. A
more natural arrangement than either would be the withdrawal of the
Endogens, placing them at the beginning, in perhaps an inverse order.
ANALYTICAL KEY TO THE ORDERS.--As stated in Dr. Gray's Preface to the
last edition, this is designed to enable the student to refer readily to
its proper Order any of our plants, upon taking the pains to ascertain
the structure of its flowers, and sometimes of the fruit, and by
following out a series of easy steps in the analysis. It is founded upon
the most obvious distinctions which will answer the purpose, and is so
contrived as to provide for all or nearly all exceptional instances and
variant cases. Referring to the Order which the Key leads him to, the
student will find its most distinctive points brought together and
printed in Italics in the first sentence of the ordinal description, and
thus can verify his results. The Synopsis which follows will then lead
him to the genus, to be verified in turn by the full generic description
in its place; and the progress thence to the species is facilitated,
when there are several to choose from, by the arrangement under
divisions and subdivisions, as already explained.
It will be seen that the Key directs the inquirer to ascertain, first,
the Class of the plant under consideration,--which, even without the
seeds, is revealed at once by the plan of the stem, as seen in a
cross-section, and usually by the veining of the leaves, and is commonly
confirmed by the numerical plan of the flower;--then, if of the first
Class, the Subclass is at once determined by the pistil, whether of the
ordinary kind, or an open scale bearing naked ovules. If the former,
then the choice between the three Divisions is determined by the
presence or absence of the petals, and whether separate or united. Each
Division is subdivided by equally obvious characters, and, finally, a
series of successively subordinated propositions,--each set more
indented upon the page than the preceding,--leads to the name of the
Order sought for, followed by the number of the page upon which it is
described in the body of the work.
The book is now submitted to those for whose benefit it has been
prepared, in the trust that its shortcomings will meet with friendly
indulgence, and with the earnest request that information be kindly
given of any corrections or additions that may appear to be necessary.
SERENO WATSON.
CAMBRIDGE, MASS., Dec. 26, 1889.
SYNOPSIS OF THE ORDERS OF PLANT
DESCRIBED IN THIS WORK.
SERIES I. PHAENOGAMOUS OR FLOWERING PLANTS: those producing real flowers
and seeds.
CLASS I. DICOTYLEDONOUS OR EXOGENOUS PLANTS.
Stems formed of bark, wood, and pith; the wood forming a zone between
the other two, and increasing, when the stem continues from year to
year, by the annual addition of a new layer to the outside, next the
bark. Leaves netted-veined. Embryo with a pair of opposite cotyledons,
or in Subclass II. often three or more in a whorl. Parts of the flower
mostly in fours or fives.
SUBCLASS I. ANGIOSPERMAE. Pistil consisting of a closed ovary which
contains the ovules and becomes the fruit. Cotyledons only two.
DIVISION I. POLYPETALOUS: the calyx and corolla both present; the latter
of _separate_ petals. (Apetalous flowers occur in various Orders, as
noted under the subdivisions.)
[A.] THALAMIFLORAE. Stamens and petals hypogynous (free both from the
calyx and from the superior ovary), upon a usually narrow receptacle
(not glandular nor discoid, except in Reseda, sometimes stipe-like).
(Stamens and petals upon the partly inferior ovary in some Nymphaeaceae.)
Apetalous flowers occur in the Ranunculaceae and Caryophyllaceae.
[*] 1. Carpels solitary or distinct (or coherent in Magnoliaceae); sepals
and petals deciduous (except in Nymphaeaceae); leaves alternate or
radical, without stipules (sometimes opposite or whorled and rarely
stipular in Ranunculaceae); embryo (except in Nelumbo) small, in fleshy
albumen.
1. Ranunculaceae (p. 34). Sepals (3 or more), petals (as many, in regular
flowers, or none), stamens (usually many), and carpels (1--many) all
distinct. Fruit achenes, follicles, or berries. Mostly herbs.
2. Magnoliaceae (p. 49). Sepals and petals colored alike, in three or
more rows of three, imbricate. Fruit cone-like, formed of the numerous
cohering pistils. Trees.
3. Anonaceae (p. 50). Sepals (3) and petals (6, in two rows) valvate.
Fruit pulpy. Shrubs or small trees.
4. Menispermaceae (p. 51). Sepals and petals in twos or threes,
imbricate. Pistils becoming 1-seeded drupes. Dioecious woody
climbers, with palmate or peltate leaves.
5. Berberidaceae (p. 52). Sepals and petals imbricate, each in two rows
of three (rarely in twos or fours). Stamens opposite the petals. Pistil
solitary, becoming a berry or pod. Shrubs or low herbs.
6. Nymphaeaceae, in part (p. 54). Sepals and petals each 3, or many in
several rows. Pistils becoming coriaceous and indehiscent. Aquatics;
floating leaves peltate.
[*] 2. Carpels (2 or more) united into a compound ovary with parietal,
often nerve-like placentae (or the seeds covering the inner surface in
Nymphaeaceae, and the placentae axile in Sarraceniaceae). Herbs (some
Cistaceae somewhat shrubby).
[+] Fruit 5--many-celled; calyx or whole perianth persistent; embryo
small, at the base of fleshy albumen.
6. Nymphaeaceae proper (p. 54). Sepals 2--6. Petals and stamens numerous,
on a thick hypogynous receptacle or inserted upon the ovary. Capsule
8--30-celled. Aquatics, with peltate or cordate leaves.
7. Sarraceniaceae (p. 57). Sepals and petals 5. Capsule 5-celled. Marsh
plants, with pitcher-shaped leaves.
[+][+] Fruit 1-celled, or spuriously 2--more-celled by partitions
connecting the placentae.
[++] Embryo minute at the base of fleshy albumen; perianth deciduous;
sepals 2.
8. Papaveraceae (p. 57.) Flowers regular. Sepals fugacious. Petals 4--12.
Stamens and seeds numerous. Capsule 2--several-valved. Juice milky or
colored.
9. Fumariaceae (p. 59.) Flowers irregular. Petals 4, in dissimilar pairs.
Stamens 6, diadelphous. Fruit 2-valved (indehiscent and 1-seeded in
Fumaria). Juice watery; leaves dissected.
[++][++] Albumen none; embryo curved or folded; perianth deciduous
(sepals persistent in Resedaceae).
10. Cruciferae (p. 61). Sepals and petals 4. Stamens mostly 6,
tetradynamous (two inserted lower and shorter). Pod 2-celled by a
transverse partition, 2-valved, or sometimes indehiscent or transversely
jointed. Bracts and stipules none.
11. Capparidaceae (p. 74). Sepals and petals 4. Stamens 6 or more, nearly
equal. Pod 1-celled, 2-valved. Embryo coiled. Leaves often palmately
divided; bracts and stipules often present.
12. Resedaceae (p. 75). Sepals and petals 4--7, irregular. Stamens
indefinite on an hypogynous disk, not covered in the bud. Pod 1-celled,
3--6-lobed, opening at the top.
[++][++][++] Embryo rather large in fleshy albumen; placentae on the
middle of the valves; calyx persistent.
13. Cistaceae (p. 76). Flowers regular; sepals and petals 5, the two
outer sepals minute. Stamens indefinite. Pod 1-celled, 3--5-valved.
Ovules orthotropous. Embryo curved. Leaves entire, the lower often
opposite.
14. Violaceae (p. 78). Flowers irregular; sepals and petals 5. Stamens 5,
with connivent introrse anthers. Style clavate. Pod 1-celled, 3-valved.
Ovules anatropous. Embryo straight. Stipules present.
[*] 3. Ovary compound, 1-celled, with central placentae; embryo curved
around mealy albumen (except in Dianthus); leaves entire; stipules
mostly none.
15. Caryophyllaceae (p. 82). Sepals (5, rarely 4) distinct or united,
persistent. Petals as many, rarely none. Stamens as many or twice as
many, rarely fewer. Styles 2--5. Leaves opposite.
16. Portulacaceae (p. 90). Sepals 2. Petals 5. Stamens 5--20. Capsule
3-valved or circumscissile. Fleshy herbs; leaves mostly alternate.
[*] 4. Calyx imbricate; stamens as many or twice as many as the petals
or often indefinite; ovary compound, 1-celled with parietal placentae or
several-celled with the placentae united in the axis; embryo straight or
slightly curved; albumen none or scanty.
17. Elatinaceae (p. 91). Small marsh annuals, with opposite leaves,
membranous stipules, minute axillary flowers, few stamens, and pod
2--5-celled.
18. Hypericaceae (p. 92). Herbs or shrubs, with opposite entire dotted
leaves and no stipules. Flowers cymose or panicled. Stamens few or many,
usually in 3 or more clusters. Pod 1-celled or 3--5-celled.
19. Ternstroemiaceae (p. 95). Trees or shrubs, with alternate leaves
and no stipules. Flowers large, axillary, solitary. Stamens numerous,
more or less united together and with the base of the petals. Pod
3--5-celled.
[*] 5. Calyx valvate; stamens numerous, usually more or less united
together and with the base of the petals; ovary 3--many-celled with the
placentae united in the axis (becoming 1-celled and 1-seeded in Tilia).
20. Malvaceae (p. 96). Stamens monadelphous; anthers 1-celled. Calyx
persistent. Seeds kidney-shaped, with curved embryo and little albumen.
Herbs or shrubs, with alternate palmately veined stipular leaves.
21. Tiliaceae (p. 101). Stamens polyadelphous or nearly distinct; anthers
2-celled. Calyx deciduous. Embryo nearly straight. Trees, with alternate
leaves and deciduous stipules.
[B.] DISCIFLORAE. Stamens as many as the petals or twice as many or
fewer, inserted upon or at the outer or inner base of a more or less
tumid hypogynous or perigynous disk, which is cushion-like or annular or
divided into glands, sometimes obscure or minute (or none in Linum,
Ilex, some Geraniaceae and Polygala); ovary superior (or half-inferior in
some Rhamnaceae); sepals more usually distinct. Petals wanting in some
Rutaceae, Rhamnaceae, and Sapindaceae.
[*] 1. Ovules (mostly 1 or 2 in each cell) pendulous, with the rhaphe
toward the axis of the ovary; disk often reduced to glands alternate
with the petals or none; ovary often lobed or the carpels nearly
distinct.
22. Linaceae (p. 101). Flowers regular, usually 5-merous. Capsule not
lobed, mostly 5-valved, spuriously 10-celled, 10-seeded. Stamens united
at base. Disk none or 5 minute glands. Herbs, with entire alternate or
opposite leaves; stipules gland-like or none.
23. Geraniaceae (p. 102). Flowers regular or irregular, 5-merous or
3-merous as to the stamens and pistils. Ovary 3--5-lobed, the cells
1--few-ovuled, and axis persistent. Disk of 5 glands or none. Herbs,
with often lobed or divided mostly alternate leaves, with or without
stipules.
24. Rutaceae (p. 106). Flowers mostly regular, 3--5-merous, dioecious
or polygamous in our genera. Ovary 2--5-lobed or the carpels nearly
distinct, upon a glandular disk; cells 2-ovuled. Mostly shrubs or trees,
with glandular-punctate compound leaves, without stipules.
[*] 2. Ovules (1 or 2) pendulous, the rhaphe away from the axis; disk
none and ovary not lobed.
25. Ilicineae (p. 107). Flowers small, dioeciously polygamous,
axillary, 4--8-merous. Fruit a 4--8-seeded berry-like drupe. Shrubs or
trees, with simple alternate leaves and no stipules.
[*] 3. Ovules (1 or 2 in each cell) erect, the rhaphe toward the axis;
disk fleshy, covering the base of the calyx; stamens as many
as the petals, at the margin of the disk; flowers perfect or
polygamo-dioecious; albumen fleshy; shrubs or trees, with simple
leaves (compound in some Vitaceae).
26. Celastraceae (p. 109). Sepals and petals imbricated, the stamens
alternate with the petals. Fruit 2--5-celled; seeds arilled.
27. Rhamnaceae (p. 111). Calyx valvate. Petals small or none. Stamens
alternate with the sepals. Fruit 2--5-celled; seeds solitary, not
arilled.
28. Vitaceae (p. 112). Calyx minute. Stamens opposite the valvate
caducous petals. Climbing by tendrils opposite the alternate leaves.
[*] 4. Ovules (1 or 2) ascending or horizontal, or pendulous from a
basal funicle; fleshy disk entire or lobed; stamens 5--10; shrubs or
trees, with compound leaves (simple in Acer) and mostly
polygamo-dioecious and often irregular flowers; petals imbricate
(sometimes none in Sapindaceae).
29. Sapindaceae (p. 115). Flowers mostly unsymmetrical or irregular.
Ovary 2--3-celled and -lobed.
30. Anacardiaceae (p. 118). Flowers regular, 5-androus. Ovary 1-celled,
becoming a small dry drupe. Leaves alternate; juice milky or resinous.
[*] 5. Ovules solitary, pendulous from the summit of the 2-celled ovary;
disk none; flowers irregular (subpapilionaceous), hypogynous; stamens
monadelphous or diadelphous; anthers 1-celled, opening by an apical
pore.
31. Polygalaceae (p. 120). Herbs, with perfect flowers and alternate or
opposite or whorled entire leaves. Stamens 6--8. Seed carunculate.
[C.] CALYCIFLORAE. Sepals rarely distinct; disk adnate to the base of the
calyx, rarely tumid or conspicuous; petals and stamens on the calyx,
perigynous or epigynous, the ovary being often inferior (hypogynous in
Drosera and Parnassia, nearly so in some Leguminosae and Crassulaceae).
Apetalous flowers in Orders 33, 35, 36, 38, 39, 41, 42, 47, and 50.
[*] 1. Ovary usually superior, the pistils solitary, or several and
distinct (sometimes more or less united but at least the styles distinct
except in some Saxifragaceae).
32. Leguminosae (p. 122). Flowers papilionaceous or regular. Stamens
usually 10, and mostly monadelphous or diadelphous. Pistil one, free,
becoming a legume; style terminal. Albumen none. Leaves mostly compound,
alternate, stipular.
33. Rosaceae (p. 150). Flowers regular, with usually numerous distinct
stamens, and 1--many pistils, distinct or (in Pomeae) united and combined
with the calyx-tube; style often lateral or basal. Calyx-lobes and
petals mostly 5. Ovules mostly 1 or 2. Albumen mostly none. Trees,
shrubs, or herbs; leaves usually alternate and stipulate, simple or
compound.
34. Calycanthaceae (p. 167). Calyx-lobes, petals, and stamens indefinite.
Pistils numerous, becoming achenes in a hollow receptacle. Albumen none.
Aromatic shrubs, with opposite entire leaves and no stipules.
35. Saxifragaceae (p. 168). Flowers regular, with 5--10 stamens (numerous
in Philadelphus), few (mostly 2) more or less united, free or partially
adnate carpels, and few--many ovules on axile or sometimes parietal
placentae. Seeds albuminous. Herbs or shrubs, with opposite or alternate
leaves, with or without stipules.
36. Crassulaceae (p. 170). Mostly fleshy herbs, with symmetrical flowers,
the usually distinct many-seeded carpels as many as the sepals. Seeds
albuminous. Leaves alternate or opposite or whorled; stipules none.
37. Droseraceae (p. 178). Glandular-haired scapose marsh herbs, with
regular 5-merous hypogynous flowers. Capsule 1-celled, with 3--5
many-seeded parietal placentae. Anthers extrorse. Leaves circinate in
vernation.
38. Hamamelideae (p. 179). Shrubs or trees; flowers often
polygamo-monoecious, in clusters, heads, or spikes; petals often none.
Seeds 2 or more, bony, in a 2-beaked woody pod opening above, the base
adnate to the calyx-tube. Stamens few or many. Leaves alternate, simple.
39. Halorageae (p. 180). Aquatic or marsh herbs; flowers perfect or
polygamo-dioecious, small, axillary or spicate; petals often none.
Stamens 1--8. Ovary inferior, the calyx-limb obsolete or very short.
Fruit small, indehiscent, 1--4-celled, 1--4-seeded. Leaves alternate or
opposite, the submersed often dissected.
[*] 2. Ovary inferior (except in Lythraceae), 1--several-celled; style
entire; flowers perfect, regular or nearly so, mostly 4-merous; herbs,
with simple and mostly entire leaves without stipules.
40. Melastomaceae (p. 183). Calyx open. Stamens definite; anthers opening
by an apical pore. Leaves opposite, 3--7-nerved; flowers cymose.
41. Lythraceae (p. 184). Calyx-lobes valvate. Pod free, but enclosed in
the calyx, membranous, 1--4-celled, many-seeded with axile placentae.
Leaves mostly opposite; flowers axillary or whorled; petals crumpled, or
none.
42. Onagraceae (p. 186). Calyx-lobes valvate. Ovary 1--4-celled, the
cells 1--many-ovuled. Stamens 2, 4, or 8. Petals 2 or 4, convolute, or
none. Leaves opposite or alternate.
[*] 3. Ovary inferior (except in Passifloraceae and Ficoideae), 1-celled
with parietal placentae or several-celled by the intrusion of the
placentae; flowers regular, perfect or unisexual; styles free or united;
herbs.
[+] Embryo straight; cotyledons foliaceous; leaves alternate, often
lobed.
43. Loasaceae (p. 193). Flowers perfect. Stamens indefinite. Style entire
or 2--3-cleft. Capsule 1-celled, with 2 or 3 many-seeded placentae.
Pubescence of hooked hairs.
44. Passifloraceae (p. 194). Climbing by tendrils. Flowers perfect.
Stamens 5, monadelphous. Ovary stalked, superior, becoming a 1-celled
many-seeded berry with 3 or 4 placentae. Styles 3, clavate.
45. Cucurbitaceae (p. 194). Tendril-bearing vines, with dioecious or
monoecious flowers. Corolla 5-lobed, often confluent with the calyx.
Stamens 3 or 5, usually more or less united and the anthers often
tortuous. Fruit fleshy or membranous, 1--5-celled, the placentae often
produced to the axis and revolute. Seeds exalbuminous.
[+][+] Embryo curved or coiled about central albumen; leaves entire.
46. Cactaceae (p. 196). Fleshy and mostly leafless prickly plants, with
solitary sessile perfect flowers. Calyx-lobes and petals indefinite,
imbricated, the numerous stamens on the tube. Fruit a 1-celled
many-seeded berry.
47. Ficoideae (p. 198). Calyx-lobes or sepals 5 and petals none in our
genera. Capsule 3--5-celled with axile placentae, loculicidal or
circumscissile, many-seeded. Often fleshy; leaves mostly opposite or
verticillate.
[*] 4. Flowers small, regular, perfect or polygamous; calyx-limb minute
or obsolete; ovary inferior, 2--several-celled, with solitary pendulous
ovules; petals and stamens mostly 4 or 5, on the margin of an epigynous
disk surrounding the styles; albumen copious.
48. Umbelliferae (p. 198). Flowers in umbels or heads. Petals (inflexed)
and stamens 5. Styles 2. Fruit of 2 dry seed-like carpels, the pericarp
usually with oil-tubes. Herbs, with alternate mostly compound leaves.
49. Araliaceae (p. 212). Flowers mostly in umbels and nearly as in
Umbelliferae; petals not inflexed and styles 2 or more. Fruit a
2--several-celled drupe. Herbs or shrubs, with alternate mostly compound
leaves.
50. Cornaceae (p. 213). Flowers not in umbels; petals (valvate, or none)
and stamens 4 or 5. Style 1. Fruit a 1--2-seeded drupe. Trees, shrubs,
or rarely herbs, with opposite or alternate simple and mostly entire
leaves.
DIVISION II. GAMOPETALOUS: calyx and corolla both present, the latter of
united petals (excepting some Ericaceae, Styracaceae, and Oleaceae, Galax,
Statice, and Lysimachia). Apetalous flowers occur in Glaux and some
Oleaceae. Stipules present only in Rubiaceae and Loganiaceae, or rarely in
Caprifoliaceae.
[*] 1. Ovary inferior; stamens borne upon the corolla, alternate with
its lobes.
[+] Stamens distinct; leaves opposite or whorled; seed albuminous except
in Valerianaceae.
51. Caprifoliaceae (p. 216). Corolla mostly 5-lobed, regular or
irregular, the stamens as many (one fewer in Linnaea, doubled in
Adoxa). Ovary 1--several-celled; fruit a berry, drupe, or pod,
1--several-seeded. Shrubs or herbs; leaves opposite, rarely stipular,
not turning black in drying.
52. Rubiaceae (p. 222). Flowers regular, 4--5-merous, the corolla mostly
valvate. Ovary 2--4-celled. Herbs or shrubs; leaves simple, entire,
opposite with stipules, or verticillate, usually turning black in
drying.
53. Valerianaceae (p. 228). Stamens (1--4) fewer than the lobes of the
somewhat irregular corolla. Ovary with two abortive or empty cells and
one containing a suspended ovule. Fruit dry and indehiscent. Herbs.
54. Dipsaceae (p. 229). Flowers mostly 4-merous and with 4 (rarely 2)
stamens, involucellate in involucrate heads; corolla-lobes imbricate.
Ovary simple, 1-celled, with a suspended ovule. Herbs.
[+][+] Anthers connate into a tube.
55. Compositae (p. 230). Stamens as many as the valvate corolla-lobes.
Ovary with a solitary erect ovule, becoming an achene. Albumen none.
Calyx-limb reduced to a pappus or none. Flowers in involucrate heads.
[*] 2. Ovary inferior (or superior in most Ericaceae and in
Diapensiaceae); stamens free from the corolla or nearly so (adnate in
some Diapensiaceae), as many as the lobes and alternate with them, or
twice as many; leaves alternate (opposite in some Ericaceae); style 1.
[+] Juice milky; capsule 2--5-celled, many-seeded; herbs.
56. Lobeliaceae (p. 305). Corolla irregular, 5-lobed. Stamens united, at
least by the anthers. Capsule 2-celled or with two placentae.
57. Campanulaceae (p. 307). Corolla regular, 5-lobed, valvate. Stamens
usually distinct. Capsule 2--several-celled.
[+][+] Juice not milky nor acrid; capsule 3--10-celled.
58. Ericaceae (p. 309). Flowers mostly regular, 4--5-merous. Stamens
distinct, more usually twice as many as the corolla-lobes or petals.
Ovary inferior or superior. Herbs or shrubs.
59. Diapensiaceae (p. 326). Flowers regular. Stamens 5, on the corolla,
or monadelphous with 5 petaloid staminodia. Ovary superior, 3-celled.
[*] 3. Ovary superior; stamens as many as the corolla-lobes and opposite
them.
60. Plumbaginaceae (p. 327). Stamens 5, on the base of the petals.
Styles 5. Fruit an achene or 1-seeded utricle. Herbs; leaves radical.
61. Primulaceae (p. 328). Stamens 4--8, perigynous. Style 1. Fruit a
capsule with several seeds on a central placenta. Herbs; leaves radical
or opposite or alternate.
62. Sapotaceae (p. 332). Flowers small, 4--5-merous. Style 1. Ovary
few--several-celled; fruit fleshy, bearing a single bony-coated seed.
Shrubs or trees, with milky juice and alternate entire leaves.
[*] 4. Ovary superior or more or less adnate to the calyx,
few--several-celled, the cells 1-ovuled; stamens twice as many as the
corolla-lobes or more; trees or shrubs, with alternate leaves.
63. Ebenaceae (p. 333). Flowers dioecious or polygamous. Stamens on the
corolla. Ovary superior. Styles distinct. Fruit fleshy, few-seeded.
64. Styracaceae (p. 333). Flowers perfect. Stamens subhypogynous. Ovary
more or less inferior. Style 1. Fruit dry or nearly so, 1--4-seeded.
[*] 5. Ovary superior, of two carpels (sometimes by division apparently
4-carpellary, sometimes of 3--5 in Polemoniaceae, Convolvulaceae, and
Solanaceae); stamens on the corolla (except in apetalous Oleaceae),
alternate with its lobes, as many or fewer.
[+] Corolla not scarious and nerveless.
[++] Corolla none, or regular and 4-cleft or -parted, the stamens fewer
than its lobes; style 1; seeds 1--3.
65. Oleaceae (p. 335). Trees or shrubs, with opposite and pinnate or
simple leaves. Flowers perfect or polygamo-dioecious. Stamens mostly
2, alternate with the usually 2-ovuled carpels.
[++][++] Corolla regular, its lobes 4--5 or rarely more; stamens as
many.
[=] Ovaries 2, becoming follicles; stigmas and sometimes the styles
united; herbs with milky juice, perfect 5-merous flowers, and simple
entire leaves.
66. Apocynaceae (p. 337). Stamens distinct or the anthers merely
connivent, with ordinary pollen. Style 1.
67. Asclepiadaceae (p. 338). Stamens monadelphous, the anthers
permanently attached to a large stigmatic body; pollen mostly in waxy
masses. Styles distinct below the stigma.
[=][=] Ovary compound (ovaries two in Dichondra), with 2 or 3 (rarely 4
or 5) cells or placentae; stamens distinct; mostly herbs.
[a.] Leaves opposite; corolla-lobes 4 or 5 or more.
68. Loganiaceae (p. 345). Leaves entire, with stipules or a stipular line
joining their bases. Capsule 2-celled, few--many-seeded. Herbs or woody
twiners (our species).
69. Gentianaceae (p. 346). Glabrous herbs; leaves entire, sessile and
simple (except in Menyanthes). Capsule 1-celled with 2 parietal placentae
or the whole inner surface ovuliferous, many-seeded.
[b.] Leaves alternate (sometimes opposite in Polemoniaceae and
Hydrophyllaceae); corolla-lobes always 5 in our species.
70. Polemoniaceae (p. 354). Capsule usually 3-celled, loculicidal; seeds
1--many in each cell on the stout placental axis. Style 3-cleft or
-lobed. Leaves opposite or alternate, simple or compound.
71. Hydrophyllaceae (p. 357). Leaves often lobed or divided, and the
inflorescence frequently scorpioid. Style 2-parted or 2-lobed. Capsule
1-celled, 2-valved with two parietal or introflexed placentae, or
sometimes 2-celled. Seeds 2 or more on each placenta.
72. Borraginaceae (p. 360). Leaves mostly entire and plants often
rough-hispid; inflorescence commonly scorpioid. Style 1. Ovary
4-ovulate, usually 4-lobed and maturing as 4 separate or separable
nutlets, or not lobed, 2--4-celled and separating when ripe into 2 or 4
nutlets.
73. Convolvulaceae (p. 367). Usually twining or trailing; flowers on
axillary peduncles or cymose-glomerate. Corolla 5-lobed or 5-plaited,
twisted in the bud. Styles 1 or 2. Ovary 2- (sometimes 3- or spuriously
4-) celled, becoming a globular 4--6-seeded capsule (or ovaries two and
distinct in Dichondra). Cotyledons broad-foliaceous.
74. Solanaceae (p. 373). Style 1. Ovary 2-celled (rarely 3--5-celled),
with numerous ovules on axillary placentae, becoming a pod or berry.
Cotyledons narrow.
[++][++][++] Corolla more or less bilabiately irregular (sometimes
nearly regular), 5-lobed. Fertile stamens 4 and didynamous, or 2. Style
1. Ovary always of two carpels.
[a.] Ovules several or many.
75. Scrophulariaceae (p. 377). Capsule 2-celled, with central placentae.
Seeds small, usually numerous. Herbs; leaves alternate or opposite.
76. Orobanchaceae (p. 393). Root-parasites with no green foliage. Capsule
1-celled, with 2 simple or double parietal placentae. Seeds many.
77. Lentibulariaceae (p. 395). Aquatic or marsh herbs, with scapes or
scape-like peduncles, sometimes nearly leafless. Corolla personate and
spurred. Capsule globular, 1-celled; placentae central, free,
many-seeded.
78. Bignoniaceae (p. 398). Large-flowered trees or often climbing shrubs,
with usually opposite simple or compound leaves. Capsule 2-celled by a
partition between the 2 parietal placentae. Seeds numerous, large, mostly
winged.
79. Pedaliaceae (p. 399). Herbs, with opposite simple leaves. Ovary
1-celled with two bilamellar parietal placentae, or 2--4-celled by their
union, becoming drupaceous or capsular. Seeds few or many, wingless.
80. Acanthaceae (p. 399). Herbs, with opposite simple leaves. Capsule
2-celled, loculicidal, with each axile placenta bearing 2--10 flattish
seeds.
[b.] Cells of the ovary 1--2-ovuled; herbs or low shrubs, with opposite
leaves.
81. Verbenaceae (p. 401). Ovary 2--4-celled, not lobed, the dry or
drupaceous fruit separating into 2 or 4 1-seeded nutlets (fruit 1-celled
and 1-seeded in Phryma). Style terminal.
82. Labiatae (p. 403). Ovary deeply 4-lobed around the style, the lobes
becoming dry seed-like nutlets. Stems square; aromatic.
[+][+] Corolla scarious and nerveless; flowers regular, 4-merous; style
1.
83. Plantaginaceae (p. 422). Scapose herbs, with perfect or
polygamo-dioecious or monoecious flowers in 1--many-flowered spikes.
Fruit a circumscissile 2-celled capsule, with one or more peltate seeds
in each cell, or an achene.
DIVISION III. APETALOUS EXOGENS. The corolla wanting (except in some
Euphorbiaceae), and sometimes also the calyx.
[*] 1. Ovary superior (though sometimes enclosed within the calyx),
1-celled with a solitary basal ovule (several-celled in Phytolaccaceae);
embryo coiled or curved (nearly straight in Polygonaceae) in or about
mealy albumen (albumen none in some Chenopodiaceae); herbs.
[+] Fruit the hardened or membranous closed base of the corolla-like
perianth enclosing a utricle.
84. Nyctaginaceae (p. 425). Perianth tubular or funnelform. Stamens
hypogynous. Fruit ribbed or winged. Leaves opposite; stipules none.
[+][+] Fruit a utricle; perianth mostly persistent, small, 4--5-lobed or
-parted, or none.
85. Illecebraceae (p. 426). Perianth herbaceous. Stamens perigynous.
Leaves opposite; stipules scarious (none in Scleranthus).
86. Amarantaceae (p. 427). Flowers sessile, bracteate, the bracts
(usually 3) more or less dry and scarious, as well as the 3--5 distinct
sepals. Stamens 1--5, hypogynous. Utricle indehiscent or circumscissile.
Embryo annular. Leaves mostly alternate, entire; stipules none.
87. Chenopodiaceae (p. 430). Flowers sessile, not scarious-bracteate.
Sepals greenish or succulent, 5 or fewer, or none. Stamens 5 or fewer,
perigynous or hypogynous. Embryo annular or spiral or conduplicate.
Leaves alternate; stipules none.
[+][+][+] Ovary of several 1-ovuled carpels, in fruit a berry (in our
genera).
88. Phytolaccaceae (p. 435). Sepals 4--5, petaloid or herbaceous. Stamens
5--30, hypogynous. Carpels 5--12. Embryo annular. Leaves alternate,
entire; stipules none.
[+][+][+][+] Fruit a triangular or lenticular achene.
89. Polygonaceae (p. 436). Flowers on jointed pedicels. Calyx 3--6-lobed
or -parted, more or less corolla-like. Stamens 4--12, on the calyx.
Embryo nearly straight. Leaves alternate, with sheathing stipules or
none.
[*] 2. Ovary compound, the cells many-ovuled (or 1-ovuled in Piperaceae);
embryo minute in copious albumen; flowers perfect.
90. Podostemaceae (p. 444). Aquatic, with the aspect of sea-weeds or
mosses, with minute naked flowers from a spathe-like involucre. Ovary
superior; pod 2--3-celled.
91. Aristolochiaceae (p. 444). Terrestrial herbs or climbing shrubs.
Calyx valvate, adnate at least at base to the 6-celled many-seeded
ovary. Stamens 6--12, more or less united with the style. Leaves
alternate, mostly cordate; stipules none.
92. Piperaceae (Sec. Saurureae), (p. 446). Marsh herb (our species). Perianth
none. Carpels 3--4, distinct, with usually a single ascending seed.
Leaves alternate, entire.
[*] 3. Ovary superior, simple, 1-celled, 1-ovuled, forming a berry or
drupe; trees or shrubs, with mostly entire leaves and no stipules.
93. Lauraceae (p. 446). Flowers perfect or dioecious. Sepals 4 or 6, in
2 rows. Stamens 9--12; anthers opening by 2 or 4 uplifted valves. Seed
suspended; albumen none. Aromatic; leaves alternate.
94. Thymelaeaceae (p. 448). Flowers perfect. Calyx corolla-like,
4--5-cleft. Stamens twice as many. Seed suspended, with little or no
albumen. Acrid shrubs with very tough bark; leaves alternate.
95. Elaeagnaceae (p. 448). Flowers mostly dioecious. Calyx-tube becoming
berry-like and enclosing the achene. Seed erect, albuminous. Leaves
silvery-scurfy, opposite.
[*] 4. Ovary inferior, 1-celled, 1--3-ovuled (but 1-seeded); albumen
without testa, bearing the embryo in a cavity at the apex; calyx-lobes
valvate.
96. Loranthaceae (p. 449). Parasitic on trees, with jointed stems and
opposite leaves. Flowers dioecious. Ovule solitary, erect. Fruit a
berry.
97. Santalaceae (p. 450). Flowers perfect. Ovules 2--4, suspended from
the apex of a central placenta. Fruit dry, indehiscent. Leaves
alternate.
[*] 5. Flowers all unisexual (polygamous in some Urticaceae and
Empetraceae, apparently perfect in Euphorbia); cells 1--2-ovuled; embryo
nearly as long as the albumen or filling the seed; calyx often wanting,
corolla-like only in some Euphorbiaceae and Empetraceae; stipules often
present.
[+] 1. Ovary superior, 3-celled (1-celled in Crotonopsis) with 1 or 2
pendulous ovules in each cell; herbs.
98. Euphorbiaceae (p. 451). Flowers monoecious or dioecious
(involucrate and apparently perfect in Euphorbia). Mostly with milky
juice, and usually alternate often stipulate leaves.
[+] 2. Ovary 1-celled, 1-seeded; trees or shrubs (except some
Urticaceae).
[++] Calyx regular, the stamens as many as the lobes and opposite them
or fewer; ovary superior.
99. Urticaceae (p. 461). Flowers monoecious, dioecious, or (in Ulmeae)
perfect. Seeds exalbuminous or nearly so. Inflorescence very various.
[++][++] Perianth mostly none; at least the staminate flowers in aments
or spikes or dense heads; albumen none.
100. Platanaceae (p. 466). Trees, with alternate palmately lobed leaves,
sheathing stipules, and monoecious flowers in separate globose heads.
Ovary superior; fruit a club-shaped nutlet.
101. Juglandaceae (p. 467). Trees, with alternate pinnate leaves, no
stipules, and monoecious flowers, the staminate in aments. Ovary
inferior; fruit a nut.
102. Myricaceae (p. 469). Shrubs, with resinous-dotted leaves, with or
without stipules, and monoecious or dioecious flowers, both kinds in
short scaly aments. Ovary superior, becoming a small drupe-like nut.
[+] 3. Ovary 2--7-celled, with 1 or 2 suspended ovules in each cell,
becoming 1-celled and 1-seeded; calyx mostly none or adherent to the
ovary; trees or shrubs with simple leaves.
103. Cupuliferae (p. 470). Flowers monoecious. Fruit a nut surrounded
by an involucre, or (in Betuleae) a small winged or angled naked nutlet
in the axils of the scales of an ament.
[+] 4. Ovary 1-celled, becoming a 2-valved pod with two parietal or
basal placentae bearing numerous small comose seeds; perianth none.
104. Salicaceae (p. 480). Dioecious trees or shrubs, with both kinds of
flowers in aments, and simple alternate stipulate leaves.
[+] 5. Ovary several-celled, becoming a drupe containing 3--9 1-seeded
nutlets; seed erect; low shrubby heath-like evergreens.
105. Empetraceae (p. 487). Flowers polygamous or dioecious,
scaly-bracted. Sepals somewhat petaloid or none. Embryo axile in
copious albumen.
[+] 6. Ovary 1-celled with a suspended ovule, becoming an achene; calyx
none; aquatic herbs, with finely dissected whorled leaves.
106. Ceratophyllaceae (p. 488). Flowers monoecious, minute, axillary
and sessile. Albumen none; the seed filled with a highly developed
embryo.
SUBCLASS II. GYMNOSPERMOUS EXOGENS. Ovules naked upon a scale, bract, or
disk. Cotyledons two or more.
107. Coniferae (p. 489). Resiniferous trees or shrubs, with mostly
awl-shaped or needle-shaped and evergreen leaves, and monoecious or
dioecious flowers.
CLASS II. MONOCOTYLEDONOUS PLANTS.
Stems without central pith or annular layers, but having the woody
fibres distributed irregularly through them (a transverse slice showing
the fibres as dots scattered through the cellular tissue). Embryo with a
single cotyledon and the early leaves always alternate. Parts of the
flower usually in threes (never in fives), and the leaves mostly
parallel-veined. Our species herbaceous, excepting Smilax.
[*] Ovary inferior (superior in Bromeliaceae, nearly so in some
Hemodoraceae); at least the inner lobes of the perianth petal-like.
[+] 1. Seeds without albumen, very numerous and minute.
108. Hydrocharidaceae (p. 495). Aquatics, with dioecious or polygamous
flowers from a spathe; outer perianth calyx-like, the inner sometimes
wanting. Stamens 3--12. Ovary 1-celled with 3 parietal placentae or
6--9-celled with axile placentae.
109. Burmanniaceae (p. 496). Terrestrial, with scale-like cauline leaves
and regular perfect triandrous flowers. Perianth corolla-like.
110. Orchidaceae (p. 497). Terrestrial, with very irregular perfect
flowers. Stamens and style connate; anthers 1 or 2. Capsule 1-celled;
placentae 3, parietal. Perianth corolla-like.
[+] 2. Seeds albuminous. (Ovary 3-celled and flowers regular in our
genera.)
111. Bromeliaceae (p. 511). Mostly epiphytes, with dry persistent scurfy
leaves. Flowers 6-androus; outer perianth calyx-like.
112. Hemodoraceae (p. 512). Fibrous-rooted, with equitant leaves and
perfect 3- or 6-androus flowers. Perianth persistent, woolly or scurfy
outside. (Ovary sometimes nearly free; leaves flat in Aletris.)
113. Iridaceae (p. 513). Root not bulbous; leaves equitant in two ranks.
Flowers from a spathe. Stamens 3, opposite the outer lobes of the
corolla-like perianth; anthers extrorse.
114. Amaryllidaceae (p. 515). Often bulbous-rooted and scapose. Perianth
corolla-like. Stamens 6; anthers introrse.
115. Dioscoreaceae (p. 517). Climbing, with net-veined leaves. Flowers
dioecious, small, 6-androus; perianth calyx-like. Ovules 1 or 2 in
each cell.
[*][*] Ovary superior (very rarely partially adnate to the calyx in
Liliaceae).
[+] 1. At least the inner perianth corolla-like; ovary compound; seeds
with copious albumen.
116. Liliaceae (p. 517). Flowers perfect, 6-androus, the regular perianth
corolla-like (dioecious in Smilax, dimerous in Maianthemum, the outer
divisions herbaceous in Trillium). Fruit a 3-celled capsule or berry.
117. Pontederiaceae (p. 535). Aquatic, with more or less irregular
perfect flowers from a spathe; perianth corolla-like. Stamens 3 or 6,
mostly unequal or dissimilar. Capsule 1-celled or imperfectly 3-celled.
118. Xyridaceae (p. 536). Rush-like, scapose. Flowers capitate, perfect,
3-androus, the calyx glumaceous. Capsule 1-celled.
119. Mayaceae (p. 537). Moss-like aquatic. Flowers perfect, axillary,
solitary, 3-androus; calyx herbaceous. Capsule 1-celled.
120. Commelinaceae (p. 538). Flowers perfect, regular or somewhat
irregular, with 3 more or less herbaceous persistent sepals and 3
fugacious petals. Stamens 6 or some sterile. Capsule 2--3-celled.
127. Eriocauleae (p. 566). Scapose aquatic or marsh plants, with linear
leaves and dense heads of monoecious (rarely dioecious) minute
flowers. Corolla tubular or none. Capsule 2--3-celled, 2--3-seeded.
[+] 2. Perianth small, of 6 equal persistent glumaceous segments;
flowers perfect; ovary compound.
121. Juncaceae (p. 539). Rush-like. Stamens 3 or 6. Capsule 1- or
3-celled, 3-valved.
[+] 3. Flowers without chaffy glumes, the perianth none or reduced to
bristles or sepal-like scales; flowers often monoecious or
dioecious; carpels solitary or united.
[++] Flowers capitate or upon a spike or spadix, with or without a
spathe.
122. Typhaceae (p. 547). Marsh or aquatic plants, with linear leaves, and
monoecious flowers without proper perianth, in heads or a naked spike.
123. Araceae (p. 548). Flowers perfect or monoecious upon the same
spadix, rarely dioecious, with 4 or 6 scale-like sepals or none.
[++][++] Flowers very minute, one or few from the margin of a floating
disk-like frond.
124. Lemnaceae (p. 551). Plants very small, green, mostly lenticular or
globose.
[+] 4. Perianth of 4 or 6 segments, the inner often petaloid, or none;
carpels solitary or distinct (coherent in Triglochin); seeds without
albumen; aquatic or marsh plants, often monoecious or dioecious.
125. Alismaceae (p. 553). Perianth of 6 segments, the inner petal-like.
126. Naiadaceae (p. 557). Perianth-segments herbaceous or none.
[+] 5. Flowers in the axils of chaffy scales or glumes arranged in
spikes or spikelets, without evident perianth; stamens 1--3; ovary
1-celled, 1-seeded; seed albuminous.
128. Cyperaceae (p. 567). Scales single. Perianth none or replaced by
bristles. Anthers basifixed. Fruit a triangular or lenticular achene.
Stem solid, often triangular, with closed sheaths.
129. Gramineae (p. 623). Glumes in pairs. Perianth replaced by minute
scales. Anthers versatile. Fruit a caryopsis. Culm usually hollow,
terete; sheaths split to the base.
SERIES II. CRYPTOGAMOUS OR FLOWERLESS PLANTS; destitute of stamens and
pistils, in fructification producing _spores_ instead of seeds.
CLASS III. ACROGENOUS PLANTS.