American species of Amelanchier

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AMERICAN SPECIES OF
AMELANCHIER

BY
GEORGE NEVILLE JONES

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THE UNIVERSITY OF ILLINOIS PRESS
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1946

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ILLINOIS BIOLOGICAL MONOGRAPHS

Vol. XX No. 2

PUBLISHED BY THE UNIVERSITY OF ILLINOIS
UNDER THE AUSPICES OF THE GRADUATE SCHOOL
UrzBana, ILLINOIS

EDITORIAL COMMITTEE

JouHN THEODORE BUCHHOLZ
FRED WILBUR TANNER
HARLEY JONES VAN CLEAVE

UNIVERSITY

800—1-46—28555 ienessi

AMERICAN SPECIES OF
AMELANCHIER

WITH 14 MAPS AND 23 PLATES

BY

GrorGE NEVILLE JONES

CONTRIBUTION FROM THE DEPARTMENT oF Botany
OF THE UNIVERSITY oF ILLINOIS

THE UNIVERSITY OF ILLINOIS PRESS
URBANA
1946

Copyright, 1946, by the University of Illinois Press.
All rights reserved. Permission for reproduction
in any form may be obiained from the publisher.

Manufactured in the
United States of America

Il.

II.

Ly.

CONTENTS

. INTRODUCTION
TAXONOMY Pot ng oo ee
Bibliography and Description of the Genus
Keys to Species
Key to Flowering Specimens .
Key to Fruiting Specimens
DESCRIPTION AND DISCUSSION OF SPECIES .

List oF NUMBERED EXSICCATAE .
PLATES

INDEX TO PLANT NAMES

LOE
25

i INTRODUCTION

AMELANCHIER is a genus of shrubs and small trees belonging to the sub-
family Pomoideae of the Rosaceae and including not more than two dozen
species widely distributed in North America, Europe, northern Africa,
and eastern Asia. Some kinds are highly ornamental and are planted for
the showy early white flowers, as well as occasionally for the more or less
edible fruits. Only the American species are included in this paper. Their
number is eighteen. Two others formerly included in Amelanchier, in-
habiting Guatemala and Mexico (and southern Texas), have been trans-
ferred to the genus Malacomeles, and are treated in another paper.

The origin of the generic name Amelanchier is not definitely known,
but probably it has been derived from the Provencal name of the Euro-
pean Amelanchier ovalis Medic. The American amelanchiers are known
by the common names serviceberry, sarviceberry, sarvis, maycherry, june-
berry, shadblow, shadbush, shadberry, shadblossom, shadflower, shad-
wood, sugar pear, wild pear, lancewood, boxwood, Canadian medlar,
bilberry, snowy mespilus, saskatoon, and perhaps some others. These
vernacular names are used as follows: serviceberry, because of the
similarity of its fruit to that of the European service tree (Sorbus tor-
minalis (L.) Crantz); juneberry because the berry-like fruits of certain
species ripen in June; in the eastern part of the United States the names
shadblow, shadberry, shadblossom, shadflower, and shadwood are used for
certain species that are in bloom when shad begin to ascend the streams.
Lancewood and boxwood have been applied to other species because their
wood has been used for handles for tools. The name saskatoon, used in
western Canada for the fruits of Amelanchier alnifolia Nutt. and the
bushes on which they grow, originated with the Blackfoot Indians, who
used the fruits either fresh or dried. The pemmican of the Indians was
composed of dried and pulverized deer or buffalo meat to which was added
saskatoon berries, the mixture then being stirred into boiling fat, and
when cooled molded into cakes. Explorers and prospectors found the
fruits a welcome addition to their food supply, and it is recorded that the
fruit of A. alnifolia was used by the members of the Lewis & Clark
Expedition when they ran short of other food. The foliage of some of the
western species is a palatable forage for grazing animals. Some of the
eastern species make a satisfactory stock on which to graft the pear
and quince.

The species of Amelanchier are closely related and are sometimes
somewhat difficult to distinguish. One or more is found in each province
of Canada, and in every state in the United States. The earlier students
of the North American flora, including Michaux, Pursh, Nuttall, Torrey,
and Gray, took the view that the genus in the western hemisphere con-

7

8 ILLINOIS BIOLOGICAL MONOGRAPHS

sisted of only one, or at the most very few, highly variable species. This
attitude characterized the understanding of the genus in the United States
during the greater part of the nineteenth century. Subsequent students
have, however, broken away from this viewpoint, and there now may be
found in botanical literature nearly two hundred binomials and trinomials
representing the species of Amelanchier in America.

In 1912 the late Professor K. M. Wiegand, of Cornell University,
presented a tentative revision of Amelanchier in eastern North America,
recognizing eight species, including three newly described by himself.
Subsequently, he described three more. His work has been followed
rather closely by almost all recent writers of manuals, floras, and
check-lists. It now appears that some species were improperly typified,
and that the assumption was made that some of the nomenclatural types
were “hybrids” and therefore to be discarded. This policy has been the
cause of a certain amount of confusion in the taxonomy and nomencla-
ture of several species. Probably this inexact typification was at least
partly responsible for the vagueness and haziness of the specific lines,
with the result that many specimens were supposed to belong to hybrid
plants rather than to true species. In later years, Wiegand (Science, n.s.
81:161-166. 1935) concluded that there are in eastern North America

. Six or seven good species with normal ranges coinciding in general with the
geographical areas in eastern North America, and fitting in with the ranges of other
plants. The hybrids were usually local. Circumstantial evidence, therefore, seemed
tc indicate that these true species had been in existence a long time, during which
they had spread over wide areas, as for instance from Newfoundland to Georgia
and Minnesota — over all the area having suitable habitat conditions. They were old
enough to have become more or less static as far as distribution was concerned.
The hybrids seem like swarms of bees, buzzing around for a time, only to disappear,
leaving the fundamental species to continue through the ages. ... It can not be
denied that species may have arisen through hybridity, quite possibly in the ways
suggested by many geneticists, but one can not become very enthusiastic. At least,
it seems evident that species are not being formed every day, or even every year,
or even every century, as some enthusiasts are inclined to think.

In the genus Amelanchier, as in many other genera of vascular plants,
hybridization between different species doubtless sometimes occurs, and
the hybrids are recognizable by those familiar with the species, but as
Wiegand has already pointed out the hybrids constitute an insignificant
element in the flora. Referring to the subject of hybridization in Rubus,
Dr. L. H. Bailey (Gentes Herbarum 2:272, 273. 1932) writes as follows:

We do not elucidate the blackberry problem by the assumption of miscellaneous
hybridity as if the species themselves were known and all the puzzles were mixed
progeny: our work takes a new direction the moment we cease to invoke crossing
as a way to escape from difficulties. The fact that certain forms are puzzling and
of doubtful specific validity does not make them hybrids. Hybrids there may be,
but the first effort is to determine the species which are supposed to spawn into
mongrels. Hybridity is to be accepted only on evidence; it can not be determined
by the examination of usual herbarium specimens.

AMERICAN SPECIES OF AMELANCHIER—JONES 9

Recent trends in taxonomic studies of vascular plants emphasize the
importance of exact typification and detailed diagnosis and description,
as well as critical examination of a large series of specimens from the total
geographical area of each species. Through the application of this method
it is usually possible to outline a much more precise concept of the
specific entities, and to interpret more accurately the boundaries between
species, than was possible in former times. The net result of this sharper
delineation is that the number of puzzling “intermediates,” “hybrids,”
“varieties,” “forms,” and other taxonomic collectanea is appreciably
diminished.

In a recent study (Gentes Herbarium 5:912, 913. 1945), Bailey says:

The office of taxonomy is to distinguish and define the units, which we call
genera and species, and then to name the units so that they may be assembled into
larger classes: the word taxonomy signifies classification. When we confuse the
definitions we obscure the program. Taxonomy is not primarily the discovery of
all detectable differences or merely the keeping of records. We are to discover the
significancies to use in schemes of arrangement. ... Marked variations which pre-
sumably have some constancy and are associated with geography or environment
may be named and formally described in a taxonomic program, but study of varia-
tion itself is not a nomenclatorial problem, and the naming of inconstant or fugi-
tive differences may obscure the undertaking. Taxonomy, or classification, stresses
agreements.

Apart from the frequent assumptions of hybridity, and the former
‘ uncertainty connected with the identity of the holotypes, one of the other
principal causes of confusion in the taxonomy of Amelanchier is the fact
that specimens showing mere variations of foliage have been not infre-
quently described as new varieties or even species. Anyone who studies
Amelanchier in the field, or who examines large series of specimens in
herbaria, is at once struck by the extraordinary variation of the foliage
that occurs even in the same species, as manifested in different stages of
development and from various habitats. There is a great deal of difference
between the appearance of specimens collected in the spring when the
leaves are of thin texture and more or less pubescent, and specimens
from the same shrub or tree in late summer or autumn when the leaves
are larger, frequently somewhat coriaceous, and, with the exception of
two western species, nearly or quite glabrous. The leaves of young shoots
are usually much larger, and of diverse shapes and types of indentation.
In the present paper no attempt has been made to describe these leaves;
probably such descriptions would be either impossible or useless; instead,
the descriptions have been drawn up from average normal leaves of the
flowering and fruiting branches. For several of the species the following
stages must be recognized and correlated before all specimens can be
correctly identified: (1) specimens with unopened flowers and very
young leaves, (2) specimens with opened flowers accompanied by
half-unfolded leaves, (3) specimens from which the petals have

10 ILLINOIS BIOLOGICAL MONOGRAPHS

dropped, but with young unfolded leaves and young fruits, (4) specimens
with mature leaves and fruits, (5) specimens with mature leaves but no
fruits, and (6) specimens consisting only of leaves of vigorous young
shoots. When placed side by side, specimens of the same species in these
different stages of development often show an almost incredible dissimi-
larity and have been not infrequently mistaken for different species.
However, collections of these different stages made over a period of years
from the same plant, as from plants in cultivation in the Arnold Arbo-
retum, and in the collections of feral plants made by critical field
botanists, show conclusively that these extraordinary variations and “in-
tergrades” are not at all, or only very rarely, due to hybridism, as has been
frequently supposed, but are well within the range of normal variation of
the species, not only among individuals in different habitats and other
ecological conditions, but among different specimens from the same
shrub or tree.

The taxonomic value of the kind of serration of the leaf-blades has
to be estimated with caution. Nearly every species has a characteristic
type of serration, but the range of variation within the species is fre-
quently greater than that found between species. The statements, e.g.,
“leaves finely toothed,” or “coarsely toothed,’ commonly used as key
characters, may be often somewhat misleading, even though attempts are
made to give them a semblance of quantitative value by recording the
number of teeth per centimeter on average leaves, or the total number of
teeth found on average blades. This is inevitable, because many species,
including A. spicata, A. florida, and A. pallida, may have the blades finely
toothed when young, while the mature leaves have a coarser indentation
with fewer or larger teeth. It is only by confining the statement of inden-
tation to the leaves of the flowering branches, or the fruiting branches,
respectively, and then only by using the most general characterizations,
that it is at all possible to use this character as a practical guide to the
specific identity of the specimen under observation.

Similarly, over-emphasis must not be placed on the habit of the plant,
which varies, of course, according to the habitat. Three American species
—A. arborea, A. interior, and A. laevis—are arborescent, and under favor-
able conditions may become small or even moderate-sized trees, but under
adverse conditions, shrubs many years of age may be only a foot or two
in height. Certain other species, e.g., A. spicata, are usually dwarf, but
under exceptionally favorable conditions may reach a height of two
meters. However, it can be safely predicted that the truly frutescent
species will never become trees, no matter how suitable the environ-
mental conditions may be. In the western part of the continent especially,
altitude is an important factor affecting the appearance of the plant, as is

AMERICAN SPECIES OF AMELANCHIER—JONES 11

also the wetness or dryness of the habitat. Certain species, e.g., 4. florida,
characteristically found at or near sea-level, may ascend the mountains to
an altitude of 5000 feet or higher, where they show corresponding varia-
tions in their foliage, habit of growth, and general appearance. All these
ecological variations must be taken into consideration in attempting to
define the species.

In Amelanchier, as in most other genera of flowering plants, by far
the most useful taxonomic characters are to be found in the morphological
peculiarities of the flowers and fruits. The number of carpels, the length,
and degree of fusion of the styles, the amount of pubescence of the
top of the ovary, the number of stamens, the shape and size of the petals,
and the shape, size, and direction of growth of the calyx-lobes, as well as
the character of the inflorescence, are the best and most reliable structural
criteria for taxonomic purposes.

Several explanations, including the theory of frequent hybridization
previously mentioned, have been advanced from time to time to account
for the peculiar variations within Amelanchier and related genera. How-
ever, certain available cytological data, although at present rather meager,
seem to provide a more plausible alternative theory. The genera of the
Pomoideae, including Amelanchier, that have been studied cytologically
have been found to have a basic chromosome number of seventeen, or are
polyploids with a basic number of seventeen. K. Sax (Journ. Arnold Arb.
12:3-21. 1931) reports that the pure species of Amelanchier that he
studied are diploids, but two natural interspecific hybrids are tetraploids.
In the same year, A. A. Moffett (Proc. Roy. Soc. Lond. ser. B, 108:423-
446, 1931) investigated four species of Amelanchier and found that they
are tetraploids. The chromosome number of the Pomoideae is a “second-
ary basic number (unbalanced relative to the primary basic number) and
the derived series of polyploids (2n = 34, 51, 68) are secondary poly-
ploids.” As previously noted, the great variation within the genus Ame-
lanchier often makes taxonomic study extremely difficult. “Such a con-
dition is to be expected from a group of complex polyploids, whose
polyploidy furnishes a mechanism for the segregation of numerous varia-
tions, but whose general character is evidently determined by their char-
acteristic balance.”’ It seems possible, therefore, that polyploidy may have
played a more important part in the differentiation of genera and species
in this subfamily than has heretofore been realized, since a change in
chromosome balance is usually accompanied by a change in the morpho-
logical characters of the plant.

The following description of the wood of Amelanchier is taken from
S. J. Record & R. W. Hess, Timbers of the New World, Yale Univer-
sity Press, 1943, pp. 447-448:

12 ILLINOIS BIOLOGICAL MONOGRAPHS

Heartwood brown or reddish brown, usually absent from small specimens;
sapwood thick, slightly brownish; appearance of lumber usually marred by numer-
ous brown lines (pith flecks). Luster medium. Odor and taste absent or not dis-
tinctive. Hard, heavy, compact, tough, and strong; sp. gr. (air-dry) 0.85; weight
53 lbs. per cu. ft.; texture fine and uniform; grain straight to irregular; rather
easily worked, taking a good polish; dark heartwood durable. Of no commercial
possibilities because of the scarcity and small size of the trees.

In the preparation of this paper, it has been necessary to study botan-
ical material in several herbaria. For the loan of specimens, and for other
courtesies, I wish to express my appreciation to the curators who have so
generously placed the necessary specimens at my disposal. For assistance
in the completion of this work I am particularly grateful to Dr. E. D.
Merrill, Professor Alfred Rehder, and Dr. A. C. Smith, all of the Arnold
Arboretum; to Dr. M.-L. Fernald and Dr. L. B. Smith, -of thesGray,
Herbarium; to Dr. J. M. Greenman, of the Missouri Botanical Garden;
to Dr. F. W. Pennell and Mr. Bayard Long, of the Academy of Natural
Sciences of Philadelphia; to Dr. H. A. Gleason and Mr. G. L. Wittrock,
of the New York Botanical Garden; to Dr. Th. Just, of the University of
Notre Dame, for the opportunity of examining material in the Greene
Herbarium; to Dr. C. O. Rosendahi and Dr. E. C. Abbe, of the University
of Minnesota; and to Dr. H. L. Mason, of the University of California,
for a few specimens from Lower California. Thanks are due Dr. Leon
Croizat of the Arnold Arboretum for nomenclatural and bibliographical
assistance.

In the citation of specimens, the name of the herbarium to which the
particular specimen belongs is indicated by the following standard abbre-
viations: (AA) Arnold Arboretum, (GH) Gray Herbarium, (NE) New
England Botanical Club, (MBG) Missouri Botanical Garden, (UI) Uni-
versity of Illinois, (Ph) Academy of Natural Sciences of Philadelphia,
(NY) New York Botanical Garden, (ND) University of Notre Dame,
(Minn.) University of Minnesota, and (UC) University of California.
Because of the large amount of material studied, it has been practicable
to cite only part of the collections.

Il. TAXONOMY
BIBLIOGRAPHY AND DESCRIPTION OF THE GENUS

Amelanchier Medicus, Phil. Bot. 1:135, 155 (1789), Geschichte 79 (1793) ; Moench,
Meth. Pl. 682 (1794); Lindley in Trans. Linn. Soc. Lond. 13:100 (1822); De
Candolle, Prodr. 2:632 (1825); Spach, Hist. Nat. Veg. Phan. 2:82 (1834);
Hooker, Fl. Bor. Am. 1:202 (1834); Endlicher, Gen. Pl. 1237, 6345 (1836) ;
Loudon, Arb. & Frut. Brit. 2:874 (1854) ; Torrey & Gray, Fl. N. Am. 1:473
(1840) ; Bentham & Hooker, Gen. Pl. 1:628 (1865); Wenzig in Linnaea 4:105
(1874) ; Decaisne in Nouv. Arch. Mus. Hist. Nat. Paris 10:133 (1874) ; Brewer
& Watson, Bot. Calif. 1:189 (1880) ; Sargent, Silva N. Am. 4:125 (1892) ; Dip-
pel, Handb. Laubh. 3:388 (1893); Focke in Engler & Prantl, Nat. Pflanzenf.
3°:26 (1894); Britton & Brown, Illustr. Fl. N. U.S. 2:237 (1897); Small, FI.
Se. U.S. 531 (1903); Britton, Man. Fl. N. States 517 (1901); Card in Bailey,
Cyclop. Am. Hort. 57 (1904) ; Sargent, Man. Tr. N. Am. 360 (1905) ; Schneider,
-Illustr. Handb. Laubh. 1:731 (1906); Dalla Torre & Harms, Gen. Siphon. 207
(1907) ; Hough, Handb. Tr. N. States & Canada 439 (1907) ; Robinson & Fern-
ald in Gray, New Man. Bot. (ed. 7) 459 (1908) ; Britton & Shafer, N. Am. Tr.
436 (1908) ; Coulter & Nelson, New Man. Rocky Mt. Bot. 265 (1909) ; Wiegand
in Rhodora 14:117 (1912) ; Britton & Brown, Illustr. Fl. N. U.S. (ed. 2) 2:291
(1913) ; Rehder in Bailey, Stand. Cyclop. Hort. 272 (1914) ; Wooton & Stand-
ley in Contr. U.S. Nat. Herb. 19:322 (1915); Farwell in Rep. Mich. Acad. Sci.
17:172 (1916); Rydberg, Fl. Rocky Mts. 446 (1917); Bailey, Man. Cult. PI.
378 (1924); Jepson, Man. Fl. Pl. Calif. 509 (1925); Wiegand & Eames, FI.
Cayuga Basin 246 (1926); Rehder, Man. Cult. Tr. & Shr. 388 (1927) ; Rosen-
dahl & Butters, Tr. & Shr. Minnesota 213 (1928); Small, Man. Se. Fl. 635
(1933) ; Munz, Man. S. Calif. Bot. 229 (1935); Marie-Victorin, Fl. Laurent.
315 (1935) ; Jepson, FI. Calif. 2:233 (1936); Nielsen in Am. Midl. Nat. 22:160
(1939) ; Rehder, Man. Cult. Tr. & Shr. (ed. 2) 386 (1940) ; Kearney & Peebles
in U.S. Dept. Agric. Miscel. Publ. 423:392 (1942); Abrams, Illustr. Fl. Pac.
States 2:470 (1944).

Amelanchus Rafinesque in Act. Soc. Linn. Bordeaux 6:265 (1834); Merrill in Proc.
Am. Phil. Soc. 86:89 (1942).

Amelancus Rafinesque, Fl. Tellur. 3:5 (1836) [1837]; F. Mueller ex Vollmann, FI.
Bayern 453 (1914).

Amelancher Bub. Fl. Pyren. 2:580 (1900).

Type Species: Mespilus Amelanchier L. = A. ovalis Medic.

Slender often scaly-barked shrubs or small trees with unarmed
branches and slender terete branchlets; wood hard, ring-porous; pith
slightly 5-sided, pale, continuous; leaves simple, deciduous, alternate,
petioled, pinnately veined, usually serrate or sometimes entire; leaf-scars
narrowly crescent-shaped; bundle-traces 3; stipules linear, caducous, free
from the petiole; stipule-scars lacking; winter-buds solitary, sessile, con-
spicuous, with several imbricate scales; flowers perfect, regular, ento-
mophilous, in racemes (rarely solitary or paired) terminating short leafy
branchlets of the season, appearing shortly in advance of the foliage, or
as the leaves unfold; pedicels bracteate at the base and bearing a second
bract at or near the middle; bracts scarious, pubescent, linear, deciduous
about the time the flowers open; hypanthium campanulate or urceolate,
more or less adnate to the carpels, becoming globose or ellipsoid in fruit;

13

14 ILLINOIS BIOLOGICAL MONOGRAPHS

disk nectariferous; calyx 5-lobed or 5-cleft, the lobes narrow, entire,
imbricate in aestivation, persistent, becoming revolute or reflexed on the
fruit, or sometimes remaining erect or ascending; petals 5, white, or rarely
pink, oblanceolate to narrowly oval; stamens 10-20, short, inserted on the
rim of the calyx; filaments subulate, more or less persistent on the fruit;
styles 2-5, free or united at the base or to the middle; carpels 2-5, more or
less united to form an inferior, compound, 2-5-loculed ovary, each locule
2-ovuled, but in fruit nearly divided by a false partition growing from the
back of each carpel, thus forming an incompletely 4-10-loculed pome
with one seed in each locule if all mature; carpel-walls of firm texture, not
bony; pome small, mealy or juicy, berry-like, edible but often insipid;
seeds small, smooth, dark brown; no endosperm.

Species and varieties of Amelanchier have been placed under a num-
ber of generic names, including Amelanchus, Amelancus, Aronia, Cra-
taegus, Malus, Mespilus, and Pyrus, and have been transferred from
one generic category to another. Amelanchier occupies a distinct position
in the Pomoideae by its usually racemose inflorescence, distinctive foliage,
and type of fruit with the peculiar carpels which have a false partition
growing from the back of each. This latter character is also present only
in the closely related monotypic genus Peraphyllum of western United
States, and in Malacomeles of Mexico and Guatemala.

KEYS TO THe AMERICAN SPECIES
KEY TO FLOWERING SPECIMENS

1. Styles normally 5 (rarely 4); stamens 20; fruit becoming purplish black at
maturity, glabrous, usually juicy and edible, mostly 10-loculed.
2. Top of the ovary glabrous.
3. Sepals more or less pubescent, at least on the ventral surface.

4. Petals 3-9 mm. long; racemes erect; young leaves densely white-tomen-
tose beneath; mature leaves elliptical or oval, serrulate acutish or obtuse
and apiculate, the base rounded or acutish; Atlantic Coastal Plain.

5. Racemes 2.5-6 cm. long, on leafy branchlets, the lower pedicels becom-
ing 5-10 mm. long; petals oblanceolate; styles 4 mm. long, united beyond
the middle; anthers 1 mm. long; fastigiately branched, alder-like
shrubs, 3-8 m. tall, forming close bushy clumps; chiefly in swampy
woods, moist thickets, or bogs............00ccee ce eee 7. A. canadensis

5. Racemes 1-3 cm. long, compact, leafless, the precocious flowers on very
short pedicels only 1-3 mm. long; petals elliptical; styles 2-3 mm. long,
separate to below the middle; anthers 0.5-0.7 mm. long; low surculose
colonial shrubs 0.2-1 m. high, forming loose colonies in open woods or
Sandy, pine Danrenss sas esse ee eee ee ace er 8. A. obovalis

4. Petals of the fully opened flowers 12-25 mm. long.

6. Leaves short-acuminate at the apex, cordate or rounded at the base,
usually ovate or obovate, finely serrate; racemes loose, becoming pen-
dent or spreading, 4-12 cm. long; trees or tall shrubs of eastern North
America.

AMERICAN SPECIES OF AMELANCHIER—JONES 15

7. Leaves at flowering time about half-grown, nearly or quite glabrous,
or only sparsely pubescent, bright green or bronze purple; sepals
lanceolate, acuminate; racemes glabrous or nearly so; fruit sweet
and juicy; lowest fruiting pedicels mostly 2.5-5 cm. long. .5. A. laevis

7. Leaves at flowering time scarcely unfolded, densely white-tomentose
beneath; sepals oval-lanceolate, abruptly pointed; racemes sericeous;
fruit somewhat dry, mealy and tasteless, falling early; lowest fruit-
AISA MEMICE]S eZ oC IONS. yates leicvaan an Sore oicie Oo See 6. A. arborea

. Leaves obtuse or acute at the apex, not acuminate; racemes erect or
ascending, 2-5 cm. long; shrubs 1-3 m. tall; western North America.

8. Petals 16-25 mm. long, 5-7 mm. wide above the middle; styles 3-4
mm. long; anthers 1-15 mm. long; leaf-blades thin, green, oval,
25-oncms lone atwmatunitye.. cakes. ct oes cevaee ser 14. A. cusicku

8. Petals 12-16 mm. long, 3-4 mm. wide above the middle; styles 2-2.5
mm. long; anthers 0.8 mm. long; leaf-blades at maturity firm, pallid,
suborpicular, 105-3 cm. lOmg..: nesses ee oes 15. A. basalticola

3. Sepals, hypanthium, and pedicels perfectly glabrous, glaucous; whole plant
perfectly glabrous; petals 8-12 mm. long; western United States..........

0010 0 ies ees ST RS CRE TEP near rt =o 16. A. pumila

2. Top of the ovary tomentose, usually densely so, rarely with only a few
trichomes.
9. Leaves permanently puberulent or finely tomentulose, at least on the lower
surface; petals 5-11 mm. long; shrubs of western United States.

10.

10.

Sepals lanceolate to deltoid-lanceolate; hypanthium and pedicels sparsely
pubescent to glabrous; leaves with 7-9 pairs of lateral veins; second-year
twigs usually brown and glabrous; top of hypanthium constricted on the
fruit; California, southern Oregon and western Nevada....17. A. pallida
Sepals linear or narrowly lanceolate to linear-spatulate; hypanthium and
pedicels usually more or less densely lanate; leaves with 11-13 pairs of
lateral veins; second-year twigs often puberulent, gray; hypanthium not
at all or only slightly constricted on the fruit; Montana to New Mexico,
westward to Lower California and eastern Oregon...... 18. A. utahensis

9. Leaves at maturity glabrous or somewhat pubescent, never finely puberu-
lent; the young leaves either glabrous or glabrescent (occasionally tardily
so), or else densely floccose-tomentose, at least on the lower surface.

11.

1

Leaves usually more or less densely tomentose or floccose beneath at
flowering time; shrubs or trees of the eastern half of North America.
(Occasional specimens of the cordilleran A. alnifolia with tardily gla-
brescent leaves might also be sought here, as well as A. arborea var.
alabamensis, whose flowers are unknown. )

12. Petals 4-10 mm. long; dwarf colonial shrubs 0.2-2 m. tall; leaves
usually with 7-9 pairs of irregular lateral veins anastomosing near
the usually rather finely serrate margins............... 9. A. spicata

12. Petals (of the fully opened flowers) 11-22 mm. long; shrubs (or
small trees) 2-6 m. tall; leaves with 11-13 pairs of regular parallel
conspicuous lateral veins, the uppermost extending into the teeth;
margins coarsely serrate-dentate...............0000. 10. A. sanguinea

Leaves not densely tomentose at flowering time, unfolded and more than

half-grown, soon becoming nearly or quite glabrous on both surfaces.

13. Leaves finely serrate, the teeth mostly 6-10 per cm., and 20-40 on
each margin of average leaves of the flowering and fruiting branches.
14. Flowers in 3-12-flowered racemes; petals oblanceolate, 8-13 mm.

long; leaf-blades mostly rounded or slightly cordate at the base.
15. Leaves usually elliptical or somewhat obovate, unfolded and
more than half-grown and dull green and glabrous at flower-
ing time, obtuse or acute at the apex, the base rounded; a

16

ILLINOIS BIOLOGICAL MONOGRAPHS

19:

low, stoloniferous, diffusely-branched shrub 0.3-1 m. tall;

western Newfoundland, St. Paul I., the Magdalen Is., Anti-

costi I., and the coast of Quebec along the Gulf of St.

Wea Wren Cee ajay mens edet aitiere oC Gee renee Cree 2. A. fernaldu

Leaves ovate or oval.

16. Racemes 2-4 cm. long; leaves ovate, shortly acuminate,
unfolded and more than half-grown and nearly or quite
glabrous at flowering time; slender shrub 1-3 m. tall;
Quebec to Massachusetts and Vermont, and adjacent New
IMOnK fey Booka sete eRe RIE ORO OEE 3. A. neglecta

16. Racemes loose, 4-7 cm. long; leaves oval, acute, unfolded
but not fully grown at flowering time, the lower surface
sparsely floccose-pubescent, varying to nearly or quite
glabrous, sometimes the pubescence remaining until the
petals have fallen; straggling or arching shrubs 2-3 m.
tall, or small trees 7-8 m. tall; Minnesota, Wisconsin, and
NOLthMeRm WH Chigatiseems sss asc eer ioe 4. A. interior

14. Flowers mostly in pairs or threes (or solitary), one terminal and
the others in the axils of the upper leaves; blades glabrous, flat,
oval, acute at each end; petals oval, 6-9 mm. long; a several-
stemmed, loosely cespitose shrub 0.5-3 m. tall; Labrador and New-
foundland to the mountains of New England and New York to
the Pocono Plateau, Pennsylvania, westward through Ontario to
northern Michigan, and northwestern Minnesota...............

eee

Sa ee eT ee tee POR Pea Aer Men een war es 1. A. bartramiana

13. Leaves more coarsely serrate, the teeth usually not more than 2-5
per cm., and only 5-20 on each margin of average blades.
17. Petals 6-10 mm. long, 2-3.5 mm. wide.

18.

18.

Petals glabrous throughout; mature leaves relatively thin,
the lower surface pale green; low shrubs 30-90 cm. tall;
Gaspé Peninsula, Quebec, westward to the region about Lake
SUNCH ORs Om aakeeeuar nee aeys odes emit aes tee 11. A. gaspensis
Petals pilosulous on the inside of the claw; mature leaves
somewhat coriaceous, glaucous beneath; shrub or small tree
2-4 m. tall; Rocky Mountains and Great Plains of western
Nort Anterica. 2,4.<195> sees ange wne se 12. A. alnifolia

17. Petals 12-25 mm. long.

19.

19.

Petals 12-15 mm. long (rarely shorter), 3-3.5 mm. wide;
sepals deltoid-lanceolate, 2-2.5 mm. long; styles 2-2.5 mm.
long; anthers 0.5-0.7 mm. long; summit of the ovary densely
tomentose; Pacific Slope, from Alaska to northern California,
west of the Cascade Mountains................ 13. A. florida
Petals 16-25 mm. long, 5-7 mm. wide; sepals lanceolate,
acuminate, 3.5-5 mm. long; styles 3-4 mm. long; anthers
1-1.5 mm. long; summit of ovary nearly glabrous or with
a ring of tomentum around the base of the styles; British
Columbia to eastern Washington, eastern Oregon, Idaho,
northern Utah, and western Montana......... 14. A. cusicku

1. Styles 4, 3, or 2 (rarely 5) ; stamens 10-15 (-18); petals 5-10 mm. long; racemes

erect or ascending, 2-4 cm. long; fruit usually small, 3-8-loculed, often drying
brownish before maturity.
20. Petals oblanceolate.
21. Leaves at maturity glabrous.
22. Hypanthium sparsely pubescent outside at flowering time; western

Washington and adjacent southern British Columbia................

Ae eee eee arc ete tate ome 13a. A. florida var. humptulipensis

AMERICAN SPECIES OF AMELANCHIER—JONES 17

22. Hypanthium glabrous; southwestern United States and adjacent Mex-
HCOMMCIADFOUS TOLIMN'S: Obit ects cena. s.asisslecleeteane eels 18. A. utahensis
21. Leaves more or less puberulent or tomentulose, at least beneath; Rocky

Mt. region, New Mexico to Montana, westward to eastern Oregon,

southeastern California and adjacent Mexico........... 18. A. utahensis
20. Petals oval or obovate; leaves puberulent (rarely glabrous) ; southwestern
Oregon, California, and adjacent Nevada..................005. 17. A. pallida

KEY TO FRUITING SPECIMENS

1. Leaves glabrous at maturity, or rarely slightly pubescent beneath, never per-
manently puberulent.
2. Styles normally 5; fruit glabrous and purplish black when ripe, usually juicy
and edible, mostly 10-loculed.
3. Top of the ovary tomentose, or tomentulose, rarely becoming nearly glabrous
with age.

4. Average leaves of the fruiting branches finely and sharply serrate or
serrulate, the lanceolate teeth usually 5-10 per cm.; species occurring
east of the 100th meridian.

5. Style-base (as seen on the young fruit) thickened and tapering into
the conical top of the densely tomentose ovary; styles on the young
fruit 4-6 mm. long; sepals 3-4 mm. long, divaricate or reflexed, subu-
late-lanceolate; hypanthium more or less constricted below on the
young fruit.

6.

Leaves oval or ovate, usually acute at each end, the petioles 4-12
mm. long; fruits solitary or in pairs or threes, or on robust speci-
mens 4 or more in a somewhat corymbose raceme, ellipsoid-ovoid
or somewhat pyriform, the larger 1-1.5 cm. in diameter when fully
TVA BUT Ceaser nen meters eacree serene seo eRe tee oe eaten 1. A. bartramiana

. Leaves mostly elliptical or somewhat obovate, obtuse or acute

at the apex, the base rounded; petioles 1-2.5 cm. long; fruits sub-
globose, 6-10 mm. in diameter at maturity, 5-10 in a slender raceme
SREY ee Eager er cRr ee oes oe Gh te Me casos chee reas 2. A. fernaldit

. Style-base not conspicuously thickened, the top of the ovary rounded

or flattish; styles on the young fruit 2-3 mm. long; sepals reflexed or
revolute, mostly oblong-lanceolate or triangular-lanceolate; leaf-base
usually rounded or subcordate.

fi:

Sepals 3-4 mm. long, reflexed; hypanthium not constricted on the

young fruit.

8. Young leaves densely white-tomentose; tree 5-10 m. tall; mature
leaves ovate or oval, acute or acuminate; Arkansas and Alabama
NRC eRe ec een reco + aan er nT Pec ee 6a. A. arborea var. alabamensis

8. Young leaves green and glabrous or nearly so from the first.

9. Leaves ovate, shortly acuminate; shrubs 1-3 m. tall; flowers
in erect or ascending racemes 2-4 cm. long; Quebec to Massa-
chusetts and Vermont, and adjacent New York..3. A. neglecta

9. Leaves oval, acute; straggling or arching shrubs 2-3 m. tall, or
small trees 7-8 m. tall; flowers in nodding racemes 4-7 cm.
long; Minnesota,. Wisconsin, and northern Michigan........
STE re IE mas ae eet eo ge a OR 4. A. interior

. Sepals 1.5-2.5 mm. long, usually revolute; hypanthium noticeably

constricted on the young fruit; leaves commonly oval, acutish or
obtuse, densely white-tomentose beneath when young; dwarf surcu-
lose colonial shrubs 0.2-2 m. tall... 0.0.00... ce eee eee 9. A. spicata

4, Average leaves usually more coarsely toothed, the ovate teeth mostly
3-6 per cm.

18

10.

10.

ILLINOIS BIOLOGICAL MONOGRAPHS

Lower surface of mature leaves often pale green, but not glaucous;
fruits 6-8 mm. in diameter at maturity; species occurring east of the
100th meridian.

11. Tall straggling shrubs or small trees 2-6 m. tall; blades coarsely
dentate-serrate, the lower surface, the petioles, and the rachis and
pedicels of the young fruiting racemes often retaining traces of
pubescence; lateral veins usually 11-13 pairs, conspicuous, parallel,
the upper ones ending in the teeth; styles on the young fruit
MOStlys3=5) MMs ONG. 0s ecw seis sac arretaee oe eee 10. A. sanguinea

11. Low shrubs 0.3-2 m. tall; principal lateral veins of the leaves
fewer and less regular; styles on the young fruit 2-3 mm. long.
12. Principal lateral leaf-veins mostly 9-11 pairs, the upper ones

usually extending to the margin and ending in the teeth, the
others anastomosing at their tips; racemes and mature leaves
quite glabrous; low, much-branched shrubs 30-90 cm. tall,
often forming dense thickets; Gaspé Peninsula, Quebec, west-
ward to the region about Lake Superior..... 11. A. gaspensis
12. Principal lateral veins mostly 7-9 pairs, somewhat irregularly
and distantly arranged, usually anastomosing before reaching
the margin; racemes, petioles, and lower part of underside of
blades often retaining traces of grayish tomentum; low
stoloniferous colonial shrubs 0.2-2 m. tall; Newfoundland to
eastern North Dakota, southward to Missouri and Georgia
Sei ths Bao AN oye Ps Pala, sha aPor snaieteacs sc aah oe SOuAN RT OR: eA Seo 9. A. spicata

Lower surface of leaves usually glaucous; the margins ordinarily

rather coarsely toothed, chiefly on the upper half of the usually

obtuse blade; fully developed fruits normally 1-1.5 cm. in diameter;
shrubs 2-4 m. tall, or small trees, of western North America.

13. Mature blades usually more or less suborbicular, subcoriaceous;
petals 6-10 mm. long; species of the Rocky Mountains and Great
Pett Savard rap cxarortievsceatrcro are sae aversoraushnna Becta ereeepeciee oe 12. A. almifolia

13. Mature blades frequently oval and of thinner texture; petals 12-
15 mm. long; species of the Pacific Slope, Alaska to northern
California, west of the Cascade Mountains......... 13. A. florida

3. Top of the ovary glabrous, rarely with a few trichomes.
14. Leaves finely and sharply serrate or serrulate almost to the base of the
blade, the teeth usually 6-11 per cm., 20-60 on each side of average
blades; eastern North America.

TD,

15.

Low colonial surculose shrubs 0.2-1.5 m. tall; fruiting pedicels

mostly 3-8 mm. in length; sepals on the fruit usually erect or

CIV ATICATE 2 serctys ernie eer eee ne eA en er 8. A. obovalis

Tall shrubs or small trees; lower fruiting pedicels becoming 1-5 cm.

long.

16. Sepals on the fruit usually erect or ascending, rarely somewhat
reflexed; fruiting racemes erect, compact, the lower pedicels 1-2
cm. long; leaves usually elliptical, varying to narrowly obovate,
oblanceolate, or oval, the apex acute, or obtuse and mucronate;
shrubs 2-8 m. tall, somewhat fastigiately branched; chiefly
Atlantic Coastal@ Plaines. asa seine sae sce 7. A. canadensis

16. Sepals on the fruit reflexed or recurved, rarely remaining erect
or divaricate; fruiting racemes loose, the lower pedicels longer;
leaves ovate or oval, short-acuminate or acute at the apex, the
base cordate or rounded; small trees 10-20 m. tall, or tail shrubs,
with ascending or spreading branches.

AMERICAN SPECIES OF AMELANCHIER—JONES 19

17. Sepals lanceolate to subulate, acuminate, 3-4.5 mm. long on
the fruit; lowest fruiting pedicels mostly 2.5-5 cm. long;
blades, and usually the petioles, completely glabrous; fruit
SWEEE ATIC “SUCEIIIEN Ese as cr cic cts teusleeaia ee tac 5. A. laevis

17. Sepals triangular-lanceolate or oblong-lanceolate, abruptly
pointed, 2-3 mm. long on the fruit; lowest pedicels 1-2.5 cm.
long; petioles, and often the underside of the blades (at least
near the base), frequently retaining some pubescence at ma-
turity; fruit somewhat dry, mealy, insipid......6. 4. arborea

14. Leaves more coarsely serrate, the teeth 3-6 per cm., 3-15 on each side of
average blades; apex obtuse or acutish; winter buds conical, acute;
shrubs 1-3 m. tall; western North America.

18. Sepals more or less pubescent within.
19. Sepals deltoid, 3-3.5 mm. long; petals 16-25 mm. long; leaves
thin, green, mostly 2.5-5 cm. long, mostly oval....14. A. cusicku
19. Sepals linear-lanceolate, 4-5 mm. long; petals 12-16 mm. long;
leaves at maturity firmer, pallid, suborbicular, 1.5-3 cm. long

a ee tree Ree ooane. CES Rivas duc arene 15. A. basalticola

18. Sepals, hypanthia, and pedicels perfectly glabrous, glaucous; whole
plant perfectly glabrous; petals 8-12 mm. long........ 16. A. pumila

2. Styles 3 or 4; petals 6-10 mm. long.
20. Calyx and ovary pubescent; western Washington and southern Vancouver

is ]aarvleentrs ever ce ots Seteleterciov ole ese letaeie Gale avert 12a. A. florida var. humptulipensis
20. Calyx and ovary glabrous; Nevada and Arizona; glabrous forms of....
ete oeen Cet sal cia Oana tee ae He Sha Sa ceoa eas 18. A. utahensis

. Leaves usually permanently puberulent or tomentulose, at least beneath, varying

to cinereous (rarely glabrous) ; petioles pubescent (except shade forms) ; styles

usually 4, 3, or 2 (rarely 5); fruit usually small, 3-6-loculed, often sparsely
pubescent at first, frequently drying brownish before maturity; shrubs of western

United States and Lower California.

20. Hypanthium more or less constricted on the young fruit; sepals lanceolate to
deltoid-lanceolate; styles 4 or 3 (rarely 5), united below; leaves entire to ser-
rate, usually with 7-9 pairs of lateral veins; southwestern Oregon, California,
ABGeAGIACEME IN CV AGA’ coo ecieicss adie acces uNiyisn oe iets Se weltea die ss t's 17. A. pallida

20. Hypanthium not at all constricted on the fruit; sepals subulate-lanceolate to
linear-spatulate; styles 2, 3, or 4, free nearly to the base; leaves usually
coarsely toothed, varying to nearly entire, usually strongly reticulate and with
9-13 pairs of lateral veins; Rocky Mountain region, New Mexico to Montana,
westward to eastern Oregon and Lower California.......... 18. A. utahensis

III, DESCRIPTION AND DISCUSSION OF SPE Giss

1, AMELANCHIER BARTRAMIANA (Tausch) M. Roemer
(Plates I and XI)

Mespilus canadensis var. § oligocarpa Michx. Fl. Bor. Am. 1:291 (1803).

Pyrus oligocarpa nana Donn ex Muhlenberg, Cat. Pl. Am. Sept. 49 (1813).

Pyrus bartramiana Tausch in Flora 21:715 (1838).

Aronia praecox Neumann ex Tausch, l.c., pro syn.

Amelanchier canadensis var. § oligocarpa Torrey & Gray, Fl. N. Am. 1:474 (1840) ;
Torrey, Fl. N.Y. 1:226 (1843); Walpers, Rep. Bot. Syst. 2:55 (1843); Gray,
Man. 131 (1848); Farwell in Rep. Mich. Acad. Sci. 17:175 (1916).

Amelancher bartramiana M. Roem. Syn. Mon. 3:145 (1847); Wiegand in Rhodora
14:158, pl. 96 (1912); Britton & Brown, Illustr. Fl. N. U.S. (ed. 2) 2:293, fig.
2334 (1913) ; Rehder in Bailey, Stand. Cyclop. Hort. 273, fig. 188 (1914) ; Hoff-
mann in Proc. Boston Soc. Nat. Hist. 36:280 (1922); Pease in ibid. 37:267
(1924) ; Rehder, Man. Cult. Tr. & Shr. 390 (1927); Rydberg, FI. Prairies &
Plains 438 (1932); Marie-Victorin, Fl. Laurent. 317, fig. 91 (1935); Nielsen
in Am. Midl. Nat. 22:189, pl. 15 (1939) ; Rehder, Man. Cult. Tr. & Shr. (ed. 2)
389 (1940). :

Amelanchier oligocarpa M. Roem. Syn. Mon. 3:145 (1847); Watson & Coulter in
Gray, Man. Bot. (ed. 6) 167 (1889); Dippel, Handb. Laubh. 3:391, fig. 196
(1893) ; Britton & Brown, Illustr. Fl. N. U.S. 2:239, fig. 1990 (1897) ; Keeler,
Our Northern Shr. 196 (1903); Schneider, Illustr. Handb. Laubh. 1:737, figs.
411, 412 (1906) ; Britton, Man. N. States 518 (1901), (ed. 3) 518 (1907) ; Card
in Bailey, Cyclop. Am. Hort. 57 (1904); Robinson & Fernald in Gray, New
Man. Bot. (ed. 7) 460 (1908); Jones & Rand in Bull. Vermont Agr. Exp. Sta.
145:101 (1909); Apgar, Ornam. Shr. U.S. 182, fig. 280 (1910); Clements,
Rosendahl, & Butters, Minnesota Tr. & Shr. 153 (1912); Bean, Tr. & Shr.
Brit. Isles 1:189 (1914).

Amelanchier sanguinea sensu Decaisne in Nouv. Arch. Mus. Hist. Paris 10:136
(1874). Non A. sanguinea (Pursh) DC.

Amelanchier arguta Nuttall ex Britton, Man. N. States (ed. 2) 1066 (1905), (ed. 3)
1076 (1907).

Amelanchier canadensis var. pauciflora Farwell in Rep. Mich. Acad. Sci. 17:175
(1916).

A several-stemmed loosely cespitose shrub 0.5-3 m. tall; stems ascend-
ing, slender; bark grayish brown; twigs slender, glabrous, brown, with
small oval lenticels; winter buds small, brown, lanceoloid, acuminate, the
scales glabrate except the ciliolate margins; leaves flat and imbricate in
the bud, glabrous from the first, about half-grown at flowering time, firm
and pale green and slightly glaucous beneath at maturity; blades of the
flowering and fruiting branches oval, varying to elliptical, obovate, or sub-
orbicular, 3-6°cm. long, 1.5-4 cm. wide, usually acutish at each end, or the
base cuneate, less commonly somewhat rounded; principal lateral veins
10-16 pairs, not prominent, irregularly spaced, curved upward, forking
and anastomosing near the margin, often with shorter intermediate ones;
margins finely and sharply serrate or serrulate or sometimes double-
serrate to below the middle or near the base of the blade, the teeth
obliquely acuminate, 6-10 per cm., 20-40 on each margin; stipules cadu-
cous, linear, sericeous, about 1 cm. long; petioles usually short, glabrous

20

AMERICAN SPECIES OF AMELANCHIER—JONES 7A

or sparingly ciliolate, those of the leaves of the fruiting branches mostly
4-12 mm. long; flowers solitary or few together, one terminal, the others
in the axils of the upper leaves, the pedicels slender, glabrous, 1-3 cm.
long; petals white, oval, obtuse, widest near the middle, 6-9 mm. long;
stamens longer than the styles; filaments glabrous; anthers 1 mm. long,
sepals lanceolate, acuminate, glabrous outside, tomentulose within; fruits
solitary or in pairs or threes, or on robust specimens 4-6 in a somewhat
corymbose raceme, but usually only one or two of them developing to
full size at maturity; hypanthium more or less constricted on the young
fruit; sepals on the fruit varying from nearly erect to divaricate or
reflexed, (3-) 4-5 mm. long, glabrous outside, tomentulose within, lanceo-
late-subulate; styles 5 (or 4), united below the middle, 4-5 mm. long,
tomentose at the base; ovary densely tomentose, tapering into the some-
what thickened style-base; mature fruits ellipsoid-ovoid or somewhat pyri-
form, glabrous, dark purple, glaucous, edible but insipid, ripening in July
or August, the larger ones 1-1.5 cm. in diameter; pedicels glabrous, 1-3
cm. long; seeds few, or only 1 maturing, dark brown, smooth, glabrous,
asymmetrically ovoid or lanceoloid, 3.5-4.5 mm. long, 2-3 mm. thick.

Type Locarity: [Northeastern] North America. Isotype in the her-
barium of the Missouri Botanical Garden. Phototype in the herbarium of
the Arnold Arboretum of Harvard University.

Rance: Mountain woods, cold swamps and bogs, or in wet rocky soil,
often at medium or higher altitudes, from Labrador and Newfoundland
to the mountains of New England and New York, to the Pocono Plateau,
Pennsylvania, westward through Ontario to northern Michigan and north-
eastern Minnesota. Flowering from May 15 to June 10; fruit ripe in July
and August.

Laprabor: Caribou I. in 1860, Chapin (UI); Petty Harbour, Bishop 371 (GH,
AA), Hopedale, Bishop 370 (GH, AA); Makkovik, Stecker 19 (GH); Red Bay,
Sandborger 20 (GH); Betchewan, Abbe 1136 (GH).

QuEBEc: East Main, east coast of James Bay, Potter 487 (GH); Mt. Sherrick,
Potter 486 (GH); L. Mistassini, Aug. 12, 1885, Macoun (GH); Orford, Sher-
brooke Co., Pease 11969 (GH); St. Agathe des Monts, June 1, 1903, Aug. 26, 1902,
Jack (AA); Brion I., Magdalen Islands, St. John 1903 (GH); Thetford, Megantic
Co., M.-Victorin 11214 (AA); Black Lake, Megantic Co., Fernald & Jackson 10107
(GH); Lac Saint-Jean, M.-Victorin 15589 (AA); St. Pierre, Arséne 311 (GH);
Anticosti I., M.-Victorin & Rolland-Germain 27901 (GH), 27899 (GH, AA);
Matamek River, Bowman 26, 237, 402 (GH); Rimouski Co., Collins & Fernald in
1904 (GH), Fernald & Collins 1100 (GH), Rousseau 26228 (GH, AA), 26771
(AA); Harrington, Abbe 1151 (GH); Saguenay Co., St. John 90526, 90527, 90528,
90529 (GH), Robinson 782 (GH), July 14, 1892, Kennedy (GH); Sept-Iles, Gaspé
Co., M.-Victorin & Rolland-Germain 18707, 18708, 18709 (GH); Mt. Albert, Col-
lins & Fernald in 1905 (GH), Fernald & Collins 233 (GH), M.-Victorin, et al.
17436 (GH, AA); Table Top Mt., Fernald & Collins 614 (GH), Rousseau &
Fortier 31441, M.-Victorin, Rolland-Germain, & Jacques 33486, 33467 (GH);
Montagne Ste.-Anne, M.-Victorin, et al. 17433 (GH, AA).

22 ILLINOIS BIOLOGICAL MONOGRAPHS

Map 1.—Range of Amelanchier bartramiana.

NEWFOUNDLAND: Avalon Bay, Fernald & Wiegand 5574, 5574a, 5576, 5578,
5755 (GH); Valley of Exploits River, Fernald & Wiegand 5597, 5600, 5602, 5736,
5737, 5739, 5742, 5743, 5744, 5749, 5751, 5752, 5754 (GH), 5738 (GH, AA); Con-
ception Bay, Howe & Lang 1203 (GH); St. John Bay, Fernald, Long, & Fogg
1789 (GH); Spruce Brook, Kennedy 17 (GH); Crabbes Station, Kennedy 257
(GH); Notre Dame Bay, Fernald & Wiegand 5535, 5745, 5746, 5756 (GH); Bay
of Islands, Fernald, Long, & Fogg 292 (GH), Fernald & Wiegand 3554, 5748, 5750
(GH), Mackenzie & Griscom 10327 (GH), Howe & Lang 1081 (GH); Bonne Bay,
Kimball 102 (GH); Hermitage Bay, W. Palmer 1340 (GH).

New Brunswick: Bass River, June 1, 1871, July 2, 1875, Fowler (GH);
Serpentine River, Hay 63 (GH); Kent Co., May 29, 1868, Fowler (GH).

Nova Scotia: Ohio, Jack 3106 (GH); Weymouth, Jack 3344 (GH); Folleigh,
Bean, White, & Linder 21459 (GH, AA); St. Paul I., Perry & Roscoe 244 (GH).

Princ—E Epwarp IsLanp: Queens Co., Fernald; Long, & St. John 7596, 7597
(GH).

Maine: Mt. Katahdin, July 1900, Fernald (GH, NE); Somerset Co., Aug. 18,
1896, Fernald (NE), Fernald & Pease 25133 (NE), St. John & Nichols 2332
(NE); Franklin Co., Furbish in 1894 (NE), Chamberlain & Knowlton in 1902
(NE); Orono, May 14, 1892, Fernald (NE); Penobscot Co., Fernald in 1897
(NE); Marshfield, Aug. 2, 1916, Knowlton (NE); Greenville, Fernald 257 (GH,
NE); Winn, Fernald & Long 13782 (NE) ; South Poland, Furbish in 1893 (NE);
Fort Kent, Fernald 2314 (GH, NE).

New HampsuirE: “White Mts., N. Hampshire,” Nuttall (type coll. of A.
arguta) (GH); Crawford Mill Pond, June 4, 1881, Faron (GH, NE); Jaffrey,
Rand & Robinson 616 (GH); Mt. Moosilanke, July 12, 1886, Faron (GH); May
11, 1895, Churchill (GH, NE); Franconia, May 31, 1892, Faron (GH, NE),
Fernald & Smiley 11720 (NE); Tuckerman’s Ravine, Sargent in 1879 (AA),
Eggleston 2369 (GH, UI), July 6, 1888, Faxon (GH, NE), July 17, 1891, Kennedy

AMERICAN SPECIES OF AMELANCHIER—JONES 23

(NE); Mt. Washington, Greenman 1058 (GH, MBG), July 12, 1855, Wm. Booitt
(NE); Mt. Ingalls, Pease 11205 (NE); Whitefield, Pease 16683 (NE, UI); Mt.
Madison, Pease 10216 (NE); Carter Notch, Pease 4090 (NE); Carroll, Pease
14372 (NE); Stewartstown, Pease 16662 (NE); Sugarloaf Mt. Pease 13482
(NE); Berlin, Pease 25275 (NE); Pittsburg, Pease 10312 (NE).

Vermont: Mt. Mansfield, July 3, 1897, Williams (GH, NE), Churchill, July 5,
1897 (NE), Greenman 936 (MBG), Pringle in 1878 (AA), Eggleston in 1893
(GH); Willoughby, July 8, 1898, Kennedy (NE); Woodford, June 20, 1925,
Carpenter, Churchill, & Knowlton (NE); Mt. Killington, June 18, 1899, Eggleston
(NE, UI), May 1913, Dutton (GH, NE); Rutland, Eggleston 1960, 1964 (GH,
NE); Barton, May 24, 1923, Knowlton (NE); Coventry, May 19, 1932, Knowlton
(NE); Craftsbury, May 19, 1932, Knowlton (NE); Greensboro, May 18, 1938,
Knowlton (NE); Lowell, May 23, 1935, Knowlton (NE); Lunenburg, July 9, 1915,
Woodward (NE); Sutton, May 20, 1932, May 18, 1933, Knowlton (NE); Sears-
burg, Pease 19561 (NE); Stratton, Eggleston 1962 (GH), Blanchard 5 (GH, AA).

MassacHuseETTs: Mt. Greylock, June 2, 1901, Churchill (GH, NE), June 16,
1901, Williams (GH), June 25, 1916, Hoffmann (NE), G. N. & F. F. Jones 16170
(UI); Florida, July 3, 1909, July 14, 1916, Hoffmann (NE); Hubbardston,
Weatherby, Smuth, et al., Pl. Exsicc. Gray. 959 (GH, UI); Ashburnham, May 19,
1924, Knowlton (NE).

New York: Canton, Phelps 1588, 1589 (GH); Essex Co., House 7264, 9468,
10227 (GH); Tug Hill Plateau, Hotchkiss 2290 (GH); Mt. McIntyre, Muenscher
& Clausen 4020 (GH).

PENNSYLVANIA: Pocono Mts., July 27, 1893, Porter (GH).

Ontario: Wingham, Morton 2691 (UI); Long Lake, Jennings 14015 (GH);
Sandy Inlet, Krotkov 5384 (GH). .

Micuican: Keweenaw Peninsula, Farwell 52d (GH), Fernald & Pease 3360
(GH) ; Gwinn, June 6, 1909, Harrison (GH); Isle Royale, Cooper 46 (GH).

MINNESOTA: Vermilion Lake, Arthur, Bailey, G Holway B407 (GH).

An examination of a part of the type collection from the Bernhardi
Herbarium, now in the Missouri Botanical Garden, leaves no doubt of
the identity of this species, or that it is the same as Michaux’s Mespilus
canadensis var. § oligocarpa, as shown by a comparison of phototypes in
the Gray Herbarium and the Arnold Arboretum.

Amelanchier bartramiana is usually to be distinguished by the small
flowers occurring mostly in pairs or threes or solitary, one terminal and
the others in the axils of the upper leaves, which are oval, acute at each
end, short-petioled, finely serrate, and quite glabrous from the first. The
styles on the young fruits are thickened at the base and taper into the
conical top of the densely tomentose ovary. It has been usually assumed
that the number of flowers and fruits never exceeds 1-3, or 4, and that
occasional robust specimens bearing a larger number in a somewhat
corymbose inflorescence must belong to plants of hybrid origin that have
resulted from crosses with either A. laevis or A. spicata. That this need
not necessarily be the case is shown by certain specimens that evidently
possess all the other diagnostic characters usually attributed to A. bar-
tramiana. It may be pointed out, however, that when the number of fruits
exceeds two or three, usually only the terminal ones attain full size at
maturity, the others remaining small and failing to develop.

24 7 ILLINOIS BIOLOGICAL MONOGRAPHS

Amelanchier arguta Nuttall ex Britton, published in 1905 without
citation of a type or any definite reference to the source of the name,
does not differ in any essential respect from A. bartramiana (Tausch)
M. Roem. One of Nuttall’s specimens, from ‘White Mts., N. Hampshire,
also on Wachusett Mt., Mass.,” has been examined in the Gray Her-
barium,

A specimen (Weatherby, Smith, & Rollins 6909) collected near Hub-
bardston, Worcester County, Massachusetts, on May 23, 1939, appears to
be a hybrid between A. bartramiana and A. canadensis. The collection
data on the label of the specimen are: “Low, loosely cespitose shrub in
bushy flats. With parents, flowering later than A. bartramiana.”

2. AMELANCHIER FERNALDII Wieg.
(Plates, l-ands Ly.)
Amelanchier fernaldii Wiegand in Rhodora 22:149 (1920).

Low surculose, straggling, diffusely branched shrubs 0.3-1 m. tall,
growing in colonies or large clumps; bark of the branches gray or brown,
smooth; twigs glabrous; winter buds small, conical, acute, glabrous or
nearly so; leaves conduplicate in the bud, unfolded and more than half-
grown, dull green and glabrous at flowering time, of rather thin texture;
mature blades firm, elliptical or somewhat obovate, glabrous on both sides,
paler beneath, darker green and often somewhat impressed-veined above,
5-8 cm. long, 1.5-4.5 cm. wide, the apex obtuse or acute, the base rounded;
lateral veins irregular, 7-13 pairs; margins sharply serrate from near the
base, the acuminate teeth 4-10 per cm., 15-35 on each side of average
leaves of the flowering and fruiting branches; stipules linear, caducous,
0.5-1 cm. long, 0.5-1 mm. wide, glabrous or pubescent; petioles slender,
glabrous or sparsely pilose, 1-2.5 cm. long on average leaves; flowers small,
in erect or ascending, loose, 3-8-flowered racemes 2-4 cm. long; rachis and
pedicels glabrous; petals 5, white, glabrous, oval or oblanceolate, obtuse,
8-11 mm. long, 5-6 mm. wide; stamens 20, the filaments glabrous, 2-4 mm.
long; anthers 1 mm. long; hypanthium campanulate, 4-5 mm. in diameter,
glabrous; sepals linear-lanceolate, acuminate, tomentulose within, glabrous
outside, 3-5 mm. long, erect or soon irregularly divaricate; styles 5,
glabrous, united near the base, 3-4 mm. long; summit of the ovary
densely tomentose; mature fruits 6-10 mm. in diameter, 10-loculed, gla-
brous, purplish black and juicy at maturity; pedicels 3-4 cm. long; seeds
brown, smooth, obliquely lanceoloid, acute at each end, 4-5 mm. long,
2-2.5 mm. wide.

Type Locatity: Grindstone, Grindstone Island, Magdalen Islands,
Quebec, Canada. Type in the Gray Herbarium of Harvard University.

AMERICAN SPECIES OF AMELANCHIER—JONES 25

RanGE: In wet woods and thickets, borders of swamps, “lime barrens
or strongly calcareous shores and swamps” ( Wiegand, l.c.), or damp hol-
lows in sand dunes, western Newfoundland, St. Paul Island, the Mag-
dalen Islands, Anticosti Island, and the coast of Quebec along the Gulf
of St. Lawrence. Flowering from the end of June to the latter part of
July; fruit ripening in July and August.

NEWFOUNDLAND: Table Mt., Port au Port Bay, Fernald & St. John 10840,
10841 (GH), 10842 (GH, AA); Old Port au Choix, St. John Bay, Fernald, Long,
& Fogg 1792 (GH).

Nova Scotia: St. Paul Island: Ethel Lake, Perry & Roscoe 243 (GH).

Prince Epwarp Istanp: Bothwell, Fernald & St. John 11083 (AA).

QuesBEc: Magdalen Islands: Grindstone Island, Fernald, Long, & St. John
7592 (GH, TYPE), 7590 (GH), Fernald, Bartram, Long, & St. John 7586 (GH),
7589 (AA, GH); Coffin Island, Fernald, Bartram, Long, & St. John 7587 (GH);
Brion Island, St. John 1909 (GH); Anticosti Island, M.-Victorin & R.-Germain
27898 (GH); Riviere Vaureal, M.-Victorin & R.-Germain 27900 (GH); Grand
River, Gaspé Co., Fernald in 1904 (GH); Cap Original, M.-Victorin, R.-Germain,
& Jacques 33216 (GH); Sept-Iles, M.-Victorin & R.-Germain 18710, 18711 (GH);
Isle-aux-Coudres, M.-Victorin 4318 (AA, GH); Grand Barachois, M.-Victorin &
R.-Germain 9507 (GH, AA); Anse Pleureuse, M.-Victorin, R.-Germain, & E. Jacques
33436, 33439 (GH).

Although A. fernaldu has some characteristics suggesting a hybrid
origin, it “seems to form a definite unit, not a fluctuating hybrid’ (Wie-
gand, l.c.), and may be regarded as a small endemic species of the region
about the Gulf of St. Lawrence. From A. bartramiana it is to be dis-
tinguished by its racemose inflorescence, elliptical leaves with longer
petioles, and longer petals and smaller fruits.

The leaves of A. fernaldu are the same general shape as those of
A. canadensis, but there is little likelihood of the two species being con-
fused. The leaves of the former are glabrous, flat, and often nearly full-
grown at flowering time, while those of the latter are folded, less than
half-grown, and densely whitish tomentose. Also, the flowers of A. fern-
aldu, with their woolly-topped ovaries, and their longer sepals, are in
nearly glabrous, looser racemes.

3. AMELANCHIER NEGLECTA Eeglest.
(Plates I and V)
Amelanchier neglecta Eggleston, in herb.

Slender-stemmed, shrubs 1-3 m. tall, with grayish brown bark and
glabrous twigs; winter buds small, glabrous, conical; leaves conduplicate
in the bud, unfolded and more than half-grown and nearly or quite
glabrous at flowering time; mature blades ovate or oval, glabrous through-
out, paler beneath, of firm texture, 5-6 cm. long, 2-3 cm. wide, the apex
usually shortly acuminate, the base rounded or subcordate; lateral veins

26 ILLINOIS BIOLOGICAL MONOGRAPHS

7-11, curved, anastomosing toward the edge of the leaf; margins finely
and sharply serrate nearly to the base, the teeth 7-9 per cm., 20-30 on
each margin of average leaves of flowering and fruiting branches;
stipules linear, fugacious, slightly pubescent; petioles of mature leaves
1-2 cm. long, glabrous; flowers small, in erect or ascending 7-10-flowered
racemes 2-4 cm. long; rachis and pedicels glabrous; petals 5, white,
glabrous, narrowly oval, mostly 8-10 mm. long, about 4 mm. wide; sta-
mens 20, the filaments glabrous, 2-3 mm. long; anthers 1 mm. long;
hypanthium campanulate, 3-4 mm. in diameter, glabrous; sepals linear-
lanceolate, acuminate, tomentulose within, glabrous outside, 3-4 mm. long,
soon recurved or reflexed; styles 5, glabrous, united near the base, 3-4
mm. long; summit of the ovary densely tomentose; mature fruits 8-10
mm. in diameter, subglobose, 10-loculed, glabrous, purplish black and
juicy at maturity; pedicels 15-25 mm. long; seeds smooth, brown, asym-
metrical, obliquely lanceoloid, acute at each end, 4-5 mm. long.

Type Locatity: Rutland, Vermont. Collected by W. W. Eggleston in
1899. Type in the Gray Herbarium of Harvard University.

RANGE: Growing at the edge of clearings, in damp thickets and open
woods that have been recently burned, or on ledges and talus, or in mossy
spruce barrens, southern Quebec, Prince Edward Island and Nova Scotia
to Massachusetts, Vermont, and adjacent New York. Flowering in May
and June; fruits ripening in July and August.

Quesec: Gaspé Bay, Aug. 23, 1897,
Jack (AA); Montmagny, Rousseau
24557 (AA, GH); Lac Long, Rousseau
24552 (GH); Montmorency Falls, Ma-
coun 66927 (GH); Ottawa River, Bro.
Rolland 7214 (GH).

New Brunswick: Woodstock, Fer-
nald & Long 13764 (GH).

Nova Scotia: Meteghan, Fernald &
Long 21450 (GH).

Prince Epwarp Is_tanp: Charlotte-
town, Fernald & St. John 7578 (GH);
Mt. Stewart, Fernald, Bartram, Long,
& St. John 7581 (GH).

MaIne: Fort Fairfield, Fernald 37
(NE); Northport, Furbish in 1891
(NE); Brownville, Parlin 1811 (GH) ;
East Livermore, Furbish in 1878
(NE); Dover, June 29, 1894, Fernald
(NE); Fort Kent, Fernald 2312 (GH, eo
NE); Oxford, Chamberlain in 1907) ~ ot
(NE) ; Moxie Mt, Aug. 25, 1902, Cole 25 A 7\/
lins & Chamberlain (NE); Cutler, Xt
July 7, 1902, Kennedy, et al. (GH);

Mt. Katahdin, July 17, 1900, Churchill Map 2—Range of Amelanchier neglecta.

AMERICAN SPECIES OF AMELANCHIER—JONES Zi

(NE); Isle au Haut, Hill 1614 (NE); Orono, Fernald, July 6, 1892, Furbish in
1891 (NE); Winn, Fernald & Long 13768, 13769 (NE) ; City Camp, July 17, 1900,
Fernald (AA, NE, GH); Jonesboro, Aug. 1, 1932, Knowlton (NE); Roque Bluffs,
July 11, 1908, Knowlton (NE).

New Hampsuire: Flume, Sargent in 1879 (AA); Gorham, Cusick in 1887
(GH); Randolph, Moore 4189 (GH), Pease 670 (NE); Crawford House, June 6,
1881, Faxon (GH); Franconia, June 26, 1895, Faxon (GH); Colebrook, Fernald
& Pease 16825 (NE); Carroll, Pease 14374 (NE); Pittsburg, Pease 10995 (NE);
Shelburne, July 12, 1882, Deane (NE); Baldface Mt., Pease 16044 (NE); Stew-
artstown, July 19, 1917, Fernaid & Pease (NE); Gilmanton, Cushman & Sanford
1217 (NE); Gilford, Harris & Pease 26551 (NE); New Hampton, Pease 26527
(NE); Plymouth, Fernald 11719 (NE); Mt. Agassiz, July 1871, Collins (NE).

VeRMonT: Manchester, Day 379 (GH, NE); Johnson, Grout, June 8, 1895
(NE); Willoughby, May 26, 1904, May 19, 1905, Kennedy (NE); Orange, May 17,
1932, Knowlton (NE); Sutton, May 20, 1932, Knowlton (NE); Brandon, May
26, 1912, Knowlton (NE); Rutland, Eggleston 1118, 1173, 1174, 1175, May 12,
June 21, July 7, 1899 (TYPE coll., GH, NE).

MASSACHUSETTS: Westminster, May 10, 1912, Hunnewell & Wiegand (NE);
Hanging Mt., July 12, 1906, Hoffmann (NE); Lanesboro, May 19, 1920, Hoffmann
(NE); Florida, June 5, 1920, July 14, 1916, Hoffmann (NE); Sheffield, May 24,
1920, Hoffmann (NE).

New York: Black Lake, St. Lawrence Co., Muenscher & Maguire 2319 (GH),
2320 (MBG) ; Washington Co., Burnham 21 (GH); Newcomb, House 10189 (GH).

Amelanchier neglecta has often been regarded as a natural hybrid
between A. bartramiana and A. laevis because in several respects it ap-
pears to be somewhat intermediate between these two species. It resem-
bles A. /aevis in the conduplicate vernation of the finely toothed, usually
acute, or shortly acuminate leaves that are quite glabrous, even at flower-
ing time. It is, however, of smaller stature than A. Jaevis, and differs in
the constantly tomentose ovary, shorter petals, and longer styles. It also
shows some resemblance to 4. bartramiana, but differs in the habit of
growth, and in the smaller, usually more numerous, constantly racemose
flowers and fruits. The usually ovate, short-acuminate leaves, which
are rounded or cordate at the base, are of different color and texture, and
are longer-petioled than those of A. bartramiana. At flowering time they are
often slightly pubescent beneath, and unfold with the flowers, whereas in
A. bartramiana the glabrous flat leaves precede the flowers. The sub-
globose fruits have shorter styles, and the top of the ovary shows a
tendency to be flattened instead of conical, and the broader and shorter
sepals are closely reflexed. Whether A. neglecta is really a hybrid, or
whether it is a species with a series of somewhat intermediate characters,
remains to be proved. Its relative abundance over a fairly extensive area,
as well as the apparent constancy of its taxonomic characters, suggests
that it is a species of equal rank with A. laevis, A. fernaldiu, A. interior,
and A. bartramiana.

28 ILLINOIS BIOLOGICAL MONOGRAPHS

4. AMELANCHIER INTERIOR Nielsen

(Plates I and VIT)
Amelanchier laevis sensu Rosendahl & Butters, Tr. & Shr. Minnesota 217 (1928),

ex p. Non Wiegand, 1912.

Amelanchier interior Nielsen in Am. Midl. Nat. 22: 185, pl. 13 (1939).
Amelanchier intermedia sensu Nielsen, op. cit. 184, pl. 12, ex p. Non Spach, 1834.
Amelanchier wiegandii Nielsen, op. Ge 180, pl. 10.

Straggling shrubs or small trees up to 8 m. tall; winter buds reddish
brown, narrowly ovoid, acute or acuminate, sometimes curved, the lateral
6-9 mm. long, the terminal 8-13 mm. long; bud-scales ciliate; leaves ovate
or broadly oval, or elliptical, conduplicate in the bud, unfolded but scarcely
fully grown at flowering time, green and sparsely pubescent beneath when
young, soon glabrous; mature blades 3-7 cm. long, 2-5 cm. wide, the apex
acute or shortly acuminate, the base subcordate or rounded, usually quite
glabrous on both surfaces when fully mature; lateral veins 8-11 pairs, not
prominent, regularly arranged, usually curved upward and becoming in-
distinct before reaching the margin; margins finely and evenly serrate or
serrulate nearly or quite to the base; teeth 5-6 per cm., 20-30 on each side
of average leaves; stipules linear, pubescent, deciduous; petioles slender,
1.5-3 cm. long, Paneee at maturity; flowers in loose, nodding, glabrous,
7-12-flowered racemes 4-7 cm. long, the lower pedicels 12-25 mm. long;
petals 5, white, oblanceolate or narrowly obovate, obtuse, 8-13 mm. long,
4-5 mm. wide; stamens about 20, the filaments glabrous; anthers 0.6-0.8
mm. long; hypanthium broadly cup-shaped, 3-5 mm. in diameter, glabrous
outside, slightly constricted on the young fruit; sepals triangular-lanceo-
late, 3-3.5 mm. long, acuminate, pubescent within, usually recurved from
the middle after anthesis; styles 5, glabrous, 3-4 mm. long, usually united
to the middle; top of the ovary densely white-tomentose; mature fruit
globose, purplish black, glaucous, glabrous, 6-8 mm. in diameter, sweet,
juicy, edible; lower pedicels 2-4 cm. long; seeds brown, smooth, obliquely
lanceoloid, somewhat flattened, about cf mm. long, 2-3 mm. wide.

Type Locatity: “East River road at junction with Seymour Avenue,
S. E. Minneapolis,’ Hennepin County, Minnesota. Collected in 1935 by
Etlar L. Nielsen. Type in the Herbarium of the University of Minnesota.

RANGE: Dry open woods, sandy wooded slopes, or on wooded bluffs
along rivers, Minnesota, Wisconsin, and northern Michigan. Flowering
in May and June; fruits maturing in July and August.

Minnesota: Caribou Lake, Nielsen 1379 (Minn.); Tofte, Breckenridge &
Nielsen 3161 (Minn.) ; Clarks Bay, Grand Portage, Breckenridge, Nielsen, & Moore
3229 (TYPE of A. wiegandti, Minn.), Moore & Nielsen 3653 (Minn.); Grand
Rapids, Nielsen 1060, 1063, 1064, 1066, 1069 (Minn.); Cohasset, Nielsen 1055
(Minn.) ; Two Harbors, Nielsen 1376 (Minn.); Finland, Moyle & Nielsen 1944
(Minn.); Baptism River, Nielsen 3170 (Minn.); Itasca Park, Nielsen 1931, 2485,
2521, 2547, 3113, 3124,-3127, 3129, 3131, 3133, 3136; 3137, 3138; Grant 2862. (GE);
Detroit Lakes, Zech 152, 161, 168, 195 (Minn.); Carlton, A. & E. Mattioli 7, 13,
20 (Minn.); Pelican Rapids, Butters 1342 (Minn.); Perham, July 13, 1926, Rosen-

AMERICAN SPECIES OF AMELANCHIER—JONES Zo

Map 3.—Range of Amelanchier interior.

dahl (GH); St. Cloud, Nielsen 2669 (Minn.); Center City, July 1892, Taylor
(GH) ; Coon Lake, Nielsen 1091, 1301 (Minn.) ; Ham Lake, Nielsen 1082 (Minn.) ;
St. Paul, Rosendahl & Butters 2578 (GH); Minneapolis, Nielsen 2961 (TYPE,
Minn.) ; Mendota, Rosendahl 5198 (Minn.) ; Hastings, Nielsen 1820 (Minn.) ; Red
Wing, Nielsen 1804 (Minn.) ; Spring Grove, Rosendahl 439, 4935 (GH), 4937 (AA),
Rosendahl & Butters 3892 (GH).

WISscoNSIN: Delton, Fassett 2825 (GH); Milford, Fassett 7128 (GH) ; Devils
Lake, Fassett 3051 (GH); Drummond, Cheney 4447 (GH).

Micuican: Foley Creek, near St. Ignace, Mackinac Co., Benner 6708 (GH).

This shadbush has the general aspect of Amelanchier laevis Wieg.,
but differs from that species in its smaller stature, wool-top ovary, some-
what shorter fruiting pedicels, and in the smaller flowers in shorter
racemes. Until it was described as a distinct species by Nielsen in 1939 it
had been variously labeled A. laevis, A. sanguinea, or A. arborea, or as a
hybrid between A. bartramiana and A. sanguinea, or between A. laevis and
A. spicata.

The leaves at flowering time are usually unfolded but not fully grown,
and in the majority of specimens the lower surface is sparsely floccose-
pubescent, varying to nearly or quite glabrous. Only rarely do the leaves
at this time remain folded and retain sufficient pubescence to give them a
whitish or grayish appearance, but by the time the petals have dropped,
this tardily-retained pubescence has disappeared.

Amelanchier wiegandii Nielsen is regarded as a synonym of A. in-
terior. In the original publication it appears in the key next to aA.
interior, from which it is said to differ in the carinate blades and acute
sinuses of the leaf-teeth. These characters are, however, quite intangible.
In a tabular comparison it was contrasted with A. sanguinea, from which
it is widely separated.

30 ILLINOIS BIOLOGICAL MONOGRAPHS

5. AMELANCHIER LAEVIS Wieg.
(Plates I and VI)

Pyrus botryapium sensu Bigelow, FI. Bost. 120 (1814), (ed. 2) 196 (1824). Non
L.f. 1781, nec Amelanchier botryapium Borkh. 1803.

Amelanchier canadensis sensu Torrey & Gray, Fl. N. Am. 1:473 (1840); Watson
& Coulter in Gray, Man. (ed. 6) 166 (1889); Sargent, Silva N. Am. 4:127, pl.
194 (1892); Britton & Brown, Illustr. Fl. N. U.S. 2:237, fig. 1985 (1897);
Clark in Bull. Vermont Agr. Exp. Sta. 73:68 (1899) ; Britton, Man. 517 (1901),
(ed. 3) 517 (1907); Small, Fl. Se. U.S. 531 (1903); Sargent, Tr. N. Am. 360
(1905); Robinson & Fernald in Gray, New Man. Bot. (ed. 7) 459 (1908);
Britton & Shafer, N. Am. Trees 437 (1908); Jones & Rand in Bull. Vermont
Agr. Exp. Sta. 145:99 (1909); Silva Tarouca, Freiland-Laubh. 139, fig. 123
(1913) ; Coker & Totten, Tr. N. Car. 60 (1916); Farwell in Rep. Mich. Acad.
Sci. 17:173 (1916). Non Mespilus canadensis L. 1753.

Amelancher botryapium sensu Emerson, Tr. Massachusetts 443 (1846); Decaisne
in Nouv. Arch. Mus. Hist. Nat. Paris 10:135 (1874). Non Borkh. 1803.

Amelanchier canadensis var. botryapium Gray, Man. Bot. 130 (1848), (ed. 2) 126
(1856).

Amelanchier lancifolia Hort. ex Hand-list Trees & Shrubs Kew 1:217 (1894), pro
syn. A. canadensis sensu Torrey & Gray.

Amelanchier stricta Hort. ex ibid.

Amelanchier laevis Wiegand in Rhodora 14:154, pl. 96 (1912); Small & Carter, FI.
Lancaster Co., Pennsylvania 154 (1913); Rehder in Bailey, Stand. Cyclop.
Hort. 273 (1914) ; Deam, Tr. Indiana 177, pl. 76 (1921); Sargent, Man. Tr. N.
Am. (ed. 2) 395, fig. 351 (1922); Silva Tarouca, Freiland-Laubh. (ed. 2) 95,
fig. 85 (1922) ; Hoffmann in Proc. Boston Soc. Nat. Hist. 36:280 (1922) ; Pease
in ibid. 37:67 (1924) ; Bailey, Man. Cult. Pl. 379 (1924); House in N.Y. State
Mus. Bull. 254:411 (1924) ; Wiegand & Eames, Fl. Cayuga Basin 248 (1926) ;
Rehder, Man. Cult. Tr. & Shr. 390 (1927) ; Sudworth in U.S. Dept. Agric. Misc.
Circular 92:134 (1927) ; Rosendahl & Butters, Tr. & Shr. Minnesota 217 (1928),
ex p.; Peattie, Fl. Indiana Dunes 219 (1930); Rydberg, Fl. Prairies & Plains
437 (1932) ; Deam, Tr. Indiana (ed. 2) 189, pl. 77 (1932); Small, Man. Se. FI.
636 (1933); Marie-Victorin, Fl. Laurent. 317, fig. 91 (1935); Coker & Totten,
Tr. Se. States 208 (1936); Nielsen in Am. Midl. Nat. 22:188, pl. 14 (1939) ;
Deam, FI. Indiana 532 (1940); Rehder, Man. Cult. Tr. & Shr. (ed. 2) 389
(1940); G. N. Jones, FI. Illinois 154 (1945); Weatherby & Adams in Contr.
Gray Herb. 158:50 (1945).

Amelanchier laevis f£. nitida Wiegand in Rhodora 13:155 (1912).

Amelanchier laevis var. cordifolia Ashe in Journ. Elisha Mitchell Sci. Soc. 34:138
(1918).

Amelanchier laevis var. nitida Fernald in Rhodora 23:267 (1922).

A small tree 10-13 m. tall, or a shrub, with spreading branches; bark
reddish brown, longitudinally fissured in age; twigs slender, glabrous;
winter buds lanceoloid, the scales ciliate; leaves of firm texture, ovate or
oval, varying to slightly obovate, conduplicate in the bud, glabrous or
nearly so from the first, often bronzy, about half-grown at flowering time,
folded lengthwise and glaucous-purplish, soon entirely glabrous, and when
mature dark green and slightly glaucous; mature blades 4-6 cm. long, 2.5-4
cm. wide, short-acuminate or acute at the apex, rounded or subcordate at
the base; lateral veins 12-17 pairs with short intermediate ones, un-
equally distant, sinuous, slightly upcurving, anastomosing near the margin,
the uppermost widely spreading; margins finely and sharply serrate nearly

AMERICAN SPECIES OF AMELANCHIER—JONES 31

to the base, the teeth subulate, callus-tipped, 6-8 per cm., 35-45 on each
margin of average leaves; stipules and bracts purplish green, deciduous,
sericeous; petioles slender, 12-25 mm. long; flowers large and showy;
racemes many-flowered, flexuous, spreading or drooping, 4-12 cm. long,
glabrous or nearly so; pedicels glabrous, slender, the lower 1.5-3 cm.
long; petals white, oblanceolate or narrowly oval, obtuse, 12-22 mm. long,
3-4 mm. wide; stamens about 20, 3-4 mm. long, the filaments glabrous;
anthers 1 mm. long; hypanthium campanulate, 3-5 mm. broad, glabrous,
very slightly or not at all constricted below; sepals lanceolate, 3-4 mm.
long, acuminate, glabrous outside, tomentulose within, becoming reflexed
on the fruit, or sometimes merely divaricate; styles 5, glabrous, 3-4 mm.
long, free to below the middle; summit of the ovary glabrous; fruit
globose, purple or nearly black, 6-8 mm. in diameter, glaucous, sweet,
succulent, edible; lower pedicels 2-5 cm. long; seeds chestnut brown,
smooth, asymmetrically ovoid or semi-ellipsoid, somewhat compressed,
4-4.5 mm. long, 2-2.5 mm. wide when fully developed.

Type Locarity: Wellesley, Massachusetts. Collected by K. M. Wie-
gand, May and June, 1911. Type in the Gray Herbarium of Harvard
University.

Rance: In dry open woodlands, roadside thickets, cool ravines and
hillsides, or damp wooded slopes and banks, from Newfoundland to
Minnesota, southward to Missouri, Indiana, and Georgia; flowering in
the southern part of the range from the middle of April, in the north
to the middle of June; fruit ripe in June and July. Common names:
serviceberry, juneberry, maycherry.

Quesec: Bolton, July 25, 1926, Knowlton (GH); Longueuil, M.-Victorin 4319
(AA, GH), 11219, 9503 (AA); Caughnawaga, May 27, 1900, Jack (AA); Bic,
Fernald & Pease 25135 (GH), Louis-Marie, et al. 34433 (GH); La Trappe, Louis-
Marie 145 (GH); Berthier-en-Bas, Rousseau 24549 (GH) ; Saint-Raphael, Rousseau
24567, 24562, 24559 (GH); Kingsmere, Macoun & Malte 88016 (GH); Shawnigan
Falls, Chamberlain & Knowlton in 1923 (GH).

NEWFOUNDLAND: Donovans, Fernald & Wiegand 5755 (GH); St.-Pierre,
Arséne 310 (GH); near St. Johns, Bishop 373 (GH), Fernald & Wiegand 5567,
5568, 5569, 5570, 5580 (GH); Whitbourne, Fernald & Wiegand 5545, 5547, 5578a
(GH); Cape St. George, Mackenzie & Griscom 11102 (GH); French Island,
Fernald, Long, & Fogg 290 (GH); Lomond, Fernald, Long, & Fogg 1791 (GH);
Grand Falls, Fernald & Wiegand 5552 (GH); Rushy Pond, Fernald & Wiegand
5542, 5599 (GH).

New Brunswick: Shediac Cape, July 25, 1914, Hubbard (GH); Kent, May
30, 1870, Fowler (GH).

_ Nova Scotra: Boylston, Hamilton in 1890 (GH); Newport, Dill in 1894
(AA); Truro, Jack 628 (AA); Middleton, Long 21447 (GH), 21448 (GH, AA);
Cape Breton Island, Macoun 19043 (GH), Nichols 557, 168 (GH); Digby, Howe
& Lang 265, 297 (GH); Marshalltown, Jack 3187, 3188 (AA); Meteghan, Fernald
& Long 21451 (GH); Weymouth, Fernald, et al. 21441 (GH), 21442 (GH, AA);
Armdale, Fernald, Bartram, & Long 23943, 23944, 24761 (GH); Halifax, Jack
680, 3648, 3671 (AA).

32 ILLINOIS BIOLOGICAL MONOGRAPHS

Map 4.—Range of Amelanchier laevis.

PrINcE Epwarp Istanp: Bothwell, Fernald & St. John 11082 (GH).

Marne: Boothbay, Fassett 446 (NE), Grover & Smith in 1922 (UI); Kenne-
bunkport, Koehler 2 (GH); North Berwick, May 1894, Parlin (GH); Vassalboro,
Chamberlain 34 (NE); South Poland, Furbish in 1895 (NE); Mt. Livermore,
Furbish in 1896 (NE); Monticello, Fernald & Long 13763 (AA, NE); Brunswick,
Furbish in 1892 (NE), May 19, 1897, Chamberlain (NE); Cumberland, Chamber-
lain 533 (NE); West Baldwin, Furbish in 1900 (NE).

New Hampsurre: Mason, May 15, 1915, Batchelder (NE); Lebanon, May 3,
1889, Kennedy (GH, NE); Peterboro, Batchelder 3 (NE); Mt. Agassiz, Collins
in 1871 (NE); Whitefield, July 4, 1896, Deane (NE); Stark, Pease 17476 (NE);
Franconia, Faxon in 1892 (GH, NE); Hooksett, July 11, 1926, Batchelder (NE) ;
Dover, Hodgdon 195, 2244, 2245 (NE); Barrington, Hodgdon & Dunn 2772 (NE);
Gorham, Pease 16010 (NE); Jaffrey, July 19, 1891, Deane (GH, NE).

VERMONT: Pownal, Floyd 845 (NE); Sunderland, May 18, 1935, Pease (NE);
Peacham, May 10, 1889, Blanchard (NE); Burlington, May 24, 1914, Knowlton
(NE); Brunswick Springs, Sanford 1066 (NE); Island Pond, Sanford 1193
(NE) ; Johnson, June 3, 1895, Grout (NE); Barton, May 24, 1923, Knowlton (NE) ;
Willoughby, May 28, 1904, Kennedy (NE); Roxbury, Winslow, July 18, 1916
(NE); Jamaica, Moldenke 9502 (UI); Stratton, June 25, 1914, Wheeler (NE);
Hartford, June 12, 1920, Eaton & St. John (NE).

MassaAcHuseETTs: Barnstable, Child, Knowlton, Bird, & Bean 16377 (NE);
Oak Bluffs, Seymour 1224 (GH, NE) ; Chilmark, Seymour 1223 (GH, NE); Mag-

AMERICAN SPECIES OF AMELANCHIER—JONES 33

nolia, Weatherby & Perry, May 12, 1936, Pl. Exsicc. Gray. 663, 664 (GH, UI);
Wellesley, Wiegand 2136 (GH, TYPE of A. laevis); Coleraine, May 11, 1912,
Batchelder, Kennedy, & Williams (NE); Whately, Harger & Fernald, May 17, 1913
(NE); Granville, Knowlton & Hunnewell in 1913 (NE); Amherst, Seymour 3501
(NE); Ayer, Ordway & Bullard, May 30, 1934 (NE); Nantucket I., Bicknell
4835 (NE); Plymouth, Fernald & Hunnewell 15192 (NE); Dorchester, May 5,
1889, Churchill (NE); Florida, Fernald & Long 9623 (NE, GH); Lanesboro, May
22, 1916, Churchill (NE); Becket, G. N. & F. F. Jones 13716, 13750, 13781, 15323
CW):

ConneEcTicuT: Oxford, Harger 3 (GH); Stratford, Eames 1 (GH); South-
ington, Bissell in 1901 (GH); Groton, Graves in 1901 (GH); Winchester, Harger
10 (GH); Waterbury, Blewitt 1512 (NE); Glastonbury, Weatherby 2861 (NE);
Oly Lyme, Woodward in 1918 (NE); Andover, Weatherby 5280 (NE); Middle-
town, Blewitt 1798 (NE); Hamden, Blewitt 1797 (NE).

Ruove Istanp: Westerly, Aug. 21, 1913, Bissell, Harger, & Weatherby (NE) ;
Little Compton, July 27, 1919, Collins (NE); Barrington, May 30, 1911, Winslow
(NE); Cumberland, Hunnewell 4129 (NE); Providence, Collins in 1902 (GH);
Hopkinton, Fernald, Woodward, & Collins in 1919 (NE); Smithfield, Chamberlain
52 (NE, AA); Prudence Island, Sanford 10215 (NE); Lincoln, Fernald 9624
(NE).

New York: Hudson Falls, April 27, 1915, Burnham (GH); French Mt.,
Burnham 12 (GH); Judd’s Falls, Wiegand 2572 (AA, GH); Newcomb, House
7961 (GH); North Harpersfield, Topping 223 (UI); Stockholm, Phelps 1591
(GH); Canton, Phelps 567 (GH).

New Jersey: Palmyra, Long 14522, 16257 (Ph); Georgia, Long 52074 (Ph);
Sharptown, Long 18378 (Ph); Green Creek, Stone 11932 (Ph).

PENNSYLVANIA: Saylorsburg, Long 6633 (GH); Lancaster Co., April 25, 1891,
Heller (GH, Ph); Pocono Plateau, Harshberger in 1904 (GH); Reitz Gap, Wahl 29
(GH); Trout Run, Wahl 298 (GH); Slateford, Long 50001 (Ph); Bake Oven
Knob, Pretz 2363a, 3246 (Ph) ; Trexlertown, Pretz 5919 (Ph) ; Alburtis, Pretz 9090
(Ph); Fleetwood, Long 12515 (Ph); Lenhartsville, Long 12867 (Ph); Huffs
Church, Wilkens 6666 (Ph); Schubert, Wilkens 5129 (Ph); Nockamixon Narrows,
Benner 2393 (Ph); Smithville, Tanger 3036 (Ph); Brickerville, Tanger 3044, 3045
(Ph); Gap, Long 30691 (Ph); Hopeland, Long 41781 (Ph); Unionville, Pennell
82 (Ph); Nottingham Barrens, Pennell & Long 7559 (Ph); Landenburg, Long
8476 (Ph); St. Peters, Long 33631 (Ph); Exton, Long 32022 (Ph); Elam, Long
32349 (Ph).

DELAWARE: Vandyke, Long 48444 (GH, Ph); Coochs Bridge, Benner 9567
(Ph); Centreville, June 13, 1898, Commons (Ph); Newark, May 11, 1922,
Meredith (Ph).

Maryann: Baltimore, May 15, 1910, Churchill (GH); Golt, Montgomery Co.,
Hunnewell 5885 (GH); Elkton, Randolph 121 (GH); Elk Neck, Pennell 24805
(Ph); North East, Long 54418, 57028 (Ph).

West VirGINIA: Canaan Valley, July 21, —, Burke (AA).

District oF CoLumBIA: Brightwood, Smiley in 1881 (GH).

VIRGINIA: Hopewell Gap, Allard 4353 (GH); White Top Mt., Britton & Vail
in 1892 (AA), June 28, 1892, Small (GH); Brushy Mt., June 17, 1892, Small (GH).

NortH CaroLina: Highlands, Harbison 2 (AA); Macon Co., Harbison 510,
913 (AA).

GrorciA: Rabun Co., Duncan 3282 (GH); Trenton, Hermann 10191 (GH).

Ontario: Cornwall, May 28, 1913, May 29, 1914, Jack (AA).

Minnesota: Mabel, Spondee in 1928 (AA); Duluth, July 1, 1914, Butters
(GH) ; near Minneapolis, Nielsen 1868 (Minn.) ; Ham Lake, Nielsen 1872 (Minn.) ;
Lake City, Rosendahl G Nielsen 1852 (Minn.).

Wisconsin: Black Falls, July 6, 1914, Lake (AA); Mud Bay, Pease 18044
(GH); Devils Lake, May 25, 1899, Cheney (GH); Trout Lake, Fassett 13745
(GH); Kilburn, June 1, 1893, Pammel (AA); Brown Co., Schuette in 1904 (AA);
Barneveld, Fassett 2826 (GH).

34 ILLINOIS BIOLOGICAL MONOGRAPHS

Micuican: Wolverine, Gleason 708 (AA); Grayling, Piper in 1922 (AA);
Port Huron, May 17, 1912, Dodge (AA); Mackinac I., July 4, 1912, Hunnewell
(GH); Sault Ste. Marie, Newins 8141 (AA); Ann Arbor, Hermann 6471 (GH);
Homestead, Hermann 7259 (GH); Gwinn, June 6, 1909, Harrison (GH) ; Lansing,
Bailey in 1887 (GH); Keweenaw Co., Farwell in 1889 (GH).

Onto: Portage Co., Webb 1180 (GH); Berea, May 1897, Watson (UI).

Kentucky: Lynch, McFarland 18 (GH).

InpIANA: Miller, Chase 197 (UI); Martin Co., Deam 12868 (GH) ; Versailles,
Deam 16116 (AA); Albion, Deam 33798 (AA).

Ittinors: Lake Forest, Harper in 1890 (AA); Oregon, May 16, 1883, Waite
(UI); Wabash Co., Schneck in 1883 (UI); Lake Zurich, Hill 321889 (UI); Bar-
rington, Chase 1047 (UI).

Iowa: Fayette, Fink in 1894 (GH); Decorah, Aug. 5, 1933, Tolstead (Minn.).

Amelanchier laevis is the most noticeably ornamental and graceful of
the eastern American serviceberries, and it is rather surprising that
such a common and conspicuous species should not have received a spe-
cific name until so treated by Wiegand in 1912. It is most closely related
to A. arborea. Besides its glabrous or nearly glabrous leaves, that are
often more or less purplish tinged when young, it differs from A. arborea
in the more spreading branches, and in the slender, flexuous, pendulous,
looser inflorescence and somewhat larger flowers. The mature leaves are
usually more abruptly pointed than those of A. arborea, rounded (scarcely
cordate) at the base, and the blades are green above, paler beneath, and
completely glabrous almost from the first. In general, it may be noted
that it has somewhat longer petals, narrower sepals, and the fruits are
longer-pedicelled, more succulent, and of a sweeter flavor. The seeds of
A. laevis are very similar to those of A. arborea, but in a large series
show a tendency to be very slightly smaller, lighter brown, and somewhat
smoother. This trend is, however, not sufficiently tangible for descriptive
purposes.

6. AMELANCHIER ARBOREA (Michx.f.) Fern.

(Plates I and VIII)

Mespilus nivea Marshall, Arbustr. Am. 90 (1785), nomen dubium.

Pyrus botryapium sensu Wangenheim, Nordam. Holzarten 90, pl. 28 (1787) ; Tausch
in Flora 21:714 (1838). Non L.f. 1781.

Amelanchier canadensis sensu Walter, Fl. Carol. 148 (1788); Darlington, Fl. Cest-
rica (ed. 3) 86 (1853); Sargent, Silva N: Am. 4:127, pl. 194 (1892), ex ‘p.;
Dippel, Handb. Laubh. 3:392 (1893); Britton & Brown, Illustr. Fl. N. U.S.
2:237, fig. 1985 (1897) ; Mohr in Contr. U.S. Nat. Herb. 6:545 (1901) ; Macken-
zie & Bush, Man. Fl. Jackson Co., Missouri 108 (1902); Keeler, Our Native
Tr. 153 (1902); Small, Fl. Se. U.S. 531° (1903).;° Card. in. Bailey; ‘Cyclops-Anr
Hort. 57 (1904) ; Dame & Brooks, Handb. Tr. New Engl. 116, pl..59 (1904) ;
Sargent, Man. Tr. N. Am. 360, fig. 283 (1905); Blanchard in Torreya 7:97
(1907) ; Hough, Handb. Tr. N. States & Canada 243, figs. 282, 283, 284 (1907) ;
Emerson & Weed, Our Tr. 191 (1908) ; Britton & Shafer, N. Am. Tr. 437, fig.
383 (1908) ; Rogers, Tree Book, pl. opp. p. 298 (1908); Apgar, Ornam. Shr.
U.S. 182, fig. 277 (1910); Blakeslee & Jarvis, in Bull. Storrs Agr. Exp. Sta.
69:492-493 (1911); Stone, Pl. So. N.J. in Rep. N.J. State Mus. 1910:488 (1911) ;

AMERICAN SPECIES OF AMELANCHIER—JONES 30

Clements, Rosendahl, & Butters, Minn. Tr. & Shr. 151 (1912); Wiegand in
Rhodora 14:150, pl. 96, fig. 6 (1912); Silva Tarouca, Freiland-Laubh. 139,
fig. 123 (1913); Britton and Brown, Illustr. Fl. N. U.S. (ed. 2) 2:292, fig. 2329
(1913); Small, Florida Tr. 30 (1913); Small, Shr. Florida 29 (1913);
Rehder in Bailey, Stand. Cyclop. Hort. 273 (1914); Bean, Tr. & Shr. Brit. Isles
1:188 (1914) ; Burns & Otis in Bull. Vermont Agr. Exp. Sta. 194:145 (1916) ;
Rydberg, Fl. Rocky Mts. 446 (1917) ; Hitchcock & Standley, Fl. Dist. Columbia
178 (1919) ; Sargent, Man. Tr. N. Am. (ed. 2) 394, fig. 350 (1922) ; Hoffmann
in Proc. Boston Soc. Nat. Hist. 36:280 (1922); House, N.Y. State Mus. Bull.
254:411 (1924); Bailey, Man. Cult. Pl. 379 (1924); Wiegand & Eames, FI.
Cayuga Basin 248 (1926); Rehder, Man. Cult. Tr. & Shr. 390 (1927); Sud-
worth in U.S. Dept. Agric. Misc. Circular 92:134 (1927) ; Pepoon, Fl. Chicago
Area 342 (1927); Rosendahl & Butters, Tr. & Shr. Minnesota 218 (1928) ;
Miller & Tehon, Native & Naturalized Tr. Illinois 203, pl. 66 (1929); Peattie,
Fl. Indiana Dunes 219 (1930); Rydberg, Fl. Prairies & Plains 436 (1932) ;
Deam, Tr. Indiana (ed. 2) 189, pl. 76 (1932); Small, Man. Se. Fl. 636 (1933) ;
Palmer & Steyermark in Ann. Missouri Bot. Gard. 22:557 (1935); Marie-
Victorin, Fl. Laurent. 316, fig. 91 (1935); Griscom in Rhodora 38:48 (1936) ;
Coker & Totten, Tr. Se. States (ed. 2) 207 (1937) ; Munns in U.S. Dept. Agric.
Misc. Publ. 287:137, pl. 133 (1938); Nielsen in Am. Midl. Nat. 22:183, pl. 11
(1939) ; Steyermark, Spring Fl. Missouri 255, pl. 68, fig. 1 (1940); Rehder,
Man. Cult. Tr. & Shr. (ed. 2) 388 (1940) ; Deam, FI. Indiana 532 (1940). Non
Mespilus canadensis L. 1753.

Mespilus amelanchier B nivea Castiglioni, Viagg. St. Uniti 2:293 (1790).

Mespilus canadensis var. B cordata Michx. Fl. Bor. Am. 1:291 (1803).

Mespilus arborea Michx.f. Hist. Arb. Am. Sept. 3:68, pl. 11 (1810), N. Am. Sylva
2:70, pl. 66 (1818), (ed. 3) 2:60, pl. 66 (1859).

Aronia arborea Barton, Comp. FI. Philadelphia 1:228 (1818).

Aroma botryapium sensu Elliott, Sketch Bot. S. Car. & Ga. 1:557 (1821). Non Pers.
1807.

Amelanchter ovalis var. B subcordata DC. Prodr. 2:632 (1825).

Aronia subcordata Rafinesque ex DC., ibid.

Malus microcarpa Rafinesque ex DC., ibid.

Aronia cordata Rafinesque, Med. Fl. 2:196 (1830) nom. nud.

Amelanchier botryapium sensu Spach, Hist. Nat. Veg. Phan. 2:85 (1834); Torrey,
Fl. N. Mid. States 203 (1826) ; Darlington, Fl. Cestrica 294 (1837); Farwell in
Rep. Mich. Acad. Sci. 17:175 (1916). Non Borkh. 1803.

Pyrus wangenheimiana Tausch in Flora 21:715 (1838).

Aronia nivea Neumann ex Tausch, ibid.

Amelanchier canadensis var. @ botryapium Torrey & Gray, Fl. N. Am. 1:473 (1840) ;
Torrey, Fl. N.Y. 1:225 (1843); Walpers, Rep. Bot. Syst. 2:55 (1843); Gray,
Man. Bot. 130 (1848); Brendel, Fl. Peoriana 47 (1887); Chapman, FI. S. U.S.
(ed. 3) 141 (1897) ; Schneider, Illustr. Handb. Laubh. 1:734, fig. 409 (1906) ;
Robinson & Fernald in Gray, New Man. Bot. (ed. 7) 460 (1908).

Amelanchier wangenheimiana M. Roem. Syn. Mon. 3:146 (1847).

Amelanchier canadensis var. tomentula Sargent, Man. Tr. N. Am. 361 (1905);
Schneider, Illustr. Handb. Laubh. 1:734 (1906); Robinson & Fernald in
Rhodora 11:47 (1909).

Amelanchier intermedia sensu Blanchard in Torreya 7:98 (1907), ex p. Non
Spach 1834.

Amelanchus canadensis Vollmann, Fl. Bayern 453 (1914).

Amelanchier canadensis {. nuda Palmer & Steyermark in Ann. Missouri Bot. Gard.
25:772 (1938); Rehder, Man. Cult. Tr. & Shr. (ed. 2) 388 (1940).

Amelanchter arborea Fernald in Rhodora 43:563, pl. 672, fig. 2 (1941); G. N.
Jones, FI. Illinois 154 (1945).

Amelanchier austromontana sensu Fernald in op. cit. 566. Non Ashe 1918.

36 ILLINOIS BIOLOGICAL MONOGRAPHS

A small tree 5-20 m. tall, with a maximum trunk diameter of 40 cm.,
or an irregularly branched shrub, the stems solitary or few together, not
growing in clumps; branches ascending; bark grayish brown, longitudi-
nally fissured on old stems, that of the young stems gray, smooth, and
often somewhat striped; twigs of the season pubescent at first, soon
glabrous, brown and slightly glossy; winter-buds lanceoloid, acuminate,
glabrous; leaves conduplicate in the bud, small, densely white-tomentose,
mostly folded at flowering time; blades 4-10 cm. long, 2-2.5 cm. wide,
ovate or oval, or slightly obovate, short-acuminate or acute at the apex,
cordate or rounded at the base; mature leaves thick, firm, dark green and
glabrous above, paler beneath and pilose at least along the midvein, be-
coming nearly or quite glabrous in age; lateral veins 11-17 pairs, un-
equally distant, sinuous, anastomosing and indistinct toward the margin;
margins sharply and finely and often somewhat doubly serrate almost or
quite to the base, the teeth ascending, incurved, slender, 6-10 per cm.,
mostly 50-60 on each side of average leaves; stipules linear, caducous,
pilose; petioles 1-2 cm. long; flowers 2-2.5 cm. in diameter, fragrant,
appearing early, usually before the leaves have unfolded; racemes spread-
ing or somewhat pendulous, 4-10 cm. long, 4-10-flowered; pedicels grayish
pubescent, the lower 8-17 mm. long; petals oblanceolate, 12-18 mm. long,
2-5 mm. wide, white, or sometimes pinkish; stamens about 20, 3-4 mm.
long, the filaments glabrous; anthers 0.6-0.8 mm. long; hypanthium cam-
panulate, 2.5-3 mm. in diameter, glabrous or somewhat floccose; sepals
triangular-lanceolate, acute, 2-3 mm. long, tomentulose on both sides,
becoming strongly reflexed from the base after anthesis; styles 5 (or 4),
about 4 mm. long, glabrous, the free portion 2-3 mm. long; summit of
ovary glabrous; fruit globose, purple (rarely whitish), 6-10 mm. in
diameter, scanty, somewhat tasteless, falling early; lower pedicels 1-2.5
cm. long; seeds 3-10, dark brown, nearly smooth, glabrous, 4.5-5 mm.
long, 2-3 mm. wide when fully developed, asymmetrically lanceoloid,
somewhat flattened.

Type Locariry: “Hab. ...a Canada ad Virginiam et in montibus
Carolinae.” Phototypes in the Gray Herbarium and the Arnold Arboretum.

RANGE: In dry woods and open ground, and on wooded hillsides, from
Maine to Minnesota, southward to Louisiana and Florida. Flowering from
the end of March to the end of May; fruits ripening in June and July.

QuEBEc: Longueuil, M.-Victorin 11222 (AA); Aylmer, Rolland 57, 58, 59
(GH); Kingsmere, Macoun & Malte in 1913 (GH); Ile Bigras, M.-Victorin 24547
(GH); Oka, M.-Victorin 24546 (GH); Grosse-isle, M.-Victorin 15586 (AA);
Montmorency Falls, Aug. 20, 1895, Jack (AA).

Marne: Monhegan Island, Jenney, Churchill, & Hill 3264 (NE); Mt. Desert
I., Rand, July 1899 (NE); Lincolnville, Rossbach 1102 (NE); Northport, Rossbach
1136 (NE); Orono, Harvey in 1898 (UI); Milford, Fernald 13767 (GH); West
Pembroke, Fernald 1885 (GH).

AMERICAN SPECIES OF AMELANCHIER—JONES 37

ae m

SDS een

Map 5.—Range of Amelanchier arborea.

New York: Pavilion, Hill in 1859 (UI); New York, Burnham 1162 (GH);
New London, House 11870 (GH); Sylvan Beach, House, May 16, 1918 (GH);
Albany, House 7787 (GH); Middlefield, Hunnewell 6889 (GH); Spencer, Eames
& Dean 4293 (GH); Canton, Phelps 1581, 1582 (GH); Black Rock Forest, Raup
7410, 7727 (GH).

VERMONT: Plainfield, Eggleston 1959 (GH); Twin Mts., Eggleston 1182, 1957
(GH) ; Pittsford, Eggleston 1183 (GH) ; Townshend, L. A. Wheeler in 1915 (GH) ;
Charlotte, May 22, 1922, Knowlton (NE); Newport, May 24, 1923, Knowlton
(NE); W. Fairlee, June 10, 1933, Anderson, et al. (NE).

New HampsuirE: Walpole, May 26, 1917, Bean & Fernald (NE); Lebanon,
Fernald, Hunnewell, & Blanchard in 1920 (NE); Wolfboro, H. E. Sargent 24
(GED):

MASSACHUSETTS: Canton, May 5, 1895 Williams (NE); Plainfield, June 14,
1913, Forbes (NE); Brimfield, May 20, 1916, Knowlton (NE); Montague, May 13,
1911, Fernald (GH, NE); Erving, Hunnewell, Macbride, & Torrey, May 14, 1915
(NE); Whately, May 17, 1913, Harger G& Fernald (NE); Shelburne Falls, May
11, 1912, Bean & Knowlton (NE); West Orange,. May 10, 1912, Hunnewell &
Wiegand (NE); Shutesbury, Tower & Seymour 3674 (NE); Northfield, May 11,
1912, Fernald & Floyd (NE); Rowe, Fernald & Long 9621 (GH); Andover,
Pease 678 (NE).

Connecticut: Waterbury, Blewitt 2038 (NE); Glastonbury, Wright in 1916
(GH); Middlebury, Apr. 23, 1896, Shepardson (NE); Danbury, Blewitt 2039
(NE); Tariffville, May 17, 1913, Winslow G Hill (NE); Southington, Blewitt
1802 (NE), Bissell in 1901 (GH); East Hartford, May 18, 1913, Briggs (NE);
Kent, Eames 8287 (NE); Barkhamsted, Blewitt 1805, 3650 (NE); Watertown,
Blewitt 2036, 3501 (NE); Winstead, Blewitt 1804 (NE); Thomaston, Weatherby
2157 CNE):

RuHopE Istanp: Portsmouth, June 10, 1911, Collins (NE); Cumberland, Humnne-
well 4128 (NE); Lincoln, May 30, 1919, Collins (NE).

38 ILLINOIS BIOLOGICAL MONOGRAPHS

New JersEY: Woodglen, Long 52340 (Ph); Penville, Long 52160 (Ph);
Charlestown, Long 46255 (Ph); Little York, Long 46808 (Ph); Tumble Falls,
Hermann 4016 (Ph) ; Drea Hook, Long 53736 (Ph) ; Bordentown, Long 3107 (Ph) ;
Smithville, Long 9610 (Ph); Cranbury, Long 51868 (Ph); Deerfield, Long 32089
(Ph); Pennington, Long 50133 (Ph); Palmyra, Long 11943, 13516, 14514, 14519
(Ph); Somerdale, Long 11605 (Ph).

PENNSYLVANIA: Safe Harbor, Apr. 18, 1889, Heller (GH); Lenhartsville,
Wilkens 1097 (GH); Ohio Pyle, May 21, 1910, Bartram (GH); Pleasant Valley,
June 9, 1923, Benner (GH); Wagontown, Apr. 25, 1925, Stone (GH); Reseca
Falls, June 9, 1918, Bartram (GH); Perkiomen Creek, Long & St. John 2479
(GH); Fairview, Wahl 34 (GH); Mercersburg, Adams & Wherry 4662 (GH);
Linfield, Long 11883 (GH); Rosscommon Creek, Long 6586 (GH); Bernville,
Stoudt & Hermann 2779, 2784 (GH).

DELAWARE: Deakyneville, Long 58228 (Ph); Mount Cuba, Long 32282 (Ph).

WEstT VirGINIA: Panther Mt., Rydberg 9052 (AA); White Sulphur Springs,
Hunnewell in 1914 (GH), Harbison 7095 (AA); Aaron’s Run, Monongalia Co.,
Myers 415 (UI); Ravenswood, May 14, 1939, Balsar (GH).

VIRGINIA: Williamsburg, Grimes 2560 (GH); Massanutten Mts., Allard 4498
(GH); Rye Valley, June 11, 1892, Small (GH); Holston River, Aug. 8, 1892, Small
(AA); Hot Springs, Hunnewell 4029 (GH) ; Wytheville, Svenson 7779 (GH) ; near
Wylliesburg, Palmer 39994 (AA); near Antioch, Allard 2819 (GH); near Aldie,
Allard 2516, 2584 (GH); Beverly, Allard 220 (GH); High Point, Allard 4342
(GH) ; Little Neck, Fernald & Long 3959 (GH); Great Neck, Fernald, Griscom,
& Long 4650 (GH); Pungo, Fernald & Griscom 4425 (GH) ; Indian Point, Fernald
& Long 11698, 11845 (GH); Clarendon, Blake 10557 (GH); Emporia, Fernald &
Long 8291 (GH, Ph); Cleopus, Fernald & Long 13040 (GH, Ph); Surry Court-
house, Fernald & Long 13039 (GH, Ph); Richmond, Fernald & Long 7063 (GH,
Ph) ; Gary Church, Fernald & Long 7064 (GH, Ph); Homeville, Fernald & Long
7065 (GH, Ph) ; Waverly, Fernald & Long 7067 (GH, Ph); South Hill, Fernald &
Long 7069 (GH, Ph); Burt, Fernald & Long 7450 (GH, Ph); Ingersoll, Fernald
& Long 11844 (GH, Ph); Scotland, Fernald & Long 13041 (GH, Ph).

MaryLanp: May 16, 1905, Hitchcock (UI); Oakwood, Tanger 3025 (Ph);
Golt, Tidestrom 11882 (GH) ; Bald Friar, St. John & Long 1009 (GH), 8059 (Ph) ;
Middle Neck, Long 37298 (GH, Ph); Elk Neck, Long 37719 (Ph); Providence
Mill, Benner 4883 (Ph).

NortH Carorina: Linville, Randolph 1200 (GH); Salisbury, Harbison 6
(AA); Raleigh, Harbison 30 (AA); Chapel Hill, Harbison 15 (AA); Ridge Crest,
Davis 1462 (UI).

SouTtH CAROLINA: Santee Canal, Ravenel, sn. (GH); Anderson, Palmer 42440
(AA).

Georcia: Holton, Harper 1806 (GH, AA); N. Georgia, Wright in 1875 (GH);
Wayne Co., Eyles 6894 (GH); Lookout Mt., July 1898, Ruth (UI).

FLoripA: River Junction, Harbison 1415 (AA); Round Lake, Harbison 4
(AA); Caryville, Mar. 30, 1927, Hume (AA); Lake Tamonia, Griscom 21578
(GH); Aspalaga Bluff, Apr. 26, 1924, Small (GH).

Ontario: Niagara, John Macoun 34296 (GH); Killaloe, May 1903, Jack (A);
Stokes Bay, Krotkov in 1934 (GH); Belleville, May 16, 1878, Macoun (GH).

WIsconsIn: Jacksonport, May 30, 1926, Kraus, et al. (GH).

Minnesota: Grand Portage, Nielsen 1630 (Minn.).

Micuican: Bridgeton, Hermann 9625 (AA).

Onto: Ironton, Werner 55 (GH); Toledo, Sanford 379 (GH); Garrettsville,
Webb 84 (GH); Sugar Grove, Horsey 323 (AA); Urbana, Apr. 22, 1838, Samples
(UI) ; Berea Co., Watson in 1897 (UI); Otway, Demaree 11260 (AA).

InDIANA: Near Attica, Deam 22524 (AA); Liverpool, Chase 2035 (UI); Dune
Park, Chase 2053 (UI); Fountain, Deam 23005 (UI); Clarke, Moffatt 190 (UI);

AMERICAN SPECIES OF AMELANCHIER—JONES 39

Bloomington, Friesner 9541 (UI); Pine, May 13, 1876, Hill (UI); Portland Arch,
G. N. Jones 13143 (UI).

Irt1no1is: Camp Grant, Mattoon 13 (UI); Marshall Co., Chase 1795 (UI);
Will Co., Hill 411912 (UI); Starved Rock, Greenman, Lansing, & Dixon 81 (GH),
Thone 172 (UI); Peoria, McDonald in 1904 (GH, UI); Adams Co., Evers 52
(UI); Urbana, Aug. 11, 1899, Clinton (GH); Mahomet, Sept. 23, 1899, Gleason
(GH) ; Homer, Sept. 16, 1899, Clinton (UI); Taylorville, Apr. 22, 1899, Andrews
(UI); Carlinville, Apr. 12, 1890, Andrews (UI); Richland Co., Ridgway 2534
(AA); Grand Tower, May 3, 1902, Gleason 2404 (GH, UI), 1695 (GH); Wabash
Co., Sept. 1886, Schneck (U1); Johnson Co., May 26, 1902, Schneck (UI); Ver-
milion River, G. N. Jones 13340 (UI); Wolf Lake, G. N. Jones, 12089 (UI);
Franklin Creek, G. N. Jones 15840 (UI) ; Sangamon River, G. N. Jones 15618 (UI) ;
Dixon Springs, G. N. Jones 11992 (UI).

Missourt: Painton, Palmer 34917 (AA); Monticello, Palmer 35890 (AA);
La Grange, Davis 2227 (AA) ; Kahoka, Palmer 25860 (AA); Livonia, Palmer 41069
(AA); Thayer, Palmer 14684 (AA); Whiteside, Davis 49 (UI); Allenton, May 20,
1882, Letterman (AA); St. Charles Co., Drouet 1421 (AA); Jefferson Co., Letter-
man 2,3 (AA); Webb City, Palmer 1602 (AA); Newton Co., Palmer 29927 (AA),
Bush 85, 3509 (GH); McDonald Co., Bush 30, 85a, 85b, 85¢ (GH); Wright
City, Davis 1259 (AA); Oronogo, Bush 1602 (GH, UI), 1602A (GH); Hannibal,
Davis 113 (AA), 704 (GH, AA), 2011 (AA, UI), 4646 (UI).

Kentucky: Louisville, Fischbach 228 (GH); Chalybeate Springs, Schacklette
261 (GH); Monticello, Smith & Hodgdon 3872 (GH); South Portsmouth, Demaree
11245 (AA).

ARKANSAS: Hardinville, Palmer 26350 (GH) ; Hamburg, Demaree 18743 (UI) ;
Baxter, Bush 15250 (AA); Hot Springs, Palmer 24470 (AA); Little Rock, G. M.
Merrill 1965 (UI); Nogo, G. M. Merrill in 1933 (UI); Van Buren, Apr. 29, 1910,
Brown (AA); Shirley, Palmer 33207 (AA); Izard Co., Palmer 35564 (AA);
Eureka Springs, Palmer 5616 (AA).

TENNESSEE: Knoxville, Ruth 317 (GH); Spence Field, Sharp & Svenson 7278
(GH); Craggie Hope, Svenson 346 (GH).

ALABAMA: Huntsville, Hubricht B2011 (MBG); Selma, “cultivated and more
or less naturalized,’ Cocks, no number or date (AA).

Mississtpp1: Rockport, Harbison 11 (AA); Tishomingo City, Harbison 40
(AA); Mooreville, Palmer 39015 (AA).

LouIsIANA: Covington, Apr. 22, 1910, Cocks (AA).

NesraskKA: Nemaha, Bates 5992 (AA).

Kansas: Baxter Springs, Bush 10380 (AA, UI).

OxKLaAHoMA: Near Page, Palmer 20761, 20764 (AA); near Ottawa, Stevens
2427 (AA); Ottawa Co., Palmer 29915 (AA).

Amelanchier arborea is an arborescent species with acuminate, cordate,
finely serrate leaves that are pubescent at flowering time, long narrow
petals, glabrous ovary, five styles, and somewhat dryish fruits, that has
been called A. canadensis by many botanists. It is closely related to
A. laevis and to A. interior. The bark is shallowly fissured and grayish
brown. When the leaves are about one-third grown the spreading or
drooping racemes of white flowers cover the tree. The flowers are there-
fore conspicuous in early spring when most other trees are leafless. The
wood is sometimes used for fishing poles, or for handles for tools. The
fruits of A. arborea, unlike those of other eastern American species are
scanty, scarcely edible, and fall early.

40 ILLINOIS BIOLOGICAL MONOGRAPHS

It has been pointed out by M. L. Fernald (Rhodora 43:562-563.
1941) that the first clear account of this species was that of Mespilus
canadensis var. B cordata Michx. 1803. “In 1810, the younger Michaux,
evidently taking the name from his father’s first word of diagnosis, ele-
vated M. canadensis var. 8B cordata to specific rank as M. arborea Michx.f.
. . . Here, then, is the first perfectly clear name for Amelanchier cana-
densis sensu Wiegand.” It is interesting to note, however, that the error
of misapplying the name A. canadensis to A. arborea (Michx.f.) Fern.
did not originate with Wiegand in 1912, for no fewer than twenty pre-
vious authors had misused the name A. canadensis in this sense. Appar-
ently the earliest binomial which might apply to this species is Mespilus
nivea Marsh. However, Marshall’s account, more of a horticultural com-
mentary than a botanical description, “is altogether too vague, unless an
original specimen of it can eventually be discovered. Its transfer into
Amelanchier would merely lead to the doubt which surrounds so many
names unfortunately taken up from Marshall’s inadequate and often
merely impressionistic accounts.”’ (Fernald, l.c.)

Pyrus wangenheimiana Tausch, based on material growing in the
Leibnitz garden at Prag, is represented by a sheet in the Bernhardi
Herbarium at the Missouri Botanical Garden, which through the kindness
of the curator, Dr. J. M. Greenman, I have had the opportunity of exam-
ining. This sheet is probably part of the original Tausch exsiccati and
may therefore be regarded as authentic. It has two specimens, one of
mature leaves, and the other consisting of a young flowering branch.
Although labeled by Wiegand in 1912 “a probable hybrid” between
A. canadensis (sensu auth.) and A. laevis Wieg., it is evidently quite
typical A. arborea (Michx.f.) Fern.

The leaves of A. arborea usually retain even at maturity some pubes-
cence on the lower surface, at least toward the base of the blades and
on the petioles. On certain specimens, however, especially those collected
from plants in exposed situations, the leaves are not infrequently glabrous
at maturity. Such specimens, which in the absence of fruits are some-
times not easily distinguished from A. laevis, have been named A. arborea
f. nuda by Palmer & Steyermark. |

The taxonomic identity of certain specimens from Westminster,
Vermont, collected by W. H. Blanchard in 1903 and 1907, and distributed
as “A. intermedia Spach” and “A. saxatilis Blanchard” is not yet de-
termined. In 1912, Wiegand was confronted with the same problem, which
he likewise was apparently unable satisfactorily to solve. The specimens
are, as Wiegand has said, difficult to understand because they do not
belong to any species now recognized. He suggested that they may have
been of hybrid origin, and concluded that they need more study in the

AMERICAN SPECIES OF AMELANCHIER—JONES 41

field. With this view I am in complete accord, but am at present unable
to add any further clarifying statement beyond the suggestion that
possibly the specimens represent hybrids between A. arborea and A.
bartramiana.

6a. AMELANCHIER ARBOREA var. alabamensis (Britt.) n. comb.

Amelanchier alabamensis Britton in Britton & Shafer, N. Am. Tr. 439, fig. 386
(1908) ; Wiegand in Rhodora 14:132, 240 (1912); Small, Man. Se. Fl. 636
(1933).

A small tree resembling A. arborea, but differing in the tomentose top
of the ovary; flowering specimens unknown.

Type Locatity: In sandy land three miles south of Auburn, Lee Co.,
Alabama, Mar. 19, and May 28, 1898, F. S. Earle & C. F. Baker 1610
(TYPE, in the herbarium of the New York Botanical Garden).

Rance: Arkansas and Alabama.

ARKANSAS: Magazine Mt., Logan Co., Palmer 23222 (AA).

ALABAMA: Auburn, Lee Co., Earle & Baker 1610 (NY); Birmingham, Jeffer-
son Co., Palmer 35329, 38948 (AA); Little Mountain, about four miles south of
Tuscumbia, Colbert Co., Harper 3322 (GH, AA).

Little has been added to the meager knowledge of this peculiar plant
since Wiegand studied it in 1912. It is here reduced to the varietal
category because it appears probable that it is not a species of equiva-
lent rank with A. arborea. Concerning the status of A. alabamensis,
Wiegand (Rhodora 14:132. 1912) commented as follows: “In 1908
Britton described Amelanchier alabamensis from material collected by
F. S. Earle and C. F. Baker three miles south of Auburn, Alabama. The
writer has seen this material which was distributed by Earle and Baker,
both the type specimen and also that which was sent to several other
herbaria, but is still unable to form any satisfactory opinion regarding it.
At flowering time the young leaves, hypanthium and sepals are like
A. canadensis [A. arborea], but the summit of the ovary is hairy. The
mature leaves are not distinctive but are more like those of A. canadensis
[A. arborea]. Nowhere in any of the herbaria studied by the writer are
there other specimens which will match these. Thus one is strongly forced
toward the opinion that A. alabamensis is not a good species of the same
grade as those with wide distribution, and that it is to be explained as a
local hybrid or as a local environmental variety. Before this conclusion is
finally reached however, a more extended search in the Southern States
should be made, for this region is one from which little material of
Amelanchier finds its way into the herbaria.” Later in the same year
Wiegand (l.c. 240) concluded that “a certain amount of woolliness on the
summit of the ovary must be admitted among the allowable variations of
A. canadensis [A. arborea] without, however, constituting a distinct

42 ILLINOIS BIOLOGICAL MONOGRAPHS

variety.” In the view of the present writer the most appropriate pro-
cedure at present is to treat the Alabama plants as a variety of A. arborea
until results of field studies are available to shed further light on the
understanding of these plants. At the present time flowering specimens
are unknown. Until these are at hand it seems inadvisable to relegate
A. alabamensis to synonymy under A. arborea, or to follow Britton &
Shafer, and Small in treating it as a distinct species.

6b. X AMELANCHIER GRANDIFLORA Rehd.

Amelanchier canadensis X laevis sensu Wiegand.
Amelanchier (?) botryapium lancifolia Simon-Louis apud Zabel, Syst. Verz.

Muenden 19 (1878), nom. nud.

Amelanchier canadensis grandiflora Zabel in Beissner, Schelle, & Zabel, Handb.

Laubh.-Ben. 191 (1903), nom. nud.

Amelanchier lancifolia hort. gall. ex Zabel, ibid., pro syn.
Amelanchier grandiflora Rehder in Journ. Arnold Arb. 2:45 (Sept. 6, 1920); Man.

Cult. Tr. & Shr. 390 (1927), (ed. 2) 388 (1940).

According to Sax (Journ. Arnold Arb. 12:9. 1931) this is a tetraploid
hybrid between A. arborea and A. laevis. In the original publication,
Rehder says that it differs from A. arborea in the larger flowers, the
longer and more slender, less pubescent racemes, and in the purplish
young leaves covered with a dense and more floccose tomentum which
soon disappears entirely; from A. Jaevis it differs in the tomentose young
leaves, the slightly villous racemes with more numerous flowers on shorter
pedicels, and in the larger, more succulent fruit. “Among the many
Amelanchiers grown at the Arboretum it is easily the most handsome and
always attracts attention by the abundance of its large flowers set off
effectively by the purplish foliage; it forms a large tree-like shrub with
wide-spreading slender branches. As I saw it in 1893 in the Botanic
Garden of the Forest Academy at Muenden it formed a well-shaped small
or medium-sized tree with spreading branches; it had been received from
the nursery of Simon-Louis near Metz as A. lancifolia, a name which
apparently was never published.” (Rehder, lc.). Accompanying the
original description are a number of citations of specimens which have
been referred to this hybrid by Wiegand. While there is little doubt that
the plants in cultivation are actually of hybrid origin, almost all the
specimens of feral plants that I have examined are clearly referable to
either A. laevis or A. arborea. The following specimens of cultivated
plants are in the herbarium of the Arnold Arboretum.

SPECIMENS OF CULTIVATED PiLants: Bot. Gard. Muenden, April 23
and July 3, 1893, A. Rehder (TYPE, AA); Arnold Arb. under no. 4406
(received from Muenden in 1892) May 11, 1900, May 3, 1902, June 22,
1903, May 11 and July 6, 1912, May 14 and July 5, 1919; Arb. Spaeth,
Berlin, May 8, 1909, H. Jensen.

AMERICAN SPECIES OF AMELANCHIER—JONES 43

6c. X AMELANCHIER GRANDIFLORA f. RUBESCENS Rehd.

A. grandiflora f£. rubescens Rehder in Journ. Arnold Arb. 2:46 (1920), Man. Cult.
Tr. & Shr. 390 (1927), (ed. 2) 388 (1940).

Flowers purple-pink in bud, tinged with pink when open.

Cultivated in the Durand-Eastman Park, Rochester, New York; speci-
mens collected: May 16, 1920, B. H. Slavin & J. Dunbar.

“This handsome form agrees in its characters with the type except
that the flowers are purple-pink in bud and suffused with pink when
open. It is a seedling from a tree of typical A. canadensis [i.e., A. arborea]
growing in Seneca Park, Rochester, and represented in our herbarium
by specimens collected by B. H. Slavin and marked No. 10. The seedling
described above, however, shows unmistakably the influence of A. laevis
which is growing at the same locality.”

7. AMELANCHIER CANADENSIS (L.) Medic.
(Plates UL TX and: Xx)

Mespilus canadensis L. Sp. Pl. 478 (1753), Syst. Veg. (ed. 13) 388 (1774); Miller,
Gard. Dict. (ed. 8) no. 6 (1768).

Mespilus virginiana Miller, Gard. Dict. ed. 8; No. 11 (1768).

Pyrus botryapium L.f. Suppl. Pl. Syst. 255 (1781); Willdenow, Sp. Pl. 2:1013
(1799), Enum. Pl. 525 (1809); Pursh, Fl. Am. Sept. 339 (1814); Sprengel,
Syst. Veg. 2:509 (1825).

Crataegus racemosa Lamarck, Encycl. Méth. Bot. 1:84 (1783); J. St. Hil. Expos.
Fam. Nat. 2:182 (1805); Poiret in Lamarck, Encycl. Méth. Bot. Suppl. 1:292
(1810).

Amelanchier canadensis (L.) Medicus, Gesch. 79 (1793); Michaux, Fl. Bor. Am.
1:291 (1803); K. Koch, Dendrol. 1:180 (1869); Fernald in Rhodora 43:566,
pl. 672, fig. 1 (1941) ; Weatherby & Adams in Contr. Gray Herb. 158:50 (1945).

Crataegus amoena Salisbury, Prodr. Stirp. Chap. Allert. 357 (1797), nomen illegit.

Amelanchier botryapium Borkhausen, Theor.-prakt. Handb. Forstb. 2:1260 (1803) ;
DC. Prodr. 2:632 (1825); Hooker, FI. Bor. Am. 1:202 (1834); Britton &
Brown, Illustr. Fl. N. U.S. 2:238, fig. 1986 (1897); Mohr in Contr. U.S. Nat.
Herb. 4:545 (1901); Card in Bailey, Cyclop. Am. Hort. 57 (1904); Britton,
Manvel IN; States 517 (1901), ied. 3). 517°(1907); Small, Fl. Se. U.S. 531
(1903) ; Keeler, Our Northern Shr. 192 (1903) ; Coker & Totten, Tr. N. Carol.
59 (1916).

Aronia botryapium Persoon, Syn. Pl. 2:39 (1807); Nuttall, Gen. Am. PI. 306
(1818); Torrey, Fl. N. & Mid. U.S. 1:479 (1824); Eaton, Man. Bot. (ed. 6)
29 (1833).

Aronia botryapium var. B racemosa Persoon, l.c.

Aronia ovalis sensu Elliott, Sketch Bot. S.C. & Ga. 1:558 (1821). Non Persoon 1807.

Pyrus ovalis sensu Bigelow, Fl. Boston. (ed. 2) 195 (1824). Non Willdenow, 1796,
nec Amelanchier ovalis Medicus 1793, nec Borkhausen 1803.

Amelanchier intermedia Spach, Hist. Nat. Vég. Phan. 2:85 (1834); M. Roemer,
Syn. Mon. 3:146 (1847); Blanchard in Torreya 7:98 (1907) ex p.; Britton &
Shafer, N. Am. Tr. 438, fig. 384 (1908) ; Stone, Pl. So. N.J. in Rep. N.J. State
Mus. 1910:488 (1911); Britton & Brown, Illustr. Fl. N. U.S. (ed. 2) 2:292,
fig. 2330 (1913) ; Wiegand in Rhodora 22:147 (1920) ; Hoffmann in Proc. Boston
Soc. Nat. Hist. 36:280 (1922) ; House in N.Y. State Mus. Bull. 254:412 (1924) ;
Wiegand & Eames, Fl. Cayuga Basin 248 (1926); Rehder, Man. Cult. Tr. &
Shr. 390 (1927).

44 ILLINOIS BIOLOGICAL MONOGRAPHS

Mespilus glabra Nuttall ex Hooker, Fl. Bor. Am. 1:202 (1834).

Pyrus neumanniana Tausch in Flora 21:76 (1838).

Aronia affinis Neum. ex Tausch, ibid.

Amelanchier canadensis var. B oblongifolia Torrey & Gray, Fl. N. Am.. 1:473
(1840) ; Torrey, Fl. N.Y. 1:225 (1843) ; Walpers, Rep. Bot. Syst. 2:55 (1843) ;
Gray, Man. Bot. 130 (1848), (ed. 2) 126 (1856); Watson & Coulter in Gray,
Man. Bot. (ed. 6) 167 (1889).

Amelanchier ovalis sensu Emerson, Tr. Massachusetts 444 (1846). Non Medicus
1793.

Amelanchier oblongifolia M. Roemer, Syn. Mon. 3:147 (1847); Robinson &
Fernald in Gray, New Man. Bot. (ed. 7) 460 (1908); Jones & Rand in Bull.
Vermont Agr. Exp. Sta. 145:99 (1909); Wiegand in Rhodora 14:147, pl. 96
(1912); Clements, Rosendahl, & Butters, Minnesota Tr. & Shr. 151 (1912);
Small, Florida Tr. 31 (1913), Shr. of Florida 29 (1913); Small & Carter, FI.
Lancaster Co. Pennsylvania 155 (1913); Bean, Tr. & Shr. Brit. Isles 1:189
(1914); Rehder in Bailey, Stand. Cyclop. Hort. 273 (1914); Hitchcock &
Standley, Fl. Distr. Columbia 178 (1919) ; Silva Tarouca, Freiland-Laubh. (ed. 2)
96, fig. 86 (1922); Bailey, Man. Cult. Pl. 378 (1924); Rehder, Man. Cult. Tr.
& Shr. 389 (1927); Rydberg, Fl. Prairies & Plains 437 (1932); Small, Man.
Se. Fl. 636 (1933) ; Rehder, Man. Cult. Tr. & Shr. (ed. 2) 388 (1940).

Amelanchier neumanniana M. Roem., l.c.

Amelanchier spicata sensu Decaisne in Nouv. Arch. Mus. Hist. Nat. Paris 10:135,
pl. 9 (1874). Non Crataegus spicata Lam.

Amelanchier canadensis var. obovalis Britton, Stern, & Poggenberg, Prelim. Cat. 17
(1888) excl. syn.; Sargent, Silva N. Am. 4:128, pl. 195 (1892), pro maxime
parte; Dippel, Handb. Laubh. 3:392 (1893) ; Schneider, Illustr. Handb. Laubh.
1:734, figs. 409, 410 (1906) ; Silva Tarouca, Freiland-Laubh. 139, fig. 124 (1913).

Amelanchier nantucketensis Bicknell in Bull. Torr. Club 38:453 (1911).

Amelanchier canadensis intermedia Ashe in Bull. Torr. Club 46:221 (1919).

Amelanchier sera Ashe in op. cit. 222; Rehder, Man. Cult. Tr. & Shr. 390 (1927),
(ed. 2) 388 (1940).

Amelanchier oblongifera Ashe, |.c. (err. in transcr.).

Amelanchier longifolia Stapf in Index Lond. 1:116 (1929) (err. in transcr.).

Amelanchier austromontana sensu Fernald in Rhodora 43:566 (1941), ex p. Non
Ashe 1918.

Shrub 2-8 m. tall, the stems slender, erect, fastigiately branched, form-
ing close bushy clumps; bark of the twigs grayish, glabrous; winter buds
small, dark brown, glabrous or nearly so; leaves thin, usually nearly
exactly elliptical, varying to oval or narrowly obovate, conduplicate in the
bud, unfolding at flowering time, the lower surface densely white-tomen-
tose, soon glabrous throughout, or the midvein and petiole often remain-
ing slightly pubescent; blades mostly 3.5-5 cm. long, 1.5-2.5 cm. wide,
acutish, or rounded and mucronate at the apex, the base rounded, rarely
subcordate or cuneate; lateral veins 9-13 pairs, irregularly and distantly
arranged, usually curved upward toward the middle and becoming ir-
regular and indistinct before reaching the margin; margins sharply serrate
with low sharp teeth, the lower part of the blade nearly or quite entire;
teeth 6-11 per cm., 25-40 on each margin on average leaves; petioles
slender, 8-15 mm. long; flowers small, appearing with the leaves; racemes
erect 2.5-6 cm. long, the rachis and pedicels whitish pubescent at first,
the lower pedicels 5-10 mm. long; petals 5, white, oval or oblanceo-

AMERICAN SPECIES OF AMELANCHIER—JONES 45

late, obtuse, 3-9 mm. long, 2-3 mm. broad; stamens about 20, the fila-
ments glabrous, 2-4 mm. long; anthers 1 mm. long; hypanthium campanu-
late, 3-5 mm. in diameter, tomentulose at the base or throughout, scarcely
constricted on the fruit; sepals triangular-lanceolate, 1.5-2.5 mm. long,
pilosulous within, mostly erect or ascending on the fruit; styles 5,
glabrous, 4-5 mm. long, fused to near the middle; summit of ovary
glabrous or nearly so; fruit globose or subglobose, 7-10 mm. in diameter,
purplish black, glaucous, sweet, juicy, edible; lowest fruiting pedicels 1-2
cm. long; seeds brown, smooth, obliquely lanceoloid, 4-5 mm. long, 2-3
mm. wide.

Type Locatity: “Habitat in Virginia, Canada.” Phototypes in the
Gray Herbarium and in the herbarium of the Arnold Arboretum. Type in
the Linnean Herbarium.

Rance: A species chiefly of swamps and bogs, and in low ground in
woods, from Newfoundland to Georgia, principally on the Coastal Plain.
Flowering from the end of March to the latter part of May; fruit ripen-
ing in June and July. Common names: shadbush, thicket shadblow,
wild pear.

NEWFOUNDLAND: Topsail Road, July 1931, Ayre (GH); Salmonier Line,
Knowlting in 1928 (GH).

New Brunswick: Grand Manan, Weatherby 7029 (GH); Bass River, June 2,
1869, Fowler (GH).

Nova Scotia: Bridgewater, Fernald & Long 23946 (GH); Dartmouth, Jack
684 (AA); Barrington, Fernald, Long, & Linder 21456 (GH); Maccan, Jack 3583
(AA); Truro, Jack 3639 (AA); Port Mouton, Bissell & Graves 21457 (GH);
Ohio, Jack 3108 (AA); Yarmouth, Bissell, Pease, Long, & Linder 21440 (GH);
Arcadia, Pease & Long 21453 (GH); Goven Lake, Fernald, Bartram, & Long
23937 (GH).

PRINCE Epwarp IsLanp: Bothwell, Fernald & St. John 11085 (GH).

Marne: Orono, May 1898, Harvey (UI); Waldo Co., Hyland 699 (GH) ; Jones-
port, Hyland 753 (GH); Stillwater, May 12, 1896, Merrill (NE); South Poland,
Furbish in 1893 (NE); Gilead, Furbish in 1897 (NE); Brunswick, Furbish in 1892
(NE); West Baldwin, Furbish in 1900 (NE); North Berwick, May 9, 1897,
Fernald (NE); Trotts Island, May 22, 1895, Fernald (NE); Alfred, Fernald &
Long 13772 (AA, NE); York, Bicknell 4840 (NE); Wells, Furbish in 1898 (NE).

New Hampsuire: Wolfeboro, Sargent 21 (GH); Winchester, May 11, 1912,
Flint (NE) ; Andover, Jack 3948 (AA); Dover, Hodgdon 2597 (NE); Barrington,
Hodgdon 2773 (NE); Merrimac Co., July 14, 1933, Bullard (NE); Bennington,
June 29, 1908, Coville (AA); Epping, Pease 24229 (NE); Derry, May 10, 1913,
Batchelder (NE); Charlestown, Woodward & Bean 17116 (NE).

VERMONT: Westminster, May 14, 1902, Blanchard (GH); Bellows Falls, May
10, 1915, Knowlton (NE).

MassacHuseETts: Plymouth, Fernald & Hunnewell 15200 (NE); Nantucket
Island, Bicknell 4815, 4847, 4851, 4858, 4862a, 4868, 4879 (NY), 4816, 4850 TYPE
collection of <A. nantucketensis (NY, NE), 4849 (NY, GH), 4857 (NE),
Day 57 (NY), 75 (NY, GH, NE); Lexington, Kennedy 3260 (NE); Concord,
Kennedy 2360 (GH); Amherst, Seymour 3493 (NE); Salem, July 4, 1925, Mackin-
tosh (NE); Chicopee, Seymour 571 (GH); Wellesley, Wiegand 2131, 2132 (GH) ;
Blue Hill, Milton, Palmer 20185 (AA); Harwich, Fernald & Long 18542 (NE);
Haverhill, Harris 423 (NE); Yarmouth, June 1916, Winslow & Sanford (NE);

46 ILLINOIS BIOLOGICAL MONOGRAPHS

Brewster, Fernald 18546 (GH, NE); Barnstable, Fernald & Long 16869, 18543
(NE); West Tisbury, Seymour 1222 (GH, NE); Revere, June 28, 1882, Young
(NE); New Bedford, May 7, 1910, Hervey (NE); Mt. Wachusett, May 11, 1897,
Bailey (NE); Douglas, Fernald 15191, 15198, 15199 (NE); Lunenburg, Fernald &
Bean 14160 (NE); Leicester, Hunnewell & Wiegand 2137, 2139, 2140, 2143 (NE).

CoNNECTICUT: Coventry,
Weatherby in 1935, Pl. Exsicc.
Gray. 662 (GH); Franklin, Wood-
ward in 1915 (NE); East Putnam,
June 10, 1922, Eaton & Fassett
(NE) ; Southington, Bissell in 1901
(GH, NE) ; Waterford, Graves in
1901 (GH); Middlebury, July 4,
1910, Blewitt (GH) ; Kent, Weath-
erby 4916 (NE) ; New Haven, Apr.
26, 1878, Allen (NE) ; Waterbury,
Blewitt 1508, 1801 (NE).

Ruope Isranp: Warwick, Apr.
17, 1910, Hope (NE); Barring-
ton, May 30, 1911, Winslow
(NE); Charlestown, Collins in
1919 (NE); South Kingston,
Gilbert, Rehder, & Smith 833
(NE); East Providence, Wiegand
991 (NE); Block Island, Fernald
& Long 9625 (GH, NE); Little
Compton, July 27, 1919, Collins
(NE).

New York: Niagara Falls,
July 8, 1898, Schneck (UI);
Waterloo, Wiegand 2541 (GH);
Oneida Lake, Muenscher & S palte-
holz 16175 (GH); Oswego, Wie-
gand 15627 (GH); Preble,
Wiegand 2539 (GH) ; Glens Falls,
ene ae an eee Mar 6.—Range of Amelanchier canadensis.
Kelvey in 1933 (AA); Brooklyn,

May 1, 1886, Stabler (GH); Ithaca, Wiegand & Eames 2515, 2517, 2518 (GH).

New Jersey: Riverton, Apr. 29, 1879, Eisele (AA); Millville, Adams 339
(GH); Merchantville, St. John & Long 1063 (GH); Lakewood, Byhouwer &
Kobuski 50 (AA); Egg Harbor City, Hunnewell 6031 (GH); Hammonton, May 27,
1923, Bassett (GH); Medford, Long 26820 (GH); Forked River, Moldenke 10580
(UI); Lakehurst, May 15, 1910, Mackenzie (GH); Island Heights, Muenscher 50
(GH).

PENNSYLVANIA: Philadelphia, May 1, 1926, McDowell (GH); Franklin Co.,
Jennings 1581 (GH); Brandamore, July 26, 1925, Stone (GH); Folsom, Long
58240 (Ph); Tinicum, Stone 6484 (Ph); Morrisville, Long 34224 (Ph); Bacton,
Long 31233 (Ph); Harmonyville, Long 33309, 33615 (Ph); Nottingham, Long
33330 (Ph); Fontaine, Long 32089 (Ph).

DELAWARE: Vandyke, Tidestrom 11947 (GH); Townsend, April 8, 1909, Long
(Ph); Wilmington, A. Commons in 1875 (Ph).

District oF CoLtumsta: North Takoma, May 2, 1897, Williams (AA); Naucks,
June 1, 1913, Steele (AA).

Marycanp: Elkton, Wherry & Adams 2775 (GH); Lanham, Blake 9324 (GH) ;
Riggs Mill, Prince Georges Co., Blake 9362 (GH); Bacon Hill, Long 57005 (Ph) ;
North East, Long 57021 (Ph); Principio Furnace, Long 54350 (Ph).

AMERICAN SPECIES OF AMELANCHIER—JONES 47

West Vircinta: Roland Park, Gabell Co., Gilbert 398 (GH).

Vireinta: Accomac, Pease 26992 (GH); Falmouth, Wiegand & Manning 1328
(GH); Richmond, April 13, 1887, Kennedy (GH); Clarendon, Allard 1229 (GH) ;
Williamsburg, Menzel 403 (GH); Sebrell, Fernald & Long 7869 (GH, Ph);
Franklin, Fernald & Long 9949, 11343 (GH, Ph); Lee’s Mill, Fernald & Long
12096 (GH, Ph); Zuni, Fernald & Long 7068 (GH); Emporia, Fernald & Long
7070 (GH, Ph); Little Texas, Fernald, Long, & Pease 11700 (GH, Ph).

NortH Carotina: Abbottsburg, Harbison 8 (AA); Pleasant Hill, Fernald &
Long 7071 (GH, Ph); Hamlet, Wiegand & Manning 1330 (GH); Blount Creek,
Godfrey, White, & Shelbourne 7038 (GH, MBG) ; Lumberton, Wiegand & Manning
1331, (GH).

SoutH CaroLina: Sumter, Rehder 955 (AA); Seneca, Palmer 35412 (MBG) ;
Manning, Stone 65 (Ph).

Georcia: Augusta, Sargent in 1900 (AA); Graymont, Harper 819 (GH);
Franklin, Hermann 10048 (GH).

In the flowering stage, this species may be recognized by the glabrous
ovary, short petals, and the erect tomentose-lanate racemes; the densely
pubescent leaves are about half unfolded. In the fruiting stage the rather
firm elliptical or slightly obovate finely serrate leaves that are soon quite
glabrous, dark green, smooth, and somewhat glossy above, paler beneath,
with rounded base and acutish or rounded, often mucronulate apex, are
distinctive. This species is apparently closely related to A. spicata, from
which it usually may be distinguished by the somewhat different habit
of growth, the elliptical leaves, glabrous top of the ovary, and the some-
what larger fruits, with erect or spreading sepals.

A peculiarity of A. canadensis has been well described by Wiegand
(Rhodora 14:149. 1912): “As the fruit matures the inflorescence expands
much less in this species than in others, the axis and pedicels remaining
short. The shoots upon which the racemes are borne remain short also,
while frequently there is a strong growth of leafy shoot beyond the in-
florescence and as a result the inflorescence often appears to have been
left far behind, and to have been lateral when really terminal. Unfortu-
nately this condition is too frequently obscure to be of use as a dis-
tinguishing characteristic.”

For some reason not apparent, the younger Linnaeus, when transfer-
ring his father’s Mespilus canadensis to the genus Pyrus, gave it a new
specific name, botryapium. However, the earliest name that applies to this
species is clearly Mespilus canadensis L., which was transferred to
Amelanchier by Medicus in 1793. In his discussion of certain eastern
North American species of Amelanchier, Fernald (Rhodora 43:560-561.
1941) observes: “Mespilus canadensis L. Sp. Pl. 1:478. 1753 was pub-
lished with unusual lack of involving references, merely the plant of
Linnaeus’s herbarium described, with a single reference to a description
of Gronovius. . . . It is, therefore, unfortunate that, when he so clearly
differentiated our species of Amelanchier and thus gave study to the genus
a new and stimulating interest, Wiegand seems to have misunderstood the

48 ILLINOIS BIOLOGICAL MONOGRAPHS

basis of A. canadensis. He had had a comparison made by a botanist not
familiar with the eastern species and he then used the Linnean name for
the largest member of the genus, the large shrub or tree with cordate,
ovate, or broadly ovate-oblong, sharply serrate leaves which, like those
of true A. canadensis (A. oblongifolia), are pubescent beneath on unfold-
ing, losing most of their pubescence with age.” This is the species now
called A. arborea (Michx.f.) Fern. For a more extended discussion of
the nomenclatural history of A. canadensis (L.) Medic., the reader should
consult the article just quoted.

Pyrus neumanniana Tausch, based on material growing in the Leibnitz
garden at Prag, is represented by a sheet in the Bernhardi Herbarium at
the Missouri Botanical Garden. This sheet is probably part of the original
Tausch exsiccatae and may therefore be regarded as authentic. Wiegand,
in 1912, labeled it “a probable hybrid” with A. bartramiana (Tausch)
M. Roem. as one of the parents. Later the sheet was annotated in pencil
by Professor Alfred Rehder: “A. Botryapium Borkh.” It contains two
specimens, a fruiting branch and a flowering one. The leaves are those
of typical A. canadensis (L.) Medic., and match perfectly those of a
photograph of the Linnean type. The small flowers, as well as the rather
characteristic inflorescence, belong unmistakably to A. canadensis.

The binomial, A. spicata Decaisne, 1874, like that of K. Koch five
years earlier, was based on the name Crataegus spicata Lam., but it is
clearly evident from Decaisne’s description (“vertice ovarii glabris, stylis
coalitis”), as well as from the floral structures shown in the accompanying
plate, and by the cited synonyms (i.e., A. ovalis Lindl., Mespilus ovalis
Willd., and M. canadensis B oblongifolia Torr. & Gray), and the state-
ment of geographical range, that Decaisne was dealing with A. canadensis
(L.) Medic. instead of the plant that Lamarck had called Crataegus
spicata. Hence, A. spicata sensu Dene. is a synonym of A. canadensis (L.)
Medic., not of A. spicata (Lam.) K. Koch, as it is sometimes cited.

In 1911, E. P. Bicknell described as A. nantucketensis Bickn. a “shrub
1.5 dm. to 2 m. high” with short petals and glabrous ovary. This was said
to be common in low grounds about the borders of swamps on Nantucket
Island, Massachusetts. From an examination of an isotype, it is evident
that Bicknell’s plants are a small form of A. canadensis (L.) Medic.
This conclusion is supported by the fact that this species is not un-
common on Nantucket, and by Bicknell’s statement that “intermediates”
between his plant and A. canadensis (L.) Medic. have been found in the
vicinity. It should be noted here that Bicknell’s plants are not the same
as those described four years earlier as A. oblongifolia var. micropetala
by B. L. Robinson. The specimens cited by Robinson are apparently a
small-flowered form of A. spicata (Lam.) K. Koch.

It may be worth noting that occasional specimens of A. canadensis

AMERICAN SPECIES OF AMELANCHIER—JONES 49

bear leaves somewhat resembling those of A. spicata. Such conditions are
the probable basis for the statement by Robinson & Fernald in Gray’s
Manual (ed. 7, p. 460) under A. oblongifolia (T. & G.) Roem.: “Highly
variable, passing into forms with broader elliptical or ovate-lanceolate
acutish leaves of deeper green color (being the A. spicata of many auth.,
not C. Koch). Apparently intergrades with other species.”

8. AMELANCHIER OBOVALIS (Michx.) Ashe
@Platess VIII Xoo, XXITT)
(?) Mespilus amelanchier sensu Walter, Fl. Carol. 148 (1788). Non L. 1753.
Mespilus canadensis var. « obovalis Michaux, Fl. Bor. Am. 1:291 (1803).
Amelanchier oblongifolia var. B walteri M. Roem. Syn. Mon. 3:147 (1847).
Amelanchier canadensis var. obovalis Sargent, Silva N. Am. 4:128 (1892), pro parte,

excl. pl. 195.

Amelanchier obovalis Ashe in Bot. Gaz. 35:434 (1903) ; Sargent, Man. Tr. N. Am.

361, fig. 284 (1905), ex parte; Fernald in Rhodora 43:566, pl. 672, fig. 3 (1941).
Crataegus canadensis obovalis Sargent ex Ashe in Lc. (err. in transcr.)
Amelanchier oblongifolia sensu Wiegand in Rhodora 14:147 (1912), ex parte.
Amelanchier stolonifera Wiegand, in op. cit. 144, ex parte.

Low shrubs 0.2-1.5 m. tall, surculose, and forming loose colonies;
winter buds conical, acute, reddish brown, dull, vernicose, the scales
ciliate; twigs slender, soon glabrous; leaves commonly oval or elliptical,
varying to slightly obovate, conduplicate in the bud, densely whitish
tomentose beneath when young, unfolding after the flowers, which are on
leafless or nearly leafless twigs; mature blades of rather firm texture, 2-5
cm. long, 1-3 cm. wide, the apex acutish, or obtuse and mucronulate, the
base acute or rounded, or less commonly slightly subcordate, the upper
surface dark green, dull, or somewhat glossy, smooth, the lateral veins not
particularly prominent, the midvein impressed, the lower surface pale
green, rather prominently veined, glabrous or nearly so at maturity, or
with lingering traces of tomentum; lateral veins 7-9 pairs, irregularly and
distantly arranged, usually curved upward and becoming irregular and
indistinct before reaching the margin; margins sharply serrulate nearly or
quite to the base, or frequently the lower third nearly or quite entire;
teeth 6-9 per cm., 20-30 on each side of average leaves; stipules linear,
pubescent, soon deciduous; petioles 5-15 mm. long, glabrous or slightly
pubescent at maturity; flowers precocious, in short, dense, compact, erect,
usually leafless, 4-10-flowered racemes 1-3 cm. long, the lower pedicels 1-3
mm. long; petals 5, white, glabrous, elliptical, minutely clawed, obtuse,
6-7 mm. long, 3-4 mm. wide; stamens 20, the filaments glabrous; anthers
0.6-0.7 mm. long; hypanthium saucer-shaped, 2-3 mm. in diameter, tomen-
tulose outside, not at all constricted on the young fruit; sepals triangular,
acute, 1-2 mm. long, pubescent within, divergent after anthesis; styles 5,
glabrous, 2-3 mm. long, usually united below the middle; top of the ovary
glabrous; mature fruits globose, purplish black, glabrous, 6-8 mm. in
diameter, sweet, juicy, edible; fruiting racemes erect, compact, 2-3 cm.

50 ILLINOIS BIOLOGICAL MONOGRAPHS

long, 2-8-fruited (average 4); sepals on the fruit erect or divaricate, less
commonly somewhat reflexed, glabrous, triangular-lanceolate, 2-3 mm.
long; fruiting pedicels usually 3-8 mm. in length, or occasionally the low-
est becoming 11 or even 14 mm. long; seeds reddish brown, smooth,
obliquely lanceoloid, somewhat flattened, obtusish at each end, 4-5 mm.
long, 2-2.5 mm. wide.

Type Locarity: “In Carolina inferiore.” Type in Michaux’s herbar-
ium in the Museum d’Histoire Naturelle, Paris. Phototype in the Gray
Herbarium.

RANGE: Open woods or sandy pine barrens along the Atlantic Coastal
Plain, from Pennsylvania to Georgia; flowering in late March and early
April; fruits ripening in May and June.

PENNSYLVANIA: Near Slatedale, Pretz 8296, 9260 (Ph); Point Pleasant, Benner
2703, Long 32883 (Ph); Naceville, Long 18759 (Ph); Yardley, Long 30300 (Ph);
Spring House, Long 33071 (Ph); Wakefield, Tanger 3351 (Ph) ; Castle Rock, June
9, 1904, Jahn (Ph); Folsom, Long 58269 (Ph).

New Jersey: Scott's, Middlesex Co., May 28, 1922, Mackenzie (Ph); Shark
River Station, May 21, 1922, Mackenzie (Ph); Clarksburg, Long 45720 (Ph);
Charleston Springs, Long 52012 (Ph); Chatsworth, Long 16451 (Ph); Clementon,
Long 21005 (Ph); Ostrom, Long 48766 (Ph); Prospertown, Long 30603 (Ph);
Friesburg, Long 35296 (Ph); Parkdale, Long & Pennell 7364, 7374 (Ph) ; Newton-
ville, Long 5915, 48268 (Ph); Fairview, Long 30773 (Ph); East Creek, Long 21590
(Ph) ; Wildwood Junction, Bartram 3220 (Ph); Browns Mills, April 30, 1905,
MacElwee (Ph).

MaryLanpb: Elkton, Long 54322 (Ph); Bacon Hill, Long 54368 (Ph); North
East, Long 54423 (Ph).

VirGINIA:. McKenney, Fernald & Long 13950 (Ph); Petersburg, Fernald, Long,
& Smart 5790, Fernald & Long 9947 (GH, Ph); Waverly, Fernald & Long 7072,
7870, 13042 (GH, Ph); Homeville, Fernald & Long 7073 (GH, Ph); Franklin,
Fernald & Long 7448 (GH, Ph); Suffolk, Fernald & Long 7074 (GH, Ph);
Whaleyville, Fernald & Long 7449 (GH, Ph); Lee’s Mill, Fernald & Long 11846
(GH, Ph); Orion, Fernald & Long 13043 (GH, Ph); Emporia, Fernald & Long
11847 (GH).

NortH CaroLtinaA: French Broad River, June 1898, Biltmore Herbarium 6706
(GH, AA).

SoutH Carottna: Near Charleston, Hunt 2969, Hunt & Martin 1408, 2526 (UI).

Amelanchier obovalis (Michx.) Ashe is a dwarf surculose shrub,
forming loose colonies, with the flowering or fruiting stems only 0.2-1.5
m. high, superficially resembling, but apparently quite distinct from, the
widespread A. spicata (Lam.) K. Koch. At flowering time it may be dis-
tinguished from that species by the precocious flowers in compact racemes
on leafless twigs, and by the glabrous ovary. It bears some structural
resemblances to A. canadensis (L.) Medic, but differs in its smaller size
and dissimilar growth-form, as well as the usually more oval leaves, some-
what shorter petals, and the shorter fruiting pedicels.

This species of dwarf shrubs was described by Michaux in 1803 as
Mespilus canadensis, “Var. a obovalis: humilior; foliis oblongiuscule
ovalibus . . . in Carolina inferiore.’”’ A phototype (Plate VIII, fig. 2)

AMERICAN SPECIES OF AMELANCHIER—JONES 51

shows Michaux’s original labels “Mespilus canadensis a obovalis. Arbriss
[eau] de deux pieds de haut. Carolines.” Wiegand in 1912 treated it tenta-
tively as a synonym of A. canadensis [oblongifolia], but it had been
raised to specific rank by W. W. Ashe in 1903. After several seasons of
field work in southeastern Virginia, Fernald & Long concluded that it is
distinct, and in a recently published study (1941) of the flora of Virginia,
Fernald has treated these plants as a separate species.

Thomas Walter’s “Mespilus Amelanchier?” is somewhat doubtfully
included here. Miss Nell Horner, Librarian of the Missouri Botanical
Garden, kindly supplied the following transcript of the original descrip-
tion: “inermis, foliis ovato-lanceolatis, crenatis, tenellis tomentosis adult-
ioribus laevibus nitidis; floribus corymbosis.”’

9 -AMELANCHIER SPICATA (Lam.) K. Koch
@Plates PL XI), XIE XLV, XV, XVI)

Crataegus spicata Lamarck, Encycl. Méth. Bot. 1:84 (1783).

Pyrus ovalis Willdenow, Berlin Baumz. 259 (1796), Sp. Pl. 2:1014 (1799), Enum.
Pl. 525 (1809); Muhlenberg, Cat. Pl. Am. Sept. 49 (1813); Pursh, Fl. Am.
Sept. 340 (1814); Sprengel, Syst. Veg. 2:509 (1825).

Amelanchier ovalis sensu. Borkhausen, Theor.-prakt. Handb. Forstbot. 2:1259
(1803); DeCandolle, Prodr. 2:632 (1825); Spach, Hist. Veg. Phan. 2:85
(1834) ; M. Roemer, Syn. Mon. 3:146 (1847); Dippel, Handb. Laubh. 3:390
(1893) ; Rehder in Bailey, Stand. Cyclop. Hort. 273 (1914). Non Medicus 1793.

Aroma ovalis Persoon, Syn. Pl. 2:40 (1807) ; Torrey, Fl. N. Middle U.S. 479 (1824).

Amelanchier spicata (Lam.) K. Koch, Dendrol. 1:182 (1869), as to name only,
excl. syn. & descr.; Koehne, Deutsche Dendr. 256 (1893); Britton & Brown,
Illustr. Fl. N. U.S. 2:238, fig. 1987 (1897); Britton, Man. Fl. N. States 517
(1901), (ed. 3) 519 (1907); Small, Fl. Se. U.S. 532 (1903); Card in Bailey,
Cyclop. Am. Hort. 57 (1904); Schneider, Illustr. Handb. Laubh. 1:737, figs.
411, 412 (1906); Apgar, Ornam. Shr. U.S. 182, fig. 279 (1910); Britton &
Brown, Illustr. Fl. N. U.S. (ed. 2) 2:292, fig. 2331 (1913); Rehder, Man. Cult.
Tr. & Shr. 389 (1927), (ed. 2) 388 (1940); G. N. Jones, FI. Illinois 154 (1945).

Amelanchier canadensis var. spicata Sargent, Silva N. Am. 4:129 (1892).

Amelanchier saxatilis Blanchard in Torreya 7:99 (1907).

Amelanchier erecta Blanchard, op. cit. 101.

Amelanchier intermedia sensu Blanchard, l.c. Non Spach 1834.

Amelanchier oblongifolia sensu Robinson & Fernald in Gray, Man. Bot. (ed. 7)
460 (1908), ex p. Non M. Roem. 1847.

Amelanchier oblongifolia var. micropetala Robinson in Rhodora 10:33 (1908) ; Rob-
inson & Fernald in Gray, New Man. Bot. (ed. 7) 460 (1908); Rehder, Man.
Cult. Tr. & Shr. 389 (1927), (ed. 2) 388 (1940).

Amelanchier humilis Wiegand in Rhodora 14:141, pl. 95 (1912) ; Clements, Rosen-
dahl, & Butters, Minnesota Tr. & Shr. 153 (1912); Rehder in Bailey, Stand.
Cyclop. Hort. 272 (1914); Rydberg, Fl. Rocky Mts. 447 (1917); House, N.Y.
State Mus. Bull. 254, 413 (1924); Deam, Shr. of Indiana 145, pl. 58 (1924) ;
Wiegand & Eames, Fl. Cayuga Basin 247 (1926) ; Rehder, Man. Cult. Tr. & Shr.
389 (1927); Rosendahl & Butters, Tr. & Shr. Minnesota 216 (1928); Peattie,
Fl. Indiana Dunes 219 (1930); Palmer & Steyermark in Ann. Missouri Bot.
Gard. 22:557 (1935) ; Marie-Victorin, Fl. Laurent. 316, fig. 91 (1935); Nielsen
in Am. Midl. Nat. 22:171, pls. 5-8 (1939) (excl. syn.); Rehder, Man. Cult.
Tr. & Shr. (ed. 2) 387 (1940) ; Steyermark, Spr. Fl. Missouri 255, pl. 68, fig. 2
(1940) ; Deam, Fl. Indiana 532 (1940).

By ILLINOIS BIOLOGICAL MONOGRAPHS

Amelancher stolonifera Wiegand in Rhodora 14:144, pl. 95 (1912), ex p.; Small &
Carter, Fl. Lancaster Co., Pa. 155 (1913); Rehder in Bailey, Stand. Cyclop.
Hort. 273 (1914) ; Hitchcock & Standley, Fl. Distr. Columbia 178 (1919) ; Hoff-
mann in Proc. Bost. Soc. Nat. Hist. 36:281 (1922) ; Pease in ibid. 37:267 (1924) ;
House in N.Y. State Mus. Bull. 254:412 (1924); Wiegand & Eames, Fl. Cayuga
Basin 247 (1926); Rehder, Man. Cult. Tr. & Shr. 389 (1927); Rosendahl &
Butters, Tr. & Shr. Minnesota 219 (1928); Small, Se. Fl. 636 (1933); Marie-
Victorin, Fl. Laurent. 316, fig. 91 (1935); Nielsen in Am. Midl. Nat. 22:177,
pl. 2, b (1939) ; Rehder, Man. Cult..Tr. & Shr. 387 (1940).

Amelancus spicata Vollman, Fl. Bayern 453 (1914).

Amelanchier botryapium var. obovalis Farwell in Rep. Mich. Acad. Sci. 17:175
(1916).

Amelanchier botryapium var. conferta Farwell, l.c.

Amelanchier botryapium var. micropetala Farwell in op. cit. 176.

Amelanchier austromontana Ashe in Journ. Elisha Mitchell Sci. Soc. 34:138 (1918).

Amelanchier beata Ashe, lc.

Amelanchier micropetala Ashe in Bull. Torr. Club 46:223 (1919).

Amelanchier micropetala var. potomacensis Ashe, l.c.

Amelanchier stolonifera var. lucida Fernald in Rhodora 23:267 (1922).

Amelanchier humilis var. typica Nielsen in Am. Midl. Nat. 22:171, pl. 5 (1939).

Amelanchier humilis var. compacta Nielsen, op. cit. 174, pl. 7.

Amelanchter humilis var. campestris Nielsen, op. cit. 176, pl. 6.

Amelanchier humilis var. exserrata Nielsen, op. cit. 177, pl. 8.

Amelanchier mucronata Nielsen, op. cit. 178, pl. 9.

Amelanchier humilis * laevis Deam, Fl. Indiana 532 (1940).

Amelanchier canadensis var. micropetala Rehder in Journ. Arnold Arb. 26:71 (1945).
Low, surculose colonial shrubs 0.3-2 m. tall; winter buds conical, acute,

reddish brown, dull, vernicose, the scales ciliate; leaves commonly oval,

varying to broadly ovate or suborbicular, conduplicate in the bud, unfold-
ing before or with the flowers and usually about half-grown at anthesis

(rarely the flowers on nearly leafless twigs), densely whitish tomentose

beneath when young; mature blades 2.5-5 cm. long, 2-3.5 cm. wide, the

apex acutish, or obtuse and more or less mucronate, the base rounded or
less commonly subcordate, usually nearly or quite glabrous on both sur-
faces; lateral veins 7-9 pairs, not prominent, irregularly and distantly
arranged, usually curved upward and becoming irregular and indistinct
before reaching the margin; margins finely and evenly serrate nearly or
quite to the base, or frequently the lower third almost or quite entire; teeth

5-8 per cm., 20-30 on each side of average leaves; stipules linear, pubescent,

deciduous; petioles slender, 1-2 cm. long, glabrous or slightly pubescent

at maturity; flowers in short, dense, erect, 4-10-flowered racemes 1.5-4

cm. long, the lower pedicels 6-18 mm. long; petals 5, white, or sometimes

pinkish, oblanceolate, obtuse, 4-10 mm. long, 3-4 mm. wide; stamens about

20, the filaments glabrous; anthers 0.6-0.9 mm. long; hypanthium saucer-

shaped, 3-4 mm. in diameter, glabrous or pubescent outside, more or less

constricted on the young fruit; sepals triangular-lanceolate, acute, 2-3 mm.

long, pubescent within, usually recurved from the middle after anthesis;

styles 5, glabrous, 2-3 mm. long, usually united only near the base; top of

AMERICAN SPECIES OF AMELANCHIER—JONES 53

the ovary densely tomentose; mature fruit globose, purplish black, glau-
cous, glabrous, 6-8 mm. in diameter, sweet, juicy, edible; lower pedicels
1-3 cm. long; seeds brown, smooth, obliquely lanceoloid, somewhat flat-
tened, about 5 mm. long, 2-3 mm. wide when well developed.

Type Locauity: Cultivated in the Jardin du Roi, Paris; said to have
come originally from Canada. Phototype and fragment of holotype in the
herbarium of the Arnold Arboretum. Type in the Museum d'Histoire
Naturelle, Paris.

Rance: On gravelly or rocky shores or river banks, sandstone or
limestone cliffs and ledges, rocky summits, in woods or thickets, pine
barrens, or sand dunes, from Newfoundland to Alabama, eastward to
Missouri and Minnesota (and eastern North Dakota). Flowering from
the beginning of April in the south to the middle of June (or somewhat
later) in the northern part of its range; fruit ripening from July to Sep-
tember. Common names; low juneberry or shadblow.

NEWFOUNDLAND: Grand Falls, Fernald & Wiegand 5557 (GH), 5558 (AA, GH),
5559565561) 5562;, 5563, 5605, 5608, 5623 (GH), 5633 (GH, AA); Rushy Pond,
Fernald & Wiegand 5627, 5630, 5635 (GH); Birchy Pond Stream, Fernald & Wie-
gand 3553 (GH); Dildo Run, Fernald & Wiegand 5565 (GH); St. Johns, Ayre in
1932 (GH); Trepassy, Fernald, Long, & Dunbar 26759 (GH).

Nova Scotia: Middleton, Fernald, Pease, & Long 21436 (GH), 21437 (GH,
TYPE of A. stolontfera var. lucida, AA), 21435 (GH, AA), Jack 3209 (AA);
Port Mouton, Jack 3479 (AA); Argyle, Fernald & White 21438 (GH), Pease &
Long 21452 (GH, AA); Kemptville, Fernald & Long 23933 (GH)-; Goven Lake,
Fernald, Bartram, & Long 23930 (GH); Gavelton, Fernald, et al. 23929 (GH);
Tusket, Jack 3357 (GH, AA), 3762 (AA); Birchtown Brook, Fernald & Long
23934 (GH); Barrington, Fernald, Long, & Linder 21439 (GH); Springhill Junc-
tion, Pease & Long 21434 (GH); Shubenacadie Grand Lake, Fernald, Bartram, &
Long 23931 (AA, GH), Fernald & Bissell 21433 (GH); Avonport, Roland 2047
(GH) ; Millville, Roland 41469 (GH); Bridgewater, Fernald & Long 23932 (GH),
Fernald, et al. 21432 (GH), Jack 3514 (AA).

Prince Epwarp Istranp: Dundee, Fernald, Long, & St. John 7593 (GH);
Southport, Fernald & St. John 7585 (GH); Mt. Stewart, Fernald, et al. 7582, 7591,
7553 (GH); Charlottetown, Fernald & St. John 11080 (GH); Cavendish, Fernald,
Long, & St. John 7595 (GH); Indian River, Fernald, Long, & St. John 7594
(GH); Tignish, Fernald, Long, & St. John 7584 (GH).

New Brunswick: Boiestown, Fernald & Pease 25134 (GH); Portage Island,
Blake 5677 (GH); Gorge of the Aroostock River, Fernald 1881 (GH); Wood-
stock, Fernald & Long 13781 (GH).

QueBEcC: Montmorency Falls, Macoun 66924 (GH); Lake Temiscaming, J/.-
Victorin 8237 (AA); Contrecoeur, M.-Victorin & R.-Germain 33110 (AA, GH);
Longueuil, M.-Victorin 9504, 11221, 11223 (AA); North Wakefield, Macoun 85506
(AA) ; Oka, M.-Victorin 24545, 2087 (AA); La Trappe, M.-Victorin & R.-Germain
33130 (AA, GH), Louts-Marie 116 (GH); Deschenes, Rolland 13033, 13035 (GH) ;
Windsor, July 25, 1923, Knowlton (GH).

Marne: Ft. Fairfield, Fernald 1888 (NE); Clifton, Fernald 2644 (NE); Winn,
Fernald & Long 13762 (AA, NE); Milford, Fernald 13760, 13773, 13774 (NE),
13775 (GH); Dover, G. B. Fernald 43 (NE); Fairfield, Fernald & Long 13766
(NE) ; Skowhegan, Chamberlain in 1903 (NE); Bald Mt., Chamberlain & Knowl-
ton in 1902 (NE); Phillips, Furbish in 1894 (NE); Oxford, Weatherby in 1914
(NE); Whitneyville, Knowlton in 1908 (NE); Mt. Desert I., Fernald in 1892

54 ILLINOIS BIOLOGICAL MONOGRAPHS

@ A. spicata a Pan Rea SZ PN e 30
® A. gaspensis ae f \ ef
\

\ ; \
= aes ie (Hie

Map 7.—Range of Amelanchier spicata and A. gaspensts.

(NE, GH), Williams in 1899 (NE), Rehder in 1936 (AA); Isle au Haut, Hill
1700 (NE) ; Matinicus, Long 320, 335 (NE); Rockland, Long 852 (NE) ; Rockport,
Fernald 9620 (NE); West Bath, Furbish in 1892 (NE) ; Standish, Fernald & Long
13777 (AA, NE); Brunswick, Chamberlain 228 (NE); Kennebunkport, Koehler 1
(GH); York, Fernald in 1900 (GH, NE).

VERMONT: Essex Junction, Eggleston 1176 (GH); Burlington, Blake 2496
(NE) ; Barnet, May 31, 1881, Blanchard (NE); Snake Mt., near Weybridge, Brain-
erd in 1878 and 1898 (GH); Rutland, Eggleston 1179, 1180 (AA, NE, GH, UI);
Twin Mts., Eggleston 185 (GH); Arlington, Schweinfurth, et al. in 1935 (NE);
Bellows Falls, Blanchard 3 (AA, GH, TYPE coll. of A. saxatilis), Blanchard 4
(TYPE coll. of A. erecta, GH); Westminster, Blanchard 4 (GH, AA); North
Westminster, Blanchard 4 (AA, GH).

New HampsHirE: Salem, May 22, 1909, Churchill & Purdie (NE, GH);
Rindge, May 31, 1913, Batchelder (NE); Jaffrey, Rand & Robinson 618 (GH);
Walpole, Fernald 105 (GH), 72 (NE, GH), Bean & Fernald 17013, 17014 (NE);
New Hampton, Pease 25912 (NE); Alton, Pease 25775 (NE) ; Gilford, Pease 26541
(NE); Strafford, Pease 24246 (NE); Rollinsford, Hodgdon 3192 (NE); Dover,
Hodgdon 2600 (NE); Durham, Hodgdon 2845 (NE); Bradford, Fernald & Sven-
son 914 (NE); Franklin, Jack 3872 (AA); Andover, Jack 3946 (AA); Shelburne,
Pease 25979 (NE); Lake Umbagog, Pease 16554 (NE); Errol, Pease 16990 (NE) ;
Ashland, Fernald 15197 (GH, NE), 15204 (NE); Bath, Pease 19680 (NE); Madi-
son, Pease 17890 (NE); Ossipee Lake, Weatherby & Smith (Pl. Exsicc. Gray
842) (GH, UI).

MassaAcHusetTts: Washington, May 31, 1909, Hoffmann (NE); Lenox, May 27,
1920, Hoffmann (NE); West Stockbridge, May 23, 1920, Hoffmann (NE); Gt.
Barrington, June 21, 1915, Hoffmann (NE) ; Sheffield, Aug. 27, 1902, July 25, 1912,
Hoffmann (NE); Montague, May 13, 1911, Fernald (NE, GH); Sunderland, Man-
ning & Seymour 3687 (NE); Shutesbury, Tower & Seymour 3668, 3671 (NE);
Agawam, Weatherby 4255 (NE); Amherst, Seymour 3512 (NE); Northampton,
Goodale & Markert 76864 (NE); Mt. Holyoke, Hubbard & Torrey T352

AMERICAN SPECIES OF AMELANCHIER—JONES 55

(NE); Russell, Fernald 9626 (NE); Chicopee, Murdoch & Torrey T391 (NE);
Lunenburg, Fernald & Bean 14132 (NE); Athol, June 19, 1935, Churchill (NE);
Ashburnham, May 19, 1924, Knowlton (NE) ; Holden, Blake & Fernald 3645 (NE) ;
Princeton, R. H. Piper 76881 (NE) ; Southbridge, May 19, 1916, Woodward (NE) ;
Gloucester, St. John 11887 (NE); Andover, Pease 683 (NE); Sherborn, Loomis
856 (NE); Tyngsboro, Pease 23232 (NE); Wellesley, Wiegand 2133 (GH, TYPE
of A. stolonifera); Blue Hill, Milton, June 10, 1900, Williams (GH), Floyd 801,
803, 980, 1008 (NE), May 11, 1903, and Sept. 22, 1900, Rehder (UI), May 7, 1899,
Kennedy & Fernald (GH, TYPE of A. oblongifolia var. micropetala), May 6,
1899, Churchill (GH, NE), Bartlett 846 (GH, UI); Mattapan, May 6, 1905, Cheever
(NE) ; Plymouth, Sanford 626 (NE) ; Chilmark, Seymour 1708, 1709 (GH); West
Tisbury, Seymour 4641 (NE); Harwich, Fernald 16867 (NE); Barnstable, Wood-
ward ¢& Fernald 15202 (NE); Sandwich, Fernald & Long 18548 (NE) ; Yarmouth,
Fernald & Long 18551 (NE); Provincetown, Greenman 3024 (GH), Fernald &
Long 18549 (AA, NE).

Connecticut: Salisbury, Blewitt 2037 (NE), Weatherby 4070, 4070a, 4070b
(NE); Sharon, Weatherby 3616 (NE); East Hartford, Weatherby 2018 (GH);
Suffield, Weatherby 5370 (NE); Stafford, Weatherby D2103 (NE); Hamden,
Blewitt 1796 (NE); Meriden, Blewitt 1795 (NE); Waterbury, Blewitt 1511, 1792,
1794 (NE); West Cheshire, Blewitt 2035 (NE).

Ruove Istanp: Barrington, May 30, 1911, Winslow (AA, NE); South Foster,
June 10, 1922, Eaton & Fassett (NE); Cumberland, Chamberlain 62 (NE).

New York: Stockholm, Phelps 1585 (GH); Canton, Phelps 1583, 1586 (GH) ;
Clare, Phelps 1584 (GH); Newcomb, House 7265 (GH); Watertown, House 8943
(GH); Forestport, Muenscher & Maguire 2322 (GH); Ledyard, Wiegand 6594
(MBG), 6603, 6587, 6593 (GH); Leroy, Hill 231895 (UI); Junius, Eames & Mac-
Daniels 4285 (GH); McKenney’s, Tompkins Co., May 1895, Wiegand (GH, TYPE
of A. humilis); Long Lake, House 10175 (GH); Hudson Falls, Burnham in 1897
(GH); Sand Lake, Wiegand 4290 (GH); Glenmont, House 17246 (MBG); Ron-
konkoma, July 5, 1908, Harper (GH); Babylon, Svenson 8012 (GH, MBG) ; Black
Rock Forest, Raup 8094 (GH); Cahoonzie, Muenscher, et al. 15609 (GH); Sulli-
van Hill, Chimung Co., Lucy 818b (GH); Monroe Co., Slavin 203 (AA).

New Jersey: Newport, June 2, 1894, Dill (AA); High Point, Mackenzie 4201
(GH) ; Chatsworth, Eames in 1894 (GH); Crowfoot, Fogg 4049 (GH, Ph) ; Char-
lotteburg, Mackenzie 3080 (GH); Elm, Fogg 1863 (Ph); Clementon, Long 20586
(Ph) ; Robbinsville, Long 51838 (Ph); Friesburg, Long 37315 (Ph); Mays Land-
ing, Pennell 12027 (Ph); Jacksons Mills, Long 52088, 52094 (Ph); Cookstown,
Long 30640 (Ph); Atsion, Long 25839 (Ph); New Gretna, Long 12504 (Ph);
Holmeson, Long 52017, 52022 (Ph); Farmingdale, Stone 12684, Brown 216 (Ph);
Spring Valley, Long 56467 (Ph).

PENNSYLVANIA: Wilkes Barre, Palmer 36296 (MBG); Easton, Porter in 1897
(GH); Fleetwood, Long 12556 (GH); Fairview, Wahl 47 (GH); State College,
Wahl 33, 73 (GH); Almont, Juiy 18, 1923, Pretzs (GH); Point Pleasant, Benner
in 1926 (GH); Beaver Meadows, Fogg 16324 (GH); Danielsville, Long 48649
(Ph); North Bangor, Long 51076 (Ph); Shimerville, Pretz 11248 (Ph) ; Schnecks-
ville, Pretz 12754 (Ph); Crackersport, Pretz 10767 (Ph); Germansville, Pretz
11762 (Ph); East Reading, Wilkens 471 (Ph); Schubert, Wilkens 5128 (Ph);
Friedensburg, Wilkens 5168 (Ph); Emilie, Benner 2926 (Ph); Bristol, Benner
7533 (Ph); Wakefield, Tanger 3065 (Ph); White Oak, Tanger 3043 (Ph) ; Spring
House, Long 32642 (Ph); Glen Riddle, Pennell 2708 (Ph).

West VIRGINIA: Ravenswood, Balser 775 (GH, MBG).

VirGINIA: Great Falls, Hunnewell 5895 (GH); Ashland, Wherry & Adams 2768
(GH) ; Loretto, Fernald & Long 14178 (Ph).

NortH CaroLina: French Broad River, June 4, 1918, Ashe (GH); Williamston,
Palmer 39808 (AA); Middlesex, Godfrey & White 7027 (GH, MBG); Raleigh,
Harbison 30 (AA); Bolton, Palmer 39833 (MBG, AA); Highlands, Harbison 9,
194, 7236, 7240 (AA), Magee in 1901 (GH).

56 ILLINOIS BIOLOGICAL MONOGRAPHS

SoutH CaroLina: Calhoun Falls, Harbison 1 (AA); Camden, Palmer 42391
(MBG).

ALABAMA: Alpine, Harbison 846 (AA); Auburn, Harbison 813 (AA).

Ontario: Thunder Bay, Pease 26323 (GH); Tobermory, Krotkov 7517 (GH) ;
Agawa Bay, Pease 18048 (GH); Britannia, M.-Victorin 15585 (AA); Dalhousie
Lake, Dunbar 10 (AA); Rockcliffe, Macoun 80733 (MBG, AA); Summerstown,
Jack in 1913 (AA); Jellicoe, Jennings 14521 (GH); Cornwall, Jack in 1914 (AA);
Belleville, Macoun in 1878 (GH); Niagara, Macoun 34298 (GH); Port Edward,
Macoun 34301 (GH); Sarnia, Dodge 57 (GH, AA).

Outro: Painesville, Werner in 1892 (GH) ; Columbus, Kellerman in 1903 (GH) ;
Georgesville, Werner 54 (GH).

Micuican: Whitefish Point, Fernald & Pease 3359 (GH); Cheboygan, Dodge
20 (GH); Burt Lake, Ehlers 1183 (GH, UI); Alpena, Wheeler in 1895 (GH);
Walhalla, Palmer 40470, 40471 (AA, MBG); Port Huron, Dodge 71, 73 (GH);
Portage Lake, Hermann 6497 (MBG), 6486 (GH).

InpIANA: Tolleston, Hill in 1894 (U1); Sheffield, Hill in 1876 (UI); Ham-
mond, 4. Chase 990 (Ph, UI); Dune Park, Hill in 1898, A. Chase 709 (UI) ; Mongo,
Deam 38196, 33770, 39083, 38194, 38195 (AA); Knox, Deam 38251 (AA); near
Rainesville, Deam 23107 (AA).

Irutnois: Sag Bridge, Hill in 1913 (UI); Lake Zurich, May 9, 1899, Hill (GH,
UI) ; Oregon, July 9, 1905, Hill (UI) ; Lombard, Moffatt 1610 (UI) ; Chicago, Hill
in 1890 (UI); Calumet, 4. Chase 702, 1745 (UI); Colehour, Hill in 1876 (UI);
Barrington, A. Chase 1048 (UI); Mississippi Palisades State Park, G. N. Jones
17143 (UI).

Wisconsin: Trout Lake, Fassett 13775 (GH); Cassian, Palmer 27795 (MBG),
27779, 27796 (AA); Marinette, Schuette in 1892 (GH); Dell Prairie, Fassett 2823
(GH); West Salem, Fassett, et al. 18359 (MBG); Dane Co., Hale in 1861 (GH,
MBG); Vermont, Fassett 2818 (GH) ; Holcombe, Fassett & Schmidt 15708 (GH) ;
Devils Lake, Fassett 2819, 2821 (GH); Delton, Fassett 2820 (GH); Barneveld,
Fassett 2822 (GH).

MinneEsoTA: Brule River, Aiton 1004 (UI); Itasca Park, Moyle 216 (MBG);
near Duluth, Lakela 2873 (MBG); Cass Lake, Pammel 52 (GH); Big Sandy Lake,
Rosendahl 4983 (GH), 4980 (AA); Perham, Rosendahl in 1926 (GH) ; Fergus Falls,
Blanchard in 1908 (AA); Center City, Taylor in 1892 (GH); Fort Snelling,
Mearns in 1891 (GH); Redwing, Sandberg 3 (UI).

NortH Dakota: Enderlin, Bergman 1376 (MBG).

SoutH Dakota: Big Stone Lake, Williams in 1894 (MBG); Warrens Woods,
Brookings Co., Thornber in 1893 (GH, MBG).

Iowa: Estherville, W/olden 1075, 1043, 1047, 1074, 1078 (GH); Armstrong,
Cratty in 1902 (MBG) ; Iowa Lake, Cratty in 1900 (MBG) ; Oak Grove State Park,
Hayden 10489 (MBG); Fayette, Fink in 1894 (GH); Ames, Pammel 14 (AA).

Mrissourt: Sedalia, Palmer 30009 (AA, MBG); Holberg, Steyermark 18635
(MBG).

The identity of Amelanchier spicata, based on Crataegus spicata Lam.,
the earliest name for the species under discussion, has long been a matter
of conjecture. Lamarck says his plant was growing in the Jardin du Roi,
and was supposed to be a native of Canada. Considering the close connec-
tion that existed between Canada and France in the eighteenth century,
there is no reason for supposing that it might have come from a more
southerly region. Several theories and speculations have been set forth in
an attempt to identify Lamarck’s plant, but the conclusion that has had the
widest influence on the choice of nomenclature is that of Wiegand, who,
in 1912, supposed it to be of hybrid origin, hence unsuitable as the type,

AMERICAN SPECIES OF AMELANCHIER—JONES o/

and therefore to be dismissed from further consideration for nomencla-
tural purposes. A few botanists, including Britton and Schneider, con-
tinued to use the name A. spicata for the common and widespread low
shrub of eastern North America with racemose flowers, tomentose ovary,
and usually finely toothed, few-veined leaves, but a number of others at-
tempted to apply in various senses half a dozen different names, the ma-
jority preferring to follow Wiegand and name their specimens either A.
humilis or A. stolonifera. |

The nomenclatural combination A. spicata was first made by Karl
Koch in 1869. Although there is some doubt whether his brief description
is applicable to Crataegus spicata Lam., and it is quite clear that the names
appended in synonymy belong elsewhere, the fact remains that he ex-
pressly cites Lamarck’s name as the basonym, hence Koch’s binomial is
the one to be used for this species.

In 1932, Professor Alfred Rehder of the Arnold Arboretum of
Harvard University obtained a photograph as well as a fragment of La-
marck’s type in the Museum d’Histoire Naturelle, Paris. This material, now
in the herbarium of the Arnold Arboretum, effectively dispels the mystery
that for a century and a half has surrounded the identity of Crataegus
spicata Lam. The photograph shows two specimens, presumably from the
same plant, one a short flowering branch, and the other a twig with ma-
ture leaves. (See Pl. XI, fig. 2.) The flowers are not quite open, but their
size and other characteristics are apparent from a microscopic examina-
tion of a fragment of the holotype that is also in the herbarium of Arnold
Arboretum. The leaf-specimen has nine leaves, some large and roundish,
others smaller and oval. The apices vary from shortly acutish to rounded
‘and mucronulate. The margins are finely serrate almost to the rounded
base. From an examination of this phototype, and of the accompanying
fragment of the type, it becomes clearly evident that Lamarck’s Crataegus
spicata is the common and widespread small serviceberry of eastern North
America with racemose small flowers, tomentose top of ovary, and usually
rather finely toothed, few-veined leaves. Recognizing the possibility that
the shrubs under consideration may comprise a somewhat polytypic
species, Wiegand in 1912 attempted to distinguish two different species
in this general cycle of affinity, chiefly on the basis of the shape, venation,
and indentation of the leaf-blades. Specimens with more coarsely toothed
and conspicuously veined blades were named A. humilis, while those with
finer indentation and fewer, less regular veins were called A. stolonifera.
Unfortunately, however, these characters are so extremely variable as
to render them practically useless. While it is true that the coarser toothed
specimens appear at first glance to be somewhat different from the finer
toothed ones, all intergradations may be found. There are no supporting
characters of flowers and fruits, or of geographical distribution. It is

58 ILLINOIS BIOLOGICAL MONOGRAPHS

perfectly evident from an examination of the types, as well as hundreds
of other specimens, including many identified and annotated by Wiegand,
that A. humilis and A. stolonifera belong to the same species, the oldest
name for which is A. spicata (Lam.) K. Koch. Even if one (or both)
of Wiegand’s proposals could be maintained as a specific entity, it would
have to be designated by one of the older names, A. erecta or A. saxatilis
of Blanchard, which were adequately published and well supported by
ample exsiccatae five years before Wiegand published A. humilis and
A. stolonifera. Although, as Wiegand has pointed out, Blanchard’s dis-
tributed specimens are a somewhat mixed lot, this fact does not exclude
his proposed species from consideration in the matter of nomenclatural
priority.

It may be of passing interest to note the historical fact that as a result
of an oversight A. humilis was published without mention of a type
specimen. Somewhat later a type was selected, but it was a specimen
collected by Wiegand in 1895, not one of those previously cited with the
original description. Unfortunately, this specimen, which closely resembles
the type of A. stolonifera, does not wholly agree with the original de-
scription and the statements in the key, which call for a plant with leaves
“coarsely dentate-serrate.”’ Although of no special importance now, the
subject is mentioned here as one of the factors which may have con-
tributed to the difficulty of accurately interpreting and delimiting the
species under discussion.

The true identity of a low, small-flowered shrub described in 1908 by
B. L. Robinson from Blue Hill, near Milton, Massachusetts, as A. ob-
longifolia var. micropetala has long been in doubt. Wiegand in 1912 dis-
cussed these small-flowered plants, and concluded that they are hybrids
between 4. canadensis and A. spicata, and designated them by the formula
A. oblongifolia X stolonifera. In 1916, after careful field studies, C. A.
Weatherby (Rhodora 18:48, 49) expressed the view that the reduced
petals may be due to the teratological condition known as staminody. In
1919, Ashe elevated Robinson’s variety to specific rank, on the view that
the plants could not be hybrids because of their general distribution and
local abundance. It now seems evident from a study of a series of speci-
mens in the Gray Herbarium and in the herbarium of the New England
Botanical Club that these plants are merely small-flowered representatives
of the common and widespread A. spicata (Lam.) K. Koch.

In 1918, W. W. Ashe described, as A. austro-montana, a “shrub not
exceeding 4 m. in height,’”’ based on specimens from the valley of French
Broad River, Transylvania County, in southwestern North Carolina. It
was said to have finely serrate leaves that are pubescent beneath when
young, but glabrous at maturity. The flowers are described as appearing
“largely” before the leaves in 7-10-flowered, nodding, pubescent racemes,

AMERICAN SPECIES OF AMELANCHIER—JONES 59

and the fruits are said to be 10-14 mm. thick, shining and almost black
when ripe, with the sepals erect or nearly so. Ashe commented that his
new species was “related to hwmulis in fruit characters.”” However, an iso-
type of A. austro-montana in the Gray Herbarium (see Pl. XVI) which
closely resembles the type of Wiegand’s A. stolonifera, does not show some
of the diagnostic characters of A. austro-montana, and in the shape, tex-
ture, serration, and indument of the leaves, the size, arrangement, and
character of the fruit, the direction of the sepals, the shape, size, and
direction of the flowering and fruiting racemes, as well as the structural
characters of the flowers themselves, can be matched almost exactly by
numerous specimens from nearly every part of the extensive geographi-
cal range of the common and variable A. spicata (Lam.) K. Koch.

10. AMELANCHIER SANGUINEA (Pursh) DC.
(Plates XVII, XVIII)

Mespilus canadensis var. Y rotundifolia Michaux, Fl. Bor. Am. 2:291 (1803).

Pyrus sanguinea Pursh, Fl. Am. Sept. 340 (1814); Bigelow, Fl. Bost. (ed. 2) 196
(1824) ; Sprengel, Syst. Veg. 2:509 (1825).

Aronia sanguinea Nuttall, Gen. Am. Pl. 306 (1818); Eaton, Man. Bot. N. Am.
(ed. 6) 29 (1833).

Amelanchier sanguinea DC. Prodr. 2:633 (1825); Spach, Hist. Nat. Veg. Phan.
2:86 (1834) ; Hooker, Fl. Bor. Am. 1:203 (1834) ex p.; Loudon, Arb. & Frut.
Brit. 2:875, figs. 630, 631 (1838); M. Roemer, Syn. Mon. 3:145 (1847) ; Britton
& Shafer, N. Am. Tr. 439, fig. 385 (1908) ; Wiegand in Rhodora 14; 138, pl. 95
(1912); Britton & Brown, Illustr. Fl. N. U.S. (ed. 2) 2:293, fig. 2332 (1913) ;
Rehder in Bailey, Stand. Cyclop. Hort. 272 (1914) ; Hoffman in Proc. Boston
Soc. Nat. Hist. 36:280 (1922); Wiegand & Eames, Fl. Cayuga Basin 247
(1926) ; Rehder, Man. Cult. Tr. & Shr. 389 (1927) ; Rosendahl & Butters, Tr.
& Shr. Minnesota 214 (1928); Peattie, Fl. Indiana Dunes 219 (1930) ; Rydberg,
Fl. Prairies & Plains 437 (1932); Small, Man. Se. Fl. 637 (1933); Marie-
Victorin, Fl. Laurent. 316, fig. 91 (1935); Nielsen in Am. Midl. Nat. 22:168,
pl. 3 (1939); Rehder, Man. Cult. Tr. & Shr. (ed. 2) 387 (1940).

Aronia latifolia Riddell, Suppl. Cat. Ohio Pl. 24 (1836).

Amelanchier canadensis var. Y rotundifolia Torrey & Gray, Fl. N. Am. 1:473 (1840),
Fl. New York 1:225 (1843) ; Walpers, Rep. Bot. Syst. 2:55 (1843); Gray, Man.
Bot. 130 (1848), (ed. 2) 126 (1856), (ed. 5) 162 (1868).

‘Amelanchier rotundifolia sensu M. Roem. Syn. Mon. 3:146 (1847); Britton &
Brown, Illustr. Fl. N. U.S. 2:238 fig. 1988 (1897); Small, Fl. Se. U.S. 532
(1903); Card in Bailey, Cyclop. Am. Hort. 57 (1904). Non (Lam.) Dum.-
Courset 1811.

Amelanchier ovalis var. willdenowiana M. Roem. l.c.

Amelanchier erecta Blanchard in Torreya 7:101 (1907), ex p.

Amelanchier spicata sensu Robinson & Fernald in Gray, New Man. Bot. (ed. 7)
460 (1908); Jones & Rand in Bull. Vermont Agr. Exp. Sta. 145:99, fig. 1,
(1909). Non Crataegus spicata Lam.

Amelanchier sanguinea f. grandiflora Wiegand in Rhodora 14:139 (1912).

Amelanchier sanguinea var. grandiflora Rehder in Bailey, Stand. Cyclop. Hort.
1:272 (1914).

Amelanchier grandiflora sensu Wiegand in Rhodora 22:149 (Oct. 29, 1920). Non
Rehder 1920 (Sept. 6).

Amelanchier huronensis Wiegand, op. cit., 150; Nielsen in Am. Midl. Nat. 22:170,
pl. 4 (1939).

60 - ILLINOIS BIOLOGICAL MONOGRAPHS

Amelanchier amabilis Wiegand in Rhodora 23:48 (1921) ; House in N.Y. State Mus.
Bull. 254:412 (1924) ; Wiegand & Eames, FI. Cayuga Basin 247 (1926) ; Rehder,
Man. Cult. Tr. & Shr. 389 (1927), (ed. 2) 378 (1940).

Straggling or arching slender shrubs 1-3 m. tall, the stems solitary, or
sometimes surculose and forming colonies; occasionally a small tree 4-6 m.
tall; young twigs reddish brown or grayish; winter buds conical, reddish
brown, dull, vernicose, or the scales ciliate, or sometimes pubescent on the
back; leaves commonly oval, varying to suborbicular, conduplicate in the
bud, unfolding before or with the flowers and usually about half-grown
or nearly full-grown at anthesis, densely whitish or somewhat yellowish
flocculent-tomentose beneath when young, tardily glabrous, or sometimes
the tomentum partially deciduous by flowering time, some of the leaves
sometimes retaining traces of tomentum at maturity, especially on the
veins beneath and on the petioles, but usually the fully mature leaves
nearly or quite glabrous on both surfaces; mature blades 2.5-7 cm. long,
2-5 cm. wide, the apex acutish or obtuse and more or less mucronate, the
base rounded subcordate; principal lateral veins of average leaves 11-13
pairs, conspicuous, rather close together, straight or nearly so, parallel,
the upper ones usually running straight to the margin and ending in the
teeth, the uppermost short and strongly curved and ascending; short
intermediate veins few or none; margins rather coarsely serrate-dentate
nearly or quite to the base with broad sharp spreading teeth, these usually
4-6 per cm., and about 20-30 on each side of average leaves of the fruiting
branches; petioles slender, 1-2 cm. long, glabrous, or remaining slightly
pubescent; flowers many, in ascending, spreading, or drooping 4-10-
flowered racemes 4-8 cm. long, the lower pedicels 6-18 mm. long; rachis
and pedicels pubescent; petals 5, white, or sometimes pinkish, oblanceo-
late, obtuse, those of the fully opened flowers 11-22 mm. long, 46 mm.
wide, more or less pilosulous on the base of the short claw; stamens about
20, the filaments glabrous; anthers 0.8-1.2 mm. long; hypanthium saucer-
shaped, 4-8 mm. in diameter, glabrous or pubescent outside, usually more
or less constricted on the very young fruit as a result of the neck of the
hypanthium being produced into a rim above the rounded summit of the
ovary; sepals triangular-lanceolate, acute, 3.5-4 mm. long, pubescent
within, usually recurved from the middle after anthesis; styles 5, glabrous,
2-3 mm. long, usually united only near the base; top of the ovary densely
tomentose; mature fruit globose or oblate-spheroidal, purplish-black,
glaucous, glabrous, 6-8 mm. in diameter, sweet, juicy, edible; pedicels 1-4
cm. long; seeds brown, smooth, obliquely lanceoloid, somewhat flattened,
about 5 mm. long and 2-3 mm. wide when well developed.

Type Locatiry: “In Canada. ..-.” Type in Michaux’s herbarium
Museum d’Histoire Naturelle, Paris. Phototype in the Gray Herbarium.

AMERICAN SPECIES.OF AMELANCHIER—JONES 61

RANGE: Woods and thickets, rocky bluffs, shores, hillsides, and ra-
vines, on various substrata, including limestone, sandstone, conglomerate,
gneiss, schist, and quartzite, from southern Quebec to Minnesota and
northern Iowa, southern Michigan, New York, and western Massachu-
setts; also in the mountains of western North Carolina. Flowering from
the beginning of May to the early part of June; fruits ripening in July
and August. Common names: round-leaved juneberry; shore shadbush.

@ A. sanguinea yy Ae fae a
ae 30
® A. fernaldii eT i JIS J

Map 8—Range of Amelanchier sanguinea and A. fernaldu.

Quesec: Isle Perrot, Jack 3928 (AA); Lake Massawippi, July 21, 1923, Knowl-
ton (GH); Aylmer, Rolland-Germain 19258 (GH) ; Chateaugay, May 28, 1901, Jack
(AA); Oka, M.-Victorin 1883 (AA), 18715 (GH); Bolton, July 25, 1926, Knowlton
(GH); Thetford, M.-Victorin 11216 (AA).

Marine: Winn, Fernald & Long 13780 (NE, AA); Orono, May 28, 1873, Scrib-
ner (NE); Veazie, June 23, 1905, Knight (UI); Milford, Fernald 13778, 13779
(GH, NE); Pembroke, Fernald 1880 (NE); Gilead, Furbish in 1897 (GH, NE);
Masardis, Sept. 8, 1897, Fernald (NE, GH), 2311 (GH, NE); Houlton, Fernald
in 1897 (GH, NE); Ashland, Fernald 2310 (GH); Fort Kent, Woodward &
Bissell in 1914 (NE); Sangerville, Fernald in 1897 (NE); Dover, Fernald 388
(GH, NE, AA).

New Hampsurre: Lebanon, Fernald, Hunnewell, & Blanchard in 1920 (NE) ;
Haverhill, Fernald 15537 (GH, NE); Bath, Pease 19794 (NE) ; Durham, Hodgdon
2998 (NE).

Vermont: Providence I., Lake Champlain, Eggleston 1178 (GH); Isle La
Motte, Cushman 871 (NE); Swanton, May 22, 1912, Knowlton (NE); Mt. Wil-
loughby, Hodgdon 2471 (NE); Canaan Falls, Eggleston 1121 (GH, UI, NE);
Charlotte, May 28, 1922, Knowlton (NE); Burlington, Pringle in 1879 (AA);

62 ILLINOIS BIOLOGICAL MONOGRAPHS

Middlebury, Brainerd in 1901 (GH) ; Williamstown, June 8, 1917, Knowlton (NE) ;
Wells, May 27, 1916, Knowlton (NE); W. Rutland, Eggleston 1127 (AA, NE),
1122 (GH); Hartland, May 8, 1921, Knowlton (NE), Eggleston 1971 (GH);
North Hartland, Drew, Hodgdon, & Taylor 2472 (NE); Hartford, Eaton & St.
John in 1920 (GH, NE); Cavendish, Fernald 449 (GH, NE); Bennington Co.,
Steyermark 7009 (MBG); Manchester, Fernald, Harris, Drew, et al. in 1932 (NE),
Cushman 4330 (MBG); Bellows Falls, May 10, 1915, Knowlton (NE); Townsend,
Wheeler in 1915 (NE); Westminster, Blanchard 4 (GH, AA).

MassacHusetTs: Sheffield, May 24, 1920, Hoffmann (NE); West Stockbridge,
Aug. 7, 1920, Hoffmann (NE); Shelburne Falls, Bean & Knowlton 12070E (NE) ;
Leverett, Tower & Seymour 3664 (NE).

New York: Canton, Phelps 568 (GH); Gouverneur, Phelps 1592 (GH);
Plattsburg, Hunnewell 4681 (GH); Depauville, Fernald, Wiegand, & Eames 14302
(GH); LeRoy, Wiegand 13976 (GH); Romulus, Eames 4287, 4288 (GH); Forest-
port, House 11201 (GH); Mt. McGregor, House 16059 (GH); Orebed Mt., June
27, 1918, Burnham (GH); Lake George, Kennedy in 1885 (GH); Lake George
Region, Aug. 10, 1916, Burnham (GH); Rochester, Wiegand 13979 (GH) ; Mendon,
Slavin 205 (AA); Ledyard, Wiegand 6589, 6592 (GH); Glenville, Svenson in 1924
& 1931 (GH); Sand Lake, Wiegand 4281 (GH) ; Middleburg, Svenson 7849 (GH) ;
Ithaca, Wiegand 2498 (GH, MBG), 2499, 2501 (GH), May 11, 1897 (GH, TYPE
of A. sanguinea f. grandiflora) ; Caroline, June 10, 1882, Dudley (AA); Dryden,
Wiegand 6582 (GH); Danby, Wiegand 2505 (GH); North Spencer, Wiegand &
Metcalf 6583 (GH).

New Jersey: Alpine, Mackenzie 5804 (Ph).

NortH CarRoLInaA: Craggy Mt., near Biltmore, Biltmore Herbarium 5664c (GH,
MBG, UI), 5664d (UI); Chimney Rock Mt., Biltmore Herbarium 5664e (MBG,
Wit, GH),

Ontario: Cloche Peninsula, Fernald & Pease 3364 (GH); Guelph, June 13,
1904, Klugh (GH); Port Franks, Lambton Co., May 26, 1904, Dodge (AA);
Thessalon, Pease & Bean 26215 (GH); Gravenhurst, Biltmore Herbarium 5664
(GH, AA); Bear Island, Krotkov 5390 (GH); Plevna, Aug. 8, 1902, Fowler (GH) ;
Nipigon, Pease & Bean 26502 (GH); Dalhousie Lake, Dunbar 12 (AA); Brock-
ville, Blanchard 4 (GH); Ottawa, Macoun 20074 (GH).

MicuicAn: West Bluff, Keweenaw Co., Fernald & Pease 3365, 3366, 3367
(GH); Copper Harbor, Hermann 7791 (MBG); Béte Grise, Fernald & Pease 3361
(GH); Gwinn, June 6, 1909, Harrison (GH); Mackinac I., July 4, 1912, Hunne-
well (GH), June 2, 1900, Schneck (UI), Sargent in 1911 (AA); Cheboygan Co.,
Gates 9544 (UI), 13914 (AA); Douglas Lake, Ehlers 323 (GH); Sand Point,
Dodge 74, 76 (GH, TYPE of A. huronensis); Sawyer, May 19, 1928, Manning
(UI); N. Manitou I., Wislizenus 937.

Wisconsin: Port Wing, Fassett 7313 (GH) ; Lake Owen, June 5, 1928, Griscom
(GH); Rhinelander, Palmer 28711 (MBG); Dyckesville, Palmer 28747 (MBG) ;
Ellison Bay, Palmer 28791 (MBG, AA); Jacksonport, Kraus 16 (GH) ; Milwaukee,
May 21, 1906, Hill (UI); Ephraim, Kraus 22 (GH); Keshena, Palmer 27740
(MBG); Port Washington, Palmer 28861 (MBG); Cedarburg, Palmer 28905
(MBG).

Minnesota: Wilmar, Palmer 36815 (AA, MBG); Falls of Minnehaha, July 16,
1886, L. H. Bailey (GH); Duluth, June 28, 1915, Stone (Ph) ; Lake of the Woods,
MacMillan & Sheldon 1747a (Minn.); Grand Portage, Rosendahl 6072 (Minn.) ;
Itasca Park, Nielsen 1967, 2500 (Minn.); Red Wing, Nielsen 1349 (Minn.).

Iowa: Estherville, Wolden 1353 (GH) ; Bluffton, July 18, 1933, Tolstead (Minn.).

The first mention of the species under discussion was made by
Michaux in 1803, who described a Mespilus canadensis var. rotundifolia
as “arborescens: foliis suborbiculato-ovalibus, utrinque rotundatis ...
in Canada.” The interpretation of Michaux’s plant is rendered quite clear

AMERICAN SPECIES OF AMELANCHIER—JONES 63

by an examination of a photograph of the type specimen in the Michaux
herbarium (Mus. Hist. Nat. Paris), which, “though not conclusive, seems
more like the large-flowered, coarse-toothed species than any other; and
this interpretation seems to have been that reached by other recent
botanists who have studied the type. The Michaux variety was raised to
specific rank and transferred to Amelanchier by Roemer in 1847. Un-
fortunately in 1814 Pursh described a Pyrus sanguinea. There has always
been doubt as to the identity of this plant; and, so far as known, no speci-
men is in existence. Pursh cites as the only synonym the M/. canadensis
y rotundifolia of Michaux, but he further says ‘tenuissime serratis .. .
A-small tree ... berries red...’ which are not the characters of
Michaux’s plant. Moreover the locality is given as, ‘In Canada and on the
banks of the Columbia.’ Since that time attempts have been made to apply
Pursh’s name, but in a most diverse manner (see Lindley in Bot. Register
t. 1171, and Loudon, Arb. et Frut. p. 875). Evidently Pursh confused at
least two plants, and evidently the only thing definite in connection with
the Pursh name is the Michaux synonym, but that is definite, and the
name need not be a source of confusion; therefore the writer is inclined
to believe, with Dr. Britton, that Pursh’s specific name should be retained
for Michaux’s plant. Even if Pursh’s name is not used, the name rotundi-
folia can not be used as a specific name for our plant, as it is an earlier
valid name’ for the native European species.” (Wiegand, in Rhodora
14125, 1912).

The phototype (Plate XVII) of Michaux’s plants depicts two fruiting
specimens, almost exactly alike, mounted over three labels. On one label
there is the statement “Lac Champlain,” on the second, “hab. in Canada,”
and on the third label, “Lac Mistassin.” Therefore, it is evident that
Michaux’s specimens came from the province of Quebec, or the northern
part of what is now the state of New York, or adjacent Vermont. Clari-
fication of the understanding of Michaux’s type is contributed by a pencil
tracing in the Gray Herbarium. This strengthens the conclusion of Wie-
gand and others that Michaux had a specimen of the taller, straggling
shrub with coarsely toothed leaves and longer fruiting pedicels that we
now call A. sanguinea (Pursh) DC.

Occasional specimens with larger flowers, and looser, often drooping
racemes, have been named A. grandiflora Wieg., A. huronensis Wieg.,
and A. amabilis Wieg., or they have been treated as a variety or form of
A. sanguinea. Since there are no evident supplementary characters of
foliage or fruit, or even any differences of geographical distribution, these
specimens are here considered to be well within the normal range of
variation found in A. sanguinea.

_ Not valid according to the current International Rules (1935) because Crataegus rotundi-
folia Lam. is an illegitimate substitute-name for Mespilus Amelanchier L. The valid name for
this European species is therefore Amelanchier ovalis Medic.

64 ILLINOIS BIOLOGICAL MONOGRAPHS

11. AMELANCHIER GASPENSIS (Wieg.) Fern. & Weatherby
(Plate XIX)

Amelanchier sanguinea sensu Lindley in Bot. Reg., pl. 1171 (1828). Non DeCandolle,
1825.
ae sanguinea var. gaspensis Wiegand in Rhodora 14:139 (1912).
Amelanchier florida sensu Wiegand in Rhodora 14:143, pl. 95 (1912); Rosendahl
& Butters, Tr. & Shr. Minnesota 216 (1928). Non Lindley, 1833.
Amelanchier gaspensis (Wiegand) Fernald & Weatherby in Rhodora 33:235 (1931).
Low, much-branched shrubs 30-90 cm. tall, often forming dense
thickets; bark grayish or brownish; winter buds conical, acute, glabrous
or more or less pubescent; leaves commonly oval, varying to suborbicular,
conduplicate in the bud, unfolding before the flowers and usually nearly
or quite full-grown at anthesis, glabrous or quickly glabrate from the
first; mature blades relatively thin, 3-6 cm. long, 1.5-4 cm. wide, the apex
usually rounded or subtruncate, or somewhat acutish, the base cordate or
subcordate, or sometimes rounded, quite glabrous on both surfaces, pale
green beneath; primary lateral veins of average leaves 6-13 pairs, prom-
inent, nearly equally distant, curved upward, the upper ones usually
extending to the margin and ending in the teeth, the others anastomosing
at their tips; margins dentate-serrate to below the middle (varying to
subentire) with broad sharp ascending teeth, these usually 3-6 per cm.,
and about 5-20 on each side of average leaves of the fruiting branches;
petioles slender, 1-2.5 (-3) cm. long, glabrous or quickly glabrate; flowers
small, in ascending or erect, 5-15-flowered racemes 3-6 cm. long, the lower
pedicels 1-2 cm. long; rachis and pedicels glabrous or barely pilose; petals
5, white, oblanceolate, glabrous throughout, 6-9 mm. long, 2-3.5 mm. wide;
stamens 20, the filaments glabrous; anthers 0.6-0.9 mm. long; hypanthium
saucer-shaped, 3-4 mm. in diameter, essentially glabrous outside, more or
less constricted on the young fruit as a result of the neck of the hypan-
thium being produced into a rim above the rounded summit of the ovary;
sepals lanceolate, acutish or acuminate, 1.5-3 mm. long, occasionally
merely divaricate but usually recurved from the middle after anthesis,
permanently glabrous on both sides, or with a small tuft of tomentum
near the tip; styles 5, glabrous, about 2 mm. long, usually united to the
middle; top of the ovary tomentose; mature fruit globose or subglobose,
purplish black, glaucous, glabrous, 8-10 mm. in diameter when fully
mature; lower pedicels 1-2 cm. long; seeds brown, smooth, obliquely
lanceoloid, somewhat flattened, 4-5 mm. long, 2-3 mm. wide.
Type Locarity: Mouth of the Bonaventure River, Bonaventure Co.,
Gaspé Peninsula, Quebec, Canada. Type in the Gray Herbarium; isotype
in the herbarium of the Arnold Arboretum.

AMERICAN SPECIES OF AMELANCHIER—JONES 65

RANGE: On cliffs, ledges, gravelly beaches, talus, or in alluvial woods,
Gaspé Peninsula, and neighboring counties of Quebec; extending north-
ward and westward to James Bay, and the region about Lake Superior.
Flowering from July to the middle of August; fruits ripe in August and
September.

Quesec: Grand River, Fernald in 1904, Richards in 1903 (GH); Percé Mt.,
Collins, Fernald, & Pease in 1904 (GH); Mt. St. Pierre, M.-Victorin, et al. 33203
(AA, GH), Fernald, Weatherby, & Stebbins 2451 (GH); Anse Pleureuse, M.-
Victorin, et al. 33440 (GH); Coin-du-Banc, M.-Victorin, et al. 17434 (GH, AA);
Riviere York, M.-Victorin, et al. 17431 (GH, AA); Mont Louis, M.-Victorin 28582
(AA, GH); Grande-Coupe, M.-Victorin, et al. 17435 (AA, GH); Cape Rosier,
Pease 20216 (GH); Lac Pleureuse, Fernald, Dodge, & Smith 25840 (GH); Mt.
Ste. Anne, Williams, Collins, & Fernald in 1905 (GH); Riviere St. Anne des
Monts, Fernald, et al. 25839 (GH, AA), Collins & Fernald in 1905 (GH); mouth
of Bonaventure River, Williams & Fernald in 1902 (TYPE, GH; AA); Maria,
M.-Victorin, et al. 33298 (GH); Bonaventure River, Collins, Fernald, & Pease
in 1904 (GH); Matapedia, M.-Victorin 28694 (GH); Sainte-Flavie, Rousseau 24537,
24554 (GH); Matane, Forbes in 1904 (GH); Milnikek, Rousseau 32418 (AA);
Cap Enrage, Rousseau 26478, 26672 (GH), 26513 (AA); Bic, Rousseau 26259
(GH), 26241 (AA, GH), Collins & Fernald in 1904 (GH), Fernald & Pease 25137
(GH), Louis-Marie, et al. 34432 (GH); Montmagny, Rousseau 24561 (GH); Ile a
Deux Tétes, M.-Victorin 24538, 24558 (GH, AA); L’Islet, Rousseau 24550 (GH) ;
Charlton I., James Bay, Potter 484 (GH); 10 miles south of East Main, Potter
485 (GH).

Ontario: 50 miles north of Jackfish, Jennings 14024c (GH); Twin Islands,
Temagami Region, Anderson & Anderson 26042 (GH); Batchawana Bay, Pease
& Ogden 25152 (GH); Awrey, Sudbury District, Fernald & Pease 3369 (GH).

MicuicaAN: Keweenaw Co., May and July 1889, Farwell (GH); Alcona Co.,
Bailey 56 (GH); Carp Lake, Ontonagon Co., Pease & Ogden 24884 (GH); rocky
shore of L. Superior, Marquette, Aug. 2, 1889, Hill (UI); Isle Royale, Cooper 122,
124, 125 (GH).

This species is related to A. sanguinea (Pursh) DC., and the cordil-
leran A. alnifolia Nutt. From the former it differs in its “glabrous or
quickly glabrate foliage, its leaves more commonly rounded or subtrun-
cate at summit, its fewer nerves with anastomosing tips, its erect racemes
with glabrous or barely pilose rachis and pedicels, its shorter and glabrous
or promptly glabrate sepals, and its shorter petals, and occupying a clearly
circumscribed area northeast of the range of A. sanguinea, A. gaspensis
seems to be quite as definite a species as any in the group.” (Fernald &
Weatherby, l.c.). From A. alnifolia it may be distinguished in the fruiting
condition by its glabrous somewhat longer sepals, the “thinner leaves with
at most pale green lower surfaces, the cordilleran series having the coria-
ceous leaves glaucous beneath,” and its different habit of growth, and
smaller fruits, as well as the wholly distinct geographical range. Although
first described as a variety of A. sanguinea, it appears to be probably as

66 ILLINOIS BIOLOGICAL MONOGRAPHS

closely related to A. alnifolia. It seems best, therefore, to follow Fernald
& Weatherby in treating the Gaspé plant as a distinct species.

Amelanchier gaspensis was apparently first illustrated (as A. san-
guinea) in 1828 by Lindley in the Botanical Register, pl. 1171, on the
basis of specimens grown in the gardens of the Royal Horticultural
Society, but which came originally from near the southern end of James
Bay, Ontario, “whence living plants were sent . . . by William Williams,
Esq., Governor of Moose Factory, in 1824.” From the beautiful colored
plate and the accompanying description, it is evident that the plant de-
scribed and illustrated is that which is now known as Amelanchier gas-
pensis. The description contains the following statements: “A handsome
hardy shrub, resembling the Snowy Mespilus [1.e., probably A. arborea]
in general appearance, but distinguished from that, and all other species
of the genus, by its young leaves being perfectly destitute of pubescence.”
(Italics mine). A. gaspensis is the only known species of Amelanchier
that grows in the region indicated and fits the description and illustration.
The only discrepancies in the plate are that the leaves are shown as being
slightly more pointed than is usual in A. gaspensis, and the flowers have
glabrous ovaries, and only 3 or 4 styles. However, these peculiarities can
be safely attributed to the merely impressionistic tendencies of the artist,
because all the eastern American species have 5 styles, and in the
description Lindley says that the calyces are “internally woolly,” and the
leaves are “obtuse at each end.”

During the time since A. gaspensis was first recognized as a distinct
entity by Wiegand in 1912, additional collections and field studies have
made possible a considerable extension of the known geographical range
of this species, which was at first thought to include only the Gaspé
Peninsula of Quebec. The plant is now known to range westward to the
region about Lake Superior. Collections of flowering specimens from Isle
Royale, and others from northern Michigan, match almost exactly those
from the type locality on the Gaspé Peninsula. They have the same kind
of inflorescence, with the short pedicels, the glabrous and somewhat
glaucous calyces, and the short petals. The obtuse leaves are fully grown
and nearly or quite glabrous at flowering time, and their margins are
toothed chiefly above the middle or toward the apex. In the light of
subsequent discoveries it is now easily seen why Wiegand, and others,
quite unable to match this material with any available specimens of the
two related and somewhat similar species of the region, e.g., A. spicata
and A. sanguinea, and apparently not suspecting that the plants belonged
to A. gaspensis, arrived at the erroneous conclusion that their plants were
identical with A. florida of the Pacific Slope of North America, whose
natural geographical area is not closer than fifteen hundred miles to the
westward.

AMERICAN SPECIES OF AMELANCHIER—JONES 67

12. AMELANCHIER ALNIFOLIA Nutt.
(Plate II)

Aroma alnifolia Nuttall, Gen. N. Am. Pl. 306 (1818).

Pyrus alnifolia Sprengel, Syst. Veg. 2:509 (1825).

Amelanchier alnifolia Nuttall in Journ. Acad. Nat. Sci. Philadelphia 7:22 (1834) ;
M. Roem. Syn. Mon. 3:147 (1847); Decaisne in Nouv. Arch. Mus. Hist. Nat.
Paris 10:135 (1874) ; Coulter, Man. Bot. Rocky Mt. Reg. 89 (1885) ; Watson in
Garden & Forest 1:185, fig. 34 (1888) ; Watson & Coulter, in Gray, Man. (ed.
6) 167 (1889) ; Sargent, Silva N. Am. 4:131, pl. 196 (1892) ; Britton & Brown,
Illustr. Fl. N. U.S. 2:239, fig. 1989 (1897); Koehne in Gartenflora 51:610,
fig. 126 (1902) ; Card in Bailey, Cyclop. Am. Hort. 57, fig. 78 (1904) ; Sargent,
Man. Tr. N. Am. 363, fig. 285 (1905) ; Rydberg, Fl. Colo. 191 (1906) ; Schneider,
Illustr. Handb. Laubh. 1:738, figs. 411, 412 (1906); Britton & Shafer, N. Am.
Tr. 440, fig. 387 (1908); Coulter & Nelson, New Man. Rocky Mt. Bot. 266
(1909) ; Apgar, Ornam. Shr. U.S. 182, fig. 278 (1910) ; Britton & Brown, Illustr.
Fl. N. U.S. (ed. 2) 2:293, fig. 2333 (1913); Bean, Tr. & Shr. Brit. Isles 1:186
(1914); Rehder in Bailey, Stand. Cyclop. Hort. 272, fig. 187 (1914); Arm-
strong, Field Book W. Wild Fl. 217 (1915); Rydberg, Fl. Rocky Mts. 447
(1917) ; Standley in Contr. U.S. Nat. Herb. 22:366 (1921); Bailey, Man. Cult.
Pl. 378 (1924); Tidestrom, in Contr. U.S. Nat. Herb. 25:283 (1925) ; Rehder,
Man. Cult. Tr. & Shr. 389 (1927); Kirkwood, N. Rocky Mt. Tr. & Shr. 188,
fig. 38 (1930); Rydberg, Fl. Prairies & Plains 437, fig. 290 (1932); Garrett,
Spr. Fl. Wasatch Reg. (ed. 5) 106 (1936); Nielsen in Am. Midl. Nat. 22:164,
208, pl. 1. (1939); Preston, Rocky Mt. Tr. 175 (1940); Rehder, Man. Cult.
Tr. & Shr. (ed. 2): 387 (1940).

Amelanchier ovalis sensu Hooker, Fl. Bor. Am. 1:202 (1834) ; Loudon, Arb. & Frut.
Brit. 2:876 (1838), ex p. Non D.C., 1825.

Amelanchier canadensis var. 6 alnifolia Torrey & Gray, Fl. N. Am. 1:473 (1840) ;
Walpers, Rep. Bot. Syst. 2:55 (1843) ; Porter & Coulter, Syn. Fl. Col. 38 (1874).

Amelanchier montana Hort. ex Hand-list Trees Kew 1:217 (1894).

Amelanchier parvifolia Hort. ex ibid.

Amelanchier lanulosa Greene, MS.

Amelanchier cuneata Piper in Bull. Torr. Club. 27:392 (1900), Contr. U.S. Nat.
Herb. 11:346 (1906) ; Schneider, Illustr. Handb. Laubh. 1:739 (1906)

Amelanchier florida sensu Piper, Contr. U.S. Nat. Herb. 11:346 (1906), ex p.;
Piper & Beattie, Fl. Se. Wash. & Adj. Idaho 133 (1914); Henshaw, Wild FI.
N. Am. Mts. 154, pl. 30 (1914) ; Sargent, Man. Tr. N. Am. (ed 2) 396, fig. 352
(1922); Raup in Contr. Arnold Arb. 6:174 (1934), Nat. Mus. Canada Bull.
74:142 (1935); Garrett, Spr. Fl. Wasatch Reg. (ed. 5) 106 (1936); St. John,
Fl. Se. Wash. & Adj. Idaho 194 (1937); Graham, Ann. Carnegie Mus. 26:231
(1937). Non Lindley 1833.

Amelanchier utahensis sensu Piper in Contr. U.S. Nat. Herb. 11:346 (1906). Non
Koehne 1890.

Amelanchier alnifolia var. typica Schneider, Illustr. Handb. Laubh. 1:739, figs. 411,
412 (1906) ; Nielsen in Am. Midl. Nat. 22:167 (1939).

Amelanchier alnifolia var. pumila Schneider, Illustr. Handb. Laubh. 1:739 (1906),
as to name only; Rehder in Bailey, Stand. Cyclop. Hort. 273 (1914).

Amelanchier macrocarpa Lunell in Am. Midl. Nat. 3:143 (1913); Rydberg, FI.
Prairies & Plains 437 (1932).

Amelanchier leptodendron Lunell in op. cit. 5:237 (1918).

Amelanchier carrii Rydberg in Brittonia 1:89 (1931), Fl. Prairies & Plains 437
(1932).

Amelanchier humilis sensu Raup in Journ. Arnold Arboretum 17:264 (1936). Non
Wiegand 1912.

Amelanchier alnifolia £. alba Nielsen in Am. Midl. Nat. 22:167 (1939).

Amelanchier alnifolia var. dakotensis Nielsen in ibid.

68 ILLINOIS BIOLOGICAL MONOGRAPHS

Shrubs or small trees 2-4 m. tall, or in exposed situations sometimes
dwarfed and prostrate; bark smooth, dark gray on the older branches,
reddish brown on the branchlets; twigs of the season more or less silky-
pubescent, soon becoming glabrous; winter-buds conical, villosulous, acute,
dark brown, 3-6 mm. long; leaves of firm texture, oval or usually sub-
orbicular or almost quadrangular, mostly flat, unfolded and more than
half-grown at flowering time, tomentose beneath when young, soon be-
coming glabrous, usually by the time the flowers are fully expanded;
upper surface dark green, smooth, quite glabrous, the lower surface pale
or glaucous, glabrous throughout, or slightly pilosulous on the lower part
of the midvein; blades 2-5 cm. long, 1.5-4 cm. wide, almost always ob-
tuse, rounded, or truncate at the apex, rounded, truncate, or subcordate
at the base, rarely somewhat tapering; lateral veins 8-13 pairs, conspicu-
ous, parallel, curving and often forked and anastomosing near the margin,
or running into the teeth, the intermediate veins none or inconspicu-
ous; margins coarsely serrate or dentate to the middle, the lower half or
third of the blade usually quite entire; sometimes the blade entire through-
out or with 1 or 2 small teeth at apex; teeth usually coarse, rigid, ovate,
acuminate, somewhat incurved, 1-3 mm. long on average leaves of the
fruiting branches, 2-5 per cm., 2-20 on each side of average leaves; stip-
ules linear, 6-18 mm. long, villous, soon deciduous; petioles 8-18 mm.
long, pilose when young, soon becoming glabrous; flowers white, fragrant,
conspicuous; racemes erect, 3-6 cm. long, 5-15-flowered, short and rather
dense, the rachis and pedicels conspicuously lanate, the lower pedicels 5-15
mm. long; petals 5, white, oblanceolate, or oval, obtuse, 6-10 mm. long, 2-3
mm. wide, more or less ciliolate and with a small tuft of hairs on the base
of the very short claw; stamens about 20, 1-2 mm. long, the filaments
glabrous; anthers 0.6-0.9 mm. long; hypanthium shallowly campanulate,
3.5-4 mm. in diameter, the outside floccose at first but soon glabrous;
sepals deltoid-lanceolate to lanceolate, 1.5-3 mm. long, more or less pilose
within, becoming reflexed in age; styles 5, rarely 4, united below, 1.5-2.5
(-3) mm. long; summit of ovary persistently tomentose; fruit globose
to obpyriform, 10-15 mm. in diameter, normally 10-loculed, and 10-seeded,
purple or nearly black when ripe, glaucous, glabrous, edible, usually sweet
and juicy, but the flavor rather insipid; fruiting pedicels glabrous, 5-15
mm. long; seeds asymmetrical, oval, flattened, brown, smooth, 4-5 mm.
long, 3 mm. wide when fully developed.

Type Locatity: “In ravines and on the elevated margins of small
streams from Fort Mandan [North Dakota] to the Northern Andes
[Rocky Mountains].” Collected by Thomas Nuttall.

AMERICAN SPECIES OF AMELANCHIER—JONES 69

RANGE: Common along streams and on moist hillsides, in woods or
thickets, or on open slopes in canyons and on mountainsides, from Yukon
to Manitoba, southward to Nebraska, Colorado, and eastern Oregon;
flowering from the middle of May to the middle of July, according to
the altitude and latitude; fruit maturing in July and August. Common
names: Western shadbush, saskatoon, serviceberry.

YuxKon: Dawson, Eastwood
269 (AA), Kusche in 1916 (GH) ;
Lewis River, Gorman 1026 (ND).

BritisH Cotumpia: Dawson
Creek; Raup & Abbe 3500, 3502,
3530 (AA); Field, Ulke S35
(AA); Kelowna, Murie 1183
(MBG); Emerald Lake, Pease
22360 (G).

WASHINGTON: Klickitat Co.,
Suksdorf 10025, 10026 (AA, UI),
10129 (AA), 10154 (GH, AA),
10234, 10247 (AA), 11841, 11859
(U1) ; Pullman, Piper 1534 (AA),
Jack 1213 (AA), G. N. Jones 1397
(GH, UI), 2057 (UI), Beattie
1819 (MBG) ; Yakima Co., Cotton
576, 571, 569, 365 (MBG); Spo-
kane Co., Suksdorf 8597 (AA),
8609 (AA, UI), 8585 (UI), Sand-
berg & Leiberg 94 (MBG),
Palmer 37835 (MBG), Jack 1453
(AA); Cle Elum, Palmer 37858
(AA); Easton, G. N. Jones 4681
(UI); Okanogan Co., Thompson
7073 (MBG); Kahlotus, Rollins,
Dillon, & Pickett 868 (MBG),
Constance & McMurray 1135
(MBG); Bishop, Constance &
Rollins 1510 (GH).

OrEGON: Strawberry Mts.,
Applegate 6219 (AA), Ferris &
Duthie 784 (AA); Canyon City,
Brown 78 (AA, MBG); Des- Map 9.—Range of Amelanchier alnifolia.
chutes River, E. Nelson 807
(MBG); Tumalo Creek, Whited
571 (MBG); Lake Co., Ferris & Duthie 411 (AA); Baker, M. E. Jones 25397
(MBG); Blue Mts., A. Nelson & Ruth Nelson 783 (MBG).

ALBERTA: Wood Buffalo Park, Raup 2645, 2647, 2648, 2650, 2652, 2053, 2654,
2656 (GH); Fort Chipewyan, Raup 6065 (GH); Fort McMurray, Raup 7078, 7089
(GH); Lake Athabaska, Raup & Abbe 4466, 4515, 4519 (GH); Waterways, Raup
2649 (GH); Jasper Park, Jack 2540, 2629, 2778 (AA); Lake Louise, Hunnewell
6169 (GH); Banff, Aug. 10, 1904, Jack (AA), Brown 23 (GH, MBG), 56 (GH);
Butters & Rosendahl 1358 (GH), Barber 90 (GH); Rosedale, Moodie 1039 (GH) ;
Calgary, Moodie 817 (GH, MBG), Barber 194 (GH).

70 ILLINOIS BIOLOGICAL MONOGRAPHS

SASKATCHEWAN: Lake Athabaska, Raup 6670, 6588, 6931, 6084, 6933 (GH);
Prince Albert, July 10, 1896, Macoun (GH); Moose Jaw, Cowles 38 (MBG, UI);
Rosthern, Munroe in 1914 (AA); Ashcroft, Cowles 38a (MBG, UI); McKague,
Breitung 523 (MBG) ; Saskatoon, July 14, 1913, Sargent (AA).

Manitospa: Winnipeg, July 22, 1913, Sargent (AA); West Selkirk, Macoun
12627 (ND).

Montana: Seeley Lake, Marsh 507 (MBG); Bigfork, Clemens in 1908 (AA);
Coram, Jack 1580 (AA); Madison River, Scribner in 1883 (AA); Wolf Creek,
Palmer 36967 (AA, MBG); Great Falls, Palmer 36950 (MBG); Bozeman, W. W.
Jones in 1901 (GH), Blankinship 135 (MBG); Gallatin Co., Suksdorf 52, 841
(AA); near Missoula, MacDougal 178 (GH), Hitchcock 2290 (MBG); near
Bonita, Muenscher 11427, 11485 (MBG); Glacier Nat. Park, Jack 1497, 1498, 1511,
1537, 1592, 2007, 2293 (AA); St. Mary’s Lake, Hunnewell 2445 (GH), G. N. Jones
5640 (UI).

IpanHo: Priest Lake, Sargent in 1896 (AA); near Moscow, Jack 1258 (AA),
G. N. Jones in 1928 (UI); Hatwai Creek, Sandberg, MacDougal, & Heller 26
(GH, AA); Lewiston, Heller 3061 (MBG); Inkom, Jack 1155 (AA); Bovill,
Jack 1359, 1374 (AA); Helmer, Jack 1361 (AA); Elk River, Jack 1336, 1379 (AA) ;
Coeur d’Alene Mts., Leiberg 1203 (AA); Tamarack, Clark 166 (MBG); Mont-
pelier, Nelson & Macbride 1052 (MBG); Salubria, M. E. Jones 6274 (MBG);
Payette National Forest, G. N. Jones 5084 (GH); Salmon, Payson & Payson 1812
(MBG); Ashton, Nelson 10086 (MBG), Cronquist & Davis 2099 (MBG); Silver
City, Macbride 926 (MBG) ; Deadwood Creek, Nelson & Macbride 1847 (MBG);
Alturas Lake, Cronquist 3778 (MBG); Ketchum, Nelson & Macbride 1273 (MBG) ;
Pocatello, Jack 1185, 1125, 1124a (AA), Cronquist 2298 (MBG), Nelson & Mac-
bride 1405 (GH, MBG) ; Lava, Nelson & Macbride 1593 (MBG).

Wyominc: Yellowstone Nat. Park, Jack in 1904 (AA), Sargent in 1896 (AA),
Dewart in 1889 (MBG, UI); Teton Mts., Merrill & Wilcox 1027a (GH); Moose,
Wiliams 1106 (MBG); Upper Prairie Dog Creek, Rollins 550 (GH, MBG);
Little Tongue River Canyon, Williams & Williams 3305 (GH, MBG); Cow Creek,
Nelson & Nelson 798 (MBG); Laramie, Jack 1051 (AA); Dayton, Jack in 1900
(AA); Bighorn, Jack in 1900 (AA); Sundance Creek, Nelson 2129 (AA); Hulett,
Ownbey 593 (MBG); Sherman, Letterman in 1884 (GH); Afton, Payson &
Armstrong 3272 (MBG, GH, UI).

CoLtorapo: Ridgway, Payson 1075 (MBG) ; Silvercliff, Horner in 1898 (GH);
San Juan Mts., Wolf 3075 (GH); Canyon City, Nelson 10530 (MBG); Tolland,
Palmer 31358 (AA, MBG); Cedaredge, Payson 1069 (MBG); Pandora, Baker 750
(MBG); Estes Park, Zobel in 1935 (MBG); Shoshone, Eggleston 14967 (AA);
Empire, Engelmann in 1874 (MBG); Sunset, Andrews in 1920 (AA); Boulder,
Cockerell in 1906 (AA), Hanson C225 (MBG).

UtaH: Logan River, Cronquist 920 (MBG); Horse Creek, Graham 9262
(MBG); Mt. Timpanogos, Garrett 3686 (AA); Timpanogos Canyon, Palmer
38097 (MBG); Eagle Creek, Harrison & Larsen 7877 (MBG); Bromide Peak,
Harrison 7473 (MBG); Thistle, Eastwood 7706 (AA); City Creek Canyon, M. E.
Jones 1447 (AA); Providence Canyon, Muenscher & Maguire 2360 (MBG).

Manitoza: Portage la Prairie, Herriot 72329 (GH).

NortH Dakota: Devil’s Lake, Palmer 36902 (AA), 36859 (MBG), Lunell in
1909 & 1910 (UI); Larimore, Palmer 36855 (AA, Ph); Butte, Lunell in 1907 and
1911 (AA, TYPE coll. of A. macrocarpa); Dunseith, Lunell in 1918 (TYPE coll.
of A. leptodendron) (GH, MBG, UI); Stump Lake, Stevens & Graves 278 (GH) ;
Pleasant Lake, Lunell in 1901 (GH); Leeds, Lunell in 1904 (UI).

SoutH Daxota: Rosebud, Williams, sn. (AA); Swan Creek, Visher 2308
(MBG); Timber Lake, Shantz 345 (UI); Deadwood, Carr 75 (TYPE coll. of A.
carrii) (GH, MBG), Palmer 37197, 37343 (MBG, AA); Redfern, Murdoch 4035
(GH); Elk Canyon, Rydberg 680 (GH); Harney Peak, Palmer 37398 (MBG,
AA); Piedmont, Palmer 37048 (MBG).

AMERICAN SPECIES OF AMELANCHIER—JONES 7A

NEBRASKA: Chadron, Pool & Folsom in 1912 (MBG); Harrison, Bates 6074
(AA); Kirkwood, Bates 1357 (AA); Valentine, Bates 5928 (AA); Johnstown,
Bates in 1898 (AA); Scottsbluff Co., Aug. 13, 1901, H. P. Baker (MBG).

The leaves of Amelanchier alnifolia are usually quite free from pu-
bescence at maturity, and are dark green above and more or less glaucous
beneath. The top of the ovary is densely tomentose, and the number of
styles is uniformly five; the petals vary from 7-10 mm. in length. It is
a small-flowered species, and therefore distinguished at once from the
larger-flowered A. cusickii, which has an overlapping range, and from
A. florida, which occupies a quite separate geographical area to the
westward.

There has been a certain amount of ambiguity concerning the identity
of A. alnifolia Nutt., caused largely, it is to be supposed, by the absence
of Nuttall’s original specimens, and on account of confusion with other
species, particularly A. florida and A. cusickii. The original material
of Nuttall was, according to his description, collected between Fort Man-
dan and the “Northern Andes.” In his “Travels and Scientific Collections
of Thomas Nuttall,’ Dr. F. W. Pennell (Bartonia, vol. 18, pp. 15, 16,
1936) comments as follows on this particular part of the Nuttall itinerary:

Fort Mandan, near the villages of the Mandan Indians, was not at the present
town of Mandan, North Dakota, but was situated on the north side of the Missouri
River in the present McLean County, and almost directly opposite the later Fort
Clark. ... All the way from the Platte River to Fort Mandan Nuttall has been
telling of plants that occur on “to the Mountains” or “to the Northern Andes.”
These terms are evidently synonymous, and the mountains indicated lie farther up
the Missouri River, but they can not be the actual Rocky Mountains for Nuttall
cites no more tributary rivers above Knife River and he could not conceivably
have crossed what is now Montana to the ranges known to Lewis and Clark... .
I suppose that Nuttall must have ascended the Missouri through this rough country,
until he came to where the plain that had seemed illimitable from the top of the
river’s bluffs at last found an end in real hills and mountains. It was no more
than the northern extension of the Bad Lands and certainly could make but feeble
claim to the imposing term “Northern Andes,” but Nuttall evidently viewed it as
an outlier of a great western mountain-system, continuous through both Americas.
In reality, it did form an obvious limit to the ranges of the species he had been
seeing.

It appears, therefore, that the type locality of Amelanchier alnifolia is
probably not very far west of Fort Mandan in what is now western
North Dakota. |

In analyzing the probable identity of A. alnifolia Nutt. two facts
stand out clearly. First, it is the only species known to occur in the area
under consideration, and second, Nuttall’s description, though brief and
incomplete, gives some of its essential distinguishing characters: “Smooth;
leaves roundish, the upper part toothed, pinnately nerved, under side
somewhat glaucous; raceme simple, elongated; fruit black and sweet.
HAB. In ravines and on the elevated margins of small streams from Fort

72 ILLINOIS BIOLOGICAL MONOGRAPHS

Mandan to the Northern Andes. OBS. A shrub 4 or 5 feet high; leaves
roundish and retuse, somewhat attenuated at the base, toothed towards
the summit; fruit dark purple, somewhat pruinose, very agreeable and
saccharine, ripening about July and August.” These statements leave no
doubt as to the identity of the shrub described by Nuttall. The comment
about the leaves being “attenuated at the base” is not incompatible with
A. alnifolia, whose leaves, although usually rounded at the base, may
vary on occasional specimens toward the condition mentioned by Nuttall.
The suggestion has been offered that Nuttall might have had specimens
of one of the eastern species, such as A. spicata, but this is highly im-
probable because that species is not known to occur so far westward.

The mature foliage of A. alnifolia, A. florida, A. spicata, and A. san-
guinea often shows under certain environmental conditions a remarkable
tendency toward parallel development, and therefore some herbarium
specimens consisting of mature leaves only are sometimes rather difficult
to separate by definite taxonomic characters. It was probably on account
of this close similarity of foliage that Wiegand and others supposed the
western A. alnifolia or A. florida to extend as far eastward as Minnesota
and Michigan, and the eastern A. humilis Wieg. [A. spicata (Lam.) K.
Koch] to occur northwestward in Canada to Mackenzie. However, flower-
ing and fruiting specimens of these species are usually readily identified.
In addition to its wholly separate westerly geographical range, A. almi-
folia is characterized by the leaf-blades being mostly rounded or truncate
at the apex, glabrous or nearly so, and almost all unfolded and more than
half-grown at flowering time, coarsely toothed only above the middle,
the lower half entire. There are also good taxonomic characters in the
flowers.

Several names, purported to be of new species related to A. alnifoha,
but based almost wholly on characters of the foliage, have been published
from time to time. These include A. macrocarpa and A. leptodendron of
Lunell, both of which appear to be based on specimens of A. alnifolia
with unusual or abnormal foliage. In 1931 Rydberg described A. carrii
from fruiting specimens collected in the Black Hills of South Dakota. It
was said to be “related to Amelanchier alnifolia but differs in the short
sepals and subentire leaves.” A series of flowering specimens collected
near the type locality in 1929 by Mr. E. J. Palmer (distributed as A.
humilis) indicate A. carrii to be merely a phase of the widespread A.
alnifolia. The name A. carrii is accordingly reduced to synonymy. Ryd-
berg’s statement that the leaves are “probably glabrous from the begin-
ning” is not supported by observations made on flowering specimens.

Several specimens are known of what is supposed to be a natural
bigeneric hybrid between Amelanchier alnifolia and a species of mountain-

AMERICAN SPECIES OF AMELANCHIER—JONES 73

ash, Sorbus scopulina Greene. When first discovered, this was thought to
be a cross between A. florida and S. sitchensis. The Sorbus-parent was
identified by the writer in 1939 as S. scopulina. It is now almost certain
that the Amelanchier-parent is A. alnifolia rather than A. florida. lf a
binary name is used for this hybrid it is written * Amelasorbus jacku
Rehder in Journ. Arnold Arb. 6:154 (1925); G. N. Jones in ibid. 20:22
(1939). Amelanchier alnifolia Nutt. X Sorbus scopulina Greene.

13. AMELANCHIER FLORIDA Lindl.
(Plate II)

Amelanchier florida Lindl., Bot. Reg. 19, pl. 1589 (1833); Spach, Hist. Nat. Veg.
Phan. 2:86 (1834); M. Roem. Syn. Mon. 3:144 (1847); Decaisne in Nouv.
Arch. Mus. Hist. Nat. Paris 10:136 (1874); Schneider, Illustr. Handb. Laubh.
1:739, fig. 412 (1906); Piper, Contr. U.S. Nat. Herb. 11:345 (1906) ; Britton
& Shafer, N. Am. Trees 441, fig. 388 (1908), excl. syn.; Bean, Tr. & Shr. Brit.
Isles 1:189 (1914); Piper & Beattie, Fl. Nw. Coast 200 (1915); Henry, FI. S.
Brit. Col. 183 (1915); Bean in Bot. Mag. 141, pl. 8611 (1915); Rehder, Man.
Cult. Tr. & Shr. 388 (1927); Sudworth in U.S. Dept. Agric. Misc. Circular
92:134 (1927); Suksdorf in Werdenda 1:21 (1927); Benson, Contr. Dudley
Herb. Stanford Univ. 2:102 (1930); G. N. Jones in Univ. Wash. Publ. Biol.
5:181 (1936); Nielsen in Madrono 4:17-21, pl. 6 (1937); G. N. Jones in Univ.
Wash. Publ. Biol. 7:108 (1938); Nielsen in Am. Midl. Nat. 22:208 (1939) ;
Applegate in Am. Midl. Nat. 22:277 (1939); Peck, Man. Higher Pl. Oregon
410 (1940) ; Rehder, Man. Cult. Tr. & Shr. (ed. 2) 387 (1940) ; Abrams, Illustr.
Fl. Pac. States 2:471, fig. 2535 (1944).

Amelanchier ovalis var. 8 semtintegrifolia Hooker, Fl. Bor. Am. 1:202 (1834) ;
Loudon, Arb. & Frut. Brit. 2:876 (1838).

Amelanchier grandiflora Douglas ex Hooker, op. cit., pro syn. A. botryapium.

Amelanchier parvifolia Hort. ex Loudon, Arb. & Frut. Brit. 2:877 (1838), (ed. 2)
1854.

Amelanchier florida var. parvifolia Loudon, l.c.

Amelanchier alnifolia sensu Sargent in Gard. & For. 5:409, fig. 69 (1892), Silva N.
Am. 4:131, pl. 196 (1892), ex p.; Dippel, Handb. Laubh. 3:389, fig. 195 (1893),
excl. syn.; Howell, Fl. Nw. Am. 165 (1898); Sudworth, For. Tr. Pac. Slope
345, fig. 162 (1908). Non Nutt. 1834.

Amelanchier gormant Greene, Pittonia 4:129 (1900); Schneider, Illustr. Handb.
Laubh. 1:739 (1906).

Amelanchier oxyodon Koehne in Gartenfl. 51:609, fig. 126b (1902).

Amelanchier alnifolia var. florida Schneider, Illustr. Handb. Laubh. 1:739, fig. 411
(1906), in Rep. Sp. Nov. Reg. Veg. 3:182 (1906); Rehder in Bailey, Stand.
Cyclop. Hort. 272 (1914).

Amelanchier canadensis var. semiintegrifolia Farwell in Rep. Mich. Acad. Sci.
7:174 (1916).

Amelancher vestita Suksdorf in Werdenda 1:20 (1927).

Amelanchier ephemerotricha Suksdorf, l.c.

Amelanchier ephemerotricha var. silvicola Suksdorf, l.c.

Amelanchier florida var. typica G. N. Jones, in Univ. Wash. Publ. Biol. 5:181
(1936).

Amelanchier florida {. tomentosa Sealy in Curtis’ Bot. Mag. 160: pl. 9496, figs. a-e
(1937) ; Rehder, Man. Cult. Tr. & Shr. (ed. 2) 387 (1940).

74 ILLINOIS BIOLOGICAL MONOGRAPHS

A slender shrub 1-5 m. tall or sometimes a small tree 10 or 12 m.
tall and 15-20 cm. in diameter; branches erect; bark brownish, becoming
gray, young twigs reddish brown, tomentose at first, soon glabrous; winter
buds usually pubescent, purplish; leaves of thin texture at flowering time
and fully expanded, oval to oblong or roundish, conduplicate in vernation,
tomentulose when young, especially beneath; at maturity becoming gla-
brous and bright green above with the midvein impressed on the upper
surface; lower surface pale green and glabrous or sparingly pubescent;
blades 3-4 cm. long, 2-3 cm. wide, rounded or subtruncate at the apex, or
occasionally varying to acutish, rounded or subcordate at the base; lateral
veins 8-12 pairs, parallel, slightly curved upward, each vein often extend-
ing into a tooth; margins mostly entire below, coarsely toothed above the
middle, or rarely below, with a few spreading deltoid teeth, the teeth 4-6
per cm., 5-20 on each margin on average leaves; blades rarely completely
entire; stipules linear, villous, caducous; petioles slender, 1-2.5 cm. long,
sparsely pubescent at first, soon becoming glabrous; flowers white, 2-3 cm.
in diameter, fragrant; racemes erect, 4-8 cm. long, 5-15-flowered, the
rachis and pedicels at first whitish pubescent but soon glabrous, the lower
pedicels 8-14 mm. long; petals oblanceolate, obtuse, 12-15 mm. long
(rarely shorter), 3-3.5 mm. wide, the apex obtuse, the base cuneate, more
or less ciliolate on the base of the very short claw; stamens about 20,
shorter than the calyx-lobes; anthers yellow, 0.5-0.7 mm. long; hypan-
thium campanulate, 4-5 mm. in diameter, tomentose when young, soon
glabrous, slightly constricted on the young fruit; sepals deltoid-lanceolate.
acute or acuminate, 2-2.5 mm. long, tomentulose at least inside at anthesis,
reflexed and glabrous on the fruit; styles 5, stout, glabrous, 2-2.5 mm.
long, united to the middle or above; summit of the ovary closely tomen-
tose in anthesis, becoming nearly glabrous in fruit; fruit globose, 10-13
mm. in diameter, glabrous, glaucous, becoming purplish black at maturity,
juicy, edible; lower pedicels 1-2 cm. long; seeds dark brown, glossy, about
5 mm. long and 2 mm. wide.

Type Loca.ity: “Northwest America,” collected by David Douglas in
1825, probably along the lower Columbia River, near the present site of
Vancouver, Washington.

RANGE: In open woods and on hillsides, usually near sea level, but
occasionally ascending the mountains to an altitude of 5000 feet; south-
eastern Alaska to western Oregon and northwestern California, flowering
from the beginning of March to the end of June, according to the latitude
and altitude.

ALASKA: Wrangell, Eastwood 995 (AA, MBG); Chilkat Valley, Walker 1075
(GH, MBG); Old Kaasan Bay, Cowles 1419 (MBG, UI); along road from Hyder
to Stewart, B.C., Whited 1197 (MBG); Yes Bay, Gorman 39 (ND).

AMERICAN SPECIES OF AMELANCHIER—JONES 75

British Cotumsia: Glacier, Brown 283 (GH, MBG); McGillivray Creek,
Macoun 93901 (AA); Mons, Macoun 93898 (GH); Skidegate, Queen Charlotte
Islands, Spreadborough 93907 (GH); Lillooet, Macoun 93897 (AA), 93899 (GH) ;
Alberni, May 25, 1917, Carter (GH); Jessie I. Spreadborough 93905 (AA) ;
Cowichan Lake, Spreadborough 93906 (AA); Elk Lake, Macoun 93881 (AA) ;
Nanaimo, Macoun 93895 (AA); Prospect Lake, Macoun 93883 (AA); Robertson
River, Spreadborough 93904 (AA) ; Renfrew, Rosendahl & Brand 88 (GH, MBG) ;
Sidney, Sargent in 1913 (AA), Macoun 93879, 93880 (GH), 93875, 93887, 93889
(AA); Victoria, Macoun 79796, Eastwood 9702, Jack 2865 (AA).

WASHINGTON: Chuckanut Bay,
Muenscher 9635 (MBG); _ Browns

x oy ’ as ~ Island, Zeller 844 (GH, MBG); Up-
<< f ' iw, = per Valley of the Nesqually, Allen 214
mee ( 7 (AA, GH, MBG); Clallam Co., Elmer
Orn i m 2512 (MBG); Seattle, Piper 84 (AA);

Snoqualmie, May 2, 1937, G. N. Jones
(GH); Yelm, Sept. 1, 1891, Piper
(AA); Montesano, Heller 3958 (AA,
GH, UI, MBG); Husum, Suksdorf
10033, 10201 (AA); Cape Horn,
Suksdorf 10395, 10360, 10361 (AA);
Bingen, Suksdorf 10154 (AA, UI,
MBG), 10194, 10195, 10382, 10455
(AA), 10494, TYPE coll. of A. ephe-
merotricha var. stlvicola (GH, UI,
AA), 11841, TYPE coll. of A. vestita,
11859, TYPE coll. of A. ephemerotricha
(GH), 11835, 11973 (GH, MBG, UI).

Orecon: Portland, Aug. 18, 1880,
Drake & Dickson (GH); Waukena
Falls, Thompson 2714 (MBG); Hood
River, Suksdorf 2138, 2139, 2153
(AA); “Oregon, Douglas’ (GH);
Sauvies I., Apr. 1881, J. Howell
(AA); Forest Grove, Thompson 616
(MBG); St. Helens, T. Howell 1132
(GH, MBG); Corvallis, Steward 230,
Epling 5623 (MBG); Salem, Nelson
1071 (GH); Goshen, Abrams 8718
(MBG); Oakland, May 3, 1914, Hunt
(AA); Sutherlin, Muenscher 15136
Map 10.—Range of Amelanchier florida. (MBG).

CaLiFoRNIA: Mendocino Co., Bo-
lander 4674 (MBG).

The identity of A. florida is securely established by Lindley’s detailed
description and excellent colored plate. It is admittedly closely related to
A. alnifolia Nutt., and at various times the two have been treated as differ-
ent species, as one species and a variety, or have been merged as a single
entity. Some botanists, including Dippel, Howell, and others, while main-
taining two species, have interchanged the names, a procedure that has
naturally increased the confusion in an already complicated subject, and
adding still further obscurity to the understanding of these species is the
existence of a third, A. cusickii Fern., discovered in 1899, which occupies

76 ILLINOIS BIOLOGICAL MONOGRAPHS

a smaller range within that of A. alnifolia. Reports of A. florida from the
region east of the Cascade Mountains, and most of the specimens so
named belong really to A. cusickii, a large-flowered species resembling
A. florida in superficial appearance, but immediately distinguished by the
usually glabrous ovary, and longer petals and sepals. A. alnifolia and
A. cusicku often grow together, but there is no evidence of hybridization
between them.

Although as now recognized by critical students of Amelanchier,
A. florida and A. alnifolia are distinct species with good distinguishing
floral characters and separate geographical ranges (the former confined
to the Pacific slope west of the Cascade Mountains, and the latter a
species of the Great Plains, and the Rocky Mountain area) leaf specimens
of these species are sometimes virtually indistinguishable from each other,
and on that account the foliage is of limited taxonomic value. Leaves of
A. alnifolia tend to be more nearly suborbicular and of somewhat thicker
texture, and often darker green, but these are by no means constant char-
acters, especially on specimens collected later in the season, or from
drier habitats. The best distinguishing morphological character is to be
found in the flowers. Flowering specimens of A. florida are at once dis-
tinguished from those of A. alnifolia by the larger flowers with longer
petals. The petals of A. alnifolia are usually 7-10 mm. long with only
occasional larger-flowered specimens. 4. florida, on the other hand, is a
relatively large-flowered plant, the petals being usually 12-15 mm. long;
however, occasional smaller-flowered specimens are found on stunted
plants, or plants of dry habitats, so that the distinction is not an absolute
one. The habit of these two species respectively is usually somewhat
different, A. florida being generally taller and more slender, but the
growth-form of each is extremely variable according to the habitat, and
can therefore scarcely serve for basic taxonomic purposes. As previously
noted, the geographical ranges of the two species are almost separate, and
it is only in a very few localities, such as in the Gorge of the Columbia
River, that the two are ever found growing near each other.

Amelanchier gormani Greene was described from specimens collected
by M. W. Gorman at Yes Bay, Alaska, in 1895, and was said to be
characterized by the calyx, whose “limb is notably dilated under the in-
sertion of the petals into a broad saucer-shaped rim; and the lanceolate
segments, either erect or somewhat spreading, are longer than all the rest
of the calyx, and are tomentulose within.’”’ However, this condition is by
no means peculiar to specimens from Alaska, but is more or less char-
acteristic of the partly ripened fruits of many specimens of both A. florida
and A. alnifolia.

AMERICAN SPECIES OF AMELANCHIER—JONES 77

13a. AMELANCHIER FLORIDA var. HUMPTULIPENSIS
G. N. Jones

Amelanchier alnifolia var. pumila sensu Schneider, Ilustr. Handb. Laubh. 1:739,
fig. 412 (1906), as to specimen cited, non A. pumila Nutt. ex Torrey & Gray
Pe hier florida var. humptulipensis G. N. Jones in Univ. Wash. Publ. Biol.

5:181 (1936).

Leaves and flowers smaller than those of A. florida var. typica; blades
of the mature summer foliage of the fruiting branches mostly oval, acutish
or obtuse at the base, acute or obtuse at the apex, 2-3 cm. long, 1-2 cm.
wide, nearly entire, or shallowly serrate near the apex with small teeth;
the racemes 2-4 (or sometimes only 1-2) cm. long, erect, 5-9-flowered,
the pilose pedicels only 2-3 mm. long; petals 6-10 mm. long, 2 mm. wide,
oblanceolate; stamens 12-15, the filaments glabrous, 1-2 mm. long;
anthers 0.6-0.8 mm. long; hypanthium shallowly campanulate, sparsely
pubescent outside at flowering time; sepals lanceolate, acuminate, soon
recurved, 1.5-2 mm. long, glabrous within, or only sparsely pilosulous;
styles 4, glabrous, unequal, 1.5 mm. long; top of the ovary tomentose,
rarely nearly glabrous; fruit 5-7 mm. in diameter, 8-loculed and 8-seeded.

Tyre Locatity: Humptulips Prairie, Grays Harbor Co., Washington.
Type: G. N. Jones 4565 (flower), 5819 (fruit) in the herbarium of the
University of Washington, Seattle.

RANGE: Olympic Peninsula, Washington, and adjacent southern Van-
couver Island, British Columbia.

British Cotumpra: Mt. Benson, July 10, 1893, Macoun (GH), 208 (GH);
near Victoria, May 15, 1893, Macoun (GH).

WASHINGTON: Humptulips, June 12, 1897, Lamb 1190 (AA, MBG); Hump-
tulips River, Murie 1122 (MBG); Kamilche, Benson 1423 (MBG); Shelton, Benson
1420 (MBG).

This appears to be a local variety of A. florida, differing from the
typical form of that species in its smaller size, usually more finely toothed
leaves, shorter inflorescences, four styles, and its restricted geographical
distribution, In 1906,-C. V. Piper (Contr. U.S. Nat.. Herb. 11:346) sug-
gested that it might be a new species, but concluded that more and better
specimens were needed. On account of the smaller flowers it might be
treated as a near relative of A. alnifolia rather than of A. florida, but the
general appearance, as well as its geographical range, suggest a closer
connection with the latter, of which it might be dismissed as merely a
stunted form, except for the fact that the flowers appear uniformly to
possess only four styles instead of five, the almost invariable number for
all other known species in this general cycle of affinity.,

78 ILLINOIS BIOLOGICAL MONOGRAPHS

14. AMELANCHIER CUSICKII Fern.
(Plates 11 XX)

Amelanchier cusickti Fernald in Erythea 7:121 (1899); Piper in Contr. U.S. Nat.
Herb. 11:346 (1906); Schneider, Illustr. Handb. Laubh. 1:735, figs. 409, 410
(1906); Piper & Beattie, Fl. Se. Wash. & Adj. Idaho 133 (1914); Henry, FI.
So. British Columbia 183 (1915); Rydberg, Fl. Rocky Mts. 446 (1917); Kirk-
wood, N. Rocky Mt. Tr. & Shr. 190 (1930); St. John, Fl. Se. Wash. & Adj.
Idaho 194 (1937) ; Peck, Man. Higher Pl. Oregon 410 (1940) ; Abrams, Illustr.
Fl. Pac. States 2:471, fig. 2533 (1944).

Shrubs 1-3 m. tall, with numerous slender, virgate, flexible branches;
bark on the young branches reddish, later becoming gray; twigs of the
season reddish brown, often somewhat glossy, quite glabrous; winter buds
reddish, nearly or quite glabrous, the inner scales more or less ciliate;
leaves commonly oval, thin, glabrous from the first, even when young,
becoming coriaceous in age, conduplicate in the bud, usually unfolded at
anthesis; mature blades 2.5-5 cm. long, 2-3 cm. wide, the apex acutish
and apiculate on some leaves, obtuse and rounded on others, the base
rounded or subcordate, perfectly glabrous on both surfaces; lateral veins
7-10 pairs, curving upward and anastomosing near the margin; margins
sharply serrate mostly above the middle, the upper teeth more prominent;
teeth 3-6 per cm., 3-15 on each side on average leaves of the fruiting
branches; stipules linear, deciduous, villous, about 1 cm. long; petioles
slender, soon glabrous, 6-22 mm. long; flowers large, in 3-8-flowered
racemes 2-5 cm. long, the lower pedicels 8-12 mm. long, glabrous; petals
5, white, glabrous, obovate or oblanceolate, obtuse, 16-25 mm. long,
usually 5-7 mm. wide above the middle, the apex obtuse or acutish;
stamens about 20, the filaments glabrous, 3-4 mm. long; anthers ellipsoid,
1-1.5 mm. long; hypanthium shallowly campanulate or saucer-shaped,
glabrous within and outside, 45 mm. in diameter; sepals lanceolate,
acuminate, or acute, 3.5-5 mm. long, pilose within at flowering time,
usually recurved from the middle during or after anthesis, the margins
very narrowly hyaline; styles 5 or 4, glabrous, 3-4 mm. long, united below
the middle, or nearly free to the base; summit of the ovary glabrous or
with a ring of tomentum around the base of the styles; mature fruit
glabrous, globose, juicy and edible, reddish at first, becoming bluish black,
about 1 cm. in diameter; fruiting pedicels 5-20 mm. long; seeds brown,
smooth, asymmetrical, 4-5 mm. long.

Type Locauity: “On stony hillsides, Union County, Oregon.” Type
(Cusick 1858), in the Gray Herbarium of Harvard University; isotype in
the herbarium of the Missouri Botanical Garden.

AMERICAN SPECIES OF AMELANCHIER—JONES 79

RaNGE: Common on basaltic ledges along rivers, British Columbia,
southward to eastern Washington, eastern Oregon, Idaho, western Mon-
tana, and northern Utah; flowering in March and April; fruit ripening
in June.

BritisH CoLuMBIA: Kamloops, June
30, 1887, Fowler (MBG).

WASHINGTON: Fort Colville, Lyall in
1861 (GH); Camden, Spiegelberg 341
(GH); Kettle Falls, Boner & Weldert
172 (GH, MBG); Spokane, Sept. 1,
1899, Piper (GH), 2694 (AA), Suks-
dorf 8575 (AA, GH, MBG, UI), Mil-
burge 258 (AA); Grand Coulee, Rollins
840 (MBG); Blewett Pass, Benson 1283
(MBG); Wenatchee R., Chelan Co.,
G. N. Jones 6573 (UI); Entiat, Thomp-
son 6002 (GH, MBG); Priest Rapids,
G. N. Jones 6349 (GH); Yakima, G. N.

Jones 1387 (UI); near Ellensburg, re

Eyerdam 1377 (MBG), Thompson 11386 te ' fi--! i
(GH, MBG); six miles s.w. of Pullman, a ; A eee
Aug. 4, 1899, Piper (GH), Elmer 135 / a me | HCA f
(MBG); Wawawai, Piper 3812 (GH), ae

G. N. Jones 1398 (UI); Almota, Con- Mar 11—Range of Amelanchier cusicku.
stance, et al. 1037 (GH, MBG) ; Bishop,
Constance & Rollins 1512 (GH);
Hooper, Constance & Rollins 1499
(MBG); Waitsburg, Horner B183 (GH); Palouse Falls, G. N. Jones 2832 (Ph).

Orecon: The Dalles, Lunell 50 (GH), Apr. 19, 1903, Lunell (UI); Canyon
Creek, Grant Co., Henderson 5147 (GH, MBG); eastern Oregon, Cusick 1858
(TYPE, GH, isotype, MBG); Wallowa Mts., G. N. Jones 7088, 7101, 7134, 7147
(UI); Jim Creek, Wallowa Co., Constance 999 (MBG); Crooked River, Whited
363 (GH, MBG).

IpaHo: Spalding, Meyer 1428 (MBG); Snake River Canyon, Idaho Co., Con-
stance, Clements, & Machlis 1008, Packard 371, Meyer 854 (MBG); Dry Buck,
Boise Co., Macbride 852 (GH, MBG); Kootenai Co., July 1890, Letberg (GH).

Montana: Near Butte, Moore in 1893 (MBG); Mt. Sentinel, Kirkwood 28,
29, 30 (AA), Barkley & Osburnson 2313 (AA, MBG, UI); near Missoula, Kirk-
wood 1182 (GH, MBG), Hughes 1185 (GH, MBG), E. C. Faust in 1915 (UI);
near Bonner, Geil 4 (MBG, UI).

UtaH: Cache Co., Logan Canyon, Maguire 3501 (GH); Blacksmith Canyon,
Maguire 3507 (GH); Sardine Canyon, Maguire 2363 (GH).

This distinctive species was named in compliment to the pioneer
Oregon botanist, William Conklin Cusick (1842-1922). It is the largest-
flowered species of Amelanchier, and thus in the flowering stage is at once
distinguishable from all others. It blooms from ten to fifteen days earlier
than A. alnifolia. Fruiting specimens are frequently less easily recognized,
but the long sepals will usually serve to separate it from A. alnifolia, the
common species occurring within its range.

80 ILLINOIS BIOLOGICAL MONOGRAPHS

15. AMELANCHIER BASALTICOLA Piper
(Plate IIT)

Amelanchier alnifolia sensu Holzinger in Contr. U.S. Nat. Herb. 3:224 (1895).

Non Nuttall, 1834.
_Amelanchier basalticola Piper, Fl. Palouse Reg. 100 (1901); Piper in Contr. U.S.

Nat. Herb. 11:346 (1906) ; Piper & Beattie, Fl. Se. Wash. Adj. Ida. 133 (1914) ;

Rydberg, Fl. Rocky Mts. 446 (1917) ; St. John, Fl. Se. Wash. & Adj. Ida. 194 (1937).

Shrubs 1-3 m. tall; bark gray; twigs of the season glabrous, brown,
smooth; winter buds small, glabrous; leaves conduplicate in vernation,
appearing before the flowers, flat and unfolded at flowering time, pale
green, glabrous, and glaucous from the first, firm in texture; blades sub-
orbicular, 1.5-3 cm. long, 1.5-2 cm. wide, rounded or truncate at apex and
base, the apex sometimes acute and mucronate; lateral veins 10-13 pairs,
curved upward, anastomosing near the margin; margins serrate above the
middle or less commonly from near the base, the teeth small, sharp, some-
what curved, 4-6 per cm., mostly 5-15 on each side on average leaves of
the flowering branches; some of the blades sometimes nearly or quite
entire; stipules linear-subulate, small, glabrous, fugacious; petioles slender,
glabrous, 7-18 mm. long, usually shorter than the blade; flowers appear-
ing after the leaves have unfolded, about 3 cm. in diameter; racemes
terminal, 2-4 cm. long, 4-8-flowered; pedicels nearly or quite glabrous, and
somewhat glaucous, 4-8 mm. long; petals 5, white, glabrous, oblanceolate,
12-16 mm. long, 3-4 mm. wide above the middle, the apex acute, obtuse,
entire or erose; stamens about 20, the filaments glabrous, 2-3 mm. long;
anthers 0.8 mm. long; hypanthium campanulate, glabrous and somewhat
glaucous, 3 mm. in diameter, sepals linear-lanceolate, attenuate-acuminate,
3.5-4 mm. long, pilosulous within, becoming reflexed after anthesis, some-
what glaucous, slightly longer than the tube; styles 5, free to below the
middle or almost to the base, 2-2.5 mm. long, summit of the ovary gla-
brous or nearly so; mature fruit globose, dark purple, juicy, glabrous,
9-12 mm. in diameter; fruiting pedicels glabrous, about 1 cm. long; well
developed seeds chestnut brown, smooth, glabrous, 5-6 mm. long, 3-3.5
mm. wide, asymmetrical, flattened-ellipsoid, bluntly pointed at each end.

Type Locatity: Bluffs of Snake River, Whitman County, Washing-
ton, opposite Clarkston.

RANGE: Southeastern Washington, adjacent Idaho, and Oregon.
Flowering in April and May; fruit ripe in June and July.

WasHIncTon: Bluffs above Wawawai, Piper 3823 (MBG, AA, GH).

IpaHo: Near Lewiston, Apr. 15, 1896, Heller (UI), 2988 (MBG, Ph); Upper
Ferry, Clearwater River, near Lewiston, Sandberg, MacDougal, & Heller 53 (MBG,
AA, GH, UI); Nez Perces Co., May 1892, Sandberg (UI, MBG); near Riggins,
G. N. Jones 972 (UI).

Orecon: Mitchell, Ferris & Duthie 656 (AA); Wallowa River, G. N. Jones
7137 (UI); Deep Creek, Snake R. Canyon, Wallowa Co., Constance, Rollins, &
Dillon 1568 (GH); Snake R. Canyon above Rogersburg, Meyer 226 (MBG).

AMERICAN SPECIES OF AMELANCHIER—JONES 81

Amelanchier basalticola Piper is a species of local distribution inhabit-
ing the basaltic cliffs and ledges of the river canyons of southeastern
Washington, and adjacent Idaho and Oregon. It is very conspicuous in the
spring when in bloom, marking the ledges on which it grows. It belongs
to the alnifolia-florida-cusicku cycle of affinity, being probably most
closely related to A. cusicku, from which it differs in its shorter and nar-
rower petals, somewhat longer sepals, and the smaller, roundish, pale
green leaves. It is distinguished from A. alnifolia by the longer petals,
longer and narrower sepals, as well as the usually smaller leaves.

16. AMELANCHIER PUMILA Nutt.
(Plates III and IX)

Amelanchier canadensis var. € pumila Nuttall ex Torrey & Gray, Fl. N. Am. 1:474
(1840) ; Walpers, Rep. Bot. Syst. 2:55 (1843) ; Dietrich, Syn. 3:158 (1843).
Amelanchier pumila Nuttall ex Torrey & Gray apud M. Roemer, Syn. Mon. 3:145

(1847) ; Rydberg, Fl. Rocky Mts. 446 (1917); Tidestrom in Contr. U.S. Nat.

Herb. 25:283 (1925) ; Graham in Ann. Carnegie Mus. 26:232 (1937).
Amelanchier glabra Greene, Fl. Franciscana 52 (1891); Schneider, [llustr. Handb.

Laubh. 1:735 (1906) ; Coulter & Nelson, New Man. Rocky Mt. Bot. 266 (1909) ;

Smiley in Univ. Calif. Publ. Bot. 9:230 (1921) ; Tidestrom in Contr. U.S. Nat.

Herb. 25:283 (1925); Abrams, Illustr. Fl. Pac. States 2:471, fig. 2532 (1944),

excl. syn.

Amelanchier polycarpa Greene, Pittonia 4:127 (1900); Schneider, Illustr. Handb.
Laubh. 1:735, fig. 409 (1906); Rydberg, Fl. Colorado 191 (1906); Wooton &
Standley in Contr. U.S. Nat. Herb. 19:323 (1915); Rydberg, Fl. Rocky Mts.
446 (1917); Tidestrom in Contr. U.S. Nat. Herb. 25:283 (1925); Tidestrom &
Kittell, Fl. Ariz. & New Mexico 251 (1941); Kearney & Peebles in U.S. Dept.
Agric. Misc. Publ. 423:393 (1942).

Amelanchier alnifolia pumila A. Nelson in Coulter & Nelson, New Man. Rocky Mt.
Bot. 266 (1909).

Shrubs 1-3 m. tall, the whole plant perfectly glabrous; bark reddish
brown, becoming gray; twigs and winter buds glabrous; leaves oval,
perfectly glabrous throughout from the beginning, thickish, somewhat
coriaceous at maturity, pale and somewhat glaucescent, at least on the
lower surface, deeper green above; conduplicate in vernation, unfolding
before or with the flowers and nearly full grown at anthesis; mature
blades suborbicular to oval, 1-5 cm. long, 1-2 cm. wide, the apex obtuse
or truncate, often somewhat mucronate, the base rounded, subcordate,
or truncate, rarely somewhat cuneate; lateral veins 7-9 pairs, curving
upward and often extending into the teeth, or anastomosing near the
margin; margins coarsely serrate to the middle, the lower third of the
biade entire, the teeth 1-2 mm. long, 3-5 per cm., 6-10 on each side of
average blades of the flowering and fruiting branches; some small-leaved
specimens with the blades nearly entire, or with only a few small teeth
near the apex; stipules linear, glabrous, fugacious; petioles rather slender,

mostly shorter than the blade, glabrous; flowers in erect or ascending

82 ILLINOIS BIOLOGICAL MONOGRAPHS

4-8-flowered racemes 2-4 cm. long, the lower pedicels about 6-12 mm.
long; hypanthium and pedicels completely glabrous, more or less glaucous;
petals 5, white, glabrous, oval, 8-12 mm. long, 3-4 mm. wide, obtuse
at the apex or sometimes acute, widest at or above the middle; stamens
12-15, the filaments glabrous, 1-2 mm. long; anthers 0.8 mm. long;
hypanthium campanulate, 3-4 mm. in diameter, completely glabrous,
somewhat constricted on the young fruit; sepals triangular-lanceolate
to linear-lanceolate, acuminate, completely glabrous on both sides, 3 mm.
long (rarely longer), soon recurved; styles 5 (or 4), united at the base,
glabrous, 1-2 mm. long; top of the ovary completely glabrous; mature
fruits depressed-globose, dark purple, glaucous, juicy, 8-9 mm. in di-
ameter; pedicels 6-12 mm. long; seeds brown, asymmetrical, 4-5 mm. long.

Type Locatity: “Near the sources of the Platte in the Rocky Moun-
tains.” Collected by Thomas Nuttall. Type in the herbarium of the Acad-
emy of Natural Sciences of Philadelphia.

RANGE: On mountain sides and plains, altitude 7,000-10,000 feet,
southwestern Montana and Idaho, to southern Oregon, Wyoming, Utah,
Colorado, and northeastern California, apparently not common, or at
least only locally abundant.

Montana: Spanish Basin, Madison Range, Flodman 545 (MBG); Helena,
May 1893, Starz (MBG).

Ipauo: Alturas Lake, Cronquist 2596
(MBG); Twin Falls, Nelson & Mac-
bride 1368 (MBG, GH); Corral, Mac-
bride & Payson 2880 (GH, MBG).

OrEGON: Cougar Peak, Lake Co.,
Eggleston 7066 (GH).

Wyvominc: Jackson Hole, June 9,
1934, Nelson & Nelson (MBG); Medi-
cine Bow Mts., Rollins 889 (MBG);
Laramie Hills, Nelson 1931 (GH,
MBG); near Leckie, Merrill & Wilcox
548 (GH), Sherman, July 29, 1884,
Letterman (MBG).

CoLtorapo: “R. Mts.” [Rocky Moun-
tains], Nuttall (TYPE in herb. Acad.
Nat. Sci. Phila.); Gunnison Canyon,
Payson 1051 (MBG); Cerro Summit,
Baker 49 (MBG); Piedra, July 10, 1899,
Baker 379 (AA, GH, MBG; ND, TYPE
of A. polycarpa); Eldora to Arapahoe
Peak, Daniels 909 (MBG); without
definite locality, Vasey in 1868 (GH).

UtaH: Logan Canyon, Mulford 192 :
(MBG, IGIB)e Milford, Hill 169 CUD): Map 12.—Range of Amelanchier
near Moon Lake, Harrison & Larsen punula.

7685 (MBG); Oak City, Harris C28699
(MBG).

CatirorNIA: Deer Park, Lake Tahoe region, Eastwood 372 (GH, AA); Donner

Lake, J. Torrey 126 (GH, Paratype of A. glabra), Heller 7176 (GH, AA, MBG).

AMERICAN SPECIES OF AMELANCHIER—JONES 83

Amelanchier pumila Nutt., included in their Flora of North America
by Torrey & Gray in 1840, was based on specimens collected by Nuttall
“Near the sources of the Platte in the Rocky Mountains.” It can usually
be distinguished from all other species by the fact that the whole plant
is completely glabrous, even on the youngest parts. It is evidently closely
related to A. cusickii and A. basalticola, from which it differs princi-
pally in its glabrous condition, and in the smaller flowers with shorter
petals, styles, and anthers.

17. AMELANCHIER PALLIDA Greene
(Plates IIT and XX1I)

Amelanchier alnifolia sensu Brewer & Watson, Bot. Calif. 1:190 (1880); Greene,
Man. Bot. Bay Reg. 110 (1894); Jepson, Fl. W. Middle Calif. 288 (1901),
(ed?) u213 911); H..M. & C.,C. Hall, Yosemite Fl. 125 (1912); Jepson,
Man. Fl. Pl. Calif. 509, fig. 508 (1925), Fl. Pl. Calif. 2:234 (1936) ; McMinn,
Illustr. Man. Calif. Shr. 216, fig. 239 (1939). Non Nuttall, 1834.

Amelanchier pallida Greene, Fl. Franciscana 53 (1891) ; Coville in Contr. U.S. Nat.
Herb. 4:97 (1893) ; Howell, Fl. Nw. Am. 165 (1898) ; Schneider, Illustr. Handb.
Laubh. 1:742, figs. 416, 417 (1906); Davidson & Moxley, Fl. S. Calif. 206
(1923) ; Tidestrom in Contr. U.S. Nat. Herb. 25:284 (1925); Benson in Contr.
Dudley Herb. Stanford Univ. 2:103 (1930); Graham in Ann. Carnegie Mus.
26:232 (1937); Peck, Man. Higher Pl. Oregon 410 (1940); Abrams, Illustr.
Fl. Pac. States 2:472, fig. 2538 (1944).

Amelanchier subintegra Greene in Pittonia 5:109 (1903) ; Schneider, Illustr. Handb.
Laubh. 1:742, figs. 416, 417 (1906).

Amelanchier gracilis Heller in Muhlenbergia 2:59 (1905) ; Schneider, Ilustr. Handb.
Laubh. 1:735 (1906); Benson in Contr. Dudley Herb. Stanford Univ. 2:103
(1930) ; Abrams, Illustr. Fl. Pac. States 2:471, fig. 2536 (1944).

Amelanchier siskiyouensis Schneider, Illustr. Handb. Laubh. 1:735, figs. 409, 410
(1906), in Fedde, Rep. Sp. Nov. 3:181 (1906); Smiley in Univ. Calif. Publ.
Bot. 9:230 (1921).

Amelanchter recurvata Abrams in Bull. Torr. Club 37:151, fig. 1 (1910) ; Davidson
‘& Moxley, FI. S. Calif. 206 (1923).

Amelanchier alnifolia var. pallida Jepson, Man. FI. Pl. Calif. 509 (1925), Fl. Calif.
2:234 (1936); Munz, Man. S. Calif. Bot. 229 (1935).

Amelanchier alnifolia var. cuyamacensis Munz in Bull. S. Calif. Acad. Sci. 31:65
(1932), Man. S. Calif. Bot. 229 (1935).

Amelanchier alnifolia var. siskiyouensis Jepson, Fl. Calif. 2:234 (1936).

Amelanchier alnifolia var. subintegra Jepson, ibid.

Amelanchier florida sensu Wynd in Am. Midl. Nat. 17:921 (1936); Applegate in
ibid. 20:277 (1939). Non Lindley 1833.

Shrubs 1-3 (-8) m. tall, with erect or ascending or divaricate branches,
these often numerous and rigid, especially in dry habitats, or rarely
spreading or more or less drooping; bark of the twigs and branches
usually reddish brown, or gray, glabrous; winter buds pubescent; leaves
oval or elliptical to suborbicular or broadly obovate, rather thick or
coriaceous, indistinctly reticulate, pale dull green on both surfaces, the
lower surface usually paler than the upper, or sometimes the upper
surface dark green, finely tomentulose or puberulent on both surfaces,

84 ILLINOIS BIOLOGICAL MONOGRAPHS

occasionally varying to nearly or quite glabrous; blades at maturity 2-4
em. long, 1.5-2.5 cm. wide, the apex usually narrow and acutish or
rounded and retuse, rarely truncate, often apiculate or cuspidate, the base
rounded; lateral veins 7-9 pairs, obscure; margins variable, entire or with
a few small teeth toward the apex, or sometimes toothed to below the
middle, both conditions often found on the same specimen; stipules
linear, villous, 1-1.5 cm. long, fugacious; petioles 6-12 mm. long, usually
more or less pubescent; flowers in short and somewhat corymbose racemes
2-4 cm. long and 4-6-flowered, on short lateral leafy spurs of the season;
pedicels rather stout, 3-5 mm. long, pubescent at anthesis; petals 5, white,
oval or obovate, 8-11 mm. long, 3-4.5 mm. wide, sparsely pilose at the
base within; stamens about 15, the filaments glabrous, 2 mm. long; anthers
0.8-1 mm. long; hypanthium shallowly campanulate, 3-4 mm. in diameter,
more or less tomentose on the outside at anthesis, or nearly glabrous,
more or less constricted on the very young fruit; sepals lanceolate, acumi-
nate 2-3 mm. long, villosulous on both sides, usually recurved from the
middle after anthesis; styles 4 or 3 (rarely 5), glabrous, about 2 mm. long,
free nearly to the base; summit of the ovary usually tomentose, sometimes
only sparsely pilose or nearly glabrous; mature fruit subglobose, purplish
black, juicy when mature, 4-6 mm. in diameter, the upper part of the
hypanthium constricted on the young fruit; fruiting pedicels 3-9 mm. long.

Type Locarity: Near Yreka, Siskiyou County, California, collected
May 12, 1876, by E. L. Greene, no. 779.

RANGE: On dry gravelly ridges and slopes, on moraines, in rocky woods
and thickets, or along streams, from sea level to 8500 feet altitude,
southern Oregon to southern California, chiefly in the Sierra Nevada and
Coast Ranges; also in northwestern Nevada. Flowering from the middle
of April to the end of June, according to the altitude; fruits ripening in
July and August. Common name: California serviceberry.

OreEconN: Reston, Peck 3530 (GH); near Klamath Falls, Wiggins 4631 (GH) ;
Crater Lake National Park, Stanford 1758 (MBG), G. N. Jones 7690 (UI); op-
posite Ashland, T. Howell in 1889 (MBG); near Ashland, Peck 9242 (GH, MBG);
near Waldo, Aug. 24, 1904, Rehder & Jack (AA), Henderson 5824 (MBG);
Wilderville, Aug. 24, 1904, Jack (AA); between Rogue R. and Union Creek,
Heller 12962 (GH, Ph, UI); Obrien, G. N. Jones 7755 (UI); Agness, Nelson 1475
(GH); Harbor, Peck 8731 (MBG, GH); Port Orford, Peck 8512 (GH, MBG).

CaLtrorNIA: Shelley Creek, Eastwood 12076 (AA); French Hill, Eastwood
72 (AA); near Mt. Eddy, Heller 13396 (MBG); Siskiyou Co., Pringle in 1882
(GH, AA, MBG); near Shasta Springs, Heller 7970 (GH, AA, MBG, TYPE coll.
of A. gracilis) ; Mt. Shasta, Brown 557 (MBG); Cantara, Eastwood 11931 (AA);
McCloud, Eastwood 1081 (GH, AA); Weed, Smith 284 (GH, AA); Yreka, Heller
8003 (GH, AA, MBG), Greene 779 (GH, MBG, TYPE coll.) ; near Alturas, C. L.
Hitchcock 6701 (MBG); Parker Creek, Ferris & Duthie 77 (AA); Hoopa Indian
Reservation, Chandler 1282 (AA, GH, MBG); Little Van Duzen R., Eastwood &
Howell 4800 (AA); Goose Valley, Eastwood 785 (AA, GH); Bennett Spring,
Tehama Co., Heller 13004 (GH, AA, MBG, UI); Fredonyer Pass, Heller 15144

AMERICAN SPECIES OF AMELANCHIER—JONES 85

(UI) ; Plumas Co., Eggleston 6235, 7596 (GH); Mt. Sanhedrin, Heller 5961 (GH,
MBG), Bacigalupi 1549 (GH, AA); Lakeport, Baker 2964 (GH, AA, MBG,
TYPE coll. of A. subintegra); near Stirling, Heller 13156 (GH, MBG, UI);
Chico Meadows, Heller 11963 (GH, AA, UI); near Donner Lake, Heller 7038
(GH, AA, MBG); Truckee, Sonne 88 (MBG) ; Emigrant Gap, Heller 12729 (GH,
UI); Fallen Leaf Lake, Abrams 4809, 4817 (GH), Smiley 214, 215 (GH); Sebas-
topol, Pammel & Davy 77 (MBG),
Heller 5794 (GH, MBG); Cazadero,
Heller 6617 (AA, GH, MBG); Napa,
Smyth in 1899 (GH) ; Silver Lake, Han-
sen 230 (GH, MBG, TYPE coll. of A.
stskiyouensis) ; San Mateo Co., Abrams
5577. (AA); Pescadero, Elmer 4659
(MBG); Santa Clara Co., Heller 7419
(GH, AA, MBG), Heller 8531 (GH,
UI); Yosemite Nat. Park, Smiley 899
(GH), Abrams 4577 (GH), 4675 (AA,
GH); Mt. Bullion, Congdon 31 (GH);
Ranch Jolon, Brewer 576 (GH, MBG) ;
General Grant Park, Culbertson 4647
(GH, MBG); Tehachapi Mts., Abrams
& McGregor 490 (GH, AA); Topatopa
Mts., Abrams & McGregor 107 (AA,
GH, TYPE ooll. of A. recurvata) ;
Cuyamaca Lake, Munz 8099 (GH,
TYPE coll. of A. alnifolia var. cuyama-
censis), Mung & Johnston 12638 (AA,
MBG), Spencer 865 (GH), Abrams 3912

(GH, MBG). .
Nevapa: Verdi, June 1, 1893, Sonne Map 13.—Range of Amelancher

(MBG), Heller 10875 (AA, GH, UI, pallida.

MBG); Kings Canyon, Ormsby Co., In oval, Amelanchier basalticola.

Baker 952 (GH, AA, MBG), 1219

(GH, AA).

It has been customary to refer nearly all the Californian Amelan-
chiers to A. alnifolia Nutt., but that species does not occur in Cali-
fornia, or indeed on the Pacific slope. The common and characteristic
species of California and southern Oregon is A. pallida Greene. It
differs from A. alnifolia in its usually smaller, narrower leaves, which
are mostly finely and inconspicuously permanently pubescent or pu-
berulent, at least beneath, the indument scarcely noticeable to the naked
eye. In fact, the original description does not mention this character.
Only rarely are the leaves quite glabrous on both surfaces at maturity.
The blades are variable in shape, texture, and degree of serration, vary-
ing apparently with the habitat and the age of the branch, those on
vigorous young shoots often showing a tendency toward being sub-
orbicular and coarsely toothed. Specimens from moist habitats have
darker green and more serrate leaves, while in drier habitats the leaves
are nearly entire, paler green, more pubescent, and subcoriaceous in
texture. These fluctuations give the specimens a somewhat different

86 ILLINOIS BIOLOGICAL MONOGRAPHS

appearance, but they seem to be well within the normal range of
variation of the species. A. pallida differs also from A. alnifola in the
fewer-flowered often somewhat corymbiform racemes, usually 15 stamens,
the styles 4 or 3 (rarely 5) free nearly to the base, and in the usually
smaller fruits.

The forms with darker green, less puberulent, and more serrate
leaves that have been described as A. gracilis Heller, and A. siskiyouensis
Schneider, present at first glance a rather distinctive appearance, and these
characters are often more prevalent on plants from higher altitudes, but
there is a complete intergradation to the form with pallid, nearly entire
leaves, and thus it is scarcely feasible to attempt to maintain more than
one species. However, special field work might yield data for possibly
varietal separation.

Regarding the Californian serviceberries as varieties of A. alnifolia
Nutt., Dr. W. L. Jepson (FI. Calif. 2:234. 1936) noted:

[They are] continuously though not highly variable as to pubescence and leaf
shape. Pubescence of the leaves or lack of it, and pubescence of the sepals are not
definitely associated with any other character. The leaf-blades are broad, elliptic,
or suborbicular and serrate at apex or a little below the apex on the sides. Plants
otherwise alike may have puberulent or glabrous leaves....In the Coast
Ranges and southward to Southern California, the leaf-blades tend to have a few
small teeth at apex or are even sometimes entire, though rarely, the teeth may be
large or the serrations extend half way to the base. ... In the Sierra and North
Coast Ranges occurs a form (var. siskiyouensis Jepson) with oblong or elliptic
leaf-blades serrate usually to the middle or nearly to the obtuse base; they are thin
at flowering time, weakly puberulent or rarely thinly arachnoid below. . . . On the
desert slopes of the Sierra Nevada and the high mountains of Southern California
or in ranges bordering the deserts a shrub is found (var. pallida Jepson) with
elliptic leaves pale or glaucous or whitish pubescent beneath: . .. All these phases,
as above indicated, are freely supplemented by intergrades representing many
indefinite fractional variations.

Amelanchier alnifolia var. cuyamacensis Munz, from Cuyamaca
Lake, San Diego Co., and A. recurvata Abrams, belong obviously to
the same species, and match exactly the typical, or oligodontous, form of
A. pallida Greene.

18. AMELANCHIER UTAHENSIS Koehne
(Eiate sii)

Amelanchier utahensis Koehne, Gattung. Pomac. in Wissen. Beil. Progr. Falk.-Real.
Berlin 95:25, pl. 2 (1890) ; Schneider, Illustr. Handb. Laubh. 1:741, figs. 415, 416
(1906) ; Rydberg, Fl. Rocky Mts. 447 (1917); Tidestrom in Contr, U.S. Nat.
Herb. 25:283 (1925); Graham in Ann. Carnegie Mus. 26:233 (1937); Peck,
Man. Higher Pl. Oregon 410 (1940); Tidestrom & Kittell, Fl. Ariz. & New
Mex. 251 (1941); Kearney & Peebles in U.S. Dept. Agric. Misc. Publ. 423:393
(1942) ; Abrams, Illustr. Fl. Pac. States, 2:471, fig. 2537 (1944).

Amelanchier alnifolia sensu Coville in Contr. U.S. Nat. Herb. 4:97 (1893), ex p.;
Parish in Pl. World 20:217 (1917) ; I. M. Johnston in ibid. 22:105 (1919). Non
Nuttall, 1834.

AMERICAN SPECIES OF AMELANCHIER—JONES 87

Amelanchier pallida var. arguta Greene in Erythea 1:221 (1893).

Amelanchier alnifolia var. utahensis M. E. Jones in Proc. Calif. Acad. Sci. 5:679
(1895).

Amelanchier prunifolia Greene in Pittonia 4:21 (1899); Schneider, Illustr. Handb.
Laubh. 1:740, figs. 413, 414 (1906); Rydberg, Fl. Col. 191 (1906); Coulter &
Nelson, New Man. Rocky Mt. Bot. 266 (1909); Rydberg, Fl. Rocky Mts. 447
(1917) ; E. H. Graham in Ann. Carnegie Mus. 26:232 (1937); Rehder, Man.
Cult. Tr. & Shr. 390 (1927), (ed. 2) 389 (1940).

Amelanchier venulosa Greene, in Pittonia 4:21 (1899); Schneider, Illustr. Handb.
Laubh. 1:741, fig. 416 (1906) ; Davidson & Moxley, Fl. S. Calif. 206 (1923).
Amelanchier bakeri Greene in Pittonia 4:128 (1900); Schneider, Illustr. Handb.

Laubh. 1:742, fig. 416 (1906); Rydberg, Fl. Colorado 191 (1906); Coulter &
Nelson, New Man. Rocky Mt. Bot. 266 (1909) ; Wooton & Standley in Contr.
U.S. Nat. Herb. 19:323 (1915) ; Rydberg, Fl. Rocky Mts. 447 (1917) ; Garrett,
Spr. Fl. Wasatch Reg. (ed. 5) 106 (1936); Kearney & Peebles in U.S. Dept.

Agric. Misc. Publ. 423:393 (1942).

Amelanchier rubescens Greene in Pittonia 4:128 (1900) ; Schneider, Illustr. Handb.
Laubh. 1:740, figs. 413, 414 (1906); Rydberg, Fl. Colorado 191 (1906); Bean
Tr. & Shr. Brit. Isles 1:190 (1914); Wooton & Standley in Contr. U.S. Nat.
Herb. 19:322 (1915); Tidestrom in ibid. 25:284 (1925); Tidestrom & Kittell,
Fl. Ariz. & N. Mex. 251 (1941).

Amelanchier crenata Greene in Pittonia 4:127 (1900); Schneider, Illustr. Handb.
Laubh. 1:742, figs. 415, 416 (1906); Wooton & Standley in Contr. U.S. Nat.
Herb. 19:323 (1915) ; Rydberg, Fl. Rocky Mts. 447 (1917).

Amelanchier rubescens var. cinerea Goodding in Bot. Gaz. 37:55 (1904).

Amelanchier elliptica A. Nelson in Bot. Gaz. 40:66 (1905) ; Rydberg, Fl. Colorado
191 (1906) ; Schneider, Illustr. Handb. Laubh. 1:739 (1906) ; Coulter & Nelson,
New Man. Rocky Mt. Bot. 266 (1909).

Amelanchier oreophila A. Nelson in Bot. Gaz. 40:65 (1905); Schneider, Illustr.
Handb. Laubh. 1:740, fig. 414 (1906) ; Rydberg, Fl. Colorado 191 (1906) ; Coul-
ter & Nelson, New Man. Rocky Mt. Bot. 266 (1909); Wooton & Standley in
Contr. U.S. Nat. Herb. 19:323 (1915); Rydberg, Fl. Rocky Mts. 447 (1917);
Tidestrom in Contr. U.S. Nat. Herb. 25:283 (1925); Kirkwood, N. Rocky Mt.
Tr. & Shr. 189 (1930); Garrett, Spr. Fl. Wasatch Reg. (ed. 5) 106 (1936) ;
Tidestrom & Kittell, Fl. Ariz. & New Mex. 251 (1941); Kearney & Peebles in
U.S. Dept. Agric. Misc. Publ. 423:394 (1942).

Amelanchier mormonica Schneider, Illustr. Handb. Laubh. 1:740, fig. 414, n-o
(1906), in Fedde, Rep. Sp. Nov. 3:182 (1906); Wooton & Standley, Contr.
U.S. Nat. Herb. 19:323 (1915); Rydberg, Fl. Rocky Mts. 447 (1917); Garrett,
Spr. Fl. Wasatch Reg. (ed. 5) 106 (1936); Kearney & Peebles in U.S. Dept.
Agric. Misc. Publ. 423:394 (1942).

Amelanchier alnifolia var. elliptica Schneider, Illustr. Handb. Laubh. 1:739 (1906).

Amelanchier jonesiana Schneider in Fedde, Rep. Sp. Nov. 3:182 (1906) ; Rydberg,
Fl. Rocky Mts. 447 (1917).

Amelanchier goldmani Wooton & Standley in Contr. U.S. Nat. Herb. 16:131 (1912).

Amelanchier australis Standley in Proc. Biol. Soc. Wash. 26:116 (1913).

Amelanchier covillei Standley in Proc. Biol. Soc. Wash. 27:198 (1914); Wooton &
Standley in Contr. U.S. Nat. Herb. 19:323 (1915); Tidestrom in ibid. 25:283
(1925) ; Abrams, Illustr. Fl. Pac. States, 2:471, fig. 2534 (1944).

Amelanchier purpusi Koehne in Engler, Bot. Jahrb. 52:278 (1915); Rehder, Man.
Cult. Tr. & Shr. 391 (1927), (ed. 2) 389 (1940).

Amelanchier plurinervis Koehne in Bot. Jahrb. 52:278 (1915); Rehder, Man. Cult.
Tr. & Shr. 391 (1927), (ed. 2) 389 (1940).

Amelanchier mtens Tidestrom in Proc. Biol. Soc. Wash. 36:182 (1923), in Contr.
WIS. Nat: Herb: 25:283 :(1925).

88 ILLINOIS BIOLOGICAL MONOGRAPHS

Amelanchier alnifolia var. covillei Jepson, Man. FI. Pl. Calif. 510 (1925), Fl. Calif.

2:234 (1936) ; Munz, Man. So. Calif. Bot. 229 (1935).

Amelanchier alnifolia var. venulosa Jepson, Man. Fl. Pl. Calif. 510 (1925), FI.

Calif. 2:234 (1936); Munz, Man. So. Calif. Bot. 229 (1935).

Amelancher alnifolia var. nitens Munz in Bull. So. Calif. Acad. Sci. 31:65 (1932).
Amelanchier utahensis subsp. covillei Clokey in Madréno 8:57 (1945).
Amelancher utahensis subsp. oreophila Clokey in ibid. 58.

Shrubs or small trees 0.5-5 m. tall, much-branched, bushy, often grow-
ing in clumps; twigs rigid, the bark ashy gray, those of the season per-
sistently copiously white-sericeous or lanate, rarely nearly or quite gla-
brous, the 2- or 3-year-old twigs also often pubescent; winter buds usually
copiously pubescent, or on occasional specimens glabrous; leaves usually
small, commonly suborbicular to oval, ovate, or obovate, grayish green, or
pale and glaucescent, finely tomentulose or cinereous on both surfaces
even at maturity, with soft, curled trichomes, varying to completely
glabrous, subcoriaceous at maturity, conduplicate in the bud, unfolding
before or with the flowers and more than half-grown at flowering time;
mature blades 0.5-3 cm. long, 0.5-2 cm. wide, the apex rounded or trun-
cate, or even emarginate, or often acute and mucronate, the base rounded
or truncate, or sometimes cuneate; lateral veins 11-13 pairs; margins
coarsely and often sharply dentate to the middle or below varying to
subentire, or crenately few-toothed toward the apex; the teeth mostly 3-5
per cm., 3-10 on each side of average leaves of the flowering and fruiting
branches; stipules linear, pubescent; petioles slender, 5-18 mm. long,
flowers small, sometimes reddish in the bud, in erect or ascending, 3-6-
flowered racemes 2-3 cm. long, sometimes subcorymbose as the result of
the elongation of the lower pedicels; rachis and pedicels lanate (rarely
glabrous), the pubescence usually persistent on the fruiting specimens;
petals 5, white, linear-oblanceolate or cuneate, 6-7 (-9) mm. long, 1.5-2.5
mm. wide; stamens 10-15 (-18), shorter than or equalling the styles, the
filaments glabrous, 1-2 mm. long; anthers 0.7-0.9 mm. long; hypanthium
shallowly campanulate or funnelform, 3-4 mm. in diameter, tomentulose,
varying to completely glabrous, not at all constricted on the young fruit;
sepals linear or linear-lanceolate, acuminate, thickish, tomentulose on both
sides (or occasionally glabrous), 3 mm. long, soon recurved from the
middle and often becoming somewhat elongated after anthesis, or remain-
ing erect or spreading on the fruit; styles 4 or 3, or only 2 (rarely 5),
glabrous, united near the base, 2.5-3 mm. long; summit of the ovary more
or less tomentulose (rarely quite glabrous), tapering into the style-base;
mature fruits few, 6-10 mm. in diameter, 3-6-loculed, usually puberulent
when young, purplish black and juicy at maturity, or often remaining
dry, insipid, pale brown and leathery, and drying on the bush without
ripening; pedicels 2-3 cm. long, usually pubescent; seeds 4-6 in each

AMERICAN SPECIES OF AMELANCHIER—JONES 89

pome, brown, microscopically puncticulate, flattened-ellipsoid, obtuse at
each end, about 5-6 mm. long and 3 mm. wide when mature and fully
developed.

Tyre Locauity: “Utah, Belleview [Pintura, Washington Co.], alt.
3000 ped., leg. Marcus E. Jones n. 1716... Vidi in herbario regio
Berolinensi.” Isotype in the herbarium of the New York Botanical Garden.

Rance: Dry rocky slopes and canyons, banks of creeks, mountain
sides, foothills, and deserts, from the Sonoran to the Transition zones,
4000 to 8000 feet altitude, New Mexico to southern Montana, westward
to eastern Oregon and Lower California. Flowering from the beginning of
April to the end of May; fruit ripening from July to September, according
to the altitude.

OrecoN: Vale, Malheur Co., Letberg
2193 (GH); Lake Abert, Eggleston
WAGON CGE),

IpAHO: Salmon, Payson & Payson
1812 (GH); Big Willow, Macbride 127
(GH, MBG); King Hill, Nelson &
Macbride 1110 (GH, MBG); Corral,
Macbride & Payson 2880a, 2880b (GH,
MBG); Soldier Mts., Macbride & Pay-
son 2904 (GH, MBG); House Creek,
Nelson & Macbride 1798 (GH, UI,
MBG) ; Twin Falls, Nelson & Macbride
1327, 1347 (GH, MBG); McCammon,
Muenscher & Maguire 2362 (MBG);
Lava, Nelson & Macbride 1593 (GH) ;
Montpelier, Macbride 13, 24, 209 (GH,
UI, MBG).

Montana: Wilsall, Suksdorf 269
(AA).

Wyominc: Chimney Rock, Green-
man ¢é Greenman 6031, 6048 (MBG);
Telephone Canyon, Porter 1059 (MBG) ;
Elk Mt., Goodding 555 (AA, GH, Map 14.—Range of Amelanchier
MBG); Rawlings, Payson & Payson utahenstis.

2540 (GH, MBG); Point of Rocks,

Merrill & Wilcox 458 (GH); Granger,

Hanna 955 (MBG); Rock Springs, Hanna 715 (MBG); Centennial, Nelson 9317
(GH), Jack 1075, 1089 (AA); Cumberland, Nelson 9010 (MBG); near Lyman,
Rollins 198 (MBG); Mountainview, Nelson 11344 (GH); Bear River, Pammel &
Blackwood 4063 (AA, GH, MBG); Rock River, Nelson 9333 (GH, MBG); Bates
Hole, Payson 4779 (MBG), Solheim 434 (UI).

CoLtorapo: Willow Creek, Goodding 1683 (GH, MBG); Camp Creek, Goodding
1456 (GH, MBG, ISOTYPE of A. oreophila); Rangeley, Graham 9074 (GH);
Buford, Hermann 5688 (GH, MBG); Breckenridge, Mackenzie 240 (MBG); Bear
Creek, June 13, 1918, Churchill (GH, MBG); Golden-Central City, Duthie &
Clokey 3795 (GH, MBG, UI); White Rocks, May 12, 1918, Andrews (AA); near
Boulder, Hanson C226 (MBG); Mesa Grande, May 1892, Purpus (AA); Paonia,
Eggleston 14572, 14959 (AA), 14574 (GH); Cedaredge, Payson 1066 (MBG) ;
Cerro Summit, Baker 47, 55, 163 (GH, MBG); near Cimarron, Baker 139, 210,

90 ILLINOIS BIOLOGICAL MONOGRAPHS

(GH, MBG); Cedar Creek, Baker 438, 439 (GH, MBG); Gunnison Mesa, East-
wood 5095 (AA); Naturita, Payson & Payson 3883 (GH, MBG), Payson 248 (GH,
MBG), Payson 982 (MBG); Nucla, Payson & Payson 4225 (GH, MBG); Sheep
Creek Canyon, Payson & Payson 3884 (GH, MBG); Bedrock, Payson & Payson
4228 (GH, MBG); Paradox, Walker 210 (GH, MBG); near Ridgway, Payson
1077 (MBG), 2307 (GH, MBG); Norwood Hill, Walker 416 (GH, MBG); Pagosa
Springs, McKelvey 4721 (AA); La Veta Pass, Hooker & Gray in 1877 (GH); Los
Pinos (Bayfield), Baker 376 (GH, MBG; ND, TYPE of A. bakeri) ; Durango, East-
wood 5273 (AA); Bob Creek, Baker, Earle, & Tracy 197 (GH, MBG) ; Mesa Verde
Nat. Park, Munz 13046 (AA); Mancos, Baker, Earle, & Tracy 665 (Minn.; ND,
TYPE of A. prunifolia).

UrtaH: Peterson, Pammel & Blackwood 3876 (GH, MBG), 3887 (AA, GH);
Goodman Ranch, Bear River Valley, Hermann 5813 (MBG); Big Cottonwood
Canyon, Rydberg & Carlton 6662 (GH); Springville, May 17, 1913, Hill (MBG);
Red Butte, May 6, 1909, Clemens (GH, AA); Salt Lake City, Palmer 38037 (AA,
MBG); Farmington Canyon, Pammel & Blackwood 3623 (GH, AA); Soldier
Summit, Eastwood 7262, 7263 (AA); Duchesne Co., Graham 9407 (GH), 8061,
9284, 9363 (MBG); Green River, Graham 9202 (GH), 9196 (MBG); Diamond Mt.,
Graham 8074 (MBG); Ashley Creek, Graham 6289 (MBG) ; near Watson, Graham
9006 (MBG); Dry Fork, Graham 8810 (MBG) ; near Vernal, Graham 8749 (MBG) ;
Dinosaur Quarry, Graham 7659 (MBG); near Helper, McKelvey 4267 (AA);
Cottonwood Canyon, Graham 9519 (MBG); Filmore Forest, Eggleston 14164
(GH); Sevier Desert, Harris C20491 (GH); Cottonwood Creek, Cutler 2372
(MBG); Stansbury Mts., Harris C20i112 (MBG); Stansbury Island, Garrett 5356
(AA); Green Lakes, Hermann 4836 (GH, MBG); Uinta Mts., Williams 577 (GH,
MBG); Milford, Goodding 1036 (AA, GH, MBG, ISOTYPES of A. elliptica) ;
La Sal Mts., Purpus 6523 (MBG), Maguire & Redd 1907 (GH); Cooper Canyon,
Cutler 2297 (MBG); Natural Bridges Nat. Mon., Maguire & Redd 1905 (GH);
Elk Ridge, Maguire & Redd 1911 (MBG); Colorado, Cutler 2651 (MBG); Abajo
Mts., Rydberg & Garrett 9274 (GH); Post Canyon, Graham 9896 (GH); Modena,
Goodding 1008 (AA, GH, MBG); Cedar City, Tidestrom 9429 (GH); 50 miles
s.w. of San Raphael, Harrison 7439 (MBG); Zion Nat. Park, Allen in 1934, Fisk
in 1930 (MBG); Virgin, Maguire 4840 (GH, MBG); Rockville, M. E. Jones 5224
(MBG); near St. George, Tidestrom 9359 (GH), Goodding 780 (MBG, TYPE
coll. of A. rubescens var. cinerea); near Glendale, M. E. Jones 25393 (MBG),
Nelson & Nelson 2890 (GH); Belleview, April 21, 1880, M. E. Jones 1716 (NY,
TYPE coll. of A. utahensis).

Nevapa: Hunter Creek, Kennedy 1866 (GH, AA); Peavine Mt., Heller 9718
(GH); near Alum Creek, Heller 9739 (AA); Fish Lake, Heller 9995 (AA, GH,
MBG), 10004 (AA, GH); W. Humboldt Mts., Heller 10630 (GH, MBG); Battle
Mts., June 1868, [Vatson 353 (GH); near Cave Creek, Mason 4725 (GH), Heller
9510 (MBG); Jarbidge, Nelson & Macbride 1939 (GH), 1935 (GH, MBG, UI),
2120, 2121 (GH, MBG), Eggleston 14105 (GH); near Deeth, Heller 10576 (GH,
MBG, AA), 10567 (GH), 9108 (AA, MBG), Eastwood & Howell 310 (AA),
Maguire 5772 (GH); Lamoille Creek, Heller 9310 (MBG); Oasis, Palmer 38025
(AA, MBG); E. Humboldt Mts., Aug. 1868, Watson 353 (GH); near Ely, Tide-
strom 11050 (AA) ; Carson City, June 2, 1897, M. E. Jones (MBG); Kings Canyon,
Baker 946 (AA, GH, MBG); Glenbrook, Baker 1002 (AA, GH, MBG); White
Mts., Duran 752 (GH, MBG); near Panaca, Tidestrom 9460 (AA); Kyle Canyon,
Charleston Mts., Tidestrom 9600 (GH), Clokey 7141, 7142 (GH), 7541, 7542 (GH,
MBG, UI); Wilson’s Ranch, Charleston Mts., Clark Co., Clokey 7970 (GH), 8236,
8237 (GH, UI), TOPOTYPES of A. nitens; Wells, July, 1893, E. L. Greene (ND,
TYPE of A. pallida var. arguta).

CaLiForNIA: Portola, Eastwood 7017 (AA); Providence Mts., Munz, et al. 4147
(GH); Panamint Mts., Mung 14820 (MBG); Argus Peak, Purpus 5376 (GH,
MBG); Old Dad Mts., WM. E. Jones 25392 (MBG); Wrightwood, Newsom in 1932
(GH, MBG); Cushenberry Springs, Bear Valley, San Bernardina Mts., Parish

AMERICAN SPECIES OF AMELANCHIER—JONES 91

453, 3383 (MBG), 1290 (GH, TYPE coll. of A. venulosa), 10882 (GH, MBG) ;
Bear Valley, Abrams 2884 (GH, MBG); Deep Creek, Abrams 2041 (AA); Sugar-
loaf Peak, Fosberg 8605 (GH); San Antonia Peak, M. E. Jones 29011 (MBG),
Parish 11969 (GH, MBG); Laguna Mts., Eastwood 9212 (AA).

Arizona: Peach Springs, Wilson in 1893 (GH); Grand Canyon, Rehder 115,
116 (AA), Eastwood 5886, 5948 (AA), Nelson & Nelson 2024 (MBG); McKelvey
4395 (AA), Eastwood 6047, 6086 (AA, GH) ; between House Rock and Jacob Lake,
Nelson & Nelson 2877 (GH, MBG); 7 miles northwest of Jacobs Lake, Peebles &
Parker 14670 (GH); 18 miles north of Inscription House Trading Post, Peebles
13920 (GH); Mormon Lake, Smith 12005, MacDougal 102 (GH, TYPE coll. of
A. mormonica); Kendrick Peak, June 22, 1911, Lowell (AA); near Flagstaff,
McKelvey 1314 (AA), Lowell in 1915 (MBG), Hanson 626 (MBG); Williams,
Nelson 10245 (MBG); between Williams and Ash Fork, McKelvey 915 (AA); Mt.
Agassiz, May 1916, Pearson (AA); between Tunitcha and Luka-Chukai Mts.,
Goodman & Payson 2797, 2846 (MBG); Navajo Indian Reservation, Voorhies 82
(GH, AA, MBG); Matazal Mts., McKelvey 1034, 1138 (AA), Collom 488 (MBG) ;
Verde R., between Payson and Pine, McKelvey (AA); Devils Canyon, between
Miami and Superior, Nelson & Nelson 1888 (MBG); Camp Grant, E. Palmer 94
(MBG).

New Mexico: Above Toadlena, McKelvey 4626, 4630 (AA); Aztec, Baker 377
(GH, AA, MBG, TYPE coll. of A. crenata), Baker 380, 381 (GH, MBG; ND,
TYPE of A. rubescens) ; Chama, Wolf 2988 (GH); Lake Burford, Wetmore 470
(AA); Tierra Amarilla, Eggleston 6575 (GH, MBG); near Gallup, McKelvey 2316,
Eastwood 5609 (AA); Pecos River, 8 miles east of Glorietta, Heller 3679 (AA,
GH, MBG); Winsor Creek, Standley 4125 (GH, MBG); Mogollon Mts., Aug. 8,
1900, Wooton (MBG); Kingston, Metcalfe 961 (GH, MBG); near Silver City,
Greene in 1880 (MBG), Metcalfe 620 (GH, MBG).

Texas: Ridge above McKittrick Canyon, Guadalupe Mts., Moore & Stevyer-
mark 3482 (GH, MBG).

Mexico: San Pedro Martin, Baja California, May 26, 1893, Wiggins & Demaree
4948 (FM, UC), T. S. Brandegee (UC).

This common Amelanchier, occurring throughout the mountainous
and desert regions of western United States, is an extremely variable
species, for which the oldest name is 4. utahensis Koehne, applied in 1890
to specimens collected near the geographical center of its range, in south-
western Utah. Its variability is indicated by the fact that during the half-
century that has elapsed since this species was first recognized, it has been
described and redescribed under approximately two dozen different names.
In its typical form, it is a twiggy shrub with rather small, permanently
pubescent, conspicuously toothed leaves that are subcoriaceous at maturity.
The branchlets may be pubescent for two or three years. Shade forms
have larger, thinner leaves, and tend to approach A. alnifolia in general
appearance, but such specimens can usually be distinguished by the
fewer styles and stamens, and the narrower, often somewhat elongated
sepals. One of the vernacular names used for this species is “red service-
berry,” because in some localities the flower buds are pink. It is some-
times abundant locally, and is said to be palatable as a browse plant in
the spring.

An Amelanchier inhabiting the arid slopes and plains of parts of
southern California, including shrubs with rather prominently veiny

92 ILLINOIS BIOLOGICAL MONOGRAPHS

leaves, was described by E. L. Greene in 1899 as A. venulosa. However,
it lacks morphological characters to distinguish it from A. utahensis.
Greene recognized the fact that it is different from A. pallida in its more
venulose leaves, but was apparently unaware that it had been published as
A. utahensis nine years earlier. The specimens described as A. rubescens
Greene, collected “In arroyos and among hills about Aztec, New Mexico,
24 April 1899, C. F. Baker” (nos. 380, 381), are also clearly conspecific
with A. utahensis. Likewise, A. crenata Greene, also from Aztec, New
Mexico, and said by its author to be “altogether peculiar in the crenate
character of its leaf indentation” is merely a phase of the widespread and
variable A. utahensis. The plants described as A. rubescens var. cinerea
by L. N. Goodding in 1904 from near the type locality of A. utahensis,
represent merely a pubescent extreme of that species. A. plurinervis
Koehne was described from plants cultivated in Berlin, but collected by
C. A. Purpus in western United States. Specimens distributed from Arb.
Spaeth., now in the herbarium of the Arnold Arboretum, belong quite
evidently to A. utahensis.

In 1905, Professor Aven Nelson, concluding that “either A. alnifolia
was unusually variable or that some segregation ought to be made,”
described the common western xerophytic shrub with puberulent leaves,
fewer styles and stamens, and smaller fruits, as A. oreophila, apparently
without knowing that this species had been described as A. utahensis,
from southwestern Utah fifteen years earlier by E. Koehne. Nelson’s
comments in Bot. Gaz. 40:66 (1895) are as follows:

After many years’ observation in the field and the study of a large series of
specimens, I am satisfied that two valid species exist and can be readily dis-
tinguished. Nuttall’s A. alnifolia is the widely distributed glabrous shrub of the
creek banks, moist canyons, and snow slopes. ... The leaves are larger, coarsely
serrate, often suborbicular or with a tendency to truncateness at base and apex.
... The fruits become much larger, are purple, with bloom, juicy and well flavored,
and are used extensively for sauces and pies, maturing during July or August,
according to the altitude. A. oreophila [A. utahensis] is a smaller shrub, scraggy-
branched, usually in dense clumps, and occurring in the driest situations (open
stony slopes, ridges, and hilltops). It is never wholly glabrous, and the fruit is of
little if any value... . Much of the material distributed from the Rocky Mountains
belongs to this species.

In 1923, Dr. Ivar Tidestrom described, as A. nitens, a glabrous shrub
with glossy foliage and yellow fruits occurring in the Charleston Moun-
tains of southwestern Nevada. In its extreme glabrous form, this is a
plant of distinctive appearance, but it intergrades completely with the
typically pubescent forms of the widespread A. utahensis, and it is there-
fore not practicable to attempt to accord it nomenclatural recognition.
Even on topotypes collected in 1938 and 1939 by Mr. I. W. Clokey (e.g.,
Clokey 8236, 8237, UI) some specimens are almost perfectly glabrous,

AMERICAN SPECIES OF AMELANCHIER—JONES 93

while others are distinctly pubescent on the blades, petioles, pedicels and
sepals. This glabrous or semi-glabrous form has no particular geographi-
cal distribution, but occurs sporadically in various intergrades, in several
parts of the range of A. utahensis. Dr. Rogers McVaugh’ reports his
field observations of these plants as follows:

In desert and semi-desert areas from Clark Co., Nevada, throughout the Grand
Canyon region, there is an abundance of a glabrous or nearly glabrous form with
small lustrous leaves, which seems to be otherwise identical with Form C [1e., A.
utahensis]. This is the plant described as A. nitens Tidestrom, which was said by
its author to have yellow fruit (“pomis maturis aureis”). In July 1941 I was able to
visit Wilson’s Ranch, the type locality of this species, and to collect mature fruits.
As in many similar localities in the southern Great Basin, most of the fruits had
dried at maturity, leaving the seeds to rattle within. On a few bushes, however, I
was able to find an occasional berry [pome] which was still soft and slightly fleshy;
all these were pale, nearly white, with a purplish-red cheek; they were in fact
identical with the fruits of A. utahensis from other parts of its range.

There seems to be no evidence that yellow or orange fruits, usually
ascribed to certain western American species, ever actually occur, except
perhaps as a result of the attacks of fungous diseases. In drier habitats, it
commonly happens that the fruits are small, dry and leathery. Frequently
they never ripen, or when they do ripen they remain on the bush and
retain a pale brown leathery appearance. Several rust fungi, particularly
Gymnosporangium, parasitize various species of Amelanchier. It may be
that fruits heavily infected with this fungus, which gives them a yellow-
ish appearance, have been the cause of the widespread misconception
regarding the color of the fruit of some western species. The fruits of all

species of Amelanchier are normally bluish black at maturity.
IMcVaugh, Rogers, Contributions Toward a Flora of Nevada, No. 22, pp. 95-96, June 15,

1942. Issued (mimeographed) by The Division of Plant Exploration and Introduction, Bureau
of Plant Industry, U.S. Department of Agriculture, Washington, D.C.

lV. List? OF NUMBERED BEXSIGGA TAw

Abbe, E. C.—1136, 1151 (1).

Abrams, LeRoy—2041, 2884 (18) ; 3912,
4577, 4675, 4809, 4817, 5577 (17);
8718 (13).

Abrams, LeRoy, & McGregor, E. A.—
107, 490 (17).

Adams, J.—339 (7).

Adams, J., & Wherry, E. T.—4662 (6).

Aiton, G. B.—1004 (9).

Allard, H. A.—220, 2516, 2584, 2819, 4342,
4498 (6); 4353 (5); 1229 (7).

Allen, O. D.—214 (13).

Anderson, E., & Anderson, D. M.—26042
(11).

Applegate, E. I—6219 (12).

Arséne, L.—310 (5); 311 (1).

Asthur, J. °C.,. Bailey;L. Hi, é& Holway,
E. W. D.—B407 (1).

Bacigalupi, R—1549 (17).

Bailey, L. H.—56 (11).

Baker, C. F—47, 55, 139, 163, 210, 376,
377, 380, 381, 438, 439, 946, 1002 (18);
AD 379 “ClOds 750. C12) = 2952-1219
2964 (17).

Baker, (C2, Earle; (FS) So Eracy,
S. M.—197, 665 (18).

Balser, G—775 (9).

Barber, M. A.—90, 194 (12).

Barkley, F. A., & Osburnson, L.— 2313
(14)

Bartlett, H. H.—846 (9).

Bartram, E. B.—3220 (8).

Batchelder, C. F—3 (5).

Bates, J. M.—5992 (6); 1357, 5928, 6074

(12).

Bean, R. C., & Fernald, M. L.—17013,
17014 (9).

Bean, R. C., & Knowlton, C. H.—12070e
Gliae

Bean, R. C., White, D., & Linder, D. H.
—21459 (1).

Beattie, R. K—1819 (12).

Benner, W. M.—4883 (6); 6708 (4);
2393* 9567 (5) 3 2703-°(8) = 2926, 7533
(9).

Benson, L.—1283 (14) ; 1420, 1423 (13a).

Bergman, H. F.—1376 (9).

Bicknell, E. P.—4835 (5); 4815, 4816,
4840, 4847, 4849, 4850, 4851, 4857,
4858, 4862a, 4868, 4879 (7).

Biltmore Herbarium—5664, 5664c, 56644d,
5664e (10) ; 6706 (9).

Bishop, H.—370, 371, 373 (1).

94

Bissell, C. H., & Graves, C. B—21457
CZ)’.

Bissell, C. H., Pease, A. S., Long, B., &
Linder, D. H—21440 (7).

Blake, S. F.—2496, 5677 (9) ; 9324, 9362
G/):-7 10557. (6).

Blake, S. F., & Fernald, M. L.—3645 (9).

Blanchard, W. H.—3, 4 (9); 4 (10);
POD

Blankinship, J. W.—135 (12).

Blewitt, A. E.—1508, 1801 (7); 1511,
1792, 1794, 1795, 1796; 20352037,
(9) ; 1512, 1797, 1798 (5) ; 1802, 1804,
1805, 2036, 2038, 2039, 3501, 3650 (6).

Bolander, H.—4674 (13).

Boner, L., & Weldert, V.—172 (14).

Bowman, P. W.—26, 237, 402 (1).

Breckenridge, W. J., & Nielsen, E. L—
3161 (4).

Breckenridge, W. J., Nielsen, E. L., &
Moore, J. W.—3229 (4).

Breitung, A. J—523 (12).

Brewer, W. H.—576 (17).

Brown, H. E.—78 (12); 557 (17).

Brown, S.—23, 56 (12); 216 (9); 283
(GlgD)e

Burnham, S. H.—11 (7); 12 (5); 21
(3) EU 1162 (6):

Bush, B. F.—30, 85, 85a, 85b, 85c, 1602,
1602a, 3509, 10380, 15250 (6).

Butters, F. K.—1342 (4).

Butters, F. K., & Rosendahl, C. O—
1358 (12).

Byhouwer, J. T. P., & Kobuski, C. E—
50 (7).

Carr, W. P.—75 (12).

Chamberlain, E. B.—34, 52, 533 (5); 62,
228 (9).

Chandler, H. P.—1282 (17).

Chase, Agnes—197, 1047 (5); 702, 709,
990, 1048, 1745 (9) ; 2035, 2053 (6).

Chase, V. H.—1795 (6).

Cheney, L. S—4447 (4).

Child, H. W.; Knowlton, GC. Hi., Bird;
F. W., & Bean, R. C—16377 (5).

Clark, J. A—166 (12).

Clokey, I. W.—7141, 7142, 7541, 7542,
7970, 8236, 8237 (18).

Collom, R. E—488 (18).

Congdon, J. W.—31 (17).

Constance, L.—999 (14).

Constance, L., Clements, H. F., & Mach-
lis, L—1008 (14).

AMERICAN SPECIES OF AMELANCHIER—JONES 95

Constance, L., Machlis, L., Rogers, B.,
& Rollins, R. C.—1037 (14).

Constance, L., & McMurray, R. L.—1135
GAs

Constance, L., & Rollins, R. C—1510
(12) ; 1499, 1512 (14).

Constance, L., Rollins, R. C., & Dillon,
L. A.—1568 (15).

Cooper, W. S.—46 (1); 122, 124, 125
(CUB!

Cotton, J. S—365, 569, 571, 576 (12).

Cowles, H. C.—38, 38a (12) ; 1419 (13).

Cronquist, A.—920, 2298, 3778 (12);
2596 (16).

Cronquist, A., & Davis, R. J.—2099 (12).

Culbertson—4647 (17).

Cushman, J. A.—871, 4330 (10).

Cushman, J. A., & Sanford, S. N. F.—
1217).

Cusick, W. C.—1858 (14).

Cuthbert, A—1 (7).

Cutler, H. C—2297, 2372, 2651 (18).

Daniels, F.—909 (16).

Davis, J—49, 113, 704, 1259, 1462, 2011,
2227, 4646 (6).

Day, M. A—57, 75 (7); 379 (3).

Deam, C. C.—12868, 16116, 33798 (5);
22524, 23005 (6) ; 23107, 33770, 38194,
38195, 38196, 38251, 39083 (9).

Demaree, D.—11245, 11260, 18743 (6).

Dodge, C. K—20, 57, 71, 73 (8); 74,
76 (10).

Drew, W. B., Hodgdon, A. R., & Taylor,
F.—2472 (10).

Drouet, F.—1421 (6).

Dudley, W. R.—56 (7).

Dunbar, J.—10 (9); 12 (10).

Duncan, W. H.—3282 (5).

Duran, V.—752 (18).

Duthie, R., & Clokey, I. W.—3795 (18).

Eames, A. J., & Dean, E—4293 (6).

Eames, A. J., & MacDaniels, L. H—4285
(9).

Eames, E. H.—1 (5); 4287, 4288 (9);
8287 (6).

Farle, F. S., & Baker, C. F—1610 (6a).

Eastwood, A.—72, 785, 1081, 11931,
12076 (17) ; 269, 7706 (12) ; 372 (16);
995, 9702 (13) ; 5095, 5273, 5609, 5886,
5948, 6047, 6086, 7017, 7262, 7263, 9212
(18).

Eastwood, A., & Howell, J. T.—310
(18) ; 4800 (17).

Eggleston, W. W.—185, 1176, 1179, 1180
CO) 8s 1173,1174,.1175 (3) : 1121,
M22, 1127; 7 1178,-1971,, (G10) ©1182)

Eggleston, W. W. (continued)
‘1183, 1957, 1959 (6); 1960, 1962, 1964,
2369 (1); -6235,-7596, (17s 7066
(16) ; 6575, 7116, 14105, 14164, 14572,
14574, 14959 (18); 14967 (12).

Ehlers, J. H.—323 (10); 1183 (9).

Eimer, A; D, E135 (14); 2512 (13);
4659 (17).

Epling, C—5623 (13).

Evers, R. A.—52 (6).

Eyerdam, W.—1377 (14).

Eyles, D.—6894 (6).

Farwell, O. A——52d (1).

Fassett, N. C—446, 2826, 13745 (5);
2818, 2819, 2820, 2821, 2822, 2823,
13775" (9)'s 2825, 3051, 7128 (4) + 7313
(10).

Fassett, N. C., & Schmidt, J. F. W—
15708 (9).

Fassett, N. C., Steyermark, J. A, &
Tryon, R. M.—18359 (9).

Fernald, G. B—43 (9).

Fernald, M. L.—37, 2312, 11719 (3);
105, 1881, 1888, 2644, 9620, 9626,
13760; 13773, 13774, 13775, 15197,
15204, 16867 (9) ; 257, 2314 (1) ; 9624
(5) ; 388, 449, 1880, 2310, 2311, 13778,
13/79, 15537 (10); 1885,, 13767 (6):
15191, 15198, 15199, 18546 (7).

Fernald, M. L., Bartram, E. B., & Long,
B.—23930, 23931 (9); 23937 (7);
23943, 23944, 24761 (5).

Fernald, M. L., Bartram, E. B., Long, B.,
& Fassett, N. C.—23929 (9).

Fernald, M. L., Bartram, E. B., Long, B.,
& St. John, H.—7553, 7582, 7591 (9);
7581 (3); 7586, 7587, 7589 (2).

Fernald, M. L., & Bean, R. C.— 14132
(9); 14160 (7).

Fernald, M. L., & Bissell, C. H.—21433
(9).

Fernald, M. L., Bissell, C. H., Graves,
C. B., Long, B., & Linder, D. H.—
21441, 21442 (5).

Fernald, M. L., Bissell, C. H., Pease,
A. S., Long, B., & Linder, D. H.—
21432 (9).

Fernald, M. L., & Collins, J. F—233, 614,
1100 (1).

Fernald, M. L., Dodge, C. W., & Smith,
L. B.—25840 (11).

Fernald, M. L., & Griscom, L.—4425 (6).

Fernald, M. L., Griscom, L., & Long, B.
—4650 (6).

Fernald, M. L., Griscom, L., Mackenzie,
K. K., & Smith, L. B—25839 (11).

96 ILLINOIS BIOLOGICAL MONOGRAPHS

Fernald, M. L., & Hunnewell, F. W.—
15192 (5); 15200 (7).

Fernald, M. L., & Jackson, H. B.—10107
Ge

Fernald, M. L., & Long, B.—3959, 7063,
7064, 7065, 7067, 7069, 7450, 8291,
9621, 11698, 11844, 11845, 13039, 13040,
13041 (6); 7068, 7070, 7071, 7869,
9625, 9949, 11343, 12096, 13772, 16869,
18542, 18543, 23946 (7); 7072, 7073,
7074, 7448, 7449, 7870, 9947, 11846,
11847, 13042, 13043, 13950 (8); 9623,
13763, 21451 (5); 13762, 13766,13777,
13781, 14178, 18548, 18549, 18551,
23932, 23933, 23934 (9); 13764, 13768,
13769, 21450 (3); 13780 (10) ; 13782
(1):

Fernald, M. L., Long, B., & Dunbar,
B. H.—26759 (9).

Fernald, M. L., Long, B., & Fogg, J. M.
—290, 1791 (5); 292, 1789 (1); 1792
(2).

Fernald, M. L., Long, B. & Linder,
D. H.—21439 (9); 21456 (7).

Fernald, M. L., Long, B., & St. John, H.
—7584, 7593, 7594, 7595 (9); 7590,
7592 (2); 7596, 7597 (1).

Fernald, M. L., Long, B., & Smart,
R. F.—5790 (8).

Fernald, M. L., & Pease, A. S—3359,
25134 <(9) > 3360; 25133. “C1 3361,
3364, 3365, 3366, 3367 (10); 3369,
2137 (ll) 216825 (3) = 25135 05),

Fernald, M. -L., Pease, A» S.,-&
Long, B.—11700 (7); 21435, 21436,
21437 (9).

Fernald, M. L., & St. John; H—7578
(3); 7585, 11080 (9); 10840, 10841,
10842, 11083 (2); 11082 (5); 11085
GZ).

Fernald, M. L., & Smiley, F. J—11720
Gl):

Fernald, M. L., & Svenson, H. K.—914
(9).

Fernald, M. L., Weatherby, C. A, &-

Stebbins, G. L.—2451 (11).

Fernald, M. L., & White, D—21438 (9).

Fernald, M. L., & Wiegand, K. M.—
3594, 5535, 5574, 5574a, 5576, 5578,
5597, 5600, 5602, 5736, 5737, 5738,
5739, 5742, 5743, 5744, 5745, 5746,
5/48, 0/49, 5/50, . 5751, 5752, _.5754,
5759,;- 9/90, (1); 5542, 5545; 5547,
5552, 5567, 5568, 5569, 5570, 5578a,
5980, 5599, 5/59 (5) + 3553, 5557, 5565,
5558, 5559, 5561, 5562, 5563, 5605,
5608, 5623, 5627, 5630, 5633, 5635 (9).

Fernald, M. L., Wiegand, K. Mei&
Eames, A. J.—14302 (10).

Ferris, R. S., & Duthie, R—77 (7);
411, 784 (12); 656 (15).

Fischbach, C. M.—228 (6).

Flodman, J. H.—545 (16).

Floyd, F. G—801, 803, 980, 1008 (9);
845 (5).

Fogg, J. M.—1863, 4049, 16324 (9).

Fosberg, F. R—8605 (18).

Friesner, R. C.—9541 (6).

Garrett, A. O.—3686 (12); 5356 (18).

Gates,. F. C., & Gates, M. T.—9544,
13914 (10).

Geil, D—4 (14).

Gilbert, F. A—398 (7).

Gilbert, F. A., Rehder, A., & Smith,
L. B.—833 (7).

Gleason, H. A.—708 (5); 1695, 2404 (6).

Godfrey, R. K., & White, R. N.—7027
(9).

Godfrey, R. K., White, R. N., & Shel-
bourne, V.—7038 (7).

Goodale, A. S., & Markert, W. C.—
76864 (9).

Goodding, L. N.—555, 780, 1008, 1036,
1456, 1683 (18).

Goodman, G. J., & Payson, L. B—2797,
2846 (18).

Gorman, M. W.—39 (13); 1026 (12).

Graham, E. H.—6289, 7659, 8061, 8074,
8749, 8810, 9006, 9074, 9196, 9202, 9284,
9363, 9407, 9519, 9896 (18); 9262
(12).

Grant, M. L.—2862 (4).

Greene, E. L.—779 (17).

Greenman, J. M.—936, 1058 (1); 3024
(9). ,

Greenman, J. M., & Greenman, M. T.—
6031, 6048 (18).

Greenman, J. M., Lansing, O. E, &
Dixon, R. A—81 (6).

Grimes, E. J—2560 (6).

Griscom, L.—21578 (6).

Hanna, L.—715, 955 (18).

Hansen, G.—230 (17).

Hanson, H. C.—C225 (12); C226, 626
(18).

Harbison, G. J.—1, 9, 30, 194, 813, 846,
7236, 7240 (9); 2, 510, 913 (5); 4, 6,
11, 15, 30, 40, 1415, 7095 (6) 3-85'((7or

Harger, E. B—3, 10 (5).

Harper, R. M—819 (7); 1806 (6);
3322 (6a).
Harris, J. A—C20112, C20491 (18);

C28699 (16).

AMERICAN SPECIES OF AMELANCHIER—JONES 97

Harris, S. K—423 (7).

Harris, S. K., & Pease, A. S—26551 (3).

Harrison, B. F.—7439 (18); 7473 (12).

Harrison, B. F., & Larsen, E.— 7685
GIG) 47877 C2).

Hay, G.—63 (1).

Hayden, A.—10489 (9).

Heller, A. A.—2988 (15); 3679, 9108,
9310, 9510, 9718, 9739, 9995, 10004,
10567, 10576, 10630 (18); 3061, 3679,
0739", (12) = 3958) (13)-;- 5794, 5961,
6617, 7038, 7419, 7970, 8003, 8531,
10875, 11963, 12729, 12962, 13004,
13156, 13396, 15144 (17); 7176 (16).

Henderson, L. F.—5147 (14); 5824,
13004 (17).

Hermann, F. J.—4836, 5688, 5813 (18) ;
6471, 7259, 10191 (5) ; 6486, 6497 (9) ;
4016, 9625 (6) ; 7791 (10) ; 10048 (7).

Herriot, W.—72329 (12).

Hill, A. F.—169 (16); 1614 (3); 1700
(9).

Hill, E. J—231895 (9); 321889 (5);
411912 (6).

Hitchcock, C. L.—2290 (12) ; 6701 (17).

Hodgdon, A. R.—195, 2244, 2245 (5);
2471, 2998 (10) ; 2600, 2845, 3192 (9) ;
25972773 (1).

Hodgdon, A. R., & Dunn, S.—2772 (5).

Horner, R. M.—B183 (14).

Horsey, R. E.—323 (6).

Hotchkiss, N.—2290 (1).

House, H. D.—7264, 9468, 10227 (1);
7265, 8943, 10175, 17246 (9); 7787,
11870 (6); 7961 (5); 10189 (3);
11201, 16059 (10).

Howe, C. D., & Lang, W. F.—265, 297
CS) 08), 1203 (>.

Howell, T.—1132 (13).

Hubbard, F. T., & Torrey, G. $S—T352
(9).

Hubricht, L—B2011 (6).

Hughes, J. A—1185 (14).

Hunnewell, F. W.—4029, 4128, 6889 (6) ;
4129, 5885 (5) ; 4681 (10) ; 5895 (9);
6031 (7); 2445, 6169 (12).

Hunnewell, F. W., & Wiegand, K. M.—
2137,. 2139, 2140, 2143° (7).

Hunt, K. W.—2969 (8).

Hunt, K. W., & Martin, F.—1408, 2526
(8).

Hyland, F.—699, 753 (7).

Jack, J. G—628, 680, 3187, 3188, 3648,
3671 (5); 684, 3108, 3583, 3639, 3948
(7); 1075, 1089 (18); 1051, 1124a,
125;6 1155, 1185.4 12135; 12589 1336;

Jack, J. G. (continued)

1359, 1361, 1374, 1379, 1453, 1497,
1498, 1511, 1537, 1580, 1592, 2007,
2293, 2540, 2629, 2778 (12); 2865
(13); 3106; #3344 (1); 3209, 3357,
3479, 3514, 3762, 3872, 3946 (9) ; 3928
(10).

Jenney, C. F., Churchill, J. R., & Hill,
A. F.—3264 (6).

Jennings, O. E—1581 (7); 14015 (1);
14024c (11); 14521 (9).

Jones, G. N.—972, 7137 (15) ; 1397, 2057,
4681, 5084, 5640 (12); 4565, 5819
(13a); 1387, 1398, 2832, 6349, 6573,
7088, 7101, 7134, 7147 (14); 7690,
7755 (17) ; 11992, 12089, 13143, 13340,
15618, 15840 (6); 17143 (9).

Jones, G. N., & Jones, F. F.—13716,
137500137815) 153237 ©) 2 16170.-(1).

Jones, M. E—1716, 5224, 29011, 25392,
25393 (18) ; 1447, 6274, 25397 (12).

Kennedy, G. G.—17, 257 (1) ; 1866 (18) ;
2360, 3260- (7).

Kimball, R..H.—102 (1).

Kirkwood, J. E—28, 29, 30, 1182 (14).

Koehler, H. J—1 (9); 2 (5).

Kraus, E. J.—16, 22 (10).

Krotkov, P. V. —5384, 5390 (10) ; 7517 (9).

Lakela, O.—2873 (9).

Lamb, F. H.—1190 (13a).

Leiberg, J. B—1203 (12); 2193 (18).

Letterman, C. W.—2, 3 (6).

Long, B.—6633, 8476, 12515, 12867, 14522,
16257, 18378, 21447, 21448, 30691,
32022, 32349, 33631, 41781, 48444,
50001, 52074, 54418, 57028 (5); 3107,
6586, 9610, 11605, 11883, 11943, 13516,
14514, 14519, 32089, 32282, 37298,
37719, 46255, 46808, 50133, 51868,
52160, 52340, 53736, 58228 (6) ; 26820,
31233;, 32089, -33309, 33330, 33615,
34224, 54350, 57005, 57021, 58240 (7) ;
5915, 16451, 18759, 21005, 21590, 30300,
30603, 30773, 32883, 33071, 35296,
45720, 48268, 48766, 52012, 54322,
54368, 54423, 58269 (8); 12504, 12556,
20586, 25839, 30640, 32642, 37315,
48649, 51076, 51838, 52017, 52022,
52088, 52094, 56467 (9).

Long, B., & Pennell, F. W.—7364, 7374 (8).

Long, B., & St. John, H.—2479 (6).

Long, C. A. E—320, 335, 852 (9).

Loomis, J. A.—856 (9).

Louis-Marie, Fr.—116 (9); 145 (5).

Lucy, T. F.—818b (9).

Lunell, J—50 (14).

98 ILLINOIS BIOLOGICAL MONOGRAPHS

Macbride, J. F—13, 24, 127, 209 (17) ;
852 (14); 926 (12).

Macbride, J. F., & Payson, E. B—2880
(16) ; 2880a, 2880b, 2904 (18).
MacDougal, D. T.—102 (18); 178 (12).
Mackenzie, K. K—240 (18) ; 3080, 4201

(9).

Mackenzie, K. K., & Griscom, L.—5804

(10) 10327~C1) = 1110215) =

MacMillan, C., & Sheldon, E. P.—1747a

(10).

Macoun, J.—208 (13a); 12627 (12);
79796, 93380, 93875, 93879, 93880,
93881, 93883, 93887, 93889, 93895,
93897, 93898, 93899, 93901 (13) ; 19043
(5); 66927 (3); 20074 (10); 34296
(6) ; 34298, 34301, 66924, 80733, 85506
(9).

Macoun, J. M., & Malte, M. O—88016

Gye

Maguire, B—2363, 3501,

4840, 5772 (18).

Maguire, B., & Redd, J. D—1905, 1907,

1911 (18).

Manning, W. E., & Seymour, F. C—
3687 (9).

Marie-Victorin, Fr.—11222, 15586, 24546,
24547 (6); 1883, 11216, 18715 (10);
2087, 8237, 9504, 11221, 11223, 15585,
24545, 24548 (9); 4318 (2); 4319,
9503, 11219 (5); 11214, 15589 (1);
24538, 24558, 28582, 28694 (11).

Marie-Victorin, Fr.; Brunel, J. B., Rol-
land-Germain, Fr., & Rousseau, Z.—
17433, 17436 (1); 17431, 17434,
17435 (11).

Marie-Victorin, Fr., & Rolland-Germain,
Fr.—9507, 18710, 18711, 27898, 27900
(2) ; 18707, 18708, 18709, 27899, 27901
CD) 33110,-33130'. ©).

Marie- Victorin, Fr., Rolland-Germain,
Fr., & Jacques, E.—33203, 33298,
33440 (11) ; 33216, 33436, 33439 (2);
33467, 33486 (1).

Marsh, V. L.—507 (12).

Mason, H. L.—4725 (18).

Mattioli, A. & E—7, 13, 20 (4).

Mattoon, E. A.—13 (6).

McFarland, F. T.—18 (5).

McKelvey, S. D.—915, 1034, 1138, 1314,
2316, 4267, 4395, 4626, 4630, 4721 (18).

Menzel, R. W.—403 (7).

Merrill, E. D., & Wilcox, E. N.—458
(18) ; 548 (16); 1027a (12).

Merrill, G. M.—1965 (6).

Metcalfe, O. B.—620, 961 (18).

Meyer, F. G—226 (15); 854, 1428 (14).

so07 C12);

Milburge, Sister M.—258 (14).

Moffatt, W. S—190 (6); 1610 (9).

Moldenke, H. N.—9502 (5); 10580 (7).

Moodie, M. E.—817, 1039 (12).

Moore, A. H.—4189 (3).

Moore, J. A., & Steyermark, J. A—
3482 (18).

Moore, J. W., & Nielsen, E. L.—3653
(4).

Morton, J. A.—2691 (1).

Moyle, J. B—216 (9).

Moyle, J. B. & Nielsen, E. L.—1944
(4).

Muenscher, W. C.—50 (7); 9635, 15136
(13) ; 11427, 11485 (12).

Muenscher, W. C., & Clausen, R. T.—
4020 (1).

Muenscher, W. C., & Maguire, B.—2319,
2320 (3); 2322 (9); 2360 (12) ; 2362
(18).

Muenscher, W. C., & Spalteholz, R.—
16175 (7).

Muenscher, W. C., Wilson, C. L., &
Foster, A. S—15609 (9).

Mulford, A. I.—192 (16).

Munz, P. A.—8099 (17); 13046, 14820
(18).

Munz, P. A., & Johnston, I. M—12638
(17).

Munz, P. A., Johnston, I. M., & Har-
wood, R. D—4147 (18).

Murdoch, J.—4035 (12).

Murdoch, J., & Torrey, G. S—T391 (9).

Murie, O. E—1122 (13a); 1183 (12).

Myers, J..C—415 (6).

Nelson, A.—1931 (16); 2129, 10086,
10530 (12); 9010, 9317, 9333, 10245,
11344 (18).

Nelson, A., & Macbride, J.
1273, 1405, 1593, 1847 (12); 1110,
1327, 1347, 1593, 1798, 1935, 1939,
2120, 2121 (18); 1368 (16).

Nelson, A., & Nelson, R—783, 798 (12) ;

1888, 2024, 2877, 2890 (18).

Nelson, E.—807 (12).

Nelson, J. C.—1071 (13); 1475 (17).

Newins, H. S—8141 (5).

Nichols, G. E—168, 557 (5).

Nielsen, E. L.—1055, 1060, 1063,
1066, 1069, 1082, 1091, 1301,

1820, 1931, 2485,

2961, 3113, 3124, 3127;

3129, 3131, 3133, 3136, 3137, 3138,

3170 (4); 1868, 1872 (5); 1630 (6);

1349, 1967, 2500 (10).

Ownbey, M.—593 (12).

F.—1052,

1064,
1376,
2521,

AMERICAN SPECIES OF AMELANCHIER—JONES 99

Packard, J.—371 (14).

Palmer, E—94 (18). ,

Palmer, E. J—1602, 5616, 14684, 20761,
20764, 24470, 25860, 26350, 29915,
29927, 33207, 34917, 35564, 35890,
39015, 39994, 41069, 42440 (6);
20185, 35412 (7); 2322, 3539, 38948
(Gadies 27740, 28711, 28747, (28791,
28861, 28905, 36815 (10) ; 27779, 27795,
27796, 30009, 36296, 39808, 39833,
40470, 40471, 42391 (9) ; 31358, 36855,
36859, 36902, 36950, 36967, 37048,
37197, 37343, 37398, 37835, 37858,
38097 (12) ; 38025, 38037 (18).

Palmer, W.—1340 (1).

Pammel, L. H.—14, 52 (9).

Pammel, L. H., & Blackwood, R. E—
3623, 3876, 3887, 4063 (18).

Pammel, L. H., & Davy, J. B—77 (17).

Parish, S. B—453, 1290, 10882, 11969
(18). ;

Payson, E. B.—248, 982, 1066, 1077, 2307,
4779 (18) ; 1069, 1075 (12) ; 1051 (16).

Payson, E. B., & Armstrong, G. M.—
3272 (12).

Payson, E. B., & Payson, L.—1812, 2540,
3883, 3884, 4225, 4228 (18) ; 1812 (12).

Pease, A. S—678 (6) ; 670, 10995, 14374,
16044, 26527 (3); 683, 16554, 16990,
17890, 18048, 19680, 23232, 24246,
25775, 25912, 25979, 26323, 26541 (9);
20216 (11) ; 4090, 10216, 10312, 11205,
11969, 13482, 14372, 16662, 16683,
19561, 25275 (1); 16010, 17476, 18044
(5); 19794 (10); 22360 (12); 24229,
26992 (7).

Pease, A. S., & Bean, R. C—26215, 26502
(10).

Pease, A. S., & Long, B.—21434, 21452
(9); 21453 (7).

Pease, A. S., & Ogden, E. C—24884,
255225011).

Peck, M. E.—3530, 8512, 8731, 9242 (17).

Peebles, R. H.—13920 (18).

Peebles, R. H., & Parker, H. W.—14670
(18).

Pennell, F. W.—82, 24805 (5); 2708,
12027 (9).

Pénnell, F. W., & Long, B.—7559 (5).

Perry, L. M., & Roscoe, M. V.—243 (2);
244 (1).

Phelps, O. P.—567, 1591 (5); 568, 1592
ClO) 1581, 1582-6) > 1583. 1584,
1585, 1586 (9); 1588, 1589 (1).

Piper, C. V.—84 (13); 1534 (12); 2694,
3812 (14); 3823 (15).

Piper, R. H.—76881 (9).

Plantae Exsiccatae Grayanae—662 (7);
663, 664 (5); 842 (9); 959 (1).

Porter, C. L.—1059 (18).

Potter, D.—484, 485 (11) ; 486, 487 (1).

Pretz, H. W.—2363a, 3246, 5919, 9090
(5); 8296, 9260 (8); 10767, 11248,
11762, 12754 (9).

Purpus, C. A.—5376, 6523 (18).

Rand, E. L., & Robinson, B. L.—616 (1) ;
618 (9).

Randolph, L. F., & Randolph, F—121
(5); 1200 (6).

Raup, H. M.—2645, 2647, 2648, 2649,
2650, 2652, 2653, 2654, 2656, 6065, 6084,
6588, 6670, 6931, 6933, 7078, 7089
(12) ; 7410, 7727 (6); 8094 (9).

Raup, H. M., & Abbe, E. C—3500, 3502,
3530, 4466, 4515, 4519 (12).

Rehder, A.—955 (7); 115, 116 (18).

Ridgway, R.—2534 (6).

Robinson, B. L.—782 (1).

Roland, A. E.—2047, 41469 (9).

Rolland, F.—57, 58, 59 (6); 7214 (3);
13033, 13035 (9).

Rolland-Germain, Fr.—19258 (10).

Rollins, R. C—198 (18); 550 (12); 840
(14) ; 889 (16).

Rollins, R. C., Dillon, L. A., & Pickett,
F, L.—868 (12).

Rosendahl, C. O.—439, 4935, 4937, 5198
(4) ; 4980, 4983 (9); 6072 (10).

osendall, ©. Ocl& Brand; Co J—-88

(3),

Rosendahl, C. O., & Butters, F. K.—2578,
3892 (4).

Rosendahl, C. O., & Nielsen, E. L—
185275),

Rossbach, G. B.—1102, 1136 (6).

Rousseau, J.—24557, 24552 (3); 24537,
24550, 24554, 24561, 26241, 26259,
26478, 26513, 26672, 32418 (11);
26228, 26771 (1); 24549, 24559, 24562,
24567 (5).

Rousseau, J., & Fortier, L—31441 (1).

Ruth, A.—317 (6).

Rydberg, P. A—680 (12); 9052 (6).

Rydberg, P. A., & Carlton, E. C.—6662
(18).

Rydberg
(18).

P. A., & Garrett, O. A.—9274

St. John, H.—11887 (9); 1903, 90526,
90527, 90528, 90529 (1); 1909 (2).
St. John, H., & Long, B—1009, 8059

(6); 1063 (7).

100 ILLINOIS BIOLOGICAL MONOGRAPHS

St. John, H., & Nichols, G. E.—2332 (1).

Sandberg, J. H.—3 (9).

Sandberg, J. H., & Leiberg, J. B—94 (12).
Sandberg, J. H., MacDougal, D. T., &
Heller, A. A—26 (12); 53 (15).

Sandborger, J. D.—20 (1).

Sanford, J. N.—379 (6).

Sanford, S. N. F—626 (9); 1066, 1193,
10215: (5):

Sargent, H. E—21 (7); 24 (6).

Schacklette, H. T.—261 (6).

Seymour, F. C—571, 1222, 3493 (7);
1223, 1224, 3501 (5); 1708, 1709, 3512,
4641 (9).

Shantz, H. L.—345 (12).

Sharp, A. J., & Svenson, H. K.—7278 (6).

Slavin, B. H.—203 (9); 205 (10).

Smiley, F. J.—214, 215, 899 (17).

Smith, E. G—12005 (18).

Smith, L. B., & Hodgdon, A. R.—3872
(6).

Smith, L. E—284 (17).

Solheim, W. G—434 (18).

Sonne, C. F.—88 (17).

Spencer, M. F.—865 (17).

Spiegelberg, C. H.—341 (14).

Spreadborough, W.—93904, 93905, 93906,
93907 (13).

Standley, P. C—4125 (18).

Stanford, E. E—1758 (17).

Stecker, A.—19 (1).

Stevens, G. W.—2427 (6).

Stevens, O. A., & Graves, H.—278 (11).

Steward, A. N.—230 (13).

Steyermark, J. A—7009 (10); 18635 (9).

Stone, W.—65, 6484 (7); 11932 (5);
12684 (9).

Stoudt, H. N., & Hermann, F. J.—2779,
2784 (6).

Suksdorf, W. N.—52, 841, 8585, 8597,
8609, 10025, 10026, 10129, 10154, 10234,
10247, 11841, 11859 (12); 269 (18);
8575 (14); 2138, 2139, 2153, 10033;
10154, 10194, 10195, 10201, 10360,
10361, 10382, 10395, 10455, 10494,
11835, 11841, 11859, 11973 (13).

Svenson, H. K.—346, 7779 (6); 7849
(10) ; 8012 (9).

Tanger, Louise F. A—3025 (6); 3036,
3044, 3045 (5); 3043, 3065 (9); 3351
(8).

Thompson, J. W.—6002, 11386 (14);
7073 (12) ; 616; 2714 (13).

- Thone, F.—172 (6).

Tidestrom, I.—9359, 9429, 9460, 9600,
11050 (18) ; 11882 (6); 11947 (7).

Topping, D. L.—223 (5).

Torrey, J.—126 (16).

Tower, A. O., & Seymour, F. C.—3664
(10) ; 3668, 3671 (9) ; 3674 (6).

Ulke, T—S35 (12).

Visher, S. S.—3308 (12).
Voorhies, C. T.—82 (18).

Wahl, H. A.—29, 298 (5); 34 (6); 33,
AT 13):

Walker, E. P.—210, 416 (18) ; 1075 (13).

Watson, S.—353 (18).

Weatherby, C. A.—2018, D2103, 3616,
4070, 4070a, 4070b, 4255, 5370 (9);
D2157 (6); 2861; 5280° (5) 4916:
7029 (7),

Weatherby, C. A., Smith, L. B., Rollins,
R. C., & Mufioz, C., Pl. Exsicc. Gray.
—959 (1).

Webb, R. J.—84 (6); 1180 (5).

Werner, W. C.—54 (9); 55 (6).

Wetmore, A—470 (18).

Wherry, E. T., & Adams, J. W.—2768
(9); 2775: (7);

Whited, K.—363 (14); 571 (12); 1197
C13):

Wiegand, K. M.—991, 2131, 2132, 2539,
2541, 15627 (7) ; |2133; 4290; 6587,
6593, 6594, 6603 (9) ; 2498, 2499, 2501,
2505, 4281, 6582, 6589, 6592, 13976,
13979 (10); 2136;°2572 26).

Wiegand, K. M., & Eames, A. J.—2515,
25175 2518. (7):

Wiegand, K. M., & Manning, W. E.—
1328,. 13305 1331 (7).

Wiegand, K. M., & Metcalf, F. P.—
6583 (10).

Wiggins, I. L—4631 (17).

Wiggins, I. L., & Demaree, D.—4948
(18).

Wilkens, H.—471, 5128, 5168 (9); 5129,
6666 (5); 1097 (6).

Williams, L. O.—577 (18); 1106 (12).

Williams, L. O., & Williams, R.—3305
(12).

Wislizenus, F.—937 (10).

Wolden, B. O.—1043, 1047, 1074, 1075,
1078 (9) ; 1353 (10).

Wolf, C. B—2988 (18) ; 3075 (12).

Woodward, R. W., & Bean, R. C—
17116-(7).

Woodward, R. W., & Fernald, M. L.—
15202 (9). :

Zech, O. F.—152, 161, 168, 195 (4).
Zeller, S. M., & Zeller, E. B—844 (13).

PLATE. 1

Leaves of species of Amelanchier from herbarium specimens.

Fic. 1—Amelanchier bartramiana (Tausch)
M.Roem.

Fic. 2—Amelanchier neglecta Egglest.

Fic. 3.—Amelanchier fernaldu Wieg.

Fic. 4—Amelanchier interior Nielsen.

Fic. 5.—Amelanchier laevis Wieg.

Fic. 6—Amelanchier arborea (Michx.f.)
Fernald.

PLATE
Leaves of species of Amelanchier
Fic. 7—Amelanchier canadensis (L.) Medic.

Fic. 8—Amelanchier spicata (Lam.) K.Koch.
Fic. 9—Amelanchier sanguinea (Pursh) DC.

103

(2 a4

II

from herbarium specimens.

Fic. 10.—Amelanchier gaspensis (Wieg.)
Fernald & Weatherby.

Fic. 11.—Amelanchier alnifolia Nutt.

Fic. 12.—Amelanchier florida Lindl.

PLATE. 11

Leaves of species of Amelanchier from herbarium specimens.

Fic. 13—Amelanchier cusicku Fernald. Fic. 16.—Amelanchier pallida Greene.
Fic. 14—Amelanchier basalticola Piper. Fic. 17—Amelanchier utahensis Koehne.
Fic. 15—Amelanchtier pumila Nutt.

104

PLATE IV

Type of Amelanchier fernaldii Wieg. in the Gray Herbarium; from Fernald, Long
& St. John 7592, Grindstone Island, Magdalen Islands, Quebec, Canada.

105

PLATE V

Type of Amelanchier neglecta Egglest. in the Gray Herbarium; from
Rutland, Vermont, May 12 and June 21, 1899, W. W. Eggleston.

106

FLORA OF EASTERN MASSACHUSETTS

EX Horariam o

PLATE VI

Type of Amelanchier laevis Wieg. in the Gray Herbarium; from Wellesley,
Massachusetts, K. M. Wiegand 2136.

107

Amelanchier interior Nielsen

j Repent; :
HERBARIUM OF TRE ?

UNIVERSITY OF MINNESOTA
MINMEAPOLIB'R. Yano 1% 35)
mail Hae yy was

=~ eppods Ke

Dike te Fos. cM
whee, duh ae 2S me, os
a 3 on erin a Buel, Ta)
z nraytd | Ss “te de at) jas bod ake
Reva Site 4 Vs

PLATE Vif

Type of Amelanchier interior Nielsen in the Herbarium of the University
of Minnesota; from Minneapolis, Minnesota, E. L. Nielsen 2961.

108

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109

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110

PLATE Xx

Photograph of Amelanchier intermedia Spach in the Gray Herbarium.

111

Land of Me afin
#3 Sy,
Cticrmed >, therecarvpa
Po G fancburhs lan

j

emcee t

aff SS oe 46 7... Par i

i dphoe : : 3 bit, Vf, : :
ce lace far ei pose fos tf ioe |
Fig. 1 Fig. 2
PLATE XI

Fic. 1—Photograph of a tracing of leaves of Mespilus canadensis var. 8 oligocarpa Michx.,
from Herb. Mus. Paris. This is Amelanchier bartramiana (Tausch) M.Roem.

Fic. 2.—Phototype of Crataegus spicata Lam. in Herb. Mus. Paris, the basis
of Amelanchter spicata (Lam.) K.Koch.

112

PLATE XII
Type of Amelanchier stolonifera Wieg. (Gray Herbarium).

113

anh KM WHEG SSD

FLORA OF EASTERN MASSACHUSETTS

Herbarium of Welleicy Cottege

No. 2.1
a eee SPs | Is IRN EF erin §
Bay Hanky 2 bak Sore ed
Ry Geta otek
Pore ix Sag Eee ee wal
ce oe Conk Watisnaes
be ape Shenete Fe eG

PLATE XIII

Photograph of the second sheet of Amelanchier stolonifera Wieg.
(Gray Herbarium).

114

FLORA OF LASTERN MASSACHUSETTS

Wellesley Cotege

PLATE XIV

Photograph of the third sheet of Amelanchier stolonifera Wieg.
(Gray Herbarium).

cis

PLATE XV

Type of Amelanchier humilis Wieg. in the Gray Herbarium.

116

UNIVERSITY OF ILORNOLS

Amelanchier spicata (Law) K.Koch
G. N. JONES mr

“Gypr rpreivie fo.
eho edly. he Ss
Gx:

K ot WHEGAND

nda dic teccates ees

(Weck rs Roche pln Vt A
i lk Qe ef
lutein 4 dpe ea

ie cv ee 1 ee
Vi age iF$y™ vag
Thar DIRS tee Y Fr Unt, 30 tay

t aie

GN, JONES

PLATE XVI

Photograph of an isotype of Amelanchier austromontana Ashe
in the Gray Herbarium.

1a?

PLATE. XV ii

Fic. 1—Phototype of Michaux’s Mespilus canadensis var. Y rotundifolia in Herb. Mus. Paris.
Fic. 2—A tracing of part of specimen shown in Fig. 1, from the original in the Gray Herbarium.

118

ie i

The Academy of Natural Sclencas of Philadatphia

Ny tere hare thee;

Inewlity, Bie bad evs

Collector, they) ations!

PLATE XVIII

Photograph of a specimen of Amelanchier sanguinea (Pursh) DC. from Minnesota.

119

PROVINCE OF QUEBEC
a hy ,

oC eg
(e029 [Pont Ceuta als Got Loe
} (

PHATE 21S

Type of Amelanchier gaspensis (Wieg.) Fernald & Weatherby
in the Gray Herbarium.

120

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wt

Eastmxn Onrcen Plants
— Cann. Wa fC. Peston
Gi PO lie eb hi -teyp

vec tl se «hace waiek ~
aach, Dra f

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64 ‘ ere of Crieriot wy

Qe euwsk ot
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PLATE XX

Type of Amelanchier cusicku Fernald in the Gray Herbarium.

{21

UNIVERSITY OF ILLINOIS
: Amelanchier pallida Greene
G. N. JONES 1942

FLORA OF N. CALIFORNIA.

~ my
No PD z a wt

a x fg #

é a é j i
oy é Ei etl 7 F ttf
COfVI2Eg lass oteier t tie oe me

§ t < f 2 J
Nee ?
Ae ff _ pee 7
Cott. E. LL, GREENE. cHlAx /G, 1876.

PLATE: AX]

Photograph of an isotype of Amelanchier pallida
Greene in the Gray Herbarium.

tay

PLOBA GF SUL THEASTELN VIRGINIA

Sussex COUNTY

Dry xaady pine teomls abost 4 ouites wurtincest of Homevieat

M.L. Fexxaep and Bavaep Lose tert §, L987

PLATE. XX11

A flowering specimen of Amelanchier obovalis (Michx.) Ashe in the herbarium
of the Academy of Natural Sciences of Philadelphia, collected
in Virginia by Fernald & Long, no. 7073,

A OF SOUTHEASERILY VIRGINES

Prexce Groras Oucn

J G0 lmelershes obecabes|Preka, jake
tH

ins bossen Eeprewion xéutheast of Petersbncg,
wit head of Boo BUN

| ML. Fasnannand Bevann Lae Mary 23. 235

PLATE XXIII

A fruiting specimen of Amelanchier obovalis (Michx.) Ashe in the herbarium
of the Academy of Natural Sciences of Philadelphia, collected
in Virginia by Fernald & Long, no. 9947.

124

INDEX TO PLANT NAMES

Amelanchier (cont'd)
carrit, 67

Amelancher, 13
Amelanchier, 13

alabamensis, 41
alnifolia, 73, 80, 83, 86
alnifolia, 67
f. alba, 67
var. covillei, 88
var. cuyamacensis, 83
var. dakotensis, 67
var. elliptica, 87
var. florida, 73
var. nitens, 88
var. pallida, 83
var. pumila, 67, 77, 81
var. siskiyouensis, 83
var. subintegra, 83
var. typica, 67
var. utahensis, 87
var. venulosa, 88

alnifolia & Sorbus scopulina, 73

amabilis, 60
arborea, 34

var. alabamensis, 41
arguta, 20, 24

couvillet, 87
crenata, 87
cuneata, 67
cusickii, 78
elliptica, 87
ephemerotricha, 73
var. silvicola, 73
erecta, 51, 59
fernaldii, 24
florida, 64, 67, 83
florida, 73
var. humptulipensis, 77
var. parvifolia, 73
f. tomentosa, 73
var. typica, 73
gaspensis, 64
glabra, 81
goldmanit, 87
gorman, 73
gracilis, 83
grandiflora, 59, 73
grandiflora, 42

Xx grandiflora f. rubescens, 43
humilis, 51, 67

var. campestris, 52

var. compacta, 52

var. exserrata, 52

Xx laevis, 52

var. typica, 52
huronensis, 59
interior, 28
intermedia, 28, 35, 40, 43, 51
jonesiana, 87
laevis, 28
laevis, 30

var. cordifolia, 30

f. nitida, 30

var. nitida, 30
lancifolia, 30, 42
lanulosa, 67
leptodendron, 67
longifolia, 44
macrocarpa, 67
mucropetala, 52

var. potomacensis, 52
montana, 67
mormonica, 87
mucronata, 52
nantucketensis, 44, 48
neglecta, 25
neumanniana, 44

australis, 87
austromontana, 35, 44, 52
bakeri, 87
bartramiana, 20
basalticola, 80
beata, 52
botryapium, 30, 35, 43
var. conferta, 52
var. micropetala, 52
var. obovalis, 52
botryapium lancifolia, 42
canadensis, 30, 34
canadensis, 43
var. 8 alnifolia, 67
var. botryapium, 30, 35
X laevis, 42
var. micropetala, 52
f. nuda, 35
var. B oblongifolia, 44
var. obovalis, 44, 49
var. £ oligocarpa, 20
var. pauciflora, 20
var. € pumila, 81
var. Y rotundifolia, 59
var. semtntegrifolia, 73
var. spicata, 51
var. tomentula, 35
canadensis grandiflora, 42
intermedia, 44

126 ILLINOIS BIOLOGICAL MONOGRAPHS

Amelanchier (cont'd)
nitens, 87
oblongtfera, 44
oblongifolia, 44, 49, 51

var. micropetala, 48, 51
var. 8 waltert, 49
obovalis, 49
oligocarpa, 20
oreophila, 87
ovalis, 44, 51, 67
ovalis, 7, 13
var. B semuntegrifoha, 73
var. B subcordata, 35
var. willdenowiana, 59
oxyodon, 73
pallida, 83
var. arguta, 87
parvifolia, 67, 73
plurinervis, 87
polycarpa, 81
prunifolta, 87
pumila, 81
purpusi, 87
recurvata, 83
rotundifolia, 59
rubescens, 87
var. cinerea, 87
sanguinea, 20, 64
sanguinea, 59
var. gaspensis, 64
f. grandiflora, 59
var. grandiflora, 59
saxatilis, 40, 51
sera, 44
siskiyouensis, 83
spicata, 44, 48, 59
spicata, 51
stolonifera, 49, 52
var. lucida, 52
stricta, 30
subintegra, 83
utahensis, 67
utahensis, 86
subsp. covillet, 88
subsp. oreophila, 88
venulosa, 87
vestita, 73

Amelanchier (cont'd)
wangenheimiana, 35
wiegandi, 28

Amelanchus, 13
canadensis, 35

Amelancus, 13
spicata, 52

Amelasorbus jackti, 73

Aronia affinis, 44
alnifolia, 67
arborea, 35
botryapium, 35, 43

var. B racemosa, 43
cordata, 35
latifolia, 59
nivea, 35
ovalis, 43, 51
praecox, 20
sanguinea, 59
subcordata, 35

Crataegus amoena, 43
canadensis obovwalis, 49
racemosa, 43
spicata, 44, 48, 51

Malus microcarpa, 35

Mespilus amelanchier, 49, 51

B nivea, 35
arborea, 35
canadensis, 43, 47
var. B cordata, 35
var. a obovalis, 49, 50
var. 9 oligocarpa, 20
var. Y rotundifolia, 59, 63
glabra, 44
nivea, 34
virginiana, 43

Pyrus alnifolia, 67
bartramiana, 20
botryapium, 30, 34, 43
neumanniana, 44, 48
oligocarpa nana, 20
ovalis, 43, 51
sanguinea, 59
wangenheimiana, 35

Sorbus scopulina, 73

Sorbus torminalis, 7

~

ILLINOIS BIOLOGICAL MONOGRAPHS
(Continued from inside front cover)

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[Continued on back cover.]

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