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ContrihTztions from the

SHAW SCHOOL OF BOTANY.

No. 4.

TRELEASE ON CEANOTHUS.

{Proc. Cal. Amd. Set, id Ser., Tol. /, Port. 1. Issued June IS, ISSS.)

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106

C. Y. P. I I-E Y,
\YcLGhin7tcn.
1 ). r.i
ca'lifukma academy of tctenCes.

2d Heb, Vol. 1.

CONTRISUnONS FROM THE SHAW SCHOOL OF BOTAHT.

No. 4.

'synoptical list op north AMERICAN SPECIES OP CEAN0THU8,

By William Trelease.

In its present limitation, Ceanothm is an exclusively Amer-
ican genus, and all but three of the species known to me
occur north of Mexico. It is quite puzzling to the student
of our flora, partly because of the variability of some spe-
cies, and partly because in some instances what are evi-
dently distinct species approach each other so closely as to
render their characterization difficult, although they may be
pretty surely recognized by the practiced eye. To the evo-
lutionist, these groups of interlocking sub-species and
barely distinguishable species, are interesting as represent-
ing different stages in the mutations of their prototypes; but
they are exceedingly trying to the systematic botanist,
whose task is to so arrange and limit them as to render
their recognition by others easy.

Aside from the monograph of the genus (in the broader
sense in which it was formerly understood) in De CandoUe's
Prodromus, which characterizes very few of the species that
are now known, and the fuller revision iu the first volume of
Torrey and Gray's Flora, it has been very judiciously and
completely treated by Mr. Watson in a revision of the en-
tire group (Proc, Amer. Acad. x. 1875, 333-9), and a separate
elaboration of the numerous and difficult Californian species
(Bot. Oal. i. 102-104; ii. 439). His views on the syno-
nymy of species are also very fully indicated in his Biblio-
graphical Index to North American Botany. Nor should
the field notes of the keen-sighted Nuttall, on a number of
the Pacific Ooast species, in Torrey and Gray's Flora, be
overlooked.

Usned June 15, 1888.

I

'.:/

NORTH AMERICAN SPECIES OP CEANOTHUS. 107

In the main, the limitation and descriptions of species in
Mr. Watson's monograph, and his references to literature,
in the Index, are so accurate as to require little alteration;
but after repeatedly going over the genus as represented in
the Gray, Engelmann, and Torrey herbaria, and in the ex-
tensive collections of the California Academy of Science,
and the United States Department of Agriculture, as well
as the fine private herbarium of Mr, J. C. Martindale, it has
seemed to me that Cemiothua is capable of subdivision into
more natural groups than have yet been proposed, and the
following is offered as a tentative arrangement of our spe-
cies, I shall be very grateful for specimens and notes eluci-
dating doubtful species, and I desire here to express my
obligation to the botanists who have already placed public
or private collections in my hands for study.

A. Leaves alternate, not spiuescent, glandular-toothed or occasionally
entire; stipules thin, often subulate, fugaoious; fruit sometimes
keeled or crested on the back of each segment, but not bearing
prominent dorsal horns.— JSiuceano<AtM.

a. Inflorescence on leafless lateral peduncles borne on the

old wood.

1. 0. SANGUINEUS, Pursh. Fl. Am. Sept. i. 167; Wat-
son, Proc. I. c. 334, Index, 166.— British Columbia to Idaho
and California.

b. Inflorescence on leafy shoots of the present season's

growth.

* Flowers white, in small simple corymbose clusters
terminating mostly leafy spineless twigs: leaves
very small (2 to 8 mm. long), 3-nerved.— Atlantic
species.

2. C. MiCBOPHYLLUS, Michx. Fl. i. 154; Watson, l. c. 336.
Georgia to Florida.

1! S.I/fePwJU^J,

4 in

""i^ft^* '<J l""*' ■•rft^^^^S"'

107

1^

II

I

108

CALIFORNIA ACADEMY OP SCIENCES.

3. C. SEBPYLLiFOLius, Nutt. Gen. i. 154; Watson, I. c.
335. — Georgia. The locality in Nuttall's Genera, and on
an original specimen in the Gray herbarium, is Florida.

* * Flowers white: inflorescjuce rather simple and

mostly compact, at the ends of slender usually
leafless or nearly leafless peduncles : twigs subte-
rete, not spinose: leaves ample or medium-sized (15
to 75 mm. long), thin, toothed, 3 -nerved: fruit
about 4 mm. in diameter. — Atlantic or Rocky
Mountain species.

4. 0. Amebioanus, L. Spec. 195; Watson, I. c. 333, In-
dex, 163. — Ontario to Manitoba, south to Florida and Texr
aa. i

5. C.0VATD8, Desf. Arb. ii. 381; Watson, l. c. 334.— Can-
ada and the lake region to Texas.

VAR. PUBE8CEN8,< Watson, Index, 166. — Rocky Mount-
ain region.

* * * Flowers blue: inflorescence cqmpound, ample, on

leafy branches : twigs conspicuously sulcate : leaves
medium-sized (25 to 50 mm. long), prominently 3-
ribbed, minutely glandular-serrulate: fruit mostly
5 or 6 mm. in diameter.

6. C. TH1B8IFLORD8, Esch. Mem. St. Petersb. Acad. x.
285; Watson, I. c. 334. — Mountains of Western Califor-
nia.

* * * * Flowers blue (often pale) or «hite: inflorescence
compound, rather loose, on lew-leaved branches:
leaves entire, mostly medium-sized. — Pacific spe-
cies.

+- Twigs (more or less spinose in the first) usually
somewhat angled and often twisted : leaves rather
firm or even coriaceous.

i

"?»■ _

NORTH AMERICAN SPECIES OF CEANOTHUS.

109

■«"•• Leaves rather broad, not at all 3-nerved: fruit 5 to
6 mm. in diameter.

7. C. HPiNosus, Nutt. in Torrey & Gray, Fl. i. 267; Wat-
son, l. c. 337.— Middle and Southern California.

8. C. Palmeri, n. sp. Glabrous throughout, or a very
few hairs on the leaves and petioles: branches greenish,
becoming brown: leaves mostly on short spurs, slender-peti-
oled, about 40 mm. long, elliptical or ovate-oblong, rounded
at both ends, mucronate or emarginate, entire, thinner
than in the last: flowering branches ascending, naked or
few-leaved: inflorescence oblong, nearly simple: exocarp of
fruit rather fleshy. — Mountains of Southern California
(Palmer, 1875, No. 42). Intermediate between G. apinoaua
and C. integcrrimus.

■t-i- ■!")• Leaves narrow, 3-nerved, the nerves often conceal-
ed by the revolute margins; fruit about 3 mm. in
diameter.

9. C. Pabryi, n. sp. Branches glabrate or sparingly
villous, strongly sulcate, more or less papillate : leaves nar-
rowly elliptical-oblong, obtuse, 15X30 mm. or less, gland-
ular-serrulate, glabrous above, the lower surface rusty-to-
mentose, at least along the veins: inflorescence oblong,
interrupted, terminating recurved - ascending slender, few-
leaved branches: flowers blue. — Known to me only from
specimens found in cultivation at Calistoga, Cal. {Parry,
1881, No. 33).

H — I- Twigs not spinose nor rigid, very slightly angled
while young: leaves thinner, at least the largest 3-
nerved: inflorescence rather dense for the group;
fruit about 4 mm. in diameter.

lU. C. INTEGERRIMUS, Hook. & Arn. Bot. Beechey, 329;
Watson. I. c. 334, Index, 165. C. thyrsiflorus, var. macro-
thyrsus, Torrey, Bot. Wilkes, 263! California to Arizona.

i

i

i*al^"jr^

110

CALIFORNIA ACADEMY OF SCIENCES.

11. C. PABVIFOLIUS. C. integerruuiis, var.f parvifoUiis, Wat-
son, I. c. 334. — California to Oregon.

***** Flowers blue or white: inHoresoence mostly com-
ponnd and ample: leaves generally medium-sized
to large 3-nerved; margin various.

•i- Twigs not spinose nor conspicuously glaucous, the
youngest angled in the first species: flowering-
branches few-leaved or leafless: leaves broad, usu-
ally large (25 to 75 mm. long): fruit about 5 mm.
in diameter.

12. C. ARBORBrs, Groene, Bull. Cal. Acad. ii. 144. C.
sorediatus, Lyon. Bot. Gaz. xi. 204, 333. — Islands off the
Californian Coast.

14. C. VELUTiNUS, Dougl. in Hook. Fl. Bot. — Am. i. 125,
pi. 45; Watson, I. r. 334. — British America to California,
Colorado an. I Nebraska, chiefly in the mountains.

VAB. LJiViGATUS, Torr. & Gray, Fl. i. 686; Watson,
Index, 167. — Range of the species.

+- -f- Twigs terete, often very divergent and rigid, some
of them ending in firm spines.

-M- Twigs short, often very glaucous: leaves large in
the first, mostly medium -sized, rather small in the
hist: fruit 4 to 6 ram. in diameter.

14. C. iNCANus, Torr. & Gray, Fl. i. 265; Watson, I c.
336. — California.

15. C. EGLANDUi.osua. C. divaricatns, var. eglandulosua,
Torrey, Pac. R. R. Rep. iv. 75. C. divaricatns, Watson,
L c. in part. — Mountains of California and Lower Cali-
fornia.

Ml -t. «I.IPI"I

IXI « '-III! — »i;

NORTH AMERICAN SPECIES OF CEANOTHUS.

Ul

16. C. DiVAUiCATUs, Nutt. in TovT. & Gray Fl. i. 266, 686;
"Wiitson, /. c. 336 in part — Californiii and Lower Cali-
fornia.

VAR. QiiOHHE - SEBIIATU8, Torrey, Pttc. R. R. Rep. iv. 75.
— California.

17. C. conDur,ATiTs, Kellogg, Proc Cal. Acad. ii. 124, f.
39; Watson, I. e. 337.— Mountains of California.

^.^ y^ Twigs slender, usually slightly if at all glaucous:
spines slender and sharp: leaves rather narrow for
the group: fruit about 4 mm. in diameter.

18. C. Fendleri, Gray, pi. Fendler29; Watson, I. c. 337.
Mountains of Colorado, New Mexico and Arizona.

VAR. VIRIDI8 Gray, in Herb. — Arizona (Lemmon.
Greene).

%%% Flowers deep blue (except sometimes in the first),
in rather compact nearly simple corymbose or ob-
long clusters, on leafless or nearly leafless (some-
times abbreviated), usually scaly peduncles: twigs
terete, mostly spineless, and not very rigid.— Pa-
cific species.
■^ Leaves medium-sized, all or nearly all 3-nerved:
twigs rather more rigid in forms of the first two
than elsewhere in this group: fruit about 4 mm. in
diameter.

19. C. SOREDIATUS, Hook & Am. Bot. Beechey, 328;
Watson, I. c. 336. —Coast Range of Southern California,
extending into Lower California.

20. C. HIRSUTUS, Nutt. in Torr. & Gray, Fl. i. 266; Wat-
son, I. c. 336. C. diversi/olius Kellogg, Proc. Cal. Acad,
i. 58, 65.— Mountains of Southwestern California.

VAR. ? GLABER, Watson, I. c. 336. —East Humboldt
mountains, Arizona. (Watson, No. 212.)

"^*^r

111

112

CALIFORNIA ACADEMY OF SCIENCES.

•1

21. C. DEruMBENs, Wivtson, l. c. SSf).— Mountains of Cen-
tral California.

-H-f- Leaves usually small, commonly only the largest
:}-nerved, not papillate: fruit 3 mm. in diameter.

22. C. DKNTATiTs, Torr. k Gray, Fl. i. 268, Watson I. c.
335; — Coast range of Southern California,

8UHHP. ? FLORinuNDUS. C. Jlonbuvdus Hook. Bot. Mag.
pi. 4806; Watson, l. c. 338.— Known certainly only
from pliints grown in European gardens, from Cali-
foruian seed.

8UBS1'. ? LoBBiANUs. C. LofManus, Hook. Bot. Mag.
pi. 4810. — Coast Range of Southern California.

23. C. LMPBESSus, n. sp. Villous, with short spreading
hairs: leaves broadly elliptical to nearly orbicular, 6 to 8
mm. long, loosely villous, esi)ecially on the veins below,
the upper surface deeply furrowed over the midrib and
several pairs of lateral nerves, the slightly glandular mar-
gin very revolute, appearing there us if crenate: pedun-
cles about 10 mm. long, scaly toward the base: inflorescence
sub-globose, compact: fruit not seen. — Santa Barbara
County, Cal.

-i- -i- -i- Leaves medium-sized, oblong, not at all 3-nerved,

papillate.

24. C. PAPILL08U8, Torr. & Gray, Fl. i. 268; Watson, I. c.
337. — Mountains of Western California.

2.5. C. Veatchiands, Hook. Bot. Mag pi. 5127; Watson,
7. ('. 338. — Decribed and known only from European
plants raised from Californian seed.

B. LeHved oppuiiite or aiturnute, pinuiitely veiued, corinceous, of-
tuu pungeutl. (onlied; stipules thick «nl spongy, taper-
pointed perfl'Htent: inflorescence densely corymbose, on short
spurs from the new woo.l; (rnit usually Urge for tho genus,
p«ch carpel commonly bearing a dorsal horn, an alternating set.
of .3 crests or horns frequently at or near the apex.— Cerastes
Species of the Pacific Coast or Southwest, mostly with rigid
almost spinose twigs.

NORTH AMERICAN SPECIES OF CBANOTIIUS. 113

a. Procumbent and radicant: flowers bright blue (ex-
cept occasional albinos or hybrids), mostly on
long, slender, colored pedicels.

26. C. PHOSTRATUS, Benth. PI. Hartweg, 302; Watson,
I. c. 339.— Washington Ter. to California and Nevada.

h. Erect or spreading, the branches mostly ligid : ped-
icels usually stouter or rather short,

* Leaves opposite.

+- Flowjrs (always?) white.

27. C. cuNEATUS, Nutt. in Torr. & Gray, Fl. i. 267; Wat-
son, I. c. 338 ^nd Index, 164, in .part. - Oregon to Lower
Calil'ornia. ,

28. C. Gbegoii, Gray, Plant. Wright, ii. 28; Watson,
L c. 338.— Utah, Arizona and New Mexico to Mexico.

29. C. CRASSiFOLius, Torr. Pac. R. R. Rep. iv. 75; Wat-
son, I c. 338 and Index, 164, excl. synonym and variety.
-Coast range of Southern California, Lower California and
Califoniian islands.

-(--1- Flowers blue (or rose-purple?).

30. C, RIGIDUS, Nutt. in Torr. & Gray, Fl. i. 268; Watson,
I g 339. — Coast region of Southern California.

VAR QRANDIF0LIU8, Torr. Pac. R.R. Rep., iv. 75; Wat-
son, Index, 164, under C. crassifoUm. G. crwisifolms,
var.' glain-atiis Gray, Cat. Bolander, 8; Watson,
Index, 164.— Range of the species.

** Leaves alternate; flowers white.

113

I

)

114 CALIFORNIA ACADEMY OF SCIENCES.

31 C. VEBBUC08U8, Nutt. in Toil-. & Gray, Fl. i. 267;
Greene, Bull. Cal. Acad. ii. 81. C caneatm Watson, U.
338, and Index, 164, in part. -Southern California and
Lower California.

32. C. MA0ROCAEPU8, Nutt. in Torr. & Gray Fl i. 267;
Greene, L c. C. cuneatm, Watson, I. c. 338. and Index. 164.
in part.-Coast Range of Middle and Southern California.

NOTES.

C. «««9ui».e«« appears to differ from all other species of
the genus in flowering from wood of the preceding season «
growth; otherwise, it is related in several respects to Nos. 4
and 5.

G. microphyUns ^uA serpyllifolim-yery closely related to
each other-show no very great aflinity for other groups
For convenience they are placed where they now stand, al-
though in the character of the inflorescence, they approach
the dentatus group. I have no fruit of either.

While a ovatus is well marked in its typical Eastern
form, it passes- gradually into var. pnbescemm he West, and
thrc. gh this approaches Americanus in its leaf-forms. The
leaLL of flowering branches is sometimes quite variable.

In a spinosHS, the firm leaves commonly turn brown in

arying. eTpecially the upper ^-^-^ '**-,, ^'^^^''^"/^f-""
Hinple Somewhat leafy loose thyrsus mostly spread at right
angles or are even recurved; and the flowers are scarcely
more than lilac-colored.

C parvifoUus appears to be distinct from irdegerrmns in
its loose low habit, smaller leaves scarcely exceeding 25
mnin length, the majority of them not 3-uerved. and in
Tsmaller oblong or (from the falling of the lower fasci-

I

V

i

NORTH AMERICAN SPECIES <)F CEANOTHUS.

115

cles) siibcoryrabose inflorescence of blue flowers — those of
bitcyerrimus ai)peaving to be always white.

C. arhoreiis is, as Professor Greene has shown, the largest
of our species. It was collected on Sta. Catalina ir-my
years ago, by Nuttall, who notes its arboreous habit on tlie
label accompanying a specimen in the herbarium of the
Philfulelphia Academy.

Numbers 14 to 17 of the list are quite difficult to dis-
tinguish.

C. iiicavKs, with large, rather thin, nearly smootli ellip-
tical leaves, often rounded in abruptly to the lateral nerves,
a short distance above ths base, usually entire, pale below;
ratiier large white flowers (often 5 or 6 mm. in expanse);
and large depressed fruit with a more fleshy exocup than
usual, is generally recognized without much difii(!nlty. The
same is true of C. eglantiiihisns, which is nearly glabious and
very glaucons, with rather small strongly ovate entire or
nearly entire leaves, usually brown above, in herbarium
specimens; and smallei I'.ngy blue flowers. But divaficatiis
and cordulatus approach each other so closely that it is hard
to draw the line between them. The specimens with smaller,
thicker leaves are commonly referred to the latter, as being
evidently what Kellogg figured, while the former species
usually has large and more flexible leaves.

C. Fendleri, with rather thin narrower leaves, silky-can-
escent in the type, green and nearly glabrous in the variety;
stands out quite well from its congeners, in geographical
distribution, also.

C sorediatus is a species which I do not at all understand.
As it is here accepted, it incluues plants with slender rather
simple twigs, and others that ax-e quite rigid and intricately
branched; the leaves of some are very broadly ovate, while
others are narrower; and the pubescence varies from silky

116

CALIFORNIA l^CADEMY OF SCIENCES.

or more or less hirsute to very densely white- or rxisty-
tomentose on the lower surface of the leaves. The inflor-
escence is intermediate between that of the preceding and
following groups of specie.. In some of the rather slender-
stemmed plants that I regard as most typical, the twigs are
very rough with crowded small warts. Probably one or
more species mav ultimately be separated out, and it is not
certain that a number of specimens now referred here do
not really belong to divoricatns on the one hand, or hirsntns
on the other.

The ,/e>,tatii, group is one of the best circumscribed, but
it is not less puzzling than others, for it is doubtful whether
the forms it comprises are best described as species or va-
rieties. Lobbianm is chiefly characterized by its conspicu-
onslv unequal leaves-those on the primary shoots 25 mm.
long", mostly acute at both ends, the larger 3-nerved.

While I have scarcely felt that this form and fiorih,i»dm
are woitliy of specific rank. I have pretty confidently sep-
arated out the plant witb rather broad hairy leaves, daeply
furrowed over the veins. It is known to me only rum two
flowering specimens in the Gray herbarium, collected at
different places.

In Cerastes the species are quite as perplexing as in
EucEANOTHUS, and the difficulty of separating them is in-
creased by the occurrence of what appear to be hybrids.
Tl.is is especially true of prostratus and caneatus, typically
very distinct in habit, foliage, flowers and fruit; but nu-
merous specimens have been collected over a large «rea,
siiowing various combinations of the characters of both. So
mu.ked are these that Mr. Thomas Howell writes me that
from his field observations he is disposed to regard i)ro8<ra-
tu8 as only a v ariety of cuneahis*

•On these supposed hybrids, aee Garden and Forest, i. 7.

t )

II

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jljl

I

NORTH AMERICAN SPECIES OF CEANOTHUS.

117

The fruit of prostrntm in usually considerably louf^er than
broad, about 8 ram. in diameter, rather Hesliy, with 3 nearly
erect prominent horns, and an accessory set of apical crests
or smaller horns.

CmwatHH produces subgloboso or very slightly elongated
fruit, about (i mm. iu diameter, with thin Hesli, and usually
3 rather small horns, an accessory set of smaller ones being
occasionally seen.

C Greygii, closely related to eanedtns, is marked by its
leaves, often white - tomentose below, raosily equally
rounded and acute at both ends, frequently with one or two
small teeth on either side, and its rather pointed fruit,
about 4 mm. in diameter, seemingly nearly or quite horn-
less, the white calyx-lobes more persistent than usual.

C crassifolius, as it is now accepted, is very heterogen-
eous, embracing plants with entire or toothed smooth green
leaves, and others with the leaves pungently toothed, revo-
lute margined, and very white-woolly below. The fruit is
as much as 8 or 10 mm. in diameter, depressed-globose,
smooth or with 3 low deeply dorsal horns, and the base (in-
vested by the adnate calyx) very prominent, and indurated.
The leaves are always firm and thick.

G. rigidus has thinner, often cuneate-obcordate, mostly
denticulate leaves, and blue or purplish flowers. Good fruit
is desirable. Some specimens suggest hybridization with
C. proatralus.

G. vei-rncosus is very similar to rigidus in its foliage char-
acters, but with slender twigs, alternate leaves and white
flowers. The figure of rlgidits in Bot. Mex. Bound, pi. 9,
is evidently this species. Its fruit is mostly 4 to 6 mm. in
diameter, with small dorsal horns, or none.

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CALIFORNIA ACADEMY OF SCIENCES.

G. mocrocarpiis approaches broad-leavod forms of cnneatm,
and especially eutire-leav.'i forms of crasslfolim, in its foli-
age, but its habit is said to be arborescent, its twigs are
slender, and its leaves opposite. Its fruit is very similar
to that of the eutire-loaved crasslfi>J!m, but with prominent
dorsal horns.