CONTENTS
BRITTON, ELIZABETH GERTRUDE. West Indian Mosses—II. Mosses of
the Danish West Indies and Virgin Islands (plate 1) -
ayaa ELIZABETH G., and Ho.iick, ARTHUR. A new hiociibate fossil
Rene, Pen: Pky iockaura phil notes on tiie Rocky Mcnecck
region—IV. Forests of the Subalpine and Montane Zone II
cic. EuGENEP. The ferns and flowering plants of Nantucket—-X1V 27
ArtTuur, J. C., and Fromme, F. D. A new North American yt’, makes
(plate 2) -
HANCE, ROBERT tT Pollen Aetiocmnant ne dheecieiide in Zebrina
pendula, with special reference to the aa cence (plates 35) - - 63
Britton, ELIZABETH G. Mosses of Bermuda (plates 6, 7) ei
Rock, JosepH F. A new Hawaiian ns (plate ’8) ee a
Maxon, Wiitti1am R. The North American species of illedvataae -
Drxon, H. N. . New and rare apne oS mosses, apa: from Mitten’s
herbarium (plate 9) - - 93
HALL, HArveEy Monroe. Notes on ‘pee ad kedchanle - - - III
RyDBERG, P. A. Notes on Rosaceae—IX - II
NIcHOLs, GEORGE E. The ia en of Connesticat2V. Piaat
. societies in lowlands I
Maxon, WILiiaM R. Pobeodian riesginellnsn iad its hinediars allie: - 219
GROSSENBACHER, J. G. Medullary spots and their cause (plates 10, 11) - 227
FARWELL, OLIVER ATxKins. Notes on the srecaacoes = of ri petty
(plates 12-18) - - 24
Evans, ALEXANDER W. The genus Plagiochasma wad its Notts Asch
ies
spec -
GRIGGS, Roset F, N ew species aad varieties s Bihai (lala 0) - - 315
BICKNELL, EUGENE P. The ferns and flowering plants of Nantucket—XV - 331
FARWELL, OLIVER ATKINS. Notes on Michigan Liliaceae (plate 20) - - 351
Britton, NATHANIEL Lorp. Studies of West Indian Plants—VI -
Witiiams, R. S. Mosses from the west coast of South America Fics
regs) - > Bt 303
MACKENZIE, KENNETH eed. Noes on ‘Corel - 405
Stout, A. B. The origin of dwarf pao as shown in a sport a Hibiscus
oculiroseus (plates 26, 27) -
Scott, Ernest L. A study of castors trees a shrubbery - : - 451
RypBerG, P. A. Notes on Rosaceae—X - 463
Britton, NATHANIEL Lorp. Studies of West Paes Plants—VII - - 487
Donce, B.O. The effect of the host on the oem of certain nage
of Gymnosporangium (plates 28, 29)
BICKNELL, EuGeNE P. The ferns and lowering eS of Nantucket—XVI .
1V CONTENTS
WiLiiaMs, R.S. Mosses of the ieee a and Hawaiian Islands collected
by the late John B. Leiberg , = 57)
Howe, R. HEBER, Jr. The genus Teloschiates in North Acai - - 579
ARTHUR, JOSEPH CHARLES. New species of Uredineae—IX - : - 585
MACKENZIE, KENNETH KENT. ot arex—IX
PitT1rER, HENRY. On the characters and aii of ‘he genus Metan
pte ruce - - 623
RYDBERG, P. A. Piyiieeierapbiesl Notes on the eae Mountath
region—V. Grasslands of the Subalpine and Montane Zones - - 62
Stosson, MarcGaret. Notes on Trichomanes—I. The iad of
richomanes selec (plates 30, 31) -
OSTENFELD, C. H. Ruppia anomala sp. nov., an cue type of the
ene (plate 32) - - 6
Harris, J. Artour. On the dikes ee rites of ee
(staminate and pistillate flowers) in the inflorescence of the aroids
Arisarum vulgare and Arisarum proboscidium -
Forrste, Auc. F. Dictyophlois reticulata gen. et sp. nov. (plate ae - oes
INDEX TO AMERICAN BOTANICAL LITERATURE - 49,
87, I61, 241, 309, 359, 423; 4st, 543, 595, ide 679
INDORE TONG UME 46 8 OBS
Dates of Publication
No. 1, for January. Pages I-54. Issued January 29, 1915.
No. 2, for February. 55-92. March 2, 1915,
No. 3, for March. 93-168. March 16, 1915-
No. 4, for April 169-246 April 27, 1915°
No. 5, for May. 247-314 May 22, 1915"
No. 6, for June 315-364 June 16, 1915
No. 7, for July. 365-428 July 29, 1915
No. 8, for August 429-486 September_22, 1915
No. 9, for September. 487-548. November 5, 1915.
No. 10, for October. 549-602. November 13, 1915,
No. 11, for November. 603-650. December 11, 1915.
No. 12, for December. 651-608. January 22, 1916.
Errata
Page 410, line 17 from bottom, for Pretz read Fretz.
Page 412, line 10, for Pretz read Fretz.
Page 418, lines 1 and 13 from bottom, for Pretz read Fretz.
Page 638, line 18, for chryptandrus read crypiandrus.
Vol. 42 : No. 1
BULLETIN
- OF THE
TORREY BOTANICAL CLUB
ee ee
JANUARY, tg15
West Indian mosses—l], Mosses of the Danish West Indies and
Virgin Islands
ELIZABETH GERTRUDE BRITTON
(WITH PLATE I)
In February, 1913, Miss Delia W. Marble and I spent four
weeks collecting on the island of St. Thomas, while Dr. Britton
and Dr. Shafer cruised around among the other islands of the
Archipelago in search of cacti and other plants. An account of
their trip will be found in the Journal of the New York Botanical
Garden.*
As far as we know, no record of any other collection of mosses
from these islands exists, excepting the brief account given by
Dr. I. Urbanf of the collections made by the Rev. Johann
Christian Breutel in 1841. Most of Breutel’s mosses, 310 numbers,
are deposited at the British Museum, but his collections included
specimens from St. Thomas, St. Croix, St. Jan, St. Kitts, and
Antigua, as well as the collections made in South Africa in 1853-
1854. This would account for the large number of specimens
preserved in his herbarium, as our experience has shown that
the Danish and Virgin Islands are not particularly rich in species
of bryophytes.
William Mitten had a few duplicates from Breutel’s collec-
tions including the following six species: Calymperes Richardt C.
Miill. (distributed as C. Afzelii Sw.), Hymenostomum Breutelu
(C. Mill.) Broth., Tortula agraria (Sw.) Sw., Philonotis tenella (C.
* Jour. N. Y. Bot. Garden 14: 99. 1913.
+ Symbolae Antill. 3: 28. I902. :
(The BuLLEtIN for December (41: 577-657- pl. 21-23) was issued Ja 8, 1915.
1
2 BRITTON: MosseEs oF DANISH WEST INDIES
Mill.) Jaeg., Stereophyllum leucostegum (Brid.) Mitt. (distributed
as Hypnum Breutelii Schimp.), and Taxithelium planum (Brid.)
Mitt. (distributed as Hypnum Antillarum Schimp. ms.).
Baron von Eggers collected a few mosses in St. Thomas, St.
Jan, and Tortola in 1887, but as far as we know no list of these _
has been published.
We collected about seventy packets, but found only twenty-
‘eight species, representing twenty-two genera, though we added
three new species, a small: Phascum, a sterile Hyophila, and a
sterile Bryum, which has since been found on Mona Island also.
A brief synonymy with localities and habitat of these species
is given in the following enumeration: Nos. 1-4 and 7-9 were
determined by Mr. R. S. Williams; Nos. 16 and 17 by M. G.
Dismier. Unless otherwise noted the specimens cited were col-
lected by Miss Marble and myself. —
1. DICRANELLA LONGIROSTRIS (Schwaegr.) Mitt. Jour. Linn. Soc.
Bot. 12:.30. 1869
St. JAN: without definite locality, Breutel, 1841.
2. LEUCOLOMA SERRULATUM Brid. Bryol. Univ. 2: 752. 1827
Leucoloma Riedlei Besch. Jour. de Bot. 5: 146. 1891.
St. THOMAS: on trees in wet woods, Riedlé.
3. OCTOBLEPHARUM ALBIDUM (L.) Hedw. Descr. 3: 15. 1791
St. THOMAS: on roots of an Anthurium, stone walls, between
Pearl and Bonne Resolution near Caret Bay, 1337.
4. LeucospryuM PoLaKkowsxkyi (C. Miill.) Cardot, Mém. Soc.
Sci. Nat. Cherbourg 32: 82. 1900
TORTOLA: on rotten wood, Sage Mt., W. C. Fishlock 83,
May, 1913. Also in Porto Rico, E. G. Britton 2518, April, 1913.*
5. FisstpeNs KEGELIANUS C. Miill. Linnaea 21: 18T. 1848
St. THomas and Sr. JAN: at base of palms and on the ground,
Breutel, 1841 (distributed as F. palmatus [Sw.] Hedw.). Sr.
JAN: wet bank, Bethania, Britton & Shafer 208a. St. Tuomas:
* Both these are additions to ranges since the publication of Part 2, Vol. 15,
of North American Flora.
Britton: Mosses or DANISH WEsT INDIES 3
moist banks near French wharf; Water Island, Cowell Point, 147;
St. Peter, 38, 98; Magin’s Bay, 191; Smith’s Bay, 1289, 1316,
T4509.
6. FIssIDENS ELEGANS Brid. Spec. Musc. 167. 1806
St. JAN; on wet rock, Bethania, Britton & Shafer 361. St.
Tuomas: Cowell Point, zoo.
7. SYRRHOPODON FLAVESCENS C. Miill. Syn. 1: 541. 1849
“St. JAN: on rotten wood, Bordeaux, 300 m. alt., Britton &
Shafer 549. TorToLa: without definite locality, W. C. Fishlock
82. May, 1913.
8. CALYMPERES RIcHARDI C. Miill. Syn. 1: 524. 1849
Calymperes Breutelii Besch. Ann. Sci. Nat. Bot. VIII. 1: 278.
1895.
Calymperes hexagonum Besch. |. c. 286.
Calymperes Hookeri Besch. |. c. 287.
St. THoMAs: without definite locality, Breutel (type of C.
Breutelii) ; without definite locality, L. C. Richard (type of C. hexa-
gonum); on rocks, summit of ridge, by roadside, north of Char-
lotte Amalia, 408; on bank, Cowell Point, ror. TORTOLA: Road-_
town to High Bush, 325 m. alt., Britton & Shafer 772.
9g. CALYMPERES LONCHOPHYLLUM Schwaegr. Suppl. 17: 133. pl. 98.-
1816 oe
TorTOLA: Sage Mt., W. C. Fishlock 85a, May, 1913.
10. HyMENOSTOMUM BrREUTELIT (C. Miill.) Broth.; E. & P.
Nat; Pil: ¥*7 486-< 1602
Weisia Breutelii C. Mill. Syn. 1: 664. 1849. Not Schimp.
Gymnostomum Breutelit Br. & Sch.; Paris, Index Bryol. 542. 1895.
Weisia senocarpa C. Miill. Syn. 2: 633. 1851.
Gymnostomum senocarpum Jaeger, Adumb. 1: 280. 1873.
Hymenostomum senocarpum Paris, Index Bryol. 597. 1895.
Weisia Pabstiana C. Mill. Bot. Zeit. 15: 382. 1857.
Weisia edentula Sull. Proc. Am. Acad. 5: 273. 1861. Not Mitt.
Hymenostomum cubense Hampe; Paris, Index Bryol. Suppl. 189.
1900.
4 Britton: Moss&és oF DANISH WeEsT INDIES
Sr. Jan: Emaus, Breutel (type locality); wet banks, Bethania,
Britton & Shafer 209; Rosenberg and Bordeaux, Britton & Shafer
288, 534, 569. ST. THOMAS: on the ground, Water Island, 155;
roadside banks, summit of ridge, Bonne Resolution, 1336;
Bordeaux, 1384; St. Peter, 1258; top of Flag Hill, Fitch & Shafer
1404. ToORTOLA: Peter Island, Britton & Shafer 860. ANAGADA:
without definite locality, Britton & Shafer 1039
11. Hyophila uliginosa E. G. Britton sp. nov.
Plants attached to rocks in stream, gregarious and matted
together by fresh water algae, soft and flaccid when moist; stems
simple, about 5 mm. high, branching at apex; leaves much dis-
colored and clogged with mud at base, green and spreading at
summit of stems, about I mm. long; base hyaline and oblong;
apex lingulate and slightly carinate, apiculate; margins plane,
entire or rarely denticulate with a few hyaline teeth at apex;
costa stout, papillose on back and smooth above ending in the
cuspidate point, in section showing one row of ducts and two
small bands of stereid cells; basal cells hyaline, oblong or square,
up to 16y long by 8yu wide, upper cells obscure up to 5y
in diameter, green and densely papillose, with several minute
papillae on each surface; dioicous; flowers and fruit unknown;
propagating by septate gemmae borne in clusters on brown fila-
ments in the axils of the leaves. [PLATE 1, FIG. I-6.]|
TYPE LOCALITY:—ST. JAN: Bethania, Britton & Shafer 367.
12. Phascum sessile E. G. Britton sp. nov.
Plants annual(?), gregarious inloose bare earth, on banks; stems
simple, or branching at base, with several rosettes from one root,
1-2 mm. high; leaves inrolled with conspicuous yellowish-white
costa when dry, bright green in color and spreading when moist,
few, 8-12, oblong at base, obovate above, I-1.25 mm. long by
0.4-0.5 mm. wide; costa percurrent or excurrent into a short
cuspidate point, terete and smooth on back, with a narrow dorsal
stereid band and 2-3 rows of large ducts; margins entire or
finely crenulate and papillose; upper cells hexagonal, up to 13 u
in diameter, densely chlorophyllose with 1-3 papillae on each
surface; lower cells hyaline, oblong, 10-12 rows, up to 40 uw long,
not papillose, occasionally curved and yellow and slightly auricu-
Jate at basal angles; paroicous, antheridia few, with paraphyses, in
small buds below or near the archegonia, of which occasionally —
several are fertilized making 2-3 fruits on one plant; calyptra
small, conic, split, slightly papilioss at apex; capsule immersed,
BRITTON: MosskEs oF DANISH WEsT INDIES 5
sessile on a small brown vaginule, globose, 0.5 mm. in diameter,
sharply apiculate, indehiscent; walls with irregular hexagonal
cells, 27-32 uw in diameter; spores brown, slightly roughened, 27-
30 w in diameter, maturing in spring. [PLATE I, FIG. 7—13.]
TYPE LOCALITY:—StT. THOMAS: Cowell Point, E. G. Britton go,
February 2, 1913.
DISTRIBUTION :—ST. THOMAS: Water Island, 150, 156.
This species belongs to the section Microbryum and is close to
P. Floerkeanum, but differs in the less acuminate and less subulate
leaves with plane margins without a yellow border, and more
chlorophyllose and papillose cells.
13. TORTULA AGRARIA (Sw.) Sw. FI. Ind. Occ. 3: 1763. 1806
St. JAN: Bethania, Breutel; Bethania, Britton & Shafer 241,
268. St. THOMAs: on limestone walls of old cemeteries, Breutel;
on damp earth, Nisky, 77; Cowell Point, 97; old walls, Crown
Estate, 450 m. alt., 1369; on rocks at waterfall, Magen’s Bay, 1315;
stone walls, Bonne Resolution School, 442.
14. Bryum micro-decurrens E. G. Britton sp. nov.
Plants gregarious, in loose soil, brown at base and also more
or less brown above, from the excurrent awns; stems dull green,
slender, erect and mostly simple, unbranched, not more than
5 mm. high, matted with brown tomentum at base; leaves
erect-appressed when dry, not twisted nor glossy; spreading when
moist, less than I mm. long by 0.2 mm. wide; costa wide at base
40 pw (at least .2 width of leaf), excurrent into a short subulate
brown awn, slightly toothed at apex and on awn; cells of blade
hexagonal, 27-40 » long X 10 » wide, basal cells shorter, oblong,
with a long, decurrent narrow wing of one row of cells, extending
down the stem to the next leaf; margins bordered by one row of
longer narrow cells 5 u X 54 u long, slightly revolute below and
serrulate above; only known from sterile specimens.
TYPE LOCALITY:—ANAGADA: rocky plain near settlement,
Britton & Shafer 1038. ;
DistTRIBUTION:—Mona Island, Britton & Hess 1751, 1753.
These specimens have been compared with type specimens of
B. decursivum C. Mill. from Porto Rico, kindly loaned to us by
Dr. Engler from the Royal Botanical Garden at Berlin, and,
though closely related by the decurrent narrow basal wing, ours
differs in its shorter leaves, serrate margins and shorter cells.
6 Britton: MosseEs oF DANISH WEsT INDIES
15. BryuM CRUEGERI Hampe; C. Miill. Syn. 1: 300. 1849
Dioicous plants bright yellowish green, shining gregarious in
loose soil in gravelly bed of stream; stems short, less than I cm.
high, upper part of stem erect, base decumbent red and radiculose;
leaves light yellowish green, glossy, spreading, lanceolate, acute
or acuminate, 2 mm. long X 0.35-0.5 mm. broad; costa narrow,
ending in the carinate, mucronate apex, margins entire below
slightly serrulate at apex, not bordered nor revolute; cells hex-
agonal 54-108 wu long X 13 broad, a few alar, shorter and
broader, square or oblong, not decurrent; sterile but often
propagating by gemmae.
St. THOMAs: in bed of stream at Tutu, 422.
Compared with Chas. Wright 63 from Cuba, distributed as
B. ovalifolium Sull., the leaves are slightly narrower and less con-
cave, but they are evidently closely related species and both
belong with the group of tropical American species having glossy
leaves, and flaccid red stems, resembling a Pohlia. | They grow
mostly along streams and form a closely related group, of iba
B. ripense C. Miill. from Jamaica is also a member.
16. PHILONOTIS SPHAEROCARPA (Sw.) Brid. Bryol. Univ. 2: 25.
1827
St. THOMAS: moist banks, Bonne Resolution 4g2z; Crown
Estate, 450 m. alt., 1368, 1458.
17. PHILONOTIS TENELLA (C. Mill.) Jaeger, Adumb. 1: 541.
1873-1874
St. JAN: on wet banks, near Corallenburg, Breutel, 1841;
Bethania, Britton & Shafer 208. TorToLtaA: near Roadtown,
325 m. alt., Britton & Shafer 773.
18. PIREELLA CYMBIFOLIA (Sull.) Cardot, Rev. Bryol. 40: 17.
1913 ;
St. JAN: on wet rock near Bethania, Britton & Shafer 350.
19. PTEROBRYUM ANGUSTIFOLIUM (C. Miill.) Mitt. Jour. Linn.
soc. BOt. 12:'426:. 1860
TortoLta: High Bush, Eggers 3240a, December, 1887; Sage
Mt., W. C. Fishlock 85, May, 1913.
BRITTON: MosseEs oF DANISH WEsT INDIES <
20. NECKERA DISTICHA (Sw.) Hedw. Descr. 3: 53. 1792
St. THOMAS: on rocks at St. Peter, 7456. TOoRTOLA: on rock in
forest, High Bush, 375 meters alt., Britton & Shafer 841.
21. NECKERA JAMAICENSE (Gmel.) E. G. Britton, Bull. Torrey
Club 40: 656. 1913
ST. JAN: on bark of trees at Bethania, Britton & Shafer 364.
22. CALLICOSTELLA BELANGERIANA (Besch.) Jaeger, Adumb. 2:
257. 1874-1875
St. JAN: on stones, Bordeaux, 300 meters alt., Britton &
Shafer 548.
23. STEREOPHYLLUM LEUCOSTEGUM (Brid.) Mitt. Jour. Linn.
Soc. Bot. 12: 543. 1869
St. Jan: Emaus, Breutel; Bethania, Britton & Shafer 1357.
St. THoMAs: on rocks in shade, ravine at Tutu, 1297; waterfall
near Magen’s Bay, 1314.
24. MITTENOTHAMNIUM DIMINUTIVUM (Hampe) E. G. Britton,
Bryologist 17: 9. 1914
St. JAN: on dead wood, Bordeaux, 400 m. ag Britton &
Shafer 595.
25. TAXITHELIUM PLANUM (Brid.) Mitt. Jour. Linn. Soc. Bot.
I2: 496. 1869
St. JAN: Breutel; Bethania, Britton & Shafer 358, 305, 300;
Bordeaux, 400 m. alt., Britton & Shafer 568, 580. St. THOMAS:
St. Peter, 1259; Crown Estate, 1360. Tortota: High Bush,
Britton & Shafer 839.
26. SEMATOPHYLLUM ADMISTUM (Sull.) Mitt. Jour. Linn. Soc.
Bot. 12: 485. 1869
St. JAN: Bordeaux, 300 m. alt., Britton & Shafer 554, 556, 578,
579. St. Tuomas: St. Peter, on rocks, 1257, 1457. TORTOLA:
High Bush, 375 m. alt., Britton & Shafer 814, 840.
27. SEMATOPHYLLUM SERICIFOLIUM Mitt. Jour. Linn. Soc. Bot.
12: 483. 1869.
TorTOLA: on logs in forest, High Bush, 375 m. alt., Britton &
Shafer 819
8 Britton: Moss&és oF DANISH WEsT INDIES
28. HAPLOCLADIUM MICROPHYLLUM (Sw.) Broth.; E. & P. Nat.
A. 2** 1067. . 1907
St. THomAs: shaded bank, Pearl to Bonne Resolution, 7335.
Explanation of plate 1
The figures were drawn by Mr. R. S, Williams from magnifications twice as
great as expressed in the numbers, which represent the magnification of the figures
as they stand in the reproduction.
HYOPHILA ULIGINOSA E. G. Britton
Fic, 1. Plant, natural size.
Fic. 2. Propagula, X 35
Fic. 3. Upper leaf, X 3
Fic. 4. Cells in upper part of leaf, X 200.
Fic. 5. Cells of leaf base, 200.
Fic. 6. Cross section in upper part of leaf, X 200.
PHASCUM SESSILE E. G. Britton
Fic. Plant, >< IT.
7
Fic. 8. Cross-section in upper part of leaf, X 200.
9. Cross-section near leaf base, X 200.
Fic. ro. Calyptra, X 35.
Fic. 11. Cells at basal angle, X 200.
Fic. 12. Cells in upper part of leaf, X 200.
Fic. 13. Upper leaf, X 20.
A new American fossil moss
ELIZABETH G. BRITTON AND ARTHUR HOLLICK
(WITH TWO TEXT FIGURES)
Some seven years ago it was our privilege to describe the
first known specimen of an American fossil moss in fruit, under
the name Glyphomitrium Cockerelleae,* found at Florissant,
iV
¥
Fr gop is Scudderi E. G. Britton & A. Hollick. Fic. 1. Photo-
graph of specimen, natural size.» Fic. 2a. 03, 14. Ja 1015. ;
Orton, C.R. The newer diseases of fruit trees and latest development
in their treatment. Proc. Adams Co. Fruit Growers Assoc. 1913:
77-89. f. I-5. 1914.
Orton, C. R. Some orchard diseases and their treatment. Proc.
State Hort. Assoc. Pennsylvania 55: 43-56. pl. 5-9. 1914.
Osterhout, W. J. V. Stetige Anderungen in den Formen von Antagonis-
mus-Kurven. Jahrb. wis. Bot. 54: 645-650. O 1914.
Otis, C. H. The transpiration of emersed water plants: its measure-
meat and its relationships. Bot. Gaz. 58: 457-494. f. 1-3 + charts
I-14. 21 D 1914.
Pease, A. S. A form of Potentilla tridentata. Rhodora 16: 194, 195
17 D 1914.
Piper, C. V., & Morse, W. J. Five oriental species of beans. U. S.
Dept. Agr. Bull. 119: 1-32. pl. 1-7. 25S 1914.
Prescott, A. Sweet flag and calla. Am. Bot. 20: 134, 135: | N 1914.
Quehl, L. Mamiliaria pseudofuscata Quehl, spec. nov. Monats.
Kakteenk. 24: 114-118. 15S. 1914. [Illust.]
Rahn, O., & Harding, H. A. Die Bemiihungen zur einheitlichen
Beschreibung der Bakterien ia America. Centralb. Bak. Zweite
Abt. 42: 385-393. 30 O 1914.
166 INDEX TO AMERICAN BOTANICAL LITERATURE
Rolfe, R.A. New Orchids. Decade 43. Kew. Mis. Inf. 1914: 372-
376.
Includes Zygopetalum Prainianum sp. nov. from Peru.
Rydberg, P. A. Carduaceae (Heleniae). N. Am. Flora 44: 1-31.
3 D 1914.
Includes paniaiaatee Pappothrix, Leptopharynx, Amauriopsis, Cephalobembix,
Trichymenia, g ov., and 35 new species in Venegazia (1), Psilostrophe (3), Baileya
(3), Perityle oy. phe tack (3), Laphamia (2), Loxothysanus (1), Bahia (2),
Hulsea (3), Tetracarpum (2), Hymenopappus (5), Othake (1), Rigiopappus (1),
Chaenactis (3).
Sears, P.B. The insect galls of Cedar Point and vicinity. Ohio Nat.
15: 377-388. pl. r8-2z. 18 D 1914.
Sherbakoff, C. D. Potato scab and sulfur disinfection. Cornell Agr.
Exp. Sta. Bull. 350: 709-743. f. 106. Au 1914.
Snow, L. M. Contributions to the knowledge of the diaphragms of
water plants. I. Scirpus validus. Bot. Gaz. 58: 495-517. f. 1-16.
21 D 1914.
Standley, P.C. The ferns of Brazos Canyon, New Mexico. Am. Fern
Jour. 4: 109-114. pl. z, 2. [D] 1914.
Stapf, O. Crataegus pubescens, forma stipulacea. Curt. Bot. Mag.
IV. ro: pl. 8589. D 1914.
A plant from Mexico.
Stewart, A. Notes on the anatomy of the Punctatus gall. Am. Jour.
Bot. 3: 531-546. pl. 51, 52. D 1914.
Tavares, J.S. Les Anonacées cultivées au Brésil. Broteria 12: 137~
147. pl. 2-6. D 1914.
Tavares, J. S. Le Goyavie: (Psidium guayava Raddi) au Brésil.
Broteria 12: 148-154. pl. 7, 8. D 1914.
Tidestrom, [, Notes on the flora of Maryland and Virginia—II.
Rhodora 16: 201-209. f. r-13. D 1914.
Trelease, W. Phoradendron. Proc. Nat. Acad. Sci. 1: 30-35. 15 Ja
IQI5.
Victorin, M. Une plante “‘fortifiée” de notre flore. Le Naturaliste
Canadien 41: 49-51. O 1914. :
Victorin, M. Une probleme biologique. Le Naturaliste Canadien qr:
68, 69. N 1914.
Visher, S. S. The biology of Harding County, northwestern South
Dakota. S. Dakota Geol. Surv. Bull. 6: 11-103. pl. 1-6. 1914.
INDEX TO AMERICAN BOTANICAL LITERATURE —- 167
Vogt, R. Notes on seedlings. Am. Mid. Nat. 3: 287-289. pl. 4-10.
17 Jl 1914. {
Vogt,R. The ecology and anatomy of Polygonatum commutatum. Am.
Mid. Nat. 4: 1-11. pl. 1-3. Ja 1915.
Waddell, J. Rubber: wild, plantation and synthetic. Pop. Sci. Mo.
85: 443-456. N 1914.
Walcott, C. D. Cambrian geology and paleontology III. No. 2.
Pre-cambrian Algonkian algal flora. Smithsonian Misc. Col. 64:
77-156. pl. 4-23. 22 Jl 1914. °
Includes descriptions of Newlandia, Kinneyia, Weedia, Greysonia, Copperia,
Collenia, Camasia, and Gallatinic, gen. nov., and ten new species.
Weatherwax, P. Aphanomyces phycophilus De Bary. Proc. Indiana
Acad. Sci. 1913: 109-111. f. 1-6. I914.
Weatherwax, P. Ecological notes on certain White River algae.
Proc. Indiana Acad. Sci. 1913: 107, 108. 1914.
Weingart, W. Phyllocactus Ruestit Weing. spec. nov. Monats.
Kakteenk. 24: 123-127. 15S 1914.
Weingart, W. Phyllocactus stenopetalus S.-D. und Phylloc. latifrons
Zucc. Monats. Kaktéenk. 24: 129, 130. 15 O 1914.
' Wheeler, W. M. Observations on the Central American Acacia ants,
Trans. Second Entomological Congress 1912: 109-139. I9I2.
Whitford, A. C. On a new fossil fungus from the Nebraska Pliocene.
Univ. Stud. 14: 1-3. pl. r, 2. 1914.
Winton, K. B. Histology of flax fruit. Bot. Gaz. 58: 445-448. f.
I-4. 16N 1914.
Wolf, F.A. Eggplantrots. Myc. Centralb. 4: 278-287. f. 1-4. 1914.
Wolf, F. A. A leaf disease of walnuts. Myc. Centralb. 4: 65-69.
f. 1-6. 1914. [Illust.]
Wolf, F. A. Leaf spot and some fruit rots of peanut (Arachis hypogoea
L.). Alabama Agr. Exp. Sta. Bull. 180: 127-150. pi. 1-5. D 1914.
Wolf, F. A., & Massey, A.B. Citrus canker. Alabama Agr. Exp. Sta.
Cire. 27: 97-101. f. 7-6. My 1914.
Woodcock, E. F. Observations on the development and germination
of the seed in certain Polygonaceae. Am. Jour. Bot. 1: 454-476.
pl. 45-48. 4D 1914.
Woodward, R. W. Station for fruiting Euphorbia Cyparissias. Rho-
dora 16: 167, 168. S 1914.
168 INDEX TO AMERICAN BOTANICAL LITERATURE
Woodward, R. W. Forms of Arenaria lateriflora. Rhodora 16: 179,
180. 23 N 1914.
Wright, C. H. Tillandsia Benthamiana, var. Andrieuxit. Curt. Bot.
Mag. IV. 10: pl. 8576. S 1914.
Found in Central America.
Wright, C. H. Agave bracteosa. Curt. Bot. Mag. IV. ro: pl. 8581.
O 1914.
A plant from northern Mexico.
Zimmermann, Ce Contribuicdo para o conhecimento das diatomaceas
da provincia de Mocambique. Broteria 12: 155-162. D 1914.
VOLUME 42, PLATE 9
But_. TORREY CLUB
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AUSTRALASIAN MOSSES
.
.
DIXON
Vol. 42 No. 4
BULLETIN
OF THE
TORREY BOTANICAL CLUB
ce my
APRIL; -4ots
The vegetation of Connecticut
IV. Plant societies in lowlands*
GEORGE E. NICHOLS
(WITH FIFTEEN TEXT FIGURES)
= term LOWLAND, as used in the present paper, embraces
primarily lakes, ponds, and swamps.f The word LAKE may be
employed to designate any body of water surrounded by land,
but more commonly this term is applied to relatively large bodies
of water, smaller water bodies being referred to as POoNDs.
Throughout the present paper the two terms are used more or less
indiscriminately. A Swamp may be defined as any area where
the ground is» saturated with water throughout a good part of the
year, but where, at least during most of the year, surface water
does not accumulate. As will be seen presently, it is not always
possible to differentiate absolutely between lakes and ponds on the
one hand and swamps on the other. Thus, the transition froma
lake to the swamp at its margin may be so gradual that one cannot
say where the one leaves off and the other begins. Similarly, a
swamp may merge by almost imperceptible gradations with the
neighboring upland, so me is likewise difficult to draw a sharp
line of demarcation between Yowlands and uplands.
* Contribution from the Osborn Botanical A ETE The previous numbers
of this series have a si me in Torrey§, 13: 89-112; 199-215. 1913, and 14:
167-194. 1914. In this paper, as in the me the as followed is that
of Gray’s Manual, seh edition
+ In earlier papers this term is also — used to designate one of the three
Sail provinces into which the state is divided, viz., Central Lowland, as con-
- trasted with Eastern and Western Highlands.
[The BULLETIN for March (42: 93-168. pl. 9) ‘was issued March 16, 1915.]
169
170 NICHOLS: THE VEGETATION OF CONNECTICUT
GEOLOGICAL RELATIONS OF LAKES AND SWAMPS
The Effect of Physiographic Factors on the Distribution of Lakes
and Swamps.—In a general way it can be said that lakes and
swamps belong to the same family, and differ only in age.* Lakes,
however, ‘‘are temporary features of a landscape, and can only
exist where the land has been recently raised above the sea or
modified by some widely acting agent’’f such as glaciers. Sooner
or later they are destined either to be drained through the activity
of streams cutting down their outlets, or else to be filled in through
the washing in of inorganic sediment from without and the ac-
cumulation of vegetable debris from within.
Considered from the standpoint of their physiographic origin,
Connecticut lakes may be grouped under three heads: GLACIAL,
RIVER, and CoastaL. The term GLACIAL LAKE is here used in
its broadest sense, to include lakes formed in any sort of a depres-
sion resulting from glacial activity. Glacial lakes may further
be subdivided as follows: (1) MoratnaL Laxes—those associated
with accumulated glacial debris of any description; and (2) ScooP
LaxEs—those which occupy depressions scoured out of the under-
lying rock by the ice. Lake Saltonstall, near New Haven, affords
a striking example of a scoop lake. Thislake is about three miles
long and hasan extreme depth of 108 feet, more than eighty feet
below sea level; yet notwithstanding that the tides rise and fall
at its very edge, its water is fresh and constitutes part of the New
Haven water supply. On the whole, however, scoop lakes are
far less abundant and of much less importance in Connecticut
than morainal lakes, and of the 1,026 lakes indicated on the topo-
graphic map of the state§ the great majority belong to this latter
class. Morainal lakes may originate under a variety of conditions.
Thus, (a) they may occupy depressions in the ground moraine;
(6) they may occur behind dams formed by the heaping up of
glacial till; (c) they may occupy depressions in stratified drift; or
(d) they may develop behind the ridges of stratified drift knoy
as eskers. Morainal lakes of the first two types here mentio
* Possible exceptions to this statement are pointed out in later paragraphs.
+ Rice, W. N., and Gregory, H. E. Manual of the wiht of Connecticu
Geol. and Nor, Hist. Surv. Connecticut, Bull. 6: 248. 190
t See Rice and Gregory, op. cit., p. 249.
§ See Rice and Gregory, op. cit., p. 247.
NICHOLS: THE VEGETATION OF CONNECTICUT 171
are common throughout the state, but are especially characteristic
of the higher, more rugged sections. Morainal lakes of the last
two types are best developed in the lower, leveler, central portion
of the state. Lakes occupying depressions in stratified drift are
well exemplified by the ‘‘kettle-hole’’ ponds, so abundant in the
sand-plains near New Haven; while Lake Compounce is a good
illustration of a lake dammed in by an esker.
RIVER LAKEs or Ponps for the most part are associated with
old-age, meandering streams. In general, they are of two sorts:
(1) Ox-Bow and (2) MarcinaL. Ox-bow ponds, originating in
flood time as a result of the abandonment of old water courses,
are common along sluggish streams throughout the state (Fic. 1).
1G. 1. Ox-bow swamp along Muddy River, North Haven. Inthe foreground
are seen Castalia odorata (in the water), Dulichium arundinaceum (along the left
bank), tufts of Carex stricta (on both banks). Elm, etc., in the background.
Marginal ponds are often caused by the deposition of sediment
along the bank of a stream at high water, thereby gradually
building up a sort of dike or ridge which cuts off the channel from
172 NicHOoLs: THE VEGETATION OF CONNECTICUT
the remoter parts of its flood plain. When the stream overflows
its banks these latter areas are flooded, and after the recession of
the water there may be left behind more or less extensive shallow
ponds. Such ponds may be seen along the Connecticut River and
elsewhere, but usually they are of a very temporary nature. Far
more common than marginal ponds are marginal swamps. These
may originate in the manner just described, but more often seepage
water is responsible in large measure for the maintenance of the
swampy condition. Swamps due to seepage will be discussed
presently.
CoastTAL LAKES, PonDs, and ae are formed mainly behind
barrier beaches, in sheltered bays and harbors, and along the
lower courses of the larger rivers. The ecological relations of
these will be considered in a later paper, in connection with plant
societies along the coast.
The Effect of Topography and Ground Water Level on the
Character of Lakes, Ponds, and Swamps.—It is evident that topog-
raphy, as moulded by physiographic forces, is an all-important
factor in governing the distribution of lakes and ponds. This is
also true with respect to swamps, but it is further true, as will be
‘pointed out presently, that swamps may be developed in areas
where bodies of standing water could not exist. Yet while their
development is dependent on suitable topographic conditions,
the presence or absence of lakes, ponds, and swamps is further
conditioned by the relation between the level of the land surface
and the water table; and this in turn is influenced by the amount
of precipitation and the character of the substratum. The
surface of the ground water rises and falls with the land surface,
but with smaller differences of elevation. Thus, while it reaches
its highest elevation in the hills it stands farthest from the surface
of the ground there. Similarly, in the depressions it stands lower,
but is nearer the surface of the ground. Wherever it reaches the
surface, lakes, ponds, streams, or swamps may be formed.*
Owing to the variation in the amount of precipitation at different
seasons of the year, the level of the water table is constantly
* For an extended discussion of the underground water resources of Connecticut,
see Gregory, H. E., and Ellis, E. E., U. S. G. S. Water-Supply — 232, Pp. 1-200,
pls: -5 + figs. 1-31. 1909.
NICHOLS: THE VEGETATION OF CONNECTICUT 173
fluctuating. It is highest during winter and spring, the seasons
of maximum precipitation; it is lowest in late summer, toward
the end of the season of minimum precipitation. In the following
paragraph, the writer has attempted to bring out, with the help
of diagrams (Fic. 2), the important bearing which these various
factors may have on the nature of lakes, ponds, and swamps.
With reference to their relation to topography and water table,
four fairly distinct types of lowland may be recognized. (1) PER-
MANENT LAKES or PonDs.—Where the relationship between the
controlling factors, i. e. topography and water table, is such that
a tract of land is submerged to an appreciable depth by standing
water throughout the year (Fic. 2, A), it is obvious that a permanent
lake or pond will exist. It need hardly be remarked that such a
water body can arise only within a completely enclosed basin.
Lakes fed entirely by subterranean waters are not uncommon,
but more frequently this source of supply is supplemented by
surface streams. A permanent water body can exist in a closed
basin only where the amount of water flowing in exceeds the
amount lost through underground drainage plus the amount lost
through evaporation. (2) INTERMITTENT LAKES or PoNDs.—
Very often the relationship between the various factors involved
is such that a basin is filled with water in spring, but later in the
season, as a result of the lowering of the water table plus increased
evaporation, the surface water vanishes, leaving the bottom of the
depression quite dry, or else merely swampy (Fic. 2,B). Shallow,
intermittent ponds of this sort are of frequent occurrence, being
especially well developed in parts of the sand plains north of New
Haven. (3) PERMANENT SwampPs.—It often happens that, in
spite of an abundant supply of underground water, topographic
relations are such as to prevent pond formation. Thus, seepage
water may keep the ground on a side hill wet throughout the year,
but as fast as the water reaches the surface it runs off to lower levels.
Similarly there may be a series of perennial springs along the base
of a hill, but in the absence of an enclosed basin of some sort to
contain the water, pond formation manifestly cannot take place,
and swampy conditions prevail. Ideal conditions for permanent
swamps are presented by the flat floors of the old glacial drainage
valleys, so common throughout central Connecticut. Lying well
&
174 NICHOLS: THE VEGETATION OF CONNECTICUT
Fic. 2. Diagrammatic vertical sections to represent the relation of topography
and level of ground water to character of lakes, ponds, and swamps. Surface of
Ground represented by continuous line (~——), Level of Ground Water in Spring
by long dashes (-— -~—), Level of Ground Water in Late Summer by short dashes
)
A
Permanent Swamp; cross section of a glacial drainage valley. rom this diagram
the relations of topography and water table might at first thought appear ceconerene
with those represented by diagram A; but a longitudinal section would show t
(C) depression to be in the nature of an open trough, rather than an enclosed basin
(asin A). D. Periodic Swamp. For further discussion, see text.
NICHOLS: THE VEGETATION OF CONNECTICUT 175
below the general level of the water table at all seasons, as they
usually do, the surface of the ground is kept constantly wet by
springs and seepage water, but the surplus is continually being
carried away by the small streams which drain these areas. A
cross section of one of these glacial-drainage-valley swamps is
shown by Fic. 2,C. (4) PErRtopic Swamps.—These bear much
the same relationship to permanent swamps that intermittent
ponds bear to permanent ponds. In other words, in an area
where the relations between topography and ground water level
are such as to preclude pond formation at any season of the year,
but where the ground is saturated with water at certain seasons and
‘dry at others, a periodic swamp may be said to exist. Swamps
of this sort are common on every hand—in low grounds, in shallow
depressions of any description (as in Fic. 2, D), and on side hills.
Between the four types of lake, pond, and swamp above de-
scribed, all intergradations exist, so that it is not always easy to
assign a given lowland area to a definite category. But while this
classification is necessarily elastic, and while improvements can
doubtless be suggested, the scheme here proposed, it is believed
by the writer, will prove serviceable in the study of the vegetation
of lowlands, not only in Connecticut but elsewhere as well.
THE ROLE OF VEGETATION IN THE CONVERSION OF LAKES INTO
SWAMPS
The important r6le commonly played by plants in the conver-
sion of lakes into swamps has long been recognized, and the manner
in which this transformation is brought about may appropriately
be outlined at this point. When the plants in a lake die, their
remains sink to the bottom where, because of insufficient oxidation,
the vegetable debris is only partially decomposed. In this way
there collects on the floor of the lake a layer of vegetable muck,
or peat; and through the continued addition of fresh layers the
deposit is gradually thickened and built upward. This con-
structive process may go on until ultimately the surface of the
deposit reaches the level of the water, when the lake gives way toa
swamp (see Fic. 3). But the rate at which the substratum is
built up and the length of time which elapses before it reaches the
water level varies greatly in different parts of a lake. As will
176 NICHOLS: THE VEGETATION OF CONNECTICUT
be shown presently, plants grow most luxuriantly in shallow water;
they may be practically absent from the deeper areas. It follows,
therefore, that the accumulation of muck or peat proceeds much
more rapidly in shallow than in deep water—so much so, in fact,
that the shoreward parts of a lake may have become completely
filled in before any appreciable accumulation has taken place in
the deeper areas. The filling in of deep lakes usually proceeds
centripetally. This is due to the fact that the shoreward zones of
vegetation, in consequence of their more vigorous growth, exhibit
a tendency to push outward into deeper water. Where this
tendency is pronounced, the shoal water zones may completely
override the deeper water zones, at the same time causing the
lakeward slope of the deposit to become much steeper.* The
filling in of the deeper parts of a lake may also be effected toa
varying degree by the accumulation of loose debris from the ad-
joining shallows or by the deposition of sediment in flood time,
while various plankton forms may contribute in a small measure
to the deposit. But asa rule, the encroachment of the shoal water
vegetation is the most important agency in bringing about the
filling in of the deeper parts of deep lakes. The most extreme
development of this method of filling is seen in connection with
floating mat formation (Fic. 10), the discussion of which is reserved
for later paragraphs.
THE SUCCESSION OF PLANT SOCIETIES IN LAKES AND SWAMPS
Succession in Lakes.—Coincident with the upbuilding of the
substratum through deposition of muck or peat, as outlined in the
preceding paragraph, transformations occur in the character of
the vegetation growing on the lake’s bottom. For, as the depth
of the water diminishes, it becomes possible for plants to develop
which were unable to grow in the deeper water. And as these
shallow water plants increase in number and abundance, they
may crowd out and eventually replace the deeper water species.
Thus there may follow one another a series of plant societies,}
* For a more detailed description of this process, see Davis, U. S. Dept. Interior,
Bur. Mines, Bull. 16: 22, 23. I9grt.
t The terms Plant Society and Plant Association, in the present series of papers,
are used indiscriminately and in their broadest sense. Except in referring to the Clit
matic Formation, the use of the term Formation has been avoided.
NICHOLS: THE VEGETATION OF CONNECTICUT 177
each one of which, by helping to raise the bottom of the lake to a
higher level, prepares the way for less hydrophytic societies, but
at the same time, by so doing, brings about its own extermination.
It is a familiar fact that the plants which fringe the edges of so
many lakes are commonly massed in more or less definite bands
or zones that tend to be concentric with respect to the deeper
parts of the lake. The floristic composition of these zones in
any given lake is determined largely by the ecological requirements
of the various species of plants which happen to be present, in
relation to the depth and clearness of the water. In progressing
from the deeper parts of a lake toward the shore (Fic. 3) the zones
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Fic. 3. Diagrammatic vertical section through the marginal portion of a
shallow lake which is being filled in through the accumulation of vegetable debris on
the bottom, showing the succession of plant societies which may accompany the
transformation of a lakeinto aswamp. Sequence of Zones (or Stages): (1) Pond-
weed Zone; (2) Water-lily Zone; (3) Bulrush—Pickerel-weed—Cat-tail Zone; (4)
Sedge Zone; (5) Shrub Zone; (6) Swamp Forest.
encountered, as characterized by the dominant plants, are usually:
(1) the PonDWEED (Potamogeion) ZONE, (2) the WATER-LILY
(Castalia, Nymphaea) ZoNE, (3) the BuLrusH (Scirpus)—
PICKEREL-WEED (Pontederia)—Cat-Tait (Typha) ZONE (or ZONES),
(4) the SEDGE (Carex, etc.) ZonE. Modifications of this arrange-
ment are frequent, but in a general way the sequence is the same
throughout the northern United States.
178 NICHOLS: THE VEGETATION OF CONNECTICUT
Reference has already been made to the succession of plant
societies which accompanies the building up of the lake bottom.
It has been found that this dynamic VERTICAL SUCCESSION corre-
sponds closely with the apparently static HORIZONTAL ZONATION
just outlined. Thus, in a hypothetical case, it may safely be
assumed that the pondweeds now growing in a given locality
are destined to be succeeded by water-lilies, the water-
lilies by pickerel-weed, and these in turn by sedges. Conversely,
sedges growing on a mucky shore have very likely been preceded
by pickerel-weed, etc. This general codrdination between
the contemporaneous horizontal sequence of zones and the his-
torical or vertical order of succession has been verified repeatedly
by the stratification of plant remains observed in peat deposits,
and is of great assistance in reconstructing the past or predicting
the future course of events in any specific locality.
As has been pointed out by C. A. Davis in his comprehensive
essays on the ecology of peat formation in Michigan,* there are
relatively few of the highly organized vascular plants which can
grow when entirely submerged in water. Thus, out of the more
than 1,900 seed plants known to occur in Connecticut, scarcely
ninety, or less than five per cent., have this ability. ‘Of these
plants which will grow in water, only a few, mainly Potamogetons or
pondweeds, can establish themselves at a depth greater than ten
feet from the surface, while the majority of submerged plants
grow in less than six feet of water, unless it is unusually clear.’’f
The maximum depth at which seed plants can grow rarely exceeds
fifteen to eighteen feet, so that the deeper parts of many lakes,
owing to insufficient light, heat, and aeration, are practically
destitute of vegetation. The first stage in the succession of plant
associations which accompanies the filling in of a lake is the Ponp-
WEED STAGE, characterized by the predominance of submersed
aquatic plants, especially of species of Potamogeton. This genus
is represented in Connecticut by no fewer than twenty-eight
species, the commonest submersed forms of which are P. Rob-
binsii, P. Richardsonii, P. pusillus, P. amplifolius, and P.
heterophyllus. The two species last named are capable of develop-
* Geol. Surv. Michigan 1906: 97-286. 1907.
+ Davis, 1907, loc. cit., p. 129.
NICHOLS: THE ‘ VEGETATION OF CONNECTICUT 179
ing two sorts of leaves, viz., submersed and floating; but the
latter type is seldom formed on plants growing in water more
than five feet deep. Flowers likewise are usually suppressed,
except in shallow water, reproduction being accomplished solely
by vegetative means. Other plants frequently present in con-
siderable abundance in the pondweed association are Najas
flexilis, Vallisneria spiralis, and Elodea canadensis. The lack
of dependence of many of these water plants on the sexual
methods of reproduction is well exemplified by the latter species.
Introduced into Europe from America about 1836, this ‘‘ Wasser-
pest’’ has become widely spread over that continent, notwithstand-
ing that it has never been observed there in fruit. Fic. 4 shows a
Fic. 4. Submersed aquatics from Twin Lakes, Salisbury e plants were
placed on papers in the bottom of a boat and Sei Paeae rh tsa shown
are as follows: 1. Potamogeton nh 2. P. lucens; 3. P. natans; 28 P. ampli-
folius; 5. P. heterophyllus; 6. Char ory page Nae ais ib 8. Elodea
Canadensis; 9. Sparganium aa tn 10. LEN Robbinsii; 11. Utricularia inter
media; 12. Najas flexilis.
representative collection of aquatic plants from a lake in northern
Connecticut. The majority of such plants have roots and grow
attached to the bottom, but there are a number of forms, notably
180 NICHOLS: THE VEGETATION OF CONNECTICUT
the bladderworts (species of Utricularia), Ceratophyllum demersum,
and sometimes Ranunculus aquatilis, which are free floating and
entirely unattached to the substratum. These usually grow near
the surface and may be very abundant in sheltered coves. The
pondweed zone, and the succeeding zones as well, may be broad
Fic. 5. Water-lilies (mostly Castalia odorata) at Round Pond, North Ston-
ington.
or narrow, according as the bottom of the lake slopes gently or
steeply.
In the absence of competition submersed aquatics develop best
in shallow water, but as soon as the bottom has been raised high
NICHOLS: THE VEGETATION OF CONNECTICUT 181
enough, say within five or six feet of the surface, a higher order
of plants—higher in an ecological sense—is able to develop, viz.,
plants whose leaves float upon the surface (Fic. 5). Most im-
portant among these are the water-lilies (Castalia odorata and
Nymphaea advena), from the almost invariable presence of which
this phase in the succession is called the WATER-LILY STAGE. Asa
rule the water-lilies come in with such force that they crowd out
most of the submersed aquatics, for the shade produced by their
large leaves is so great that nothing can live underneath them.
One of the most remarkable structural peculiarities of the water-
lilies is seen in their enormous rhizomes. These organs persist
year after year, growing in length, sending out roots into the soft
muck, and branching at intervals. In Nymphaea advena they
Fic. 6. Bulrushes (Scirpus validus) at Twin Lakes, Salisbury.
frequently acquire a length of fully eight feet, with a diameter of
from two to three inches. When present in abundance they fulfil
an important ecological rdle by holding together the loose deposits
at the bottom of the lake. This same end is accomplished, though
probably less efficiently, by the weak, slender roots and rhizomes
of various submersed aquatics. Frequently other plants with
floating leaves, such as Brasenia Schreberi and Nymphoides lac-
unosum, together with Potamogeton epihydrus and similar hetero-
182 NICHOLS: THE VEGETATION OF CONNECTICUT
phyllous pondweeds, grow in company with the water-lilies. The
duckweeds (Spirodela and Lemna) and other free-floating forms
also commonly occur here, but these, as might be expected, are
not definitely restricted to any particular association.
With the continued shoaling of the water it becomes possible
for plants to develop which root at the bottom and are partly sub-
merged but whose foliage is raised above the surface of the water.
As these increase in number, and as more and more light is inter-
cepted by their aerial photosynthetic organs, species with floating
leaves become scarcer and may be completely crowded out. This
stage, because of the widespread prominence of the lake bulrush
(Scirpus validus), the pickerel-weed (Pontederia cordata), and the
cat-tail (Typha latifolia), may be referred to in a general way as the
BULRUSH—PICKEREL-WEED—CAT-TAIL STAGE. But quite as often
as not only one or at the most two of the character species will be
conspicuous in a given lake or pond. Furthermore, where more
than one is present, there is a marked tendency for one or another
form to dominate locally. It would frequently seem as though the
one first to arrive on the scene gained control. For these and other
reasons it is convenient to divide this stage into the BuLRusg, the
PICKEREL-WEED, and the CaT-TAIL SuBSTAGES. Roughly speak-
ing, these three substages may be regarded as parallels. Thus in
a given pond the pickerel-weed may fill the place occupied by the
bulrush in another. Yet these substages cannot be regarded as
absolute ecological equivalents. This is evidenced by the fre-
quently observed zonal arrangement of the species concerned where
two or three are present in the same pond. The cat-tail grows best
in water only a few inches deep; the pickerel-weed thrives in water
from six inches to nearly two feet deep; while the bulrush, although
it develops best in shallow water, can growin water more than five
feet deep. Of the three species, the pickerel-weed (Fic. 7) is the
most widely distributed and from an ecological standpoint the most
important in Connecticut lakes and ponds. Scarcely a pond is
encountered from which this plant is absent, and more often than
not it forms a conspicuous fringe in the shoal water along the shore.
The bulrush is abundant in many lakes, as at Twin Lakes, Salis-
bury (Fic. 6), but quite as often as not it is absent or else too poorly
developed to be of any ecological significance. The cat-tails are
NICHOLS: THE VEGETATION OF CONNECTICUT 183
conspicuous marginal plants in some ponds, but on the whole
cat-tail swamps are more characteristic of low flood plains along
rivers than of pond margins. Very often a luxuriant development
of cat-tails follows upon the removal of the original vegetation
from a swampy tract by drought or some similar disaster. In
Fic. 7. Pickerel-weed (Pontederia cordata) at Twin Lakes, Salisbury. In the
background is an extensive growth of cat-tails and bulrushes.
the subjoined list are given a number of other aquatics with aerial
foliage which are found in greater or less abundance growing in
shallow water along mucky shores in many lakes and ponds.
Alisma Plantago-aquatica Pontederia cordaia
Cladium mariscoides Sagittaria latifolia
Dulichium arundinaceum Sparganium americanum
Orontium aquaticum var. androcladum
Peltandra virginica Sparganium eurycarpum
To these should perhaps be added the curious AMPHIBIOUS
PLants, Proserpinaca palustris, Sium cicutaefolium, and Cicuia
bulbifera,—species adapted to alternate submergence and emer-
gence.
184 NICHOLS: THE VEGETATION OF CONNECTICUT
Most of theplantscl teristic of the bulrush—pickerel-weed—
cat-tail stage in the lake-swamp succession are provided with more
or less extensive rhizome systems, which ramify near the surface
of the mucky substratum, thereby tending to make the bottom
more solid. The building up of the substratum proceeds much
more rapidly as the depth of the water decreases, due partly to
the denser plant population of the shoreward zones, as compared
with those farther lakeward, and partly to the fact that mechanical
tissues, which naturally resist decay, are much more highly de-
veloped in plants with aerial organs than in plants which are
entirely or for the most part submerged.
Succession in Lake Swamps—As soon as the bottom of a
pond has been built up high enough so that the substratum is
exposed during part of the year, aquatic plants begin to give way
to terrestrial species. For the sake of convenience, the various
stages in the succession up to this point may be referred to col-
lectively as the LAKE SERIES, and the subsequent stages as the
SWAMP SERIES. But it is manifestly impossible to draw a sharp
line of demarcation between the two; for not only are various
species of the bulrush—pickerel-weed—cat-tail association quite
characteristic of wet swamps, but it is often easy to observe within
this zone, along the shore of a lake, a gradual transition from lake
on the one hand to swamp on the other. The bulrush—pickerel-
weed—cat-tail stage, then, represents a connecting link between
the vegetation of lakes and that of swamps. Swamps which
have originated through the filling in of a lake or pond, after the
manner above described, may be termed LAKE SwWAmpPs.
Almost always, in a succession like the one under discussion,
the first true land plants to appear in the succession are sedges;
hence, the first swamp stage is known as the SEDGE STAGE. Toa
greater degree than any of the plants that have preceded them,
the sedges are equipped with tough rhizomes and copious slender
roots, through the interweaving of which the soil is more firmly
bound together and converted into a compact turf. By far the
commonest, and ecologically the most important, sedge of Con-
necticut swamps is Carex stricta, a species which is readily recog-
nized from its tussock-forming habit (Fic. 8); but frequently other
plants with grass-like foliage are equally or even more prominent.
NICHOLS: THE VEGETATION OF CONNECTICUT 185
Among these latter, special mention should be made of the spike
rush Eleocharis tenuis, the grass Glyceria canadensis, and the rush
Juncus effusus. A swamp with predominantly grass-like vegeta-
tion is commonly referred to as a MARSH.
The sedge association is much richer in species than any of the
Fic. 8. Carex stricta swamp along the Farmington River, Simsbury, showing the
tussock-forming habit of this sedge. Picture taken in winter.
preceding stages in the succession, but the plants are still almost
exclusively herbaceous. In the subjoined list are given a number
of additional vascular species commonly found here.
Acorus Calamus Hypericum virginicum
Asclepias incarnata Impatiens biflora
Aspidium Thelypteris* Iris versicolor
Aster novi-belgii Juncus acuminatus
Bidens frondosa Juncus canadensis
Calamagrostis canadensis* Lycopus americanus
Campanula aparinoides* Lysimachia terrestris
Carex filiformis Mimulus ringens
Carex lurida* Osmunda regalis
Carex stipata* Pogonia ophioglossoides
186 NICHOLS: THE VEGETATION OF CONNECTICUT
Cicuta maculata Ranunculus septentrionalis
Cyperus strigosus* Saxifraga pennsylvanica
Epilobium densum Scirpus cyperinus
Eriophorum virginicum Scutellaria galericulata
Eupatorium perfoliatum* _ Solidago neglecta*
Eupatorium purpureum* Thalictrum polygamum*
Geum rivale Verbena hastata*
Woodwardia virginica
A Carex stricta swamp might well be said to represent a com-
posite habitat. For while the low wet areas between the tussocks
afford a suitable substratum for markedly hydrophytic species
like Glyceria canadensis, Iris versicolor, Lysimachia terrestris,
and Scirpus cyperinus, the tussocks, higher and drier than the
intervening spaces, offer a foothold for plants like Aspidiwm
Thelypteris, Calamagrostis canadensis, Eupatorium purpureum, and
Verbena hastata, which thrive best in soil that is not permanently
saturated.* In sedge swamps where tussocks of Carex stricta are
absent, the more mesophytic species, such as Aspidium Thelypteris
and Calamagrostis canadensis, are for the most part restricted to
the higher, drier areas, and frequently form a distinct zone around
the margin of the swamp.
The sedge stage may be of short or long duration. The
building up of the substratum continues, but with less speed than
heretofore, due to the accelerated and more thorough oxidation
of plant remains which results from exposure to the air. And
just as earlier in the series submersed aquatics were shaded out, so
to speak, by water-lilies, so in the competition for light the low
herbaceous species of the sedge association are destined in the
course of time to be superseded as the dominant type of vege-
tation by shrubs, and these in turn by trees. Many of the plants
of the sedge stage persist into the SHruB STAGE, but they come to
occupy a more and more subordinate position. The commonest
shrubs of open swamps are the alders (Alnus rugosa [Fic. 9] and
Alnus incana). Indeed, an alderless swamp is a rarity. Along
with the alders occur many other shrubs, the most important of
which are indicated below.
* All species starred (*) in the above list belong to this latter class. Some of
those not starred should perhaps also be included here, but it is very difficult to
draw sharp lines.
NICHOLS: THE VEGETATION OF CONNECTICUT 187
Cephalanthus occidentalis Rosa carolina
Clethra alnifolia Rubus hispidus
Cornus Amomum Salix discolor
Ilex verticillata Salix sericea
Lyonia ligustrina Spiraea latifolia
Pyrus melanocarpa Spiraea tomentosa
Rhododendron viscosum Vaccinium corymbosum
Rhus vernix Viburnum dentatum
Fic.9. Alders (Al )inaswamp, East Haven. View taken in January.
During the early has of the shrub stage, certain of these
shrubs may be more conspicuous than the alders. This is espe-
cially true of the Cephalanthus, which frequently comes in so early
in the succession and occupies the territory invaded in such force
that the sedge stage may be almost, if not quite, suppressed. But
ultimately, because of its greater height growth, its relative toler-
ance of shade, and its ability to adapt itself to a wide range of
soil conditions, the alder usually becomes the predominant form.
The terminal member of the lake-swamp succession is a
188 NICHOLS: THE VEGETATION OF CONNECTICUT
SwAmP Forest, in which red maple (Acer rubrum) is invariably
present and more often than not predominates. Commonly
growing with it are Ulmus americana, Betula lutea, and Fraxinus
nigra. In northern Connecticut the tamarack (Larix laricina) is
an important swamp tree,* while the coast white cedar (Chamae-
cyparis thyoides) is similarly common in the eastern part of the
state. Sometimes other trees, to be mentioned elsewhere (p. 193),
occur in lake swamps. Several of the shrubs of open swamps are
equally characteristic of wooded swamps, but the herbaceous
vegetation is composed very largely of forms not heretofore noted.
The following lists of species commonly represented in the under-
growth of swamp forests have been compiled from observations
made in more than twenty swamps—exclusive of bogs—in various
sections of the state.
Wooby PLANTS
Alnus incana
Alnus rugosa
Amelanchier oblongifolia
Benzoin aestivale
Clethra alnifoha
Cornus Amomum
Ilex verticillata
Lyonia ligustrina
Psedera quinquefolia
Rhododendron maximum}.
Rhus Toxicodendron
Rubus hispidus
Rubus triflorus
Sambucus canadensis
Vaccinium corymbosum
Viburnum dentatum
Viburnum Lentago
HERBACEOUS PLANTS
Arisaema triphyllum
Aspidium cristatum
Aspidium Thelypteris
Aster puniceus
Boehmerta cylindrica
Caltha palustris
Carex crinita
Carex stricta
Chrysosplenium americanum
Impatiens biflora
Lobelia cardinalis
Lycopodium lucidulum
- Maianthemum canadense
Onoclea sensibilis
Osmunda cinnamomea
Osmunda regalis
Polygonum arifolium
Solidago patula —
* Thuja occidentalis, one of the characteristic swamp trees farther north, occurs
in swamps a t Salisbury and Canaan.
yA prominent shrub in many a the Chomoscyberis swamps of southeastern
Connecticut; found also in a few swamps in Tolland and Litchfield Counties.
NICHOLS: THE VEGETATION OF CONNECTICUT 189
Cinna arundinacea Symplocarpus foetidus
Coptis trifolia Thalictrum polygamum
Galium Claytont Veratrum viride
Glyceria Torreyana Viola cucullata
Hydrocotyle americana Viola pallens
Just as was pointed out in connection with Carex stricta swamps,
the presence in wooded swamps of elevations, formed here by
stumps, fallen trunks, roots, etc., makes possible the existence of
many species which would be unable to grow in the oS wetter
situations.
The Réle of Liverworts and Mosses in Lowland Successions.—
A short digression may be made at this point to consider the part
played in the succession by liverworts and mosses. These are
sometimes present in great profusion and may contribute in some
measure to the accumulation of humus. In the subaquatic stages
of the succession they are often represented by submersed or
floating species, such as Ricciella fluitans, Ricciocarpus natans,
Fontinalis antipyretica var. gigantea, Fontinalis novae-angliae,
and various forms of Drepanociadus. Species of Sphagnum may
be present here as well as in the earlier swamp stages, but they are
often absent and seldom develop as luxuriantly as in the bog
succession which will be described presently. Common mosses
of open sedge or shrub swamps are Aulacomnium palustre, Bryum
ventricosum, Philonotis foniana, Elodium paludosum, Chrysohypnum
stellatum, Acrocladium cuspidatum, and Polytrichum commune,
while the liverworts Anthoceros laevis and Pellia epiphylla often
grow on the perpendicular sides of Carex stricta hummocks. In
wooded swamps the bryophytes are usually well represented, the
following species being perhaps as characteristic as any.
Growing in moist or wet depressions:
Pallavicinia Lyellii- Mnium punctatum
Chiloscyphus rivularis Mnium cinclidiotdes ;
_ Sphagnum sp. Brachythecium novae-angliae
— Bryum bimum | Calliergon yleo.coccmidll
Climacium americanum
Growing on stumps, logs, etc.: ; y oe
Cephalozia connivens == ~—SdDiicranum flagellare
~ Bazzania trilobata - Leucobryum glaucum
190 NicHOLS: THE VEGETATION OF CONNECTICUT
Trichocolea tomentella Fissidens adiantoides
Dicranum scoparium Thuidium delicatulum
Georgia pellucida
The Relation of Swamp Forests to Upland Forests.—In relation
to the various plant societies of the lake-swamp series, the swamp
forest may be regarded as at least a temporary climax, although it
may not always represent the ultimate condition. So long as the
ground remains saturated with water throughout a great portion
of the year, it is manifestly difficult for a mesophytic flora to de-
velop. Yet very frequently seeds of white pine (Pinus Strobus)
and other trees from an adjoining upland germinate in a swamp
during periods of protracted drought, and once firmly established
these are sometimes able to persist in spite of unfavorable soil
conditions. Moreover, many upland mesophytes, like Tsuga
canadensis, Hamamelis virginiana, and Kalmia latifolia, are
commonly present in such swamps, growing on elevations of
various kinds. Through the extension of these elevations and the
formation of new ones, it is apparent that the general level of the
ground may become raised above the zone of saturation, so that
mesophytes will become more numerous and the swamp plants cor-
respondingly restricted. In this fashion it is perhaps conceivable
that a swamp forest might ultimately be replaced by a forest es-
sentially similar to that which marks the climax of succession on
uplands. Theoretically, at least, such a forest represents the
culmination of all hydrarch, as well as xerarch, successions in this
region. But, parenthetically, it must be conceded that while
upland mesophytes undoubtedly tend to become more and more
abundant in a lowland forest, and while the tendency for such a
forest to approach the upland type is unmistakable, nevertheless,
so rapid is decomposition under subaerial conditions, that it is
doubtful whether the substratum is ever raised sufficiently to
produce a truly mesophytic habitat through the operation of
biotic factors alone.
Variations in the Rapidity of Succession in Lakes.—In the
foregoing paragraphs it has been shown that there is a tendency for
lakes and ponds to become filled in and converted into swamps
through the activity of vegetation; but there are great differences
in the rate at which this transformation is brought about. Thus
NICHOLS: THE VEGETATION OF CONNECTICUT 191
while many ponds have become completely clogged up through
the accumulation of muck or peat, there are others in which scarcely
any such accumulation would appear to have taken place. And
between these two extremes are ponds which exhibit various
intermediate degrees of filling. Although the reason for this
discrepancy between different ponds is not clear at the present
time, it may be due merely to the chance failure of certain plants to
become established in the “‘ barren’’ lakes as soon as in the “‘fertile’”’
ones. Or it is quite conceivable that a paucity of organic debris
may be due to the existence of environmental conditions unfavor-
able to the luxuriant development of aquatic plants. Certain it
is that in ponds with a sandy, gravelly, or rocky bottom, i. e.,
in ponds where muck or peat has failed to accumulate, the
vegetation invariably is sparser than in ponds with a mucky
bottom. Particularly is there a dearth here of those species most
active in peat formation, such as the larger pondweeds, the water-
lilies, the pickerel-weed, and the cat-tail. The vegetation along
the sandy or gravelly shores of such a pond is quite different
from that along mucky shores. Representative species are Erio-
caulon septangulare, Najas flextlis, and Lobelia Dortmanna—growing
in shallow water; and Panicum agrostoides, Cyperus dentatus,
Cyperus strigosus, Eleocharis acicularis, Juncus brevicaudatus,
Hypericum mutilum, Lysimachia terrestris, Gratiola aurea, and
Gnaphalium uliginosum—growing on the shore. The factors
directly or indirectly responsible for the assumed adverse envi-
ronmental conditions referred to above may possibly be many.
Among those that may enter in are the depth of the water body,
the temperature and clearness of the water, the steepness of the
shores, the amount of inorganic detritus washed in by streams or
otherwise, and the source of watersupply. That the degree of pro-
tection from or exposure to wind, and the nature of the water
currents may also be important factors is suggested by the fact
that mucky and sandy shores often occur around the margin of
the same lake, the former in the more sheltered, the latter in the
more exposed situations. Failure of peat to accumulate might also
of course result from any conditions which would promote rapid
and complete decomposition; or locally it might be effected through
the removal of plant remains by water currents.
192 NicHoLs: THE VEGETATION OF CONNECTICUT
Succession in Spring Swamps.—Many swamps have arisen
through the filling in of lakes and ponds in the manner described.
Buta large proportion of Connecticut swamps, as already pointed out
(pp. 173-175), owe their origin to the relation between topography
and ground water level. The former type of swamp has been
referred toasalakeswamp. In contrast to this, the latter type of
swamp, by reason of the fact that its existence depends on the
presence of seepage or spring water, may be called a seepage or
SPRING Swamp. Ina spring swamp there of course are never any
truly aquatic stages in the succession. Terrestrial plants are
present from the very outset. In a PERMANENT SPRING SWAMP
the pioneer association is commonly dominated by sedges, rushes,
and grasses, and in general its composition conforms closely to
that described for the sedge stage of the lake-swamp succession.
But in a PERIODIC SPRING SWAMP the pioneer association may be
quite different, both floristically and in its general aspect, from
that of a permanent swamp. To be sure, many of the plants
characteristic of permanent swamps are represented here also,
but on the whole the vegetation is much more mesophytic. Per-
haps as good an illustration of a periodic swamp as can be selected
is afforded by a Low MrEapow. Such an area may be very wet in
winter and spring, but later in the season the surface layers of
the soil usually become relatively dry. As in permanent swamps,
the predominant vegetation here is grass-like, but many other
herbaceous plants are also conspicuous. The list of species given
below may be considered as fairly representative of meadows.
Agrostis alba — Gentiana crinita
Anthoxanthum odoratum Glyceria nervata
Calamagrostis canadensis
Carex lurida
Carex scoparia
Carex stellulata
Carex stipata
Carex stricta
Castilleja coccinea
Cyperus strigosus
Eupatorium perfoliatum
Eupatorium purpureum
Juncus effusus
Poa pratensis
Rhexia virginica
Rynchospora glomerata
Sanguisorba canadensis
Selaginella apus
Thalictrum polygamum
Verbena hastata
Vernonia noveboracensis
Veronica virginica
NICHOLS: THE VEGETATION OF CONNECTICUT 193
Two types of meadows might be distinguished, viz., wet meadows
and dry meadows. But the two are not sharply delimited from
one another, and it is hardly worth while here to attempt to treat
them separately. Owing to their suitability for the growth of
grasses, meadows, especially the drier ones, are commonly culti-
vated and cut over annually for hay, and much of their natural
vegetation is thereby eliminated.
The shrub stage in a permanent spring swamp closely approxi-
mates the corresponding stage in a lake swamp. The tree stage
likewise is quite similar, although some of the trees to be mentioned
presently as characteristic of periodic swamps frequently occur
here also. In periodic spring swamps the shrub stage in the
succession scarcely differs from the shrub stage in permanent spring
swamps, except for the scarcity or absence of markedly hydro-
phytic species like Cephalanthus, the greater abundance of the
spiraeas and some others of the less hydrophytic shrubs, and the
frequent occurrence of mesophytic shrubs. In northwestern
Connecticut Potentilla fruticosa is very characteristic of such areas.
But the tree stage in periodic swamps exhibits noteworthy differ-
ences as compared with the tree stage in permanentswamps. For,
while any of the trees listed as characteristic of permanent swamps
may be found here also, and while here as there red maple and elm
may be the predominant species, the forests of periodic swamps
are characterized by the presence of a number of trees which
require an abundant water supply, but which do not thrive well in
a permanently saturated soil. Among these may be mentioned
Tsuga canadensis, Carya cordiformis, Carpinus caroliniana, Quercus
bicolor, Quercus palustris, Liriodendron Tulipifera, Nyssa sylvatica,
Fraxinus pennsylvanica and Platanus occidentalis. The herbaceous
vegetation of such swamps includes species which are seldom found
in permanent swamps, such as Erythronium americanum, Dentaria
diphylla, and Panax trifolium.
As concerns the building up of the substratum to a higher level
through the accumulation of plant remains, a state of equilibrium
somewhat analogous to that observed in many lakes is very com-
monly encountered in clayey or sandy swamps where, for various
reasons, dead organic matter oxidizes so rapidly that humus fails to
collect. Particularly is this true of periodic swamps. Ina colder,
194 NICHOLS: THE VEGETATION OF CONNECTICUT
more humid climate, peat might develop on such sites; but in Con-
necticut the mineral soil is frequently covered by little more than
the thin mantle of living vegetation and there is little humus.
Humus, as is well known, possesses a very high capacity for
retaining water, so that a soil rich in humus dries out very slowly.
A soil poor in humus may be equally wet in spring, but with the
lowering of the water table it dries out much more quickly. Such
a swamp, in consequence of the better drainage and aeration of the
soil, must necessarily afford a habitat quite different from that in
a swamp underlain by a mucky soil. It seems probable, there-
fore, that the abundance or scarcity of humus may be an important
factor in restricting the distribution of the trees mentioned above
as characteristic of periodic swamps.
Intermitient Ponds.—From an ecological standpoint these
should be classed as swamps rather than ponds, since the seasons
when their basins contain water coincide for the most part with
the period of vegetative inactivity. During the growing season,
except for a short time in spring and early summer, the bottom is
merely muddy, and very often in late summer it is quite dry.
With the exception of species having more or less amphibious pro-
clivities, like Proserpinaca palustris, Ludvigia palustris, Alisma
Plantago-aquatica, Sium cicutaefolium, and occasionally Nymphaea
advena, aquatic plants are absent. Those parts of the bottom which
remain longest submerged may be almost destitute of vascular
plants and frequently support a rich growth of algae, like Vauche-
ria and Botrydium, or of bryophytes, like Riccia, Fossombronia, and
Ephemerum. On parts of the bottom which are exposed during a
somewhat longer period commonly grow Eleocharis acicularis,
Hypericum virginicum, Lysimachia terrestris, etc.; while very often
there isa rank growth of various species of Bidens and Polygonum,
together with other rapid-growing, fall-flowering annuals. Fringing
the margin of the “pond” may occur almost any of the herbs,
shrubs, or trees characteristic of swamps. It should be noted, per-
haps, that Cephalanthus is an almost omnipresent marginal shrub.
PLANTS IN RELATION TO THE FORMATION OF MARL
There is one other important phase of plant activity in con-
nection with the filling in of lakes and ponds to which no ref-
NICHOLS: THE VEGETATION OF CONNECTICUT 195
erence has yet been made, namely, the formation of marl. Marl
deposits are always associated with calcareous regions, and in
Connecticut are confined to the limestone areas in the western
part of the state. At Twin Lakes, Salisbury, there are quite exten-
sive deposits, estimated in places to exceed a hundred feet in depth,
and there are similar deposits near Danbury and elsewhere. Chem-
ically, pure marl contains about ninety-five per cent. of calcium
carbonate. Formerly such deposits were supposed either to rep-
resent shell remains, or else to have originated through mechanical
sedimentation or chemical precipitation from calcium-containing
waters.* But while the presence in marl beds of Chara and other
plant ‘‘fossils’” had frequently caused comment, and although it
is a common observation that in lakes where marl is being de-
posited, submersed aquatic plants are usually coated with a thin
crust of calcium carbonate, it remained for Davis} to demonstrate
the important réle played by plants, particularly by the alga
Chara, in marl formation.
In a general way it may be asserted that plants are
able to bring about the formation of marl by causing the
precipitation of various calcium salts which may occur dis-
solved in the waters of lakes and ponds. Two methods by which
plants may effect this result are easily conceived. (1) The ab-
straction of carbon dioxide from water in which calcium is present
in excess, held in solution by free carbon dioxide, would cause the
precipitation of the calcium salts. (2) The combination of the
oxygen liberated by plants with easily soluble salts like calcium
bicarbonate, thereby converting them into less soluble salts like
calcium carbonate, would result in precipitation. The existence
of still a third, and possibly even more effective, method of calcium
concentration has been pretty conclusively demonstrated by Davis
in the case of Chara. (3) The calcium accumulates in the cells of
this plant in the form of calcium succinate, one of the few water-
soluble salts of calcium. Later on — oat how is not as yet
wholly understood, the dissolved salts are excr: nd deposited in
* For further discussion, see Hale, D. J., Theories of origin of bog lime or marl.
Geol, Surv. Michigan 8: 41-64. 1903.
' +A contribution to the natural history of marl. Geol. Surv. Michigan 83: 65-
96. 1903. Also earlier papers in Journal of Geology.
196 NICHOLS: THE VEGETATION OF CONNECTICUT
an insoluble form on the surfaces of the cells. At Twin Lakes this
alga is extremely abundant, more so than in any other lake or pond
in the state, so far as the writer’s observations have extended.
Here, not only Chara, but the leaves and stems of Najas, Elodea,
and various species of Potamogeton are incrusted by a thin, white,
flaky deposit of marl. Some of the blue green algae, notably
Zonotrichia, are likewise instrumental in causing marl formation.*
Because of the inability of most of the plants active in its pre-
cipitation to survive in competition with shallow water species,
the production of marl is largely restricted to the deeper parts of
lakes.
THE FLOATING Mart
Occurrence and Importance of the Floating Mat—lIn an earlier
paragraph the writer has alluded to the manner in which the shal-
low-water zones of vegetation in a lake tend to push out from the
shore and to encroach upon the deeper areas. In most lakes and
a
Sos
Why thy diy tf
LL WNT Y) /
TILA fd hy fit phy fe
VP AYL A PLL YAY Vf Tha
CA PETETITIA RSS EES 6s iM
Yh SALT SG Lt GE) ates
LLL DTT LLLL TEAL LLL ALITA AYA LL, (eee
Fic. 10. Diagrammatic vertical section through the marginal portion of a
lake which is being filled in through the intervention of a floating mat, showing the
usual succession of plant societies. Sequence of Zones: 1. Pondweed Zone; 2-
Water-lily Zone; 3. Decodon Zone; 4. Cassandra—Sphagnum Zone; 5. Cassandra—
Sphagnum—Sedge—Low Spruce Zone; 6. Tall Shrub Zone; 7. Bog Forest. F =
Marginal Ditch or ‘‘ Fosse”. er ee
* For detailed discussion of the relation of Chara and Zonotrichia to marl forma-
tion, see Davis, 1903, op. cit., especially pp. 87-02. bia a
NICHOLS: THE VEGETATION OF CONNECTICUT 197
ponds where rapid peat accumulation is taking place the filling
in of the deeper waters is brought about in this manner. Very
often the growth of certain marginal swamp plants is so vigorous
as to give rise to a raft-like zone of vegetation which spreads out
from the shore over the surface of the water, and which is com-
monly referred to as a FLOATING Mat (Fic. 10). There will be
described next, then, the manner in which this mat is developed,
together with the succession of plant associations which accom-
panies the process. And since in Connecticut the floating mat
succession is commonly associated with bogs, a type of swamp not
yet treated, the two will be considered together, i. e., the suc-
cession is assumed to culminate in the formation of a bog.
Mai Formation due to Sedges.—The lake stages in such a suc-
cession are identical with those already described. There may
thus appear, in order, submersed aquatics, aquatics with floating
leaves, and aquatics with aerial foliage; but, due to the encroach-
ment of the mat, these stages may be quite abbreviated, and one
or all may be eliminated. It is in the transition from lake to
swamp that the floating mat comes into play. According to the
conception which has been widely circulated, especially in geolog-
ical literature previous to 1907, this mat is formed in the following
manner. ‘‘Certain mosses, particularly those of the genus Sphag-
num, have a habit of growing out on the water surface and forming
a mat of intertwined stems connected with the shore. At this
stage the lake is an open water body with a border of vegetation
floating near the rim. Gradually this rim of moss creeps toward
the center of the pond until it is completely closed in and covered
over with a layer of vegetation. The lake is now a swamp; and
such a swamp, with a floating layer of aquatic plants, is known as
a ‘quaking bog’; it is possible in some cases to walk across an old
lake on a mat of vegetation while the water remains below. These
mosses have a habit of growing at the top while the old stems are
dying below, and the rotted fragments drop to the bottom of the
pond and help to fill it up.’’*
_ Except for the stress laid on the rdle of Sphagnum and other
mosses, the description above quoted portrays the origin and
behavior of the floating mat fairly accurately; and for cold,
* Rice, W. N., and Gregory, H. E., op. cit., p. 249.
198 NICHOLS: THE VEGETATION OF CONNECTICUT
humid regions like parts of eastern Canada the importance of
the mosses would hardly be overestimated. But, as_indi-
cated by Transeau* and conclusively proved by Davis,f in
milder, less humid regions the mosses play at best a very subsid-
iary part in mat formation. In Michigan, according to these
authorities, the most important mat-forming plants are the sedges,
particularly Carex filiformis. This species is capable of spreading
rapidly by means of rhizomes. As pointed out by Davis, these
subterranean or subaquatic stems will often grow horizontally a
foot or more in length during a single season, producing at the
nodes an abundance of tough, slender roots, and bearing at their
tips terminal buds “from which new plants rise to send out in
turn a new series of horizontal stems. When conditions are un-
Fic. 11. Floating sedge mat at Twin Lakes. The plants along the margin
of the mat are bulrushes.
favorable for the rhizomes to grow outward into open water, they
sometimes grow diagonally downward over the edge of the mat,
and thus the mat is strengthened as well as extended by the growth
of the plant.’’ On account of its low specific gravity, the tangle
of roots and rhizomes thus produced floats on the surface of the
* Bot. Gaz. 40: 363. 1906.
+ 1907, loc. cit., pp. 135-138.
NICHOLS: THE VEGETATION OF CONNECTICUT 199
water like a raft. Along the western shore of one of the Twin
Lakes is a beautiful example of a floating mat which thus owes its
origin to Carex filiformis (Fic. 11); and similar mats are frequently
encountered in other sections of the state. Toward its lakeward
margin such a mat is usually thin, but farther away from the edge
it reaches a thickness of two feet or more and becomes firm enough
to support considerable weight. For some distance from its outer
margin, in cases where it is typically developed, the mat is actually
floating and is underlain by clear water. But in proceeding from
the margin shoreward the water underneath is ‘‘found to become
more and more full of finely divided matter until, at a variable
distance from the water’s edge, the deposit is nearly solid and the
mat no longer floats”’.* The filling up of the open water under-
neath, and the ultimate grounding of the mat is brought about
very largely in the manner already suggested, namely, by the
dropping down of debris from the lower surface of the raft. Often,
owing to local conditions, the mat grows outward from the shore
so slowly that the water beneath the floating substratum becomes
filled out to its very margin. By the time the mat has grounded,
if not before, shrubs have asserted themselves as the controlling
element of surface vegetation; and these in turn, as in the lake-
swamp succession already described, ultimately give way to trees.
Mat Formation due to Decodon.—But, on the whole, in Connecti-
cut the sedges are overshadowed as pioneer mat formers by certain
shrubs, notably Decodon verticillatus. The manner in which
this latter shrub not only brings about the formation of a floating
mat but accelerates the upbuilding of the surface as well, has been
admirably described by Davis.t Growing at the water’s edge it
sends up slender, arching stems, many of which reach out over the
open lake, curving gracefully downward toward their tips until
they enter the water (Fic. 12). Here the stem becomes greatly
swollen, due to the formation of aerenchyma, and develops numer-
ous adventitious roots. The aerial part of the plant is herbaceous
and dies at the end of the growing season. But the swollen,
submerged portion of the stem becomes woody and perennial,
lives through the winter, moored in place, as it were, by the other-
* Davis, 1907, loc. cit., p. 138
7 1907, loc. cit., pp. I51, 152, 204—206.
200 NICHOLS: THE VEGETATION OF CONNECTICUT
wise functionless, dead aerial part of the stem, and the following
season sends up a fresh series of aerial stems. Some of these grow
outward over the open water, thus extending the margin of the
mat lakeward; others reach shoreward, thereby more effectually
uniting the newer with the older portions of the mat; while yet
others grow out in lateral directions, in this way binding together
Fic. 12. Decodon verticillat i ing int ter al in of a pond,
x S t—3
Southington. View taken in winter.
more firmly the integral parts of the still rudimentary structure.
At this stage the mat is little more than a framework, consisting
of a loose aggregation of the Decodon plants. But upon the stools
formed by the Decodon, and about its roots, other plants soon
appear, and as these increase in number, reaching across from one
stool to another, the skeleton gradually becomes sresmalormed into
a continuous, compact body. :
Importance of Other Plants in Mat Formation—Often one of
the earliest plants to appear among the Decodon stools is the
Sphagnum. But while this moss may act as a space filler, it adds
little to the strength of the mat; for its stems, weak and nearly
NICHOLS: THE VEGETATION OF CONNECTICUT 201
devoid of mechanical tissue, could hardly be expected to possess
any great degree of tensile strength. Moreover, as a rule, Sphag-
num does not develop in any abundance until a later stage in the
succession. Various species of Drepanocladus may also grow in the
water about the Decodon roots, but their rdle in mat formation is in-
consequential. As in the case of the sedge mat earlier described,
it is the vascular plants with their tough stems and roots, partic-
ularly species like the cranberry (Vaccinium macrocarpon) and
cassandra (Chamaedaphne calyculata), which are primarily re-
sponsible for the increase in the continuity and compactness of
the substratum. Decodon is primarily a pioneer, and in competi-
tion with other plants is speedily exterminated. When present,
it usually occupies a zone from six to a dozen feet wide along the
water front, but, except for scattered specimens, is absent else-
where. Inside the Decodon zone there is commonly a zone dom-
inated by cassandra, along with which, in increasing abundance,
grows the Sphagnum.
It might naturally be anticipated that where the pioneer
plants are shrubs, the sedges would be eliminated from the
succession, but this is rarely the case. Almost always sedges
and other herbaceous bog plants occur in among the shrubs,
and quite frequently the cassandra zone is bordered on the land-
ward side by a more or less continuous zone in which the domi-
nant plants are Sphagnum, cranberries, and such sedges as -
Rynchospora alba and Eriophorum virginicum. The exact ex-
planation of this last mentioned phase in the succession is puzzling.
It seems to represent a RETROGRESSIVE CONDITION due to some
disturbance which has caused the disappearance of the cassandra
and has thus created an opening for a more primitive association to
become established. Such a result might be effected either by a
sinking of the mat, or by a temporary elevation of the water level
which might flood the grounded portions of the mat, and, if of
sufficient duration, might cause the death of the cassandra. Pro-
ceeding farther landward from this zone, shrubs once more assume
control, and under normal conditions, as in other cases, trees
ultimately become the predominating element of the vegetation.
There are other ways in which the formation of a floating mat
may be inaugurated. At Bingham Pond, Salisbury (Fic. 14), and
202 NICHOLS: THE VEGETATION OF CONNECTICUT
elsewhere,(Fic. 15) the plant which fringes the water’s edge and forms
theskeleton for the advancing mat is thecassandra. The extension
outward into the open water is slow when dependent on this
plant, and the mat is usually thick and firm out to its very edge.
At Bailey Pond, Voluntown, is a remarkable submersed mat,
some ten feet wide, composed entirely of the rhizomes of the fern
Woodwardia virginica. Floating logs, sticks and similar debris
doubtless contribute their quota to the formation of the raft;
while sometimes great patches of rhizome-permeated muck break
loose from: the bottom of a pond, rise to its surface, and furnish a
substratum favorable for the rapid spread of sedges and other mat
forming plants.*
ECOLOGICAL RELATIONS, ORIGIN, AND DISTRIBUTION OF Bocs
Bogs Compared with Ordinary Swamps.—A Boc may be defined
as a fresh water swamp characterized, especially in the shrub stage,
by an abundance of xerophytic plants. It might well be desig-
nated a XEROPHYTIC Swamp. In Connecticut, so far as known,
bogs are always developed in morainal depressions. This type of
swamp, by reason of its unique vegetation, the ecological problems
involved, and the economic value of the frequently underlying peat
deposits, has probably received more attention at the hands of in-
vestigators than all other swamp types put together. In com-
paring the plant associations encountered in a lake-bog succession
with those in an ordinary lake-swamp series, the first appreciable
differences are perceived in the sedge stage, while the departure
of the two types from one another becomes very pronounced in
the shrub and tree stages. From a floristic standpoint, Connec-
ticut bog vegetation is unique because of the predominance of
ericaceous shrubs, the prevalence of black spruce (Picea mariana)
and other boreal species otherwise absent from this region, the
presence of bizarre forms like the pitcher plant (Sarracenia pur-
purea) and the sundews (Drosera rotundifolia and Drosera longi-
folia), and the usually luxuriant development of Sphagna. Eco-
logically, the most interesting problems relate to the phenomenon
of bog xerophytism.
* Such masses often rise to the surface during the summer, only to sink to the
bottom again in the fall. In this connection see S. Powers’s paper on “Floating
Islands”’, Bull. Geog. Soc. Philadelphia 12: 1-26. f. 1-9. 1914.
NICHOLS: THE VEGETATION OF CONNECTICUT 203
Succession of Plant Societies in Bogs.—As suggested in earlier
paragraphs, succession in bogs, in Connecticut at any rate, is
usually associated with floating mat formation, and several of .
the phenomena of a bog succession have already been discussed
under that head. Where a bog still borders an- open body
of water the various plant associations mentioned in connection
with the floating mat succession are often clearly distinguishable.
But quite as often, especially where the bog occupies a basin that
has become completely filled in (Fic. 13), the three associations are
Fic. 13. Spruce bog, Willington. Picea mariana and Chamaedaphne calyculata
the dominant forms; small, scattered specimens of Pinus rigida in the foreground.
more or less completely merged into one, i. e., they are telescoped.
The shrubs here, along with Sphagnum, tend to form elevated
patches; sedges, with Sphagnum, dominate on the intervening lower
ground; while the spruces—seldom over fifteen feet high, mostly
under eight, and commonly less than four—for the most part grow
scattered about among the shrubs, here and there developing
colonies of sufficient size and density to shade out the more in-
204 NICHOLS: THE VEGETATION OF CONNECTICUT
tolerant undergrowth. In a bog of this sort the more depressed,
wetter spots may be occupied largely by various bryophytes, e. g.,
Cephalozia fluitans, Mylia anomala, Scapania trrigua, Aulacom-
nium palustre, Acrocladium cuspidatum, Drepanocladus fluitans,
and certain species of Sphagnum. The following herbaceous seed
plants also commonly occur in like situations.
Arethusa bulbosa Hypericum virginicum
Calla palustris Menyanthes trifohata
Drosera longifolia Rynchospora alba
Drosera rotundifolia Scheuchzeria palustris
Dulichium arundinaceum Smilacina trifolia
Eleocharis palustris Utricularia cornuta
Xyris caroliniana
The vegetation of these depressions may represent a relict of a
primitive stage in the succession, but it seems more likely that it
represents a retrogressive phase like the one already described.
Characteristic also of open bogs, but mostly growing in less wet
places, are the following herbaceous species.
Asclepias incarnata Epilobium sp.
Aspidium Thelypteris Eriophorum sp.
Calamagrostis canadensis Habenaria blephariglotiis
Carex rostrata Lysimachia terrestris
Carex trisperma Pogonia ophioglossoides
var. Billingsit Woodwardia virginica
From these lists are omitted purposely a number of species, like
Carex pauciflora and Carex paupercula, which are peculiar to bogs
but which have been recorded from only one or two localities in
the state.
The shrubs of open bogs may conveniently be divided into two
groups, somewhat as follows. :
First Group
Andromeda glaucophylla Gaylussacia baccata
Chamaedaphne calyculata Kalmia angustifolia
Chiogenes hispidula Kalmia polifolia
Gaultheria procumbens Ledum groenlandicum
Gaylussacia dumosa Vaccinium macrocarpon
var. Bigeloviana* Vaccinium Oxycoccus
* This northern variety of Gaylussacia dumosa was described by Fernald in
Rhodora 13: 95-99. I9QII.
NICHOLS: THE VEGETATION OF CONNECTICUT 205
SECOND GROUP
Alnus rugosa (or A. incana) Pyrus melanocarpa
Amelanchier oblongifolia Rhododendron viscosum
Ilex verticillata Rhus vernix
Myrica carolinensis Vaccinium corymbosum
Nemopanthus mucronata Viburnum cassinoides
Most of theshrubs peculiar to bogs areincluded in thefirst group.
It will be noted that all of the species there listed are ericaceous,
and that practically all of them are distinctly northern in their
range. The shrubs in the second group are not so strictly confined
to bogs, but, with one or two exceptions, are frequent or common in
swamps of the ordinary type. The majority of them are equally
well developed far to the south. It should be added that the
species in the first group are comparatively low growing—mostly
less than three feet high when mature—while those in the second
group average well above three feet. In studying their distribu-
tion in an open bog, it is noticeable that the lower shrubs compris-
ing the first group are pretty uniformly distributed throughout,
while the taller shrubs which comprise the second group are best
developed toward the landward margin, or in slightly elevated
situations. In the case of a bog bordering an open pond, as at
Bingham Pond (Fic. 14), there are usually four rather definite
bands of woody vegetation, paralleling the-water’s edge: first, a
zone of low shrubs; then, a zone of low shrubs intermixed with
young spruces and tamaracks;* next, a zone of taller shrubs along
with older trees; and finally, farthest removed from the pond, a
zone of large spruces together with some of the more tolerant of
the taller shrubs. Undoubtedly some correlation exists between
the position usually occupied by shrubs of the second group in bogs
and the fact that most of them quite commonly occur in swamps of
the ordinary type, a correlation which almost certainly relates to
soil aeration and drainage. There very likely is also some con-
nection between this local distribution of these plants in bogs and
swamps and their geographic distribution.
Sphagnum and its Relations.—Occasionally, as for example at
Twin Lakes, a bog is encountered from which apparently Sphagnum
is entirely missing. In the particular case cited the bog is under-
* Ret 4h Aé+h as, £. + Snémdl éive <0
¥ £
cranberry—sedge zone already mentioned.
206 NICHOLS: THE VEGETATION OF CONNECTICUT
lain by marl, and since Sphagnum is reputedly calciphobous, it
was at first thought that here, at least, its absence could be easily
accounted for. But in a nearby swamp which likewise overlies a
calcareous subsoil Sphagnum is abundant. Similar observations
by Davis* in Michigan, together with the experimental evidence
advanced by Transeau and Weber} would make it appear highly
improbable that the presence or absence of Sphagnum can be cor-
related with the presence or absence of lime. As a rule, in the
floating mat succession, Sphagnum seldom precedes the shrubs
and sedges, except to an inconsequential degree. There are a
few ponds in southeastern Connecticut where Sphagnum macro-
phyllum, a remarkable southern coastal-plain species with leaves
fully half an inch long, constitutes an important part of the free-
floating vegetation. But this condition is exceptional. Ina bog
the Sphagnum usually appears first as a superficial layer growing
on the surface of the shrub or sedge mat; but once established it
often develops so rapidly and luxuriantly as to exert a profound
influence on the character of the bog vegetation.{ Growing
upward in dense masses around and among the stems of various
shrubs, etc., this plant commonly forms cushions which sometimes
rise as much as two feet above the original water level. Ante-
cedent plants which are unable to accommodate themselves to this
change in environmental conditions are gradually eliminated.
Broadly speaking, the ability of a plant to exist in a bed of
rapidly growing Sphagnum may be said to depend onits ability to
keep pace with the upward growth of the moss. The necessity of
keeping the foliage above the surface of the substratum is too
obvious to require more than passing mention. But it seems very
likely that it is equally important that the roots of the plant be
kept above the zone of permanent saturation. For as the
Sphagnum plants grow upward, dying away below, the lower part
of the cushions becomes more and more compact and, due largely
to the well-known capacity of Sphagnum stems and leaves for
sucking in water, acting like a sponge, it becomes completely
saturated. As the Sphagnum cushions increase in height, then,
* 1907, loc. cit., p. 276.
t+ See Transeau, 1906, loc. cit., at 32.
¢ In this connection see also W. S. Cooper’s discussion of the behavior of Sphag-
num in Isle Royale bogs. Bot. Gaz. 55: 200-206. I912.
NICHOLS: THE VEGETATION OF CONNECTICUT 207
the water level rises correspondingly, with the result that the roots
of plants originally growing on the mat’s surface are submerged.
The characteristic plants of SPHAGNUM Boas are those which are
able in some way to develop root systems above this zone of
saturation. Species like Aspidium Thelypteris and many sedges
accomplish this result through the upward growth of their rhi-
zomes. Sprawling vines like Chiogenes and Vaccinium macrocar-
pon similarly have little difficulty in establishing fresh footholds
from time to time. The pitcher plant possesses little power of
stem elongation and individual plants are buried quite rapidly,
but like some other bog plants it reproduces itself with sufficient
rapidity by means of seeds to enable it to hold its own. Among
the shrubs the most successful species are those which freely
produce adventitious roots from the old upright stems above the
zone of permanent saturation. Among others, Chamaedaphne,
Kalmia polifolia, and Ledum are noteworthy in this respect. In
a specimen of Ledum, for example, which the writer collected in a
Sphagnum bogat Norfolk, theleafy shoot rose buteight inches above
the surface, the remaining partof the stem, more than three feet long,
being imbedded in the moss. Adventitious roots had developed
throughout the length of the stem, those on the first foot of stem
below the surface being vigorous and obviously the only ones
functional at the time of collection. It is interesting to note in
this connection that Cowles,* in his study of the vegetation on the
sand dunes of Lake Michigan, found that the ability of many
woody plants to withstand burial by sand likewise depends on their
power of stem elongation plus the development of adventitious
roots. The upward growth of a Sphagnum cushion proceeds
_ de aged at first; eventually it attains a height where the
tion and decreasing water supply
is such that further upward igcowitls 3 is no longer possible. When
this stage is reached species of the lichen Cladonia and relatively
xerophytic mosses like Polytrichum commune and Hypnum
Schreberi frequently establish themselves on top of the Sphagnum
cushions.
The Significance of the Marginal DitchSeparating the bog
from the adjoining upland, more often than not, there is a broad,
-* Bot. Gaz. 27: 384. 1890.
208 NICHOLS: THE VEGETATION OF CONNECTICUT
open ditch, or fosse, which at certain seasons of the year may
be filled with water to a depth of one or two feet. This ditch is
a widespread feature of bogs and its significance is not yet wholly
clear. But of the various explanations offered, that suggested by
Davis* seems to the writer most plausible. Davis attributes the
formation of the fosse to ‘‘ the fluctuation of the water level through
rather brief intervals and the constant recurrence of these fluctu-
ations. These are attendant upon the variations in the rainfall,
and the water level in the lakes and depressions may vary one, two,
or more feet every few years, and may remain at the low water
stage for several years in succession. . . . It is also a matter of
observation that, during dry times, the water does disappear from
these marginal ditches for long periods during the summer and
autumn, the bottoms becoming quite dry, and this has the effect of
destroying much of the hydrophytic vegetation which has estab-
lished itself and also of thoroughly decomposing and disintegrating
the organic matter which has accumulated during periods of high
water.’’ In this way the surface is lowered ‘‘ below that of the area
directly above the zone of permanent water, which, being covered
by a thicker layer of vegetable debris, is kept wet by the upward
capillary movement of the water from below its surface,”
The Tree Stage in Bogs.—The most distinctive feature of the
tree stage in Connecticut bogs, considered collectively, is the
presence of the black spruce. Commonly associated with this,
and sometimes forming considerable forests in the older, better
drained portions of the bog, grows the tamarack. The tamarack
is not so tolerant of shade as the black spruce, but its more rapid
and greater height growth enable it to maintain its own in the
competition for light. More or less extensive tamarack forests
formerly occurred in bogs at New Fairfield, Suffield, and else-
where. Due to the rather open character of such forests, and the
slight shade produced by the tamarack, the spruce, together with
much of the undergrowth present in the adjoining parts of the open
bog persists here. The black spruce itself seldom forms a stand of
sufficient height and density to merit the title of forest; but in the
cool highlands of northwestern Connecticut, about Bingham Pond
* 1007, loc. cit., p. I51. For references to other explanations see Cooper, loc. cit.,
p. 203.
NICHOLS: THE VEGETATION OF CONNECTICUT 209
(Fic. 14), there is a magnificent forest of spruce. Here, however,
the black spruce of the opener parts of the bog seems to have been
largely superseded by the red spruce (Picea rubra), a species which
attains a much larger size, but which is absent from most sections
of the state.* Along with the spruce in this Bingham Pond forest
Fic. 14. Spruce bog surrounding Bingham Pond, Salisbury. Behind the
marginal fringe of Chamaedaphne, etc., is seen the bog forest, largely composed of
Picea rubra. Water of pond somewhat lowered through artificial drainage.
grow the mountain ash (Pyrus americana), a species sometimes
found in bogs in other sections of the state, and commonly in
bogs farther north. There are also scattered yellow birches, but
the spruce predominates, casting a shade so dense as to effectually
exclude nearly all of the herbs and shrubs characteristic of the
open bog. The ground beneath in many places is carpeted with
various mosses and liverworts, e. g., Bazzania trilobata, Dicranum
scoparium, Dicranum undulatum, Leucobryum glaucum, Hyloco-
mium splendens, Ptilium Crista-castrensis, Stereodon tmponens,
* The correctness of the identification of these trees as Picea rubra is possibly
open to question. At the time this forest was studied, the writer assumed them to
be Picea mariana. Since then, however, study of these two species in the Canadian
woods has led to the conviction that they must be Picea rubra, an opinion which was
first expressed and has recently been reaffirmed by Dr. C. A. Davis. These trees
also grow on the mountain slopes in the vicinity of the pond.
210 NICHOLS: THE VEGETATION OF CONNECTICUT
Hypnum Schreberi, and Georgia pellucida, and there are occasional
patches of Sphagnum. Characteristic herbaceous plants here are
Aralia nudicaulis, Clintonia borealis, Coptis trifolia, Cypripedium
acaule, and Maianthemum canadense; these grow for the most
part in the least shaded spots. The commonest shrub is Kalmia
latifolia.
In eastern Connecticut, the coast white cedar largely supplants
the black spruce as the characteristic bog conifer, while throughout
the state bogs are frequently encountered from which conifers are
entirely absent. At Berlin, for example, there is a large bog»
perhaps sixteen acres in extent, whose surface, except for a peri-
pheral band of red maple and taller shrubs, is entirely covered with
heaths and Sphagnum. In Beaver Swamp, a bog within the New
Haven city limits, red maple is likewise the only arborescent species
present. The omnipresence of this latter tree in swamps of the
ordinary type has already been commented on. This observation
may be extended to include bogs; but here, except where the
original vegetation has been disturbed, it is usually conspicuous
only toward the landward margin.
The Problem of Bog Xerophytism.—In attempting to explain
the differences between bog vegetation and that of ordinary
swamps, two sets of factors must be taken into account, namely,
factors which are active at the present time, and factors which have
operated during the past. The immediate effect of the first set
is seen in the phenomenon of bog xerophytism; the effect of the’
second set may be reflected in the restriction to bogs of so many
boreal plants. That, despite their physical wetness, the soil
conditions in bogs are conducive to xerophytism has long been
recognized. This xerophytism is evidenced by the fact that
many of the commonest bog plants are likewise characteristic of
habitats which are physically dry. Thus, in Connecticut,
Gaylussacia baccata and Gaultheria procumbens grow abundantly in
dry woods and clearings; Kalmia angustifolia and Myrica caroli-
nensts flourish on dry, open hillsides; while Pyrus melanocarpa and
Vaccinium corymbosum are among the commonest shrubs on ex-
posed, rocky ledges along the coast. Moreover, many of the
more distinctive bog plants, in contrast with most plants of or-
dinary swamps, exhibit structural peculiarities which are char-
NICHOLS: THE VEGETATION OF CONNECTICUT Zit
acteristic of xerophytes; while analogous xerophytic modifications
may frequently be observed in the leaves of bog-grown individuals
of such a species as Alnus rugosa, which occurs in both types of
swamps. The xerophytic peculiarities of the leaves of bog ever-
greens, e. g., the cassandra, however, need not necessarily be
attributed to the nature of the habitat, but may rather, to a certain
extent at any rate, be associated with protection from excessive
transpiration during the winter.*
Many theories have been advanced with a view to explaining
the condition of physiological xerophytism prevalent in bogs. It
has been variously ascribed to the acidity of the soil, to the low
temperature of the soil, to the insufficient aeration of the soil, and
to the accumulation in the soil of root excretions or various toxic
substances.t| Doubtless there are still other factors whose effect
may be equally potent. For example, in view of the low diffusion
properties of humus soils, which to a great extent would exclude
salts derived from the subjacent mineral soil, together with the
only partial decomposition of vegetable remains in bogs, it is
conceivable that an actual scarcity in the bog substratum of the
requisite mineral elements in form suitable for absorption and
assimilation by most plants may be a factor of some significance.
Indirectly, the topographic relations of bogs are considered im-
portant. Bogs usually develop in undrained or poorly drained
depressions, and almost invariably in connection with a water
body of considerable depth. In Salisbury, Norfolk, Kent, Litch-
field, and elsewhere there are bog-bordered lakes, but in many
cases, the depressions occupied by bogs, although quite deep, are®
only a few acres in extent. In two small bogs at Southington, for
example, the writer was unable to touch bottom with a twenty-five
foot peat sampler. One of these bogs isshown in Fic.15. That the
lack of drainage in such basins would tend to promote soil acidity,
poor aeration, and the accumulation of deleterious substances
seems obvious, while their depth would favor low soil temperatures.
Regarding the relative importance of these different factors, how-
See Gates, F. C. Winter as a factor in the xerophily of certain evergreen
plants. Bot. Gaz. 57: 445-489. f. I-12. I914.
+ For references to the literature relating to these various theories, see Dach-
nowski, Bot. Gaz. 52: I-3. I9II.
212 NICHOLS: THE VEGETATION OF CONNECTICUT
ever, no definite statement can yet be made, although it seems
more than likely that the observed conditions may result from a
combination of several causes. Whatever the direct cause or
causes, the prevailing consensus of opinion among ecologists seems
to be that their ultimate effect on the plants concerned is such as
Fic. 15. Spruce bog surrounding a small pond, Southington. Marginal fringe
of Chamaedaphne, with small black spruces and tall shrubs in the rear. On the
@snow-covered hill in the background can be made out oaks, chestnut, and red cedar.
either to directly hinder absorption, or else to indirectly reduce it
by retarding root development.
The Origin of the Northern Flora in Bogs.—While soil conditions
satisfactorily account for the phenomenon of bog xerophytism,
it is doubtful whether the remarkable assemblage of northern
plants found in bogs is to be explained on this basis alone. In
considering this problem some years ago, Transeau* arrived at the
conclusion that the explanation must be sought in the physio-
graphic history of the areas concerned. According to his con-
* On th hic distributi d ecological relations of the bog plant societies
of North Anecion. "Bot. Gaz. 36: 401-420. f. 1-3. 1903; also 1906, loc. cit
NICHOLS: THE VEGETATION OF CONNECTICUT 213
ception: ‘‘Where the habitat dates back to Pleistocene times and
has remained undisturbed, we find today the bog flora. Where
the habitat is of recent origin or has been recently disturbed, we
find the swamp flora, or mixtures of swamp and bog species.”
The plausibility of some such explanation for the presence of
northern plants in bogs was re-impressed on the writer during the
past summer in the course of ecological investigations conducted
on Cape Breton Island. Therethe bog is the common swamp type,
and its vegetation seems wholly in harmony with the forests of
fir and spruce that clothe the surrounding uplands; the transition
from one to the other is gradual; bog vegetation there is much less
restricted than here, as is evidenced by the fact that species which
in Connecticut are confined to bogs commonly occur there in
shallow depressions along streams, while a much larger proportion
of bog species grow on the uplands. Yet notwithstanding the
marked contrast in the upland vegetation there and here, the
resemblance of bog vegetation in the two regions, not only in
general aspect but in specific composition, particularly with ref-
erence to sedges, ericaceous shrubs, and trees, is remarkable.
Here in Connecticut, however, in contrast to the conditions farther
north, the bog is a comparatively rare swamp type, and its vege-
tation seems utterly out of harmony with the deciduous forests of
the surrounding uplands; the transition from one to the other is
abrupt, while the characteristic northern bog plants for the most
part are restricted to bogs.
In the first paper of the present series* the writer suggested
the probability that subsequent upon the final retreat of the con-
tinental ice sheet, this region for a long time was populated by
northern types of vegetation which, having originally been forced
southward by the advance of the glaciers, once more migrated
northward in the wake of the retreating ice-front. During the
time that has elapsed since the recession of the ice, variously
estimated at from 20,000 to more than 50,000 years, there un-
doubtedly has been a gradual readjustment of the climate, and as
a result of this, coupled with the invasion of plants from the south
and west, our present vegetation has been evolved. In the his-
torical sequence of vegetative formations which have occupied
* Torreya 13: 93. I913.
214 NICHOLS: THE VEGETATION OF CONNECTICUT
the region under discussion during this period, there doubtless
have been represented many of the types of vegetation which at
the present time are characteristic of regions farther north. Thus
the first vegetation to seize upon the freshly exposed land areas
may have been quite similar to the tundra formation which today
is restricted to the far north. But the dominance of the tundra
must have been relatively ephemeral, for except as it may possibly
be represented by isolated species of plants, all traces of it have
vanished. Following the tundra there presumably was gradually
developed a type of vegetation essentially similar to that which
today prevails in northern Maine and in the eastern maritime
provinces of Canada. Forests of fir and spruce dominated the
landscape. At this time, associated perhaps with a moister, cooler
climate, the relationship between lowland and upland vegetation
must have been quite similar to what it now is farther north. The
bog was a common swamp type, and bog plants were by no means
confined to bogs. Just how long this condition prevailed is of
- course largely a matter of speculation. It is perhaps significant,
however, that red spruce and fir still persist as upland species in
certain localities in northwestern Connecticut, while the southern
boundaries of the great fir and spruce forests of Maine are scarcely
two hundred miles north of Long Island Sound. From such facts
as this it seems probable that within very recent geological time
spruce and fir comprised an important element in the forests of
this region. When in their northward march the deciduous trees,
which at present pred te the forests of the state, commenced to
invade this region, they doubtless became established first in the
more favorable sites, so that there thus arose a mixture of conifer-
ous and deciduous types of forest. Such conditions actually
exist in the north-woods today. In northern Cape Breton Island,
for example, forests of beech (Fagus grandifolia), sugar maple
(Acer saccharum), and yellow birch (Betula lutea), together with
hemlock (Tsuga canadensis), almost identical in aspect with virgin
forests in northern Connecticut,* occupy the intervales and many
other favorable sites; elsewhere fir and spruce predominate. Such
a condition may have continued in Connecticut for a very long
period. Ultimately, however, perhaps to the accompaniment of
* Described by the writer in Torreya 13: 199-215. 1913.
NICHOLS: THE VEGETATION OF CONNECTICUT 215
climatic changes, the more vigorous southern species gained the
upper hand, so that spruce and fir have largely been eliminated
from this region.
In the present connection the discussion set forth in the pre-
ceding paragraph is of course significant only as it bears on the
question as to the origin of Connecticut bog vegetation. In the
opinion of the writer the answer to this question may be stated
somewhat as follows. From the above discussion it seems fairly
certain that within very recent geological time the vegetation of
this region has been quite like that of northern Cape Breton
Island today, i. e., the upland forests included a mixture of broad-
leaf and coniferous forest types and bog vegetation was common in
allswamps. As the conquest of the region by the southern types
of vegetation progressed further, the coniferous type was first
vanquished on the uplands, since here environmental conditions
proved most congenial to the invaders. It was not, perhaps,
until these uplands had become pretty completely occupied that
the last strongholds of the northern plants—the swamps—were
stormed.* The northern vegetation in spring swamps was first
to succumb, since soil conditions here proved quite suitable for
certain of the southern invaders. Similarly the northern vegeta-
tion in most lake swamps was gradually replaced by southern
types. But the conquest of some of these lake swamps proved a
difficult undertaking, owing to the peculiar soil conditions which
have been described in connection with the discussion of bog
xerophytism. For to these adverse soil conditions the northern
defenders proved singularly well adapted—so well adapted, in
fact, that many of these isolated plant communities still survive
the siege. A bog, therefore, is to be regarded as a relict swamp
type whose vegetation represents vestigial remnants of a more
northern type of flora which has been much more widely repre-
sented in this region within very recent geological time. It does not,
however, seem advisable to assume that “the habitat dates back
again the serine ca condition of the vegetation in northern
Cape ten Island affords an instructive analogy. Four of the most characteristic
Connecticut swamp trees, viz., Acer oa Ulmus americana, Fraxinus nigra,
and Betula lutea, are likewise found there; but they are restricted to the uplands,
almost never occurring in swamps. This is the condition which the writer presumes
obtained in this state at the period in question.
216 NICHOLS: THE VEGETATION OF CONNECTICUT
to Pleistocene times,’’* since from a physiographic standpoint it is
doubtful if such an assumption is tenable.j In the opinion of the
writer the bog habitats at present in existence may be of much
more recent origin.
As has been mentioned elsewhere,{ there is abundant evidence
tending to show that this invasion of areas populated by northern
bog plants is still going on. At several localities in the state are
bogs in which black spruce and coast white cedar occur together,
and in every instance the cedar seems unmistakably to be gaining
the ascendancy. In Southington is a large, boggy swamp now
overgrown with red maple and elm. Scattered about in parts of
the swamp, however, are numerous specimens of black spruce,
most of which are either dead or dying. Similar relations may be
observed elsewhere, and it is of importance to note that wherever
* Transeau, E. N., Dp: 37:
+ One of the bat to an assumption is pointed out by Dr. C. A. Davis in
a recent letter to the writer. ‘It is probably a conservative estimate ... that
the end of the Wisconsin glaciation in Connecticut occurred from 20,000 to 50,000
years ago. If even the lower estimate be taken, and we assume that any given bog
started 20,000 years ago and grew at its present rate of development, say with the
peat accumulating at the rate of one foot per century, it is obvious that in the two
hundred centuries which have elapsed, the physiography of the habitat would be
entirely changed, and that we would have no bogs at the present day.”” He cites the
occurrence of a “‘cassandra bog at sea level on the coast of North eeaaigaieos hue ” cer-
tainly g tent of the t, and
hich app tl bel level not 1 ” He further
states as his opinion — S seas aie: = the theory of vierial isan, kere so
far he ire ch +
a p of bogs,
practically untenable.” Nor does he ‘elie that we need this patents since
our present climatic conditions closely approximate those of glacial times.
probably near enough to the borders of the zone of boreal influences so that our eal
is a mixed one, not from influences exerted in the past, but from those of the present
sea and the fact that we find on all our peat deposits overlapping northern and
rn floras, as is shown by every list of plants pe from associations a aontied
ony in a way supports this contention. It that our
should be that the northern plants are present in bogs because in eee places ieey
find conditions favorable for their development and growth, and that they are dis-
placed only when these conditions become unfavorable. The fact bat most of the
northern plants of bogs have fruits that are attractive to birds, or else very smal}
light seeds—and we know that migrating birds make bogs a part of their southern
migration route—would seem to help account for the occurrence of northern plants
which, after they are established, spread rapidly.”’ It will be seen that Dr. Davis
is not wholly in accord with the views of the writer, much less so with those of
Transeau.
t Torreya 13: 99. 1913.
NICHOLS: THE VEGETATION OF CONNECTICUT 2i¢
the spruce is cut off it becomes largely, if not entirely, replaced by
red maple. While, therefore, the writer’s conception as to the
origin of spruce bogs is based entirely on circumstantial evidence,
the explanation set forth seems to account in a satisfactory manner
for the observed facts.
Occurrence of Spruce Bogs in Connecticut——So far as the writer
has been able to ascertain from his own field work and from inquiry
among local botanists, fewer than thirty spruce bogs are known to
exist within the state. These are distributed as follows: Salis-
bury*, Norfolk (3)*, Kent (2), Litchfield*, Suffield (6)*, South
Windsor (2), Willington*, East Thompson (2), New Fairfield*,
Monroe*, Middlebury (2), Southington (2)*. Bogs containing an
admixture of spruce and coast white cedar occur at Bethany*,
Plainfield (2)*, Willington and Windham; while spruce grows in a
red maple swamp at Southington*.
ACKNOWLEDGMENTS
The manuscript of this paper has been examined by Professor
H. E. Gregory of the Geological Faculty of Yale University,
Dr. C. A. Davis of the United States Bureau of Mines, Professor A.
W. Evans of the Botanical Faculty of Yale University, Mr. C. A.
Weatherby of Hartford, and Mr. C. H. Bissell of Southington,
to all of whom the writer is greatly indebted for helpful sug-
gestions and criticism.
SHEFFIELD SCIENTIFIC SCHOOL OF YALE UNIVERSITY
* Bogs so marked have been visited by the writer.
Polypodium marginellum and its immediate allies*
WILLIAM R. Maxon
Among the numerous groups of species comprising the section
Eupolypodium of the genus Polypodium, as represented in tropical
America, there are few if any which have a general structure so
simple and unusual as that exhibited by Polypodium marginellum
and several closely related species. This group, of which there are
at least five American members, is represented also in Africa, as
discussed below. The several species, which apparently are all
epiphytes of moist mountain forests, are closely similar in general
form, the fronds being of a narrowly linear type, simple, 5-25
cm. long, pinnately veined, and having the entire or slightly un-
dulate margin bordered with a capillary or flattish, lustrous, dark
brown or ebeneous band of sclerotic tissue, which from its firm
structure is not readily perishable and may even persist long after
the green tissue of the frond has disintegrated and disappeared.
This sclerotic band has, so far as can be observed, no connection
whatever with the fibrovascular conducting system; that is, the
midvein and veins, the latter terminating at a point relatively
remote from the margin. It seems rather to function primarily as
a mechanical device to give strength and rigidity to the fronds,
though not improbably it meets also some physiological need that
is not at once apparent.
The marginal sclerotic structure just described is apparently
unique in Eupolypodium. What appears at first to be a similar
case is seen in Polypodium gramineum Swartz, a West Indian plant
with small, very narrow, simple, entire, darkish-margined fronds;
but in this species the dusky border is actually composed of con-
ducting tissue, consisting of a marginal vein which connects the
excurrent ends of the once or twice-branched lateral veins. Just
outside of the marginal vein may be observed a delicate line of
greenish tissue (as is not the case in P. marginellum and its allies),
* Published by permission of the Secretary of the Smithsonian Institution.
: eee
220 Maxon: POLYPODIUM MARGINELLUM AND ITS ALLIES
which suggests that reduction in the leaf expanse has taken place
inward to the boundary of the longitudinal series of simple areoles
constituting the venation of the leaf. The venation of P. grami-
neum is not precisely matched in any other species of Eupoly-
podium, but its peculiar character is certainly not of sufficient
weight to warrant giving more than minor rank, in classification,
to this species which is the type of Swartz’s genus Grammiutis.
In any event the actual structure is fundamentally different from
that of P. marginellum and its immediate allies, as may be noted
readily upon an examination of specimens, and clearly places
P. gramineum outside of this group.
Polypodium marginellum Swartz, 1788, was the first species of
its own group to be described. It has since been very generally
misunderstood and has been regarded mistakenly in so broad a
sense as to include most of its allies, at one time or another, largely
because of insufficient material for study. Without tracing in
detail the varying treatment of the several forms, which is of
historical interest merely, it seems worth while to offer the follow-
ing key and brief synopsis, by means of which it is believed the
several species may be distinguished.
KEY TO THE SPECIES
Fertile and sterile veins both mostly once forked, the branches
produced.
linear, long-ettenuate downward from the middle
stipe usually short; rhizome scales entire or nearly so.. 1. P. marginellum.
ina oblanceolate, attenuate downward to the slender
greatly OPN stipe, deciduously ciliate with lax, deli-
cate, gland-tipped, mostly simple hairs, a few similar
hairs evident at first along the midvein beneath; rhi-
zome scales evenly sinuate-dentate
Sterile veins simple; fertile veins nearly simple, the elliptical
receptacle partially overlying the vein or wholly diver-
gent as a short, oblique, translucent distal spur.
Rhizome scales rather 7 ligulate, yellowish brown in
mass; sori _— mi upon the veins, mostly apart
from the mifiveitl., : 3. 265.2254 2 3. P. limbatum.
Rhizome scales aa subulate to acicular, reddish brown
in mass; sori distinctly inframedtal, NP thrust
against the midvein or even conc
geaninionns eee pee ot 0 to 35°.
Lamina rigidly coriaceous, rere with a
relatively heavy ead marginal band........ 4. P. nigrolimbatum.
NS
. P. leptopodon.
Maxon: POLYPODIUM MARGINELLUM AND ITS ALLIES 22]
Lamina membrano-chartaceous, translucent;. mar-
ginal sclerotic line very slender, delicate........ 5. Pf. Hesstt.
Veins arising at an angle of about 20°, closer; plants
more delicate than the preceding, with smaller,
usually much narrower Tronds;. ...5.:....20.-...6+. 6. P. ebeninum.
1. POLYPODIUM MARGINELLUM Swartz, Prodr. Veg. Ind. Occ. 130.
1788
Grammitis marginella Swartz, Jour. Bot. Schrad. 18007: 17. 1801.
Mecosorus marginellus Klotzsch, Linnaea 20: 405. 1847, in part.
TYPE LocaALity: High mountains of Jamaica.
DIsTRIBUTION: Blue Mountains, Jamaica, at 1,500 to 2,100
meters elevation.
_ In_ustration: Schkuhr, Krypt. Gew. 1: pl. 7 (as Grammitis
marginella).
As indicated in the key, Polypodium marginellum is very
strongly characterized by its long-forked veins and persistently
hirtellous surfaces, characters which distinguish it at once from
those allied species which have commonly been referred to it as
the same or as only subspecifically different. Jenman, who clearly
indicated its distinctness,* had seen no continental specimens; nor
has the writer, though there are several mainland records, in-
cluding a recent one for Costa Rica.t These probably relate to
P. nigrolimbatum Jenman. Thus, Wright, in writing of the ferns
of Mount Roraima,t has listed certain specimens as P. marginellum,
with the remark: “Swartz says this species has ‘‘venis bifidis’’;
subsequent authors describe the veins as simple, which is the case
with all the specimens at Kew.”’ Itis remarkable that true P. mar-
ginellum should be lacking at Kew, for it is decidedly the commoner
of the two Jamaican species. Of the four Roraima specimens cited
by Wright only one (McConnell & Quelch 568) has been seen; this
is P. nigrolimbatum.
The following specimens of P. marginellum, all from the Blue
Mountains, Jamaica, are in the National Herbarium: Hart 72;
Maxon 1335, 1476, 2676, 2709; Underwood 1518, 2494, 3190, 3200.
* Bull. Bot. Dept. Jamaica II. 4: 69. 1897.
+ Christ, Bull. Herb. Boiss. II. 4: 1101. 1904.
t Trans. Linn. Soc. II. Bot. 6: 83. 1901.
222 Maxon: POLYPODIUM MARGINELLUM AND ITS ALLIES
2. POLYPODIUM LEPTOPODON Wright, Trans. Linn. Soc. II. Bot.
6: 83. I901
TYPE LOCALITY: Summit of Mt. Roraima, British Guiana, alti- ,
tude about 2,620 meters.
DisTRIBUTION: Known only from the original collection (Mc-
Connell & Quelch 569).
Polypodium leptopodon is a well-marked species, related to true
P. marginellum Swartz, with which it is contrasted in the preceding
key. In general form it is simulated somewhat by an occasional
specimen of P. marginellum in which the fronds are very numerous
and through crowding have developed uncommonly long and
slender stipes; but it differs constantly in characters of pubescence
and rhizome scales. The sori, as usually also in P. marginellum,
are borne on the base of the elongate branch, the two veinlets
being subequal and terminating well short of the margin.
Besides a specimen of the type collection sent from Kew the
writer has examined several plants of the same number in the
Underwood Herbarium of the New York Botanical Garden. No
other specimens have been seen, the species apparently being very
rare.
3. Polypodium limbatum (Fée) Maxon
Grammitis limbata Fée, Gen. Fil. 233. 1852; Mém. Foug. 6: 6.
1853.
TYPE LOCALITY: Guadeloupe.
DISTRIBUTION: Guadeloupe, at 750 to 1,300 meters elevation;
also St. Vincent, at 600 meters.
ILLUSTRATION: Fée, Mém. Foug. 6: pi. 5, f. 1.
This species, described originally upon specimens collected by
Perrotet, has been very imperfectly known and_ understood,
probably because of its rarity. Excellent Guadeloupe specimens
of the Duss collection in the Underwood Herbarium show it to be
amply distinct, however. Fée’s illustration is excellent in all
particulars save one; namely, that the sori, while nearly medial
upon the veins, are not borne midway between the midvein and
margin, owing to the basal curvature of the veins. The relatively
pale scales, ligulate form of the exstipitate fronds, broad marginal
band, and position of the sori at once distinguish it from its nearest
ally, P. Hessit.
MAXON: POLYPODIUM MARGINELLUM AND ITS ALLIES 223
4. POLYPODIUM NIGROLIMBATUM Jenman, Bull. Bot. Dept.
Jamaica IT. 4:69. 1897
Grammitis fluminensis Fée, Crypt. Vasc. Brés. 85. 1869. Not
Polypodium fluminense Vell. Fl. Flum. 11: pl. 66. 1827; Mus.
Nac. Rio Janeiro 5: 447. 1881.
TYPE LOCALITY: Jamaica.
DIsTRIBUTION: Blue Mountains, Jamaica, at 1,500 to 1,800
meters elevation; also in Brazil, British Guiana, and Bolivia,
ascending to 2,600 meters.
ILLUSTRATION: Fée, op. cit. pl. 19, f. 3 (as Grammitis flumi-
nensis).
Jenman in writing of the ferns of Jamaica well distinguished
this species from P. marginellum, its only near relative in Jamaica,
but erroneously cited Grammitis limbata Fée as a synonym. The
fronds are numerous, fasciculate, acute, and rigidly coriaceous,
and narrower and more stiffly erect than in P. marginellum, which,
though widely different in venation and minute characters, it
somewhat resembles otherwise. The continental specimens are
often larger than the Jamaican, but seem no different in other
respects. Fée’s species, Grammitis fluminensis, is apparently the
same, judging by a fragment of the type collection in the Under- °
wood Herbarium, but the specimen figured by him is certainly
atypical in having obtuse apices.
The largest and most perfect Jamaican material studied is
that collected by the writer from mossy tree trunks on the upper
forested slopes of Monkey Hill (above New Haven Gap), altitude
about 1,800 meters, June 22, 1904 (Maxon 2724, 2748).
5. Polypodium Hessii Maxon, sp. nov. |
Plants epiphytic, 8-14 cm. high, the fronds few, ascending,
subfasciculate. Rhizome decumbent or short-creeping, I-I.5
cm. long, 2-3 mm. thick, sparingly radicose beneath, the small
apical portion slightly exposed, inconspicuously paleaceous;
scales dark reddish brown, rigidly acicular, subfalcate, subflexuous
toward the tip, 2-4 mm. long, 0.2-0.3 mm. broad, the cell walls
richly colored; stipe slender, about I cm. long, brownish, alate,
passing gradually into the lamina, deciduously pubescent with
minute branched glandular hairs; lamina 7-13 cm. long, 5-9 mm
broad, linear, narrowly attenuate in the basal third, a little more
224. Maxon: PoLYPODIUM MARGINELLUM AND ITS ALLIES
abruptly attenuate in the apical part (the extreme apex sub-
caudate), the margins delicately repand, appearing undulate in
drying, bordered by a delicate filiform line of lustrous blackish-
brown sclerotic tissue, this bearing a few minute caducous branched
glandular hairs, similar hairs extending sparingly to the leaf
tissue; veins free, arising at an angle of about 35°, slender, con-
cealed, but their course easily evident, the sterile ones simple; sori
oval, extending from below the apex to the middle of the lamina,
seated partially or sometimes wholly upon a very short divergent
translucent distal spur of the vein, far below its middle, thus
borne close to the concealed but sharply elevated slender costa,
separate or at maturity subconfluent. Leaf tissue firmly mem-
brano-chartaceous, dark green, paler beneath, the veins. easily
evident without transmitted light.
Type in the U. S. National Herbarium, no. 694419, collected
from mossy trunks of forest trees, Sierra de Naguabo, eastern
Porto Rico, March 8, 1914, by Mr. W. E. Hess (no. 312). Mr.
J. A. Shafer’s no, 2328, collected at the same time and place, is
identical. Other specimens studied, all from Porto Rico, are as
follows: Hioram 334; Blauner 272; Sintenis 1791; Shafer 3320, 3516,
3660
Polypodium Hessii is closely related to P. limbatum of the
Lesser Antilles, but differs sufficiently in its more rigid and darker
thizome scales, in its nearly basal sori, and in its distinctly stipitate
(rather than nearly or quite exstipitate), narrower, attenuate
fronds. The marginal band of sclerotic tissue is, moreover, very
delicate and threadlike, as opposed to the glistening, broadish,
almost flat sclerotic band of P. limbatum. P. Hessii is not likely
to be mistaken for the widespread P. nigrolimbatum, with which it
is contrasted in the preceding key.
6.. Polypodium ebeninum Maxon, sp. nov.
Plants apparently epiphytic; fronds numerous, 5-10 cm. long,
ascending, imbricate-fasciculate in a small crown. Rhizome erect
or ascending, 7-10 mm. long, I-1.5 mm. in diameter, paleaceous,
the apical portion partially concealed; scales acicular, 2-2.3 mm.
long, 0.17—-0.25 mm. broad, subflexuous toward the slender gland-
tipped apex, rich castaneous from a yellowish brown base; stipe
proper nearly wanting, alate, passing gradually into the narrow
lamina from a brownish base; lamina 5-9 cm. long, 2-4 mm
broad, straight or slightly arcuate, narrowly linear, acutish or
obtuse, broadest in the apical third, thence very gradually long-
attenuate to the base, entire, slightly repand, the margin consisting
Maxon: POLYPODIUM MARGINELLUM AND ITS ALLIES 225
of a slender band of rigid lustrous ebeneous sclerotic tissue, gla-
brate; veins numerous, free, very oblique, arising at an angle of
about 20°, the sterile ones simple, the fertile ones essentially so, the
sori (usually confined to the apical half of the lamina) distinctly
inframedial in attachment, seated upon an elliptical translucent
receptacle lying upon the distal side of the vein and scarcely di-
vergent from it, relatively large, crowded against the costa, the
two lines of sori more or less confluent at maturity. Leaf tissue
glabrous, rigidly herbaceous, translucent, dark green above, paler
beneath; costa slender, dark brown, elevated beneath; veins readily
seen beneath without transmitted light.
Type in the U. S. National Herbarium, no. 370659, collected
upon the island of St. Helena, 1885, by W. H. King.
Reported commonly from St. Helena, as P. marginellum, but in
the National Herbarium represented only by the specimens of Mr.
King. It is the smallest member of the group and readily recog-
nizable as distinct from its American relatives in both gross and
minute characters. This ora very closely related form is mentioned
from Guinea by Link,* as P. marginellum. There is also a single
Canary Island specimen, collected long ago by L. von Buch, which
may possibly be the same. This was first referred to the West
Indian P. gramineum (Grammitis linearis Swartz), but was after-
ward described as a new species, Grammitis quaerenda Bolle.t
Still later it was discussed by Kuhn,ft who mentions the “char-
acteristic deep black margin”’ and refers it to P. marginellum,
adding that it is identical with specimens collected on St. Helena
by d’Urville and apparently misidentified by Mettenius as P. gram-
ineum. Kuhn’s unsupported statement as to the identity of the
Canary Island and St. Helena plants does not, however, seem at all
conclusive, in view of the several very obvious errors of fact which
occur elsewhere in his paper and considering also the circumstance
that under the name P. marginellum, with a single variety, he
lists a total of only six specimens (apparently all that were known
to him) which certainly pertain to no less than three species. It
seems proper to describe the St. Helena plant as new, for it is not
at all improbable that the Canary Island form, if eventually found
in sufficient quantity for study, will prove to be a distinct species.
* Fil, Afric. 149. 1868.
+ Zeitschr. Allgem. Erdk. II. = 324. 1863.
t Oesterr, Bot. Zeitschr. 16: 866.
Medullary spots and their cause
J. G. GROSSENBACHER
(WITH PLATES IO AND 11)
INTRODUCTION
It is practically an established fact now that the common types
of medullary spots or “pith flecks” so frequently found in the
wood of trees and shrubs, are due to the mining of insect larvae
in the cambium. The reliability of such marks as aids in deter-
mining the wood of forest trees therefore depends upon the
relative distribution of the different species of cambium miners and
the plants they infest.
The injurious effect of the mining on the health of the host
usually depends upon the relation of the mined or eaten area of
cambium to the area of the portion that is uninjured. In the case
of large trees the effect on the host is usually slight while on a small
tree or shrub a very heavy infestation may result in marked stunt-
ing of the plant’s growth. Some woods may become worthless
for finishing purposes owing to the presence of numerous occluded
mines. Perhaps the chief pathological interest in this matter lies
in following the response of the affected regions of the bark which
results in proliferations to occlude the channels left by the foraging
miners, and also in the development of new cambium over the
outside of the mines.
This article is an addition to and a continuation of a short
paper* written a few years ago, in that more of the details of the
host’s response as well as the complete life history of the miner
discussed in the latter part of the earlier paper (pp. 63-65) are
given. It is based on notes and specimens accumulated during
the years 1910-12.
SOME ADDITIONAL AND MORE RECENT LITERATURE
All of the important earlier literature on the botanical side of
the subject of medullary spots was reviewed in some detail in the
* Medullary spots: a contribution to the life history of some cambium miners.
N. Y. Agr. Exp. Sta. Tech. Bull. 15: 49-65. 1910.
227
9928 GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE
above cited paper before giving the life history of the Ribes miner
(Opostega nonstrigella). A little later Record* published a very
similar review of the same literature and added some observations
on the occurrence of medullary spots in other hosts. Another
review of the early literature was given in a later paper by Brownt
in which he also records additional hosts and gives some details
regarding the mines and miners. Five very excellent plates give
interesting information about the mines. However, in his dis-
cussion of the occlusion of the channels made by the insects, occurs
an error that should not be left unmentioned: ‘“ The larva destroys
only those cells in its immediate path through the inner bark. . .
As the cambium layer moves outward radially, the passage left
by the larva increases in diameter. For this reason, when growth
is very rapid, the pith-fleck spots are larger than when it is slow.”’
The error in these statements may be readily shown by quoting
from a few lines below out of the same paper: ‘‘ The healing process
proceeds mainly from the bast pith-ray cells in the bark.”” This
shows that Brown’s observations were correct and that the chan-
nels once made are not moved outward radially by further growth
of cells on the wood-side of the wound. In other words, it is
unlikely that subsequent growth could widen a channel made by a
cambium miner and it is very apparent that the mines are made
narrower by the enlargement of the normal cells on both sides.
Plate 4 of his paper shows how the channels are filled mainly by
proliferations from the rays on the bark side.
The first to determine the most important stages in the life
history of a cambium miner was Nielsen.t He studied the larva
in all its stages, the pupa as well as the imago, though the egg was
not found. The miner proved to be the larva of a fly, Agromyza
carbonaria. He found that the larvae fed in the cambium until
autumn and then entered the ground, where they passed the winter
as pupae. The adult flies emerge in May.
* Record, S.J. Pith flecks or medullary spots in wood. For. Quar. 9: 244-251.
Ig1r.
t Brown, H. P. Pith-ray flecks in wood. Forest Ser. U. S. Dept. Agr. Cir.
215: 5-15. pl. 1-5. 1913.
t Nielsen, J. Cc. Uber die Entwicklung von Agromyza carbonaria Zett., der
Urheber der ‘“Markflecken.” Zool. Anzeiger 29: 221-222. 1905; Zoologische
Studien tiber die Markflecke. Zool. Jahrb. 23: 725-737. 1906.
GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE 229
At first the larva has but one hook and well-developed girdles
of plates which increase in number per segment from the first to
the last: the first segment having one, and the last or twelfth, from
seven to nine. This is also characteristic of the Prunus miner as
given in my former paper. The second hook eventually appears
on the left of the first. It is very small and lies in the curve of the
large one as is characteristic of an Agromyza larva. The length of
the larva increases from about three millimeters to twenty milli-
meters before emerging from the bark to pupate in the ground. A
girdle of small spines is developed on the second segment. They
project caudad. Before the end of larval life the number of plate
girdles is much reduced; the skin becomes thicker and wrinkles
crosswise. The pupa is three to four millimeters long, barrel-
shaped, and has the spiracles at the ends on the dorsal side.
Some of Nielsen’s figures of the causal insect were later re-
produced by Tubeuf* in an article on medullary spots. He made
the interesting observation, also frequently noted by the writer,
that the miners are present only in such trees of a species which
have active cambium throughout most of the growing season. In
the autumn they were found most numerous in trees growing in
wet places. He secured only the larval stages of the miners.
Greenef recently gave an account of the pupation of a single
larva of the river birch miner discussed by Brown and described
the adult that emerged from it. The fly was thought to be
Agromyza pruinosa, and the opinion is expressed that possibly it
is the only species in America producing medullary spots. The
latter, of course, is a premature and very unlikely assumption even
as regards the dipterous insects that mine in the cambium of woody.
plants; besides it has long since been shown to be incorrect as
regards cambium miners in general. At least one other cambium
miner is a lepidopterous insect (Opostega nonstrigella). Owing
to the fact that our knowledge of the subject is so very fragmentary
and incomplete it is evident that we are not in position to generalize
as yet. In fact if Greene’s statement on page 474 of the above
.* Tubeuf, K. von. Uber die Zellginge der Birke und anderer Laubhdlzer.
Naturwiss. Zeit. Forst- und Landwirt. 6: 235-241. 1908.
reene, C. T. The cambium miner in river birch. Jour. Agr. Research 1:
471-474. 1914.
230 GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE
paper, to the effect that “‘Agromyza pruinosa remains in the pupal
stage in the ground during winter”’ applies to the fly which he and
Brown studied it certainly differs from the one discussed in this
paper. The Prunus miner, as noted above, was found to hibernate
in the cambium of the host during the winters of 1910-11 and I91I-—
12in the larval stage and to pupate early the following summers.
LIFE HISTORY OF THE PRUNUS MINER
On pages 63-65 of my paper on medullary spots cited above
are described the early stages of a cambium miner that feeds in
Prunus and Crataegus but does not attack Salix. The observa-
tions upon which that discussion was based extended from August
15 to November 15, 1910; or to the time when the cambium had
dormancy forced upon it by the early cold weather of western
New York. The larvae continued mining until nearly mid-
November but were mostly still under six millimeters in length
when they ceased feeding.
In 1911 the first careful examination and the first large collec-
tion of miners was made on May 21. They had apparently
been active for some time. The mines were most numerous in the
thicker parts of the infested shrubs. The channels were larger
than the ones made by the same larvae in the 1910 growth, as was
readily seen by tracing them back to the previous year’s growth.
The larvae still fed with their lateral sides toward the wood and
bark, but the end turns in their mines were both clockwise and
counter-clockwise. In the stem near the ground the return trips
were often only a decimeter or two in length resulting in a zigzag
course, but some of them still were making trips 1.6 meters in
length. About a fourth of the larvae had left their hosts by boring
their way out through the bark. The holes were still open and could
be readily seen once they had been found. The larvae usually
came out through the bark a centimeter or more after making a
lower turn. Some exit holes were found as high as a meter above
but most of them were within two to four decimeters of the ground;
others entered the cambium of the crown and roots before mining
their way out. In many instances mines had been made downward
to about the surface of the ground and then turned distad or upward
a decimeter or more before making an exit. Several larvae were
GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE 231
found in their channels mining their way out above the ground
during the early forenoon of May 21. Again late in the afternoon
some were found with their heads protruding from exit holes.
Many larvae were collected into fifty per cent. alcohol. They were
from eleven to thirteen millimeters in length.
A large bundle of sprouts and seedling stems of Prunus avium
and P. domestica 0.7-4 cm. in diameter was cut; and the ends of the
sticks were stuck into vessels of soil. The soil in one vessel was
made very wet and that in the other was about in condition for
good tilth.
Several P. domestica and P. avium ccediicns I-4 cm. in diam-
eter were found which had the cambium near the ground almost
entirely destroyed by the numerous short-trip zigzag mines
crossing and recrossing each other in that region.
On May 25 only about a sxith of the larvae were still in the host
and by May 30 none were found under the bark; the mines were
found occluded or closed. After a careful examination of the
surface soil under clumps of Prunus sprouts and seedlings quite a
number of small yellowish, somewhat fusiform and barrel-shaped
pupae were found. Some were collected into alcohol. They
were about 2.7 mm. in length and about 0.9 mm. in diameter
(PLATE 10, FIG. 1). On examination of the surface soil in the
vessels into which the Prunus stems had been inserted large num-
bers of the same pupae were found, indicating that they must be the
pupae of the cambium mining larvae. In the vessel where the
soil had been kept excessively wet a number of dead larvae were
present, showing that excessive moisture prevents pupation in
some cases. The stems were carefully removed and examin
but no larvae could be found in them, while mines and exit holes
were numerous. One of the vessels containing the pupae in its
soil was placed in a cheese-cloth cage, and cheese-cloth was tied
over the tops of the other vessels.
Pupae collected under Prunus shrubs on June 4 sank at once
when dropped into alcohol while those collected on May 25 usually
floated several minutes before sinking. The puparia seemed more
tightly filled by the pupae on June 4 than before. On holding the
vial containing pupae up to the light fully formed fly-like imagines
could be distinctly seen. Twenty-five of the pupae collected under
232 GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE
Prunus shrubs were placed in soil in a breeding cage. On June 6
some more pupae were collected under both Prunus and Crataegus
shrubs and placed in soil in the cheese-cloth cage. On June 14
they looked greyish black, apparently owing to the dark color of
the insect inside.
On the morning of June 17 ten very small black flies were
found flying about in the air space of a vessel over which cheese-
cloth had been tied, and some were found walking about on the soil.
Three were found dead and were placed in alcohol. They were
about 2.5 mm. long. See Fics.2and3. Five flies (three females
and two males) were placed into a glass tube containing the distal
end of a P. avium shoot and the others into a quart fruit-jar which
was inverted over a short green branch of P. avium.
Later in the day a search was made for the flies in the Prunus
and Crataegus thickets but none could be found. However, quite
a number of empty puparia were obtained from the soil beneath
the shrubs. On June 21 all but one of the flies in the glass tube
had died and it was placed in the fruit jar. The shoot in the
tube was found to have groups of from one tofour whitish eggs.
inserted under the periderm at the edge of a number of lenticels.
Twenty eggs had been deposited under the periderm around the
lenticels of this small twig by the three females caged with it.
A few flies emerged from the soil in the breeding cages each
day until June 21 and were placed in vessels containing Prunus
shoots; after that date all of the puparia were found empty. By
June 25 most of the flies in the fruit-jar had died. ‘The shoots were
carefully examined and were also found to have eggs inserted under
the periderm at many of the lenticels. Numerous. larvae had
already emerged from the eggs and were mining in the cortex while
some had even mined as far as three centimeters in the cambium.
This shows that oviposition and hatching of the eggs took less than
a week. The eggs measured about 0.2 X0.07 mm.
Seedlings and sprouts of P. avium and P. domestica ina thicket
were found to have eggs inserted about numerous lenticels on
June 21. In many cases the larvae had emerged from the eggs.
and were mining in through the cortex and cambium. But in no
instance could more than one egg be found ata lenticel. It appears
therefore that the deposition of eggs in the caged Prunus shoots.
GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE 233
was so frequently found to be in groups because of the large number
of flies present as compared to the few lenticels in which oviposition
could occur. Both in nature and in the cages the deposition of the
eggs seemed to have been accomplished by the insertion of the
ovipositor through the lenticel; as may be inferred from Fic. 4.
This is a photograph of a shoot from one of the breeding jars
after removing the periderm from a portion of it.
The larvae always emerged from the end of the egg turned
away from the lenticel and fed first in the cortex just under the
periderm, mining parallel with the surface of the bark from half a
millimeter to three millimeters; then straight in toward the
cambium, where they continued their mines up or down depend-
ing upon how the eggs had been inserted. Most of the eggs
were inserted so that their long axes coincided with that of the
plant. In cases where the eggs had been inserted at an angle with
the long axis the larvae always turned in the direction of the lesser
angle with the stem, irrespective of whether it was up or down
the plant. The direction of the first mine therefore depends upon
oviposition and perhaps the direction of the long axes of the
cambium cells encountered by the larva and is not uniformly up
or down as suggested by Kienitz, Greene, and others. For
example, in the case of those shown in the above cited figure there
is a group of two eggs from which the larvae would have mined first
toward the distal end of the shoot, while from the group of three
they would have mined toward the base of the plant first. Some
larvae were found which had mined as far as four decimeters in
the cambium in one direction; others had mined only about a deci-
meter and then reversed the course and mined in the opposite direc-
tion, after having veered to one side from the old course. The
emerging larvae were about 2-2.5 mm. long and not more than
0.2 mm. in diameter. They were comparatively numerous: four
empty egg cases were found at lenticels of a piece of a P. avium twig
which was only about six millimeters in diameter and three deci-
meters long. The larvae were collected into alcohol.
The egg stage of these cambium miners is very short, apparently
less than three days; while the larval stage lasts at least eleven
months. The pupal stage lasts perhaps about three weeks, and
the flies apparently oviposit within two days after emerging from
the puparia.
934 GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE
On July 16 the new generation of cambium miners was found
to be rather few in number as compared with the large number
of eggs and young that were present in late June. Possibly the
excessive and very long drought in connection with the un-
precedented hot weather in western New York during the latter
part of June and the first part of July (1911) reduced cambial
activity to such an extent as to make mining difficult and resulted
in the death of many larvae.
COMPARISON OF THE PRUNUS AND OTHER DIPTEROUS CAMBIUM
MINERS
When the life history of this Prunus miner iscompared with that
recorded by Nielsen for A gromyza carbonaria and with that given
by Greene for A. pruinosa, as well as with the fragmentary history
published by Kientz (discussed in my earlier paper), it becomes
evident that though these miners are similar in many ways they
must be different species. A. carbonaria and A. pruinosa pass the
_ winter in the pupal stage in the ground while the Prunus miner
goes through winter in the cambium of the host asa larva. The
fact that the Prunus miner was never found in willows growing
among infested Prunus, as recorded in my earlier paper, also goes
to show that the insect so commonly present in Salix and Betula
is not identical with the Prunus miner. In addition to the above
noted and irreconcilable differences in the life histories and hosts
of the miners there is a marked difference in the size of all the
stages of the Agromyza pruinosa discussed by Greene and others
and those of the one described in this paper. The most striking
difference is to be found in the length of the larvae just before
pupation: The measurements given by Greene and others for the
fully grown larvae studied by them are 20-30 mm. while those
of the fly under consideration here are only 11-13 mm. in length.
The larvae of A. carbonaria also measure about 20 mm. in length;
and besides, Nielsen calls especial attention to the reduction in
the number of plate-girdles preceding pupation which does not
hold of the miner of Prunus. The Prunus miner retains all the
girdles at least to within a day of pupation. The adults of A. car-
bonaria and A. pruinosa as given by Nielsen and by Greene are
also larger than these. The venation of the wings figured by
GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE 235
Nielsen for A. carbonaria and of A. pruinosa by Greene also differs
from that of the wings of this species, as may be seen by comparing
a, b, and c of Fic. 5. However, Brown’s observations, published
in the paper cited above, show beyond a doubt that the miner of
the river birch also passes the winter as a larva in the cambium of
the host, since on April 25 it was found feeding and measured 15
mm. in length. The Prunus miner appears to differ so materially
from the other species in both its life history and morphology
that I consider it best, at least provisionally, to name it Agromyza
Pruni sp. nov.
THE OCCLUSION OF THE MINES AND THE NATURE OF THE WOUND-
TISSUES PRODUCED
The configuration of the mines was described in some detail in
my former paper but their occlusion and the reformation of the
cambium over the streak-wound were only incidentally mentioned.
In sections showing mines near the cambium or within the un-
lignified zone one is able to see that a distinct new cambium de-
velops over the channels by the division of the first to the third
cell of the phloem outside the mine. Fics. 6 and 7 give some
idea of later but closely related stages. This new cambium is often
found formed over the wounds before the regular cambial line is
advanced that far, so that the cambium arches over the channels.
Fic. 8 illustrates this point. The first noticeable change taking
place in the cells surrounding a mine after the larva has passed is
the rapid growth of the uninjured cells on the radial and bark sides
of the channel. The phloem-ray cells usually enlarge more rapidly
than the others and give rise to bladder-like proliferations which,
after attaining considerable size, are cut off by septa from the cells
giving rise to them (Fic. 9). The cells arising in this manner con-
tinue the growth in the same direction if space permits, and, if
other proliferations do not crowd on the sides, their diameter
becomes so great that septa soon form also parallel with the original
direction of growth. See Fics. 10 and 11. After the proliferating
outgrowths from the rays encounter the opposite side of the mine
the further enlargements are diverted in the directions of least
resistance that are most nearly in line with their former direction
of growth. In many instances of this type the cell back of the
236 GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE
terminal one undergoes lateral enlargement and proliferates into
the available space, cutting off the outgrowth by a septum. An
early stage of this process may be noted near the left of Fic. 9.
During the development of the substitute cambium over the
streak-wound the growth of the phloem cells usually reduces the
radial diameter of the mines from forty to sixty per cent. In the
case shown in FIG. 12'this compression growth was so pronounced as
to wholly prevent the usual form of occlusion. The phloem-ray
cells appear to retard this general growth that brings about such a
marked reduction of the channels, as may be seen from FIGs. 6
and 7 which show the status of things in an early stage. The
broader rays have a more decided influence than the mono- or
biseriate rays. Asa matter of fact, the same characteristics may
be noted in all the figures to a varying degree. The bark side of an
occluded mine nearly always has a marked undulatory boundary
line owing to the more active growth of the interray portions of
the bounding phloem that occurs during the development of the
substitute cambium over the wound. In some instances where
channels were made through a region in which the rays are all
monoseriate and at a time when radial growth is especially pro-
nounced, they may be entirely closed by general growth from the
bark and radial sides, thus leaving only the irregular lines of com-
pressed waste and some irregularities in the arrangement of the
cells to show the location of the medullary spot. That is, the
growth of the interray portions dominates the healing process and
thereby prevents proliferation from the rays, as shown in Fic. 12.
The boundary line along the pith side of the occluded ‘mine is
usually fairly straight. This is especially noticeable in Fics. 13
and 14. The tangential diameter of the occluded mine is from
one to four or more times the diameter of the miner. This greater
or tangential diameter of the original channel is also considerably
reduced during the early stages of the healing process, as may be
seen by the incurving of the bounding ray-cells, as well as by their
bulging enlargements encroaching on the mined side. Fics. 11,
15, and 16 show results under consideration. However, it is very
unusual for the uninjured ray-cells on the sides to give rise to pro-
liferations into the channels: the neighboring cells of the inter-
rupted rays seem to respond more quickly, apparently owing to the
GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE 237
fact that longitudinal resistance or compression has been elimi-
nated while in case of the others only the lateral pressure has been
reduced. Neither diameter of a mine is increased by subsequent
growth as suggested by Greene but all available evidence indi-
cates that a considerable decrease occurs in both diameters during
the occlusion of a cambium mine.
It appears that ordinarily the radial diameter of a channel is
but little greater than that of the miner while the tangential diame-
ter may be several times that of the larva. The principal reasons
for this seem to be that the portion of the cambium most desired
by the larvae for food is a sheath only a few cells in thickness and
that they feed with the lateral sides toward the bark and wood.
In that position the hooks with which the cell-walls are ripped
open also lend themselves more readily to foraging a wide path in
the plane of the cambium mantle than at right angles to it. The
tangential widths of mines made by larvae of a certain size seem
therefore to depend upon the relation of the movement and feeding
impulses dominating them. When the cambial cells about a miner
are in just the right stage to be most agreeable for food and the
larva is hungry, and besides has no special desire to travel, it makes
a very wide path; while if the cells encountered afford less suitable
food or the insect is dominated by an impulse for movement rather
than for feeding, the mine is made only large enough to permit its
passage. That is, the mines have a greater tangential diameter
at times chiefly because the larvae feed more voraciously and
browse more widely at some times than at others. Compare the
two medullary spots of Fic. 14: the lower resulted from a mine
made in early summer, and the upper from one made in mid-
summer. Both appeared to have been made by the same larva,
the lower one while the insect was younger and smaller.
The irregular mass of cells making up the tissue that occludes
or fills up a channel left by a miner has the general appearance of.
pith. Like pith these occlusion masses come to serve as places
for storing elaborated food. The chief difference between the
appearance of sections from the two places lies in the greater
variation in the size and shape of the medullary spot cells, as
shown especially in Fic. 9. Sections of material collected in sum-
mer show that in general the cells occluding mines do not lignify
238 GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE
as soon as those of the normal tissue at an equal distance from the
cambium; they also show that the cells making up medullary spots
are not all of the same age nor produced under equal and like pres-
sures: some of them lignify before others, some are angular while
others are nearly circular in cross section. Fics. 9,13,and 14 show
practically all of the types usually found.
The study of the development of such substitute tissues has
much of indirect interest to plant pathology, and the wounds,
when numerous, may sometimes not only prove very harmful to
the host but incidentally afford entrance to parasitic organisms.
In the case of Prunus no such connection with a fungous disease
was established; a very striking example, however, was found in con-
nection with the study of the Ribes cambium miner discussed in
my earlier paper. In that case many of the shoots, vacated by a
miner that it might pupate in the ground, were invaded by a fungus
through the exit hole and killed back.
Description of plates 10and 11
PLATE 10
Fic. 1. Pupae and ese of the Prunus cambium miner (Agromyza Pruni).
Upper right collected May upper left collected June 4; lower right collected
June 11; lower left valucina ae emergence of fly.
Fic. 2. Adult fly of Prunus miner (Agromyza ews from breeding cage, IQrt.
ae 3. Female with ovipositor extruded. eeding cage, IgII
G. Twig of Prunus avium from breeding cage. The periderm conaved to
Pine sgh oo of two groups of eggs; the upper group had been inserted downward
from ray aseydi and the lower one upward from lenticel below.
gs of various species of Agromyza. a, Wing of A. carbonaria, from
eiailies “ Wits of A. pruinosa, from Greene; c, Photographed wing of A. Pruni.
Fic. 6 edullary spot in Prunus domestica, near cambium; showing an early
stage in the development of a new cambium from inner phloem cells.
Fic. 7. Medullary spot in an domestica near cambium, after the production
of a few xylem cells outside the spo
Fic. 8. Medullary spot oka new cambium just developed outside the wound;
being in advance of the other cambium as shown at right of figure.
Fic. 9. Medullary spot in which the cells are not all mature, showing a marked
variation in the size and shape of the cells,
Fic. 10. Medullary spot, made by a young larva and filled by proliferations from
the rays and by the incurving of the radial rows on the sides as well as by the gen-
eral growth on the bark side. (Upper.)
Fic. 11. Medullary spot, made by a young larva, filled by proliferation from
ray on bark side and by bulging of ray from left side. The cells of the bulging
ray are shown also to have undergone longitudinal division.
GROSSENBACHER: MEDULLARY SPOTS AND THEIR CAUSE 239
Fic. 12. Medullary spot shown only by the presence of irregular lines of com-
pressed waste. This channel was occluded at this point by general growth and not
by proliferations from rays as is usually the case.
Fic. 13. Medullary spot, showing that bark rays not only proliferate freely
into mines but that they thereby prevent or retard the general ern growth,
that is especially pronounced at the left where large rays are abse
PLATE 15
Fic. 14. Two medullary sp made by tl t also showi ing
well marked aro hal aan lines on the bark and pith sides of the
sorry chann
es
eae spot showing incurving of bounding rays at right and left,
as eae as a bulging of the ray at right
Fic. 16. Medullary spot showing incurving of rays and of other radial rows.
INDEX TO AMERICAN BOTANICAL LITERATURE
1903-1915
The aim of this Index is to penn ‘all current botanical literature written by
Americans, published in ae. or based upon’ American material ; the word Amer-
ica being used in the broadest se
Reviews, and papers that ie exclusively to ‘Gunty: agriculture, horticulture,
manufactured products of vegetable origin, or laboratory methods are n d nd
no attempt is made to index the sheretain: of ee: An occasional exception is
made in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not men iets unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
subscriber must ee all cards published during the term of his subscription, Corre
spondence re to the card issue should be —— to the Treasurer of the ocr:
Botanical Clu
Allard, H. A. Effect of dilution upon the infectivity of the virus of
the ae disease of tobacco. Jour. Agr. Research 3: 295-299.
15 Jal
Anderson, 4 ” Retention of chlorophyll through the cate tin process.
Bot. Gaz. 58: 528. 21 D 1914.
Anderson, P. J. The effect of dust from cement mills on the setting
of fruit. Plant World 17: 57-68. Mr 1914.
Andrews, E. F. The galax odor. Torreya 15: 16-18. f. 2. 25 Ja 1915.
Arthur, J. C., & Fromme, F. D. The taxonomic value of pore char-
acters in the grass and sedge rusts. Mycologia 7: 28-33. f. I-
Atkinson, G. F. The development of Armillaria mellea. Myc.
Centralb. 4: 113-121. pl. 1, 2. 21 Ap 1914.
Baker, C. F. First supplement to the list of the lower fungi of the
Philippine Islands. Leaflets Philip. Bot. '7: 2417-2542. 14 N 1914:
Barbour, E. H. Carboniferous plani tissue. Am. Jour. Sci. 39: 173,
74. 7. 2: F I9%5.
Belling, J. The chromosome hypothesis of heredity. Jour. Heredity
6: 67. 25 Ja 1915.
Berger, E.W. History of Citrus canker. Florida Agr. Exp. Sta. Bull.
124: 27-30. O 1914.
Berry, E.W. Notes on the geologic history of Platanus. Plant World
17: 1-8. f. 1-5. Ja 1915.
241
242 INDEX TO AMERICAN BOTANICAL LITERATURE
Berry, E. W. Notes on the geological history of the walnuts and
hickories. Smithsonian Rept. 1913: 319-331. f. I-4. 1914.
Britton, E.G. West Indian mosses—II. Mosses of the Danish West
Indies and Virgin Islands. Bull. Torrey Club 42: 1-8. pl. 2. 29
Ja 19t5.
Includes Hyophila uliginosa, Phascum sessile, and B icrod: spp. nov
Britton, E. G., & Hollick, A. A new American fossil moss. Bull.
Torrey Club 42: 9-10. f. r, 2. 29 Ja 1915.
Plagiopodopsis Scudderi gen. et sp. nov.
Brown, H. P. A timber rot accompanying Hymenochaete rubiginosa
(Schrad.) Lev. Mycologia 7: 1-20. pl. rgg-151. Ja 1915.
Bryan, G. S. The archegonium of Sphagnum subsecundum. Bot.
Gaz. 59: 40-56. pl. 4-7. 27 Ja 1915.
Burlingame, L. L. The morphology of Araucaria brasiliensis—III.
Fertilization, the embryo, and the seed. Bot. Gaz. 59: 1-39. fi.
I-3. 27 Ja 1915.
Burt, E. A. The Thelephoraceae of North America—III. Craterellus
borealis and Cyphella. Ann. Missouri Bot. Gard. 1: 357-382. pl. 19.
30 Ja 1915.
Includes descriptions of 6 new species.
Campbell, D. H. Botanizing excursions in Borneo. Pop. Sci. Mo.
86: 193-203. F 1915.
Campbell, D. H. The present and future of botany in America.
Science II. 41: 185-191. 5 F 1915.
Cannon, W. A. A note on the reversibility of the water reaction in a
desert liverwort. Plant World 17: 261-265. S 191
Cannon, W. A. On the density of the cell sap in some desert plants.
Plant World 17: 209-212. Jl 1914.
Cannon, W. A. Specialization in vegetation and in environment in
California. Plant World 17: 223-237. f. 1-3. Au 1914.
Chapman, R. N. Observations on the life history of A an bilineatus.
Jour. Agr. Research 3: 283-293. pl. 38, 39. 15 Jai19g
Includes a brief account of the coma sagas pleads uation and the
chestnut borer in causing the death of ce
Christensen, C. Index Filicum. Supplementum 1906-1912. I-132.
1913
Cockeceil, T. D. A. Characters of Helianthus. Torreya 15: 11-16
25 Ja 1915.
Cocks, R. S. Notes on the flora of Louisiana, I. Plant World 17:
186-191. Je 1914.
Cook, O. F. Two classes of hybrids. Jour. Heredity 6: 55, 56. 25
Ja 1915.
INDEX TO AMERICAN BOTANICAL LITERATURE 243
Couch, E. B. Notes on the ecology of sand dune plants. Plant World
17: 204-208. f. 1-4. Jl 1914.
Cowles, H.C. The economic trend of botany. Science II. 41: 223-
220. -12:F39ts:
Criddle, N. A note on the colours of tumbling mustard seed. Ottawa
Nat. 28: 138, 139. 26 Ja 1915.
Dean, A. F. The Myxomycetes of Wisconsin. Trans. Wisconsin
Acad. Sci. 17: 1221-1299. O 1914.
Duggar, B. M., & Davis, A.R. Enzyme action in Fucus vesiculosus L.
Ann. Missouri Bot. Gard. 1: 419-426. 30 Ja 1915.
East, E. M. The phenomenon of self-sterility. Am. Nat. 49: 77-87.
F 1915.
Fallis, A. L. Growth of the fronds of Nereocystis Lnetkeana. Puget
Sound Marine Sta. Publ. 1: 1-8. pl. r. 15 Ja 1915.
Fawcett, H.S. Citrus canker in Florida and the Gulf States. Monthly
Bull. State Comm. Hort. California 3: 512, 513. D 1914.
Fawcett, W., & Rendle, A. B. Flora of Jamaica. 3: i-xxiv + 1-280.
pl. 1-5 +f. I-113. 1914.
Fitzpatrick, H. M. A parasitic species of Claudopus. Mycologia 7:
34-37. pl. 153 +f. 1. Ja 1915.
Claudopus subdepluens sp. nov. is described.
Garner, W. W., Allard, H. A., & Foubert, C. L. Oil content of seeds
as affected by the nutrition of the plant. Jour. Agr. Research 3:
227-249. 15 D 1914.
Gates, R. R. Some Oenotheras from Cheshire and Lancashire. Ann.
Missouri Bot. Gard. 1: 383-400. pl. 20-22. 30 Ja 1915.
Gates, R. R. A Texan species of Megapterium. Ann. Missouri Bot.
Gard. 1: 401-404. pl. 23. 30 Ja 1915.
Megapterium argyrophyllum sp. nov
Giddings, L. A. A eadaptata of Silphium laciniatum L. Plant
World 17: 309-328. f. 1-10. N 1914.
Gortner, R. A., & Harris, J. A. Notes on the technique of the deter-
mination of the depression of the freezing point of vegetable saps.
Plant World 17: 49-53. F 1914.
Greenman, J. M., & Thompson, C. H. Diagnoses of flowering plants,
chiefly from the southwestern United States and Mexico. Ann.
Missouri Bot. Gard. 1: 405-418. of 24-26. 30Jai19g15
Includes Oecopetalum mexicanum gen nov. and eine Chandleri,
Siphonoglossa Greggii, Zephyranthes cco, Randia Gaumeri, R. Purpusii, R,
truncata, spp. nov., and several new combina
Greenwood, H. E. Revised list of Fisisc collected in and near
Worcester, Massachusetts. Bryologist 18: 6-9; 28,29. I915.
244 INDEX TO AMERICAN BOTANICAL LITERATURE
Grout, A. J. Leptobryum pyriforme (L.) Wilson, with gemmae.
Bryologist 18: 9, 10. f. 7. Ja 1915.
Halsted, B. D., & others. Report of the Botanical Department. Rep.
Bot. Dept. New Jersey Agr. Exp. Sta. 1913: 537-629. pl. I-19.
1914.
Hamblin, S. F., & Bodfish, E. H. Identifying plants without a key.
1-25. f. 1-50. Boston. 20 Ja 1914.
Harper, R.M. Phytogéographical notes on the coastal plain of Arkan-
sas. Plant World 17: 36-48. f. 1-3. ;
Harris, J. A., & Gortner, R.A. On the influence of the order of develop-
ment of the fruits of Passiflora gracilis upon the frequency of
teratological variations. Plant World 17: 199-203. Jl 1914.
Harvey, J. C. Orchids in South Mexico, Orchid Rev. 23: 12-18.
fa 36 F9is.
Hasselbring, H., & Hawkins, L. A. Physiological changes in sweet
potatoes during storage. Jour. Agr. Research 3: 331-342. 15 Ja
1915.
Hayes, H. K. Tobacco mutations. Jour. Heredity 6: 73-78. f. 12,
13 + frontispiece. 25 Ja 1915.
Hill, E. J. Fontinalis Umbachiit Cardot. Bryologist 18: 10-12. Ja
IQI5.
Hitchcock, A. S. A text book of grasses. i-xvii + 1-276. f. 1-63 +
frontispiece. New York. 1914.
Hooker, H. D., Jr. Thermotropism in roots. Plant World 17: 135-
153- My 1914.
House, H. D. The sand dunes of Coos Bay, Oregon. Plant World 17:
238-243. f. r, 2. Au 1914.
Howe, R. H. A list of lichens collected in Newfoundland, with critical
notes on the family Usneaceae. Plant World 17: 154-160. f. 1.
y 1914.
Illick, J. S. Pennsylvania trees. Penn. Dept. Forestry Bull. 11:
1-231. pl. 1-129 +f. I-103. Je 1914.
Jones, B. J. The natural modes of distribution of pear blight in
California. Monthly Bull. State Comm. Hort. California 3: 505-
sri. f. 810,777. Dig
Lewis, I. F. The seasonal life-cycle of some red algae at Woods Hole.
Plant World 17: 31-35. F 1914.
Lipman, C. B. Antagonism between anions as related to nitrogen
transformation in soils. Plant World 17: 295-305. O 1914.
Lipman, C. B., & Fowler, L. W. Isolation of Bacillus radicola from
soil. Science II. 41: 256-259. 12 F 1915.
INDEX TO AMERICAN BOTANICAL LITERATURE 245
Loeb, J., & Wasteneys, H. On the identity of heliotropism in animals
and plants. Proc. Nat. Acad. Sci. 1: 44-47. Ja 1915.
Marshall, C. G. Perjugate cotton hybrids. Jotr. Heredity 6: 57-64.
f. 7-5. 25 Ja 1915.
Mason, D. T. The life history of lodgepole pine in the Rocky Moun-
tains. U.S. Dept. Agr. Bull. 154: 1-35. pl. 1-5. 14 Ja 1915.
Memminger, E.R. A list of plants growing spontaneously in Hender-
son County, N. C. Jour. Elisha Mitchell Sci. Soc. 30: 126-149.
Ja 1915.
Murrill, W. A. A new Bolete from California. Mycologia 7: 44.
a 1915.
Rostkovites californicus sp. nov.
Murrill,W.A. Southern polypores. i-iv + 1-66. NewYork. 1915.
O’Gara, P. J. New species of Colletotrichum and Phoma. Mycologia
7: 38-41. Ja 1915
Four new species are described.
O’Gara, P. J. Occurrence of silver scurf of potatoes in the Salt Lake
Valley, Utah. Science II. 41: 131, 132. 22 Ja 1915.
Orton, C. R. Structural parallelism between spore-forms in the
Ascomycetes. Mycologia 7: 21-27. pl. 152. Ja 1915.
Osterhout, W. J. V. The value of antagonism. Science IJ. 41: 255,
256.12: F 191s.
Parish, S. B. Sketches of the Colorado desert. Plant World 17: 122-
130. Ap 1914.
Pearson, G. A. The rdle of aspen in the reforestation of mountain
burns in Arizona and New Mexico. Plant World 17: 249-260.
1.25 oS TOTAL
Rigg, G. B. Notes on the flora of some Alaskan sphagnum bogs
Plant World 17: 167-182. f. 1-3. Je 1914
Rosendahl, C. O. Experiments in forcing native plants to blossom
during the winter winter months. Plant World 17: 354-361. D
1914.
Rumbold, C. A summer’s record of evaporation and precipitation in
Lancaster County, Pennsylvania. Plant World 17: 213-215. Jj]
1914.
Sharp, S.S. Notes on the determination of Rocky Mountain conifers.
Torreya 15: 1-8. f. r. 25 Ja 1915.
Shive, J. W. The freezing points of Toidnehan' s nutrient solutions.
Plant World 17: 345-353. D 1914.
Shive, J. W., & Livingston, B. E. The relation of atmospheric evap-
orating power to soil moisture content at permanent wilting in
plants. Plant World 17: 81-121. f. 7-5. Ap 1914.
246 INDEX TO AMERICAN BOTANICAL LITERATURE
Shreve, F. A montane rain-forest. 1-110. pil. 1-129 +f. 1-18.
Washington, D. C. 1914.
Carnegie Inst. Wash. Publ. 199
Shreve, F. Rainfall as a — of soil moisture. Plant World
9) 9-26:f- 1-3. Ja
Shull, G. H. The Tae of submerged seeds. Plant World 17:
329-337. f. I, 2. N 1914.
Smith, G. M. The organization of the colony in certain four-celled
algae. Trans. Wis. Acad. Sci. 17: 1165-1220. pl. 85-91 +f. I-6.
Snow, J. W. The plankton algae of Lake Erie, with special reference
to the Chlorophyceae. U.S. Fish Com. Bull. 1902: 371-394. Pl.
I-4. 1903.
Includes descriptions of 1 new genus and 13 new species.
Standley, P. C. Two plants new to the flora of Louisiana. Torreya
15: 9-11. f. I. 25 Ja 1915
Stevens, H.E. Studies of Citrus canker. Florida Agr. Exp. Sta. Bull.
124: 31-43: f. 6-11. O 1914.
Stuckert, T. Beitrage zur Kenntniss der Flora Aseeatiileen: 1... Ao
Conserv. & Jard. Bot. Genéve 17: 219-234. 1914. II. Ann. Con-
serv. & Jard. Bot. Genéve 17: 278-309. 1914.
Stuckert, T., & Heimerl, A. Die Nyctaginaceen Argentiaiens. In
Stuckert, T., Beitrage zur Kenntniss der Flora Argentiens. Ann.
Conserv. & Jard. Bot. Genéve 17: 220-234. 1914.
Stuckey, H. P. Tomatoes. Georgia Agr. Exp. Sta. Bull. 112: 211-
248. f. 1-14.) Ja tots.
Includes a chapter on “ Blossom-end rot.”
Taubenhouse, J. J. Diseases of the sweet pea. Delaware Agr. Exp.
Sta. Bull. 106: 1-93. f. 1-43. N 1914.
Taylor, N. Flora of the vicinity of New York. Mem. N. Y. Bot.
Gard. 5: I-vi + 1-683. pl. 1-9. 30 Ja 1915.
Thom, C., & Shaw, R.H. Moldinessin butter. Jour. Agr. Research 3:
301-310. f. I. 15 Ja 1915.
Watson, J. R. Whitefly control, 1914. Florida Agr. Exp. Sta. Bull.
123: 3-24. f. 1-5. S 1914.
Includes notes on the fungous diseases of the larvae.
Weaver, J. E. Evaporation and plant succession in southeastern
Washington and adjacent Idaho. Plant World 17: 273-294. f. 1-
ro. O19!
Wolf, F. A. SS caiehginy leaf-blight. Proc. Alabama State Hort. Soc.
" 11: 56-58. 1914.
-
Butt. TORREY CLUB VOLUME 42, PLATE 10
GROSSENBACHER: MEDULLARY SPOTS
BULL. TORREY CLUB VOLUME 42, PLATE II
GROSSENBACHER: MEDULLARY SPOTS
Vol. 42 No. 5
BULLETIN
OF THE
TORREY BOTANICAL CLUB
ee ee
MAY, 1915
Notes on the Michigan species of Polygonatum*
OLIVER ATKINS FARWELL
(WITH PLATES 12-18)
Much difficulty has been experienced, during the past few
years, in trying satisfactorily to place all our forms of Polygonatum,
known as Solomon’s Seal, within the prescribed limits of the two
species accredited in our manuals to North America. There area
number of characters which seem to be common to all. The stem
is. finely many-striate, cylindrical and smooth when green but
channelled more or less deeply when dry; the leaves of all are
glaucous beneath and have minutely pubescent or papillose mar-
gins, the papillae of which range from 85 w in P. cuneatum to 7 u
in P. biflorum; they are indefinitely nerved, the smallest having not
less than fifty-five nerves, which can be definitely distinguished
by the unaided eye; and they are never acute but end in a short,
narrow or broad, very obtuse or rounded apex; the peduncles are
compressed, channelled on one side, and arching; the pedicels,
at least at the flowering season, are bracteate near their junction
with the peduncle; the perianth-lobes are deltoid, dark green,
and barbellate within; the filaments are partly adnate, partly
free; the anthers are sagittate; the fruits are black with a bluish
bloom, as in many species of Vaccinium, and if collected before
maturity appear, in the dried specimens, to be indehiscent, but
at full maturity they are dehiscent, rupturing with the slightest
pressure; the bony, whitish or greenish white seeds are turgid, sub-
reniform, and smooth. The forms can be sped separated into
* Contributions to the Botany of Michigan, No.
[The BULLETIN for April (42: 169-246. fl. 10, Ae was issued April 27, 1915.)
247
%
248 FARWELL: MICHIGAN SPECIES OF POLYGONATUM
two groups based upon pubescence of foliage or lack of it, size
of flowers, and form of filaments, the two groups corresponding
to the two species usually recognized. But it is next to impossible
to include under one species forms with yellow flowers and those
with green; forms that are constantly seven or eight feet in height
with those that never exceed one or two feet; forms that have
elongated very narrow leaves with those that have very short
and broad ones; forms with slender flexuous peduncles with those
having coarse, rigidly wiry ones. Differences in size alone are not
characters upon which to base new forms of plants but when
correlated with other constant differences there is pretty good
evidence for their recognition. An effort to place these forms led
to an investigation of the publications of earlier botanists with some
surprising and unexpected results.
It seems that neither Linnaeus, Miller, the elder Aiton, Murray,
nor any of their contemporaries had recognized any American
species of this group of plants nor even included American
plants in the European species with which they dealt. Walter
was the first to describe an American species, the Convallaria
biflora, a glabrous plant with three-nerved leaves, which has since
had a rather varied career. Bosc came next, describing C. hirta,
a species with hispid peduncles and stems, otherwise glabrous.
Michaux and Persoon referred the American species to the Euro-
pean C. multiflora. Wildenow followed with two new species,
C. pubescens and C. canaliculata. Besides recognizing the above
species under Polygonatum, now considered distinct from Con-
vallaria, Pursh, suppressing Walter’s specific name biflora and
substituting for it his own name angustifolium, admitted P.
latifolium, a European species, to a place in our flora. Poiret
came next with Convallaria parviflora. The two Schultes de-
scribed C. commutata and, in 1835, Dietrich added still another,
P. giganteum. Thus in the first half century after the first
American species had been described eight others were recognized
by the earlier students of the American flora, seven being con-
sidered endemic and the other two identical with European species.
It would thus seem that the botanists of a century ago had a more
accurate conception of the genus in America than those of the
present day. More than seventy years later Dr. Greene described
FARWELL: MICHIGAN SPECIES OF POLYGONATUM 249
three other species, P. virginicum, P. boreale, and P. cuneatum,
bringing the total number of species described for America up to
twelve; but these, like most of the others, have not been generally
accepted as valid. Dr. Hooker, in 1840, united the nine species
then known and placed them all under P. multiflorum as var.
americanum. Dr. Torrey followed the lead of Hooker and went
even further by uniting them all under P. multiflorum, not even
giving them varietal rank. In the first edition of the Manual,
Asa Gray recognized P. pubescens Pursh and P. canaliculatum
Pursh. In the second edition, the former becomes P. biflorum
Ell., the latter, P. gigantewm Dietr., and a third species, P. lati-
folium Desf., is admitted. In later editions P. commutatum super-
sedes P. giganteum and P. latifolium is eliminated.
As stated above, Walter was the first to describe an American
species under Convallaria, and his description is as follows:
biflora 2. foliis semiamplexicaulibus trinervis laevibus, oblongis,
acutis, altnernis; caule tereti laevi; pedunculis solitariis bi-
floris axillar, floribus flavescentibus.
Willdenow’s descriptions, in part, of C..pubescens and of C.
canaliculata are as follows:
Convallaria pubescens
Convallaria foliis alternis amplexicaulibus ovatis subtus pubes-
centibus, caule teretiusculo sulco exarto, pedunculis axillaribus
subbifloris. . . .
Folia alterna ovata secunda basi leviter amplexicaulia integerrima
nervosa, nervis 7 majoribus, supra saturate viridia glabra,
subtus albicantia et ad nervos pilis albis obsita. Pedunculi
uni- vel biflori secundi nutantes axillares. Corolla... .
albido-flavescens ....
Convallaria canaliculata foliis alternis amplexicaulibus oblongis —
margine pubescentibus, caule canaliculato, pedunculis bifloris
axillaribus.
Differt a Convallaria pubescenti: foliis oblongis glabris margine
tantum tenuissime pubescentibus, coralla magnitudine et facie
Convallariae Polygonati. Hanc sub nomine Convallariae multi-
florae in Flora boreali-Americana indicavit Michaux p. 202.
250 FARWELL: MICHIGAN SPECIES OF POLYGONATUM
A very careful study of these descriptions never should have per-
mitted any confusion as to the application of the names; if there
are but two species in North America, then the proper names for |
them are Polygonatum pubescens (Willd.) Pursh, for the plant with
pubescent leaves, and P. biflorum Walt., for the plant with glabrous
leaves. Willdenow’s plate illustrating the former (see PLATE 12)
is excellent and is exactly matched by plants in Michigan. Both
C. biflora and C. canaliculata are described as having oblong
leaves and two-flowered peduncles; in the former the flowers are
described as yellow and in the latter as of the size and appearance of
those of C. Polygonatum, which are green and about seven lines in
length; the leaves in the former are said to be three-nerved but
Willdenow makes no mention of this feature; Kunth, however, in
his description of Willdenow’s plant says the leaves are ovate-
oblong, about three inches long by sixteen or seventeen lines in
width and striately many-nerved. The two cannot therefore be
of the same species. Pursh described P. angustifolium with
elliptical-lanceolate leaves; Elliott described P. biflorum Walt.
likewise; in other respects these descriptions are essentially the
same as that of Walter’s C. biflora, each quoting the latter as a
synonym. But an oblong leaf that is acute may be essentially
the same as a leaf that is elliptical-lanceolate,and these three names
and descriptions may, therefore, be considered as synonymous,
Elliott restoring Walter’s specific name, which had been suppressed
by Pursh for one of his own coining.
It seems scarcely possible by the widest stretch of the imagina-
tion to include, under the above description of Walter, the pubes-
cent-leaved plants with small greenish or greenish white flowers,
yet that is not only what Gray and subsequent authors have done
but they have finally come to make the latter form the principal
element of the species to which Walter’s name has been applied
or rather misapplied. The confusion of these species was started
by Dr. Hooker, continued by Dr. Torrey, and finally completed
by Dr. Gray, when he ultimately and inexcusably transferred the
name of Walter from the species described by that author to an
entirely different one and not of very close relationship. And this
interpretation has been blindly followed by subsequent authors
for nearly half a century.
FARWELL: MICHIGAN SPECIES OF POLYGONATUM 251
The Michigan species naturally fall into two groups—one
containing low, slender plants having pubescent chartaceous
foliage and small flowers with slender filaments; and the other
containing robust plants having glabrous, membranaceous leaves
and large flowers with large, stout filaments. Both are to be
placed under Baker’s section ALTERNIFOLIA. The first group
may be known as the Pubescentes and the second, as the Glabrata.
Under the Pubescentes three species have been described, P.
pubescens, P. cuneatum, and P. boreale. In all these forms the
filaments are papillate. Dr. Greene describes the peduncles of
P. boreale as filiform and flexuous; this can scarcely be said of
the specimens that have come under the author’s observation,
except as to the two or three uppermost, the others being too
stout, compressed, and channelled to be called filiform; otherwise
the specimens agree with Dr. Greene’s descriptions. There is
another form with the perianth pale green throughout or with the
lobes just noticeably of a darker green than the tube.
In the Glabrata seven species have been described, P. biflorum,
P.hirtum, P.canaliculatum, P. parviflorum, P.commutatum (all orig-
inally as species of Convallaria), P. giganteum, and P. virginicum;
two others have been referred to the European P. Jatifolium and
P. multiflorum, making nine in all. These species will fall into
two series, one with yellowish white flowers and relatively narrow
leaves and the other with greenish flowers and relatively broad
leaves. Several forms can readily be recognized; also one form with
ancipital stems has been found, i. e., in cross-section the outline
is elliptical. P. commutatum and P. giganteum are but varying
forms of P. canaliculatum and may better be considered as varieties
of it. P. virginicum appears to be a broad-leaved form of P.
biflorum. In Michigan another form is found which is analogous
to P. canaliculatum but the flowers are yellowish. In this group
it would be natural to look for P. parviflorum Dietr. (Convallaria
parviflora Poir.). Such descriptions (not the original) of this
species which the author has seen point unmistakably to some
species of the Glabrata notwithstanding it is said to have flowers of
the size of those of the Pubescentes. Probably the description was
drawn from an immature dried plant and the flowers had not yet
opened at the time of collection. Oftentimes the pressure exerted
252 FARWELL: MICHIGAN SPECIES OF POLYGONATUM
in course of pressing will burst open the buds so that they appear to
be matured flowers in the dried specimens. P. hirtum, a species
with glabrous leaves, but with the upper part of the stem and the
peduncles hispidly pubescent, would also be sought here but
nothing answering to the description of either has been found in
Michigan. No intermediate forms have come under the author’s
observation, i. e., none of the large-leaved, large-flowered forms
has had pubescent leaves or small flowers nor none of the small-
leaved, small-flowered forms has had glabrous leaves or large
flowers. There appears to be no intergradation between the
Pubescentes and the Glabrata; Walter’s P. biflorum is typical of
the latter and Pursh’s P. pubescens, of the former.
Key to the species
Leaves pubescent on the veins underneath; flowers
6—10 mm. in length; filaments filiform, papillate;
plants up to 4.5 dm. in height (Pubescentes).
Flowers with a yellowish white tube.
Leaves ovate. P. pubescens.
Leaves elliptical, cuneate. P. pubescens cuneatum.
Flowers green, leaves elliptical.
Leaves large, flowers dark green. P. boreale,
Leaves small, flowers light green. P. boreale australe.
Leaves glabrous; flowers 12-23 mm. in length, fila-
ments more or less flattened; plants taller and
stouter, 6-24 dm. in height (Glabrata).
Flowers yellowish, free part of filament smooth,
onger than “ng anthers
Stems cylindric
aves ee te. P. biflorum.
Leaves ovate- to oblong-elliptical, flowers
12-14 mm. long. P. biflorum virginicum.
Leaves ovate to lance-ovate, flowers 16-18
: P. biflorum ovatum.
Stems ancipital. P. ellipticum.
Flowers spriae filaments aeons or :
art shorter than the
Petunce two-flowered.
ves lance-ovate, flowers 14 mm. long. P. canaliculatum.
nee two- to eight-flowered.
Leaves ovate, flowers 12 mm. long. P. canaliculatum americanum.
lish e, fl 4 long. P. canaliculatum giganteum.
FARWELL: MICHIGAN SPECIES OF POLYGONATUM 252
- POLYGONATUM PUBESCENS (Willd.) Pursh, Fl. Am. Sept. 234.
1814
Convallaria pubescens Willd. Hort. Berol. 45. pl. 45. 1816.
P. multiflorum B americanum (in part) Hook. Flor. Bor. Amer.
2: 176. 1840.
P. biflorum Ell. in A. Gray, ont Ed 2, 466, 1856, mostly and
of subsequent authors. Not Walter in Elliott, Bot. S.C. & Ga.
02.308. 1817.
Salomonia biflora Farwell, Rep. Com. Parks Detroit 11: 53. 1900.
Stems glabrous, up to 4.5 dm. high, 2-4 mm. in diameter, leaf-
less part the longer; leaves nine to thirteen, ovate to lance-ovate,
17-33 mm. wide by 5-7.5 cm. long, amplexicaul, subsessile, or short-
petioled, green above, pubescent on the nerves below; three to
seven of the nerves more or less prominent; peduncles short, about
12 mm. long, one- or two-flowered; pedicels shorter than the
peduncle; flowers 8-10 mm. long by 2-3 mm. wide, occasionally
sessile, cylindrical, tube yellowish white or white in dry specimens;
the free part of the filament filiform, papillate, and shorter
than the anthers; mature fruit globular, 8-10 mm. in diameter;
seeds three to nine, 4:mm. wide. [PLATEs 12, 13, A.]
Generally in rather dry woods and copses of oak and maple.
MICHIGAN: Keweenaw Peninsula, June, 1886, Farwell 380;
Rochester, May, 1914, Farwell 3624. MASSACHUSETTS: Russell,
May, 1873, H. H. Rusby (distributed as P. biflorum). NEw
Jersey: Franklin, August, 1879, H. H. Rusby (likewise distributed
as P. biflorum).
v Polygonatum pubescens cuneatum (Greene) comb. nov.
P. cuneatum Greene, Leaflets 1: 181. 1906.
Differs from the species only in having longer and relatively
narrower leaves, cuneate at the base (18-35 mm. wide by 6-12
cm. long). [PLATE 13, B.]
MICHIGAN: near Marquette, Greene ; Detroit, May, 1895,
Farwell 380c; Keweenaw Peninsula, October, 1914, Farwell 3900.
POLYGONATUM BOREALE Greene, Leaflets 1: 181. 1906
Leaves larger, elliptical, 33-45 mm. wide by 7.5-10 cm. long,
generally sessile; flowers green, darker than the foliage, otherwise
as in P. pubescens. [PLATE 14, A.]
Copses of oak and maple. MINNESOTA: Winona, Greene.
254 FARWELL: MICHIGAN SPECIES OF POLYGONATUM
MicwiGAN: Keweenaw Peninsula, June, 1886, Farwell 380a;
Ypsilanti, May, 1891, Farwell 38ob.
‘ Polygonatum boreale australe var. nov.
Leaves small (10-22 mm. wide by 2.5—5 cm. in length), elliptical
or the lower inclined to ovate-elliptical; peduncles filiform, com-
pressed only below the arch, one-flowered, flowers water-green
(Ridgway’s nomenclature), paler than the foliage. [PLATE 14, B.]
Sparsely wooded fields. MuicHiGANn: Detroit, May, 1895,
Farwell 380d.
POLYGONATUM BIFLORUM Walt. in Elliott, Bot. S. C. & Ga. 1: 393.
1817
Convallaria biflora Walt. Fl. Car. 122. 1788.
Polygonatum angustifolium Pursh, Fl. Am. Sept. 234. 1814.
Polygonatum multiflorum 8B americanum (in part) Hook. Flor
Bor. Am. 2: 176. 1840.
Whole plant glabrous, with nine to thirteen, secund elliptical-
lanceolate leaves (18 mm. wide by 10-12.5 cm. long), rounded or
somewhat cuneately narrowed to a sessile base, and with three to
@frve more or less prominent nerves; peduncles slender, about 12-7,
mm. long, two-flowered; flowers yellow; stems about 4.5 dm. high
and 2-3 mm. indiameter. [PLATE 15, A.]
Atlantic seaboard, not typically represented in Michigan.
¥ Polygonatum biflorum virginicum (Greene) var. nov.
Polygonatum virginicum Greene, Leaflets 1: 181. 1906.
Salomonia commutata (in part) Farwell, Rep. Com. Parks Detroit
TI: $34. 1000:
Differing from the typical form of the species in its =a
ovate-elliptical to oblong-elliptical leaves, which are 25-55 m
wide by 10-18 cm. long, with fifteen to twenty-three of the nerves
more or less prominent; in its larger stems, 6-8 mm. in diameter,
6-9 dm. high, with the naked lower portion equal to or longer
than the upper leafy part; in its larger flowers which are funnel-
shaped to linear-cylindric (4-6 mm. wide by 14-25 mm. long, even
on the same plant), borne on longer (often three-flowered) pedun-
cles (25-37 mm. in length), with subequal pedicels 6-10 mm.
in length; fruit ovoid to globular, 12-16 mm. in diameter; seeds
eight to ten. [PLATE 15, B.|
FARWELL: MICHIGAN SPECIES OF POLYGONATUM 255
Moist grounds in woods or in shady places. VIRGINIA:
Greene. MIcuIGAN: Belle Isle, Detroit River, June, 1893, Farwell
1150b; Parkdale Farm, June, 1914, Farwell 3676, 3677, 3682,
3683. The most developed form of the species.
“ Polygonatum biflorum ovatum var. nov.
Salomonia commutata (in part) Farwell, Rep. Com. Parks Detroit
II: 53. 1900.
Differs from the above in having smaller ovate to elliptical
leaves (18-37 mm. wide by 7.5-10 cm. long), with fewer conspicu-
ous nerves (seven to nine) and short stems (4-6 mm. in diameter
and 4.5-6 dm. in length), often with a foliaceous, sheathing bract
10-12 cm. below the first leaf, and generally smaller flowers (4-8
mm. wide by 14-18 mm. long). [PLATE 16, A.
Open, rather dry woods and fields. MiIcHIGAN: Belle Isle,
Detroit River, June, 1893, Farwell 1150a; near Rochester, June,
1914, Farwell 3674, 3678, 3670.
“ Polygonatum ellipticum sp. nov.
Whole plant glabrous; stems about 9 dm. in height, the naked
lower portion (3.5 dm.) shorter than the leafy part (5.2 dm.) and
ancipital, the cross-section being elliptical in outline (6 X I0 mm.),
whence the specific name; leaves about twenty, ovate-lanceolate
(25-37 mm. wide by 7.5-10 cm. long), five to nine more or less
prominent nerves, semi-amplexicaul, sessile; peduncles slender, up
to 60 mm. in length with very unequal pedicels up to 25 mm.
from four-flowered in the lower to one-flowered in the uppermost;
flowers yellowish white, 16-20 mm. long by 4-5 mm. wide.
[PLATE 16, B.]
Dry fields and pastures. MICHIGAN: Rochester, June, 1914,
Farwell 3673.
POLYGONATUM CANALICULATUM (Muhl.) Pursh, Fl. Amer. Sept.
234. 1814; Kunth, Enumeration 5: 134. 1850
Conccilaria canatteulats Mahl. in Willd: Hak. Geel a5. 1816. F
Polygonatum multiflorum 8B americanum (in part) Hook. FI. Bor.
Am, 2: 176. 1840.
Whole plant glabrous; stems cylindrical, 6-10 mm. in diam-
eter, about 10 dm. high, equally divided between the naked lower
portion and the upper leafy part; leaves about fourteen, ovate to
elliptical, 30-55 mm. wide by 10-14 cm., with nine to thirty-one
more or less conspicuous nerves, semi-amplexicaul, subsessile or
ii
256 FARWELL: MICHIGAN SPECIES OF POLYGONATUM
sessile; peduncles up to 30 mm. in length, with equal pedicels 12
mm. or less, two-flowered; flowers green, broadly tubular, 12-14
mm. long by 4-6 mm. wide. [PLATE 17, A.|
Moist elder thickets. MiIcHIGAN: Rochester, June, 1914,
Farwell 367714.
v ‘Polygonatum canaliculatum americanum (Hook.) comb. nov.
Polygonatum latifolium Desf. in Pursh, Flor. Amer. Sept. 235.
1814. ,
Convallaria commutata J. A. & J. H. Scult. Syst. 7: 1671. 1830.
Polygonatum commutatum Dietr. in Otto & Dietr. Gartenz. 3:
27s. 5 VERE.
Polygonatum multiflorum B americanum (in part) Hook. FI. Bor.
Amer. 2: 176. 1840.
Polygonatum latifolium var. commutatum Baker, Jour. Linn. Soc.
Bot. 14: 555. 1875
Polygonatum biflorum commutatum Morong, Mem. Torrey Club
S835. 2806,
Stems 9-12 dm. in height, cylindrical, about 10 mm. in
diameter, only the lower third naked; leaves eighteen to twenty-
eight, ovate-lanceolate, 25-60 mm. wide by 7.5-14 cm., with nine
to thirty-one more or less conspicuous nerves, peduncles often 5 cm.
in length with subequal pedicels, 18 mm. or less, two- to eight-
flowered (generally six- or seven-flowered on the lower peduncles) ;
flowers 14 mm. long by 4 mm. wide. [PLATE 17, B.]
Low wet situations in open fields and thin woods. MICHIGAN:
Ypsilanti, June, 1893, Farwell 1150; near Rochester, June, 1914,
Farwell 3675. NeW JeRSEy: Franklin, August, 1875, H. H.
Rusby (distributed as P. gigantewm). ONTARIO: Walkerville,
June, 1891, A. L. Johnson (likewise distributed as P. giganteum).
¥ Polygonatum canaliculatum giganteum (Dietr.) comb. nov.
Convallaria multiflora L. in Michx. Fl. Bor. Amer. 1: 202. 1803.
Polygonatum multiflorum Desf. in Pursh, Fl. Amer. Sept. 234.
1814.
Polygonatum giganteum Dietr. in Otto & Dietr. Gartenz. 3: 222.
183
Polygonatum multiflorum B americanum (in part) Hook. FI. Bor.
Am. 2: 176. 1840.
Polygonatum biflorum y giganteum Wood, Bot. & Flo. 346. 1870.
FARWELL: MICHIGAN SPECIES OF POLYGONATUM 257
Stems 20 dm. or more in height, 18 mm. in diameter near the
base, the lower naked portion about 9 dm. high; upper portion
carrying about twenty-eight sessile or subsessile leaves, which are
round-ovate or oval (30-37 mm. wide by 60-90 mm. long), with
about fifty-three more or less conspicuous nerves; peduncles flat,
2 mm. wide by 90 mm. in length, rigid; pedicels 25 mm. or less,
subequal; flowers usually four (two to five), 14-16 mm. long by
4-6 mm. wide; fruit 20 mm. in diameter ripening fifteen seeds, 5
mm. wide. [PLATE 18.]
Moist elder thickets. MicHIGAN: Rochester, June, 1914, Far-
well 3473. The hyaline subulate bracts of the pedicels, which in
the other species are small (2 mm. long) and early deciduous, are
in this variety large (4-8 mm. long) and persistent, at least in
some Cases.
The author wishes to thank Miss Mary A. Day, of the Gray
Herbarium, most heartily for transcripts of the original descrip-
tions of Walter and Willdenow and for a sketch of Willdenow’s
plate of C. pubescens which is herewith reproduced for comparison;
for a like purpose a photograph of the narrow-leaved plant (P.
biflorum) collected by Dr. Rusby is also reproduced.
DEPARTMENT OF BOTANY,
ParKE, Davis & COMPANY,
DetrRoIT, MICHIGAN
258 FARWELL: MICHIGAN SPECIES OF POLYGONATUM
Explanation of plates 12-18
PLATE 12
Polygonatum pubescens (Willd.) Pursh. From photograph of Willdenow’s
plate 45 in Hort. Berol
PLATE 13
eDppeadeaae pubescens (Willd.) Pursh. Photographs of specimens from
Keweenaw Peninsula, Farwell 380, and from near Rochester, Farwell
ll 3624,
Michigan. B. Pore 'ygonatum pubescens cuneatum (Greene) Farwell. Photographs of
specimens from near Detroit, Farwell 380c, and from the Keweenaw Peninsula, Far-
well, 3909, Michigan.
PLATE 14
A. Polygonatum boreale Greene. Photo s of specimens from Keweenaw
Dupage Farwell 380a, and from near Sudha pane 3806, Michigan. B. Poly-
m boreale australe Farwell. Photograph of specimens from near Detroit,
oo. Farwell 480
PLATE 15
A. Polygonatum biflorum Walt. Photograph of a specimen from Franklin,
New Jersey, H. H. Rusby. B. Polygonatum biflorum virginicum (Greene) Farwell.
Photograph of specimens from Parkedale Farm, Michigan, Farwell 3682.
PLATE 1
Polygonatum biflorum ovatum Farwell. Photograph of specimens from near
Rochester, Michigan, Farwell 3674. B. Polygonatum ellipticum Farwell. Photo-
graph of specimens from near Rochester, Michigan, Farwell 3673.
PLATE 17
A. Polygonatum canaliculatum (Willd.) Pursh. Photograph of specimens from
Parkedale Farm, Michigan, Farwell 36777/2. B. Polygonatum canaliculatum ameri-
canum (Hook.) Farwell. Sides of specimens from near Rochester, Michigan,
Farwell 3675
PLATE 18
Polygonatum canaliculatum giganteum (Dietr.) Farwell. Photographs of speci-
mens from Parkedale Farm, Michigan, Farwell 3473.
-
The genus Plagiochasma and its North American species *
ALEXANDER W. EVANS
(WITH EIGHT TEXT FIGURES)
HISTORICAL INTRODUCTION
Of the various Marchantiaceae occurring in North America the
genus Plagiochasma has long been in especial need of critical study
and revision. It is hoped that the present paper may clear up some
of the confusion and misunderstanding regarding it, although the
writer admits that further work will be necessary before our knowl-
edge of the species can be considered at all complete. Prolonged
observation in the field under varying conditions of light and
moisture is especially to be recommended, while further collection
of material, particularly in fruiting condition, is much to be desired.
Fortunately dried specimens are more satisfactory for study than
in most genera of the thallose liverworts, even if they can never
take the place of material properly fixed and preserved.
The genus Plagiochasma Lehm. & Lindenb., as originally pro-
posed by its authors in 1832 (17, p. 13), contained two undescribed
species from Nepal, P. cordatum and P.appendiculatum. The first
of these should be regarded as the type of the genus. Four years
later Lehmann (16) published another genus, Anthrocephalus, which
was likewise based on an undescribed species from Nepal, A.
nepalensis. In 1838 Nees von Esenbeck (26, pp. 33, 40) accepted
both of these genera. He pointed out the fact, however, that
Aytonia rupestris Forst. was referable to Plagiochasma, and he
added a new species, P. peruvianum Nees & Mont., to this genus.
Aytonia rupestris, which was published in 1776, was based on
material collected by its authors on the island of Madeira, but
Nees von Esenbeck was able to extend its known range to the
island of Corfu, in Greece, citing specimens collected by Spathys.
Even at the time he wrote the species had received many names,
but he discarded them all and renamed the plant P. Aitonia
Lindenb. & Nees.
3 * Contribution from the Osborn Botanical Laboratory.
260 EvANs: THE GENUS PLAGIOCHASMA
Between 1838 and 1846 Sassi (27) proposed Anthrocephalus
italicus as a new species, his material having been collected near
Finale in Liguria, and the following species of Plagiochasma were
published: P. Rousselianum Mont. (23, p. 334, pl. 9, f. 1), of
western Algeria; P. chlorocarbum (Nees & Mont.) Mont. (24, p.
59), of Chile; and P. validum Bisch. (3, p. 56, pl. 56, f. 2753), also
of Chile. The number of species was thus increased to two in
Anthrocephalus and to seven in Plagiochasma.
In 1846 the part of the Synopsis Hepaticarum which treated
the Marchantieae- appeared (11, pp. 511-521) and the authors of
this important work showed that the genus Anthrocephalus was
based on poorly developed specimens of a Plagiochasma, which
they renamed P. Colsmannianum Lehm. & Gottsche, thus dis-
carding Lehmann’s specific name nepalensis altogether. For some
reason they make no mention of A. italicus, possibly because Sassi’s
work was unknown to them. In addition to P. Colsmannianum
and the seven species of Plagiochasma already known they include
the following species in the genus: P. ‘intermedium Lindenb. &
Gottsche, of Mexico; P. australe (Tayl.) Nees, of New Zealand;
P. limbatum (Tayl.) Neés, of Ascension Island; P. elongatum
Lindenb. & Gottsche, of Mexico; and P. mexicanum Lindenb. &
Gottsche, also of Mexico. They thus increased the total number
of species to thirteen.
In 1849 a posthumous work of Griffith was published in which
four news species of Plagiochasma from India were described (12,
PP. 330, 331, 337, pl. 65, 68, 69e), namely: P. paradoxum Griff.,
P. pedicellatum Griff., Anthrocephalus polycarpon Griff., and A.
megacarpon Griff. In 1856 Sullivant (38, p. 688, pl. 6) published
P. Wrightii, a new species from Texas. In 1859 DeNotaris (5,
p. 476, pl. 1) showed that Anthrocephalus italicus was clearly
a species of Plagiochasma and transferred it to the latter genus
under the name P. ttalicum. In 1863 Gottsche (10, p. 266)
described a new species of Plagiochasma, P. crenulatum, from
Mexico. In 1869 Austin (1, p. 229) described, as a new species,
P. erythrospermum Sulliv., from the Rocky Mountains.
In 1868 Lindberg (20, p. 291) revived the generic name
Aytonia of Forster and secured a number of followers in both
Europe and America. He spelled the name “ Aitonta,’”’ which was
Evans: THE GENUS PLAGIOCHASMA 261
perhaps justifiable, both spellings having been used by the original
authors. In 1874, Dumortier (7, p. 148) discarded both Plagio-
chasma and Aytonia and applied the name Otiona of Corda to the
genus. In 1877 Trevisan (39, p. 436) revived the name Rupinia
of the younger Linnaeus. In 1894 Le Jolis (19, pp. 109-116) gave
excellent reasons for discarding the names Aytonia, Otiona, and
Rupinia, and for retaining the name Plagiochasma. This course
is now advocated by both Stephani and Schiffner, who included
Plagtochasma in their proposed lists of nomina conservanda
presented at the International Botanical Congress of Brussels in
1910. The name Plagiochasma is therefore used in the present
paper.
In 1881 Leitgeb (18, p. 51) showed that P. Rousselianum
belonged in the genus Clevea, proposed by Lindberg in 1868, and
the species is now known as C. Rousseliana (Mont.) Leitg. In
1899 Howe (14, p. 39) showed that P. erythrospermum was a
synonym of Clevea hyalina (Sommerf.) Lindb. Between 1881
and 1898 the following species of Plagiochasma were published
(some of them under the name Aitonia): A. subplana Spruce and
A. lanigera Spruce (32, pp. 567, 568), both of Ecuador; A. extensa
Steph., of Australia (33, p. 17); A. eximia Schiffn., A. Fischeriana
Steph., and A. microcephala Steph. (34, pp. 300, 301) all three of
Africa; A. fissisquama Steph. of China (35, p. 209); A. japonica
Steph., of Japan (36, p. 54); and P. pterospermum Massal., of
China (21, p. 46, pl. 13, f. 18). The various species added to the
genus since the Synopsis Hepaticarum increased the total number
to twenty-eight.
In 1898 Stephani (37) published the last general account of
Plagiochasma in his Species Hepaticarum. With the exception of
Griffith’s species, three of which he mentions but does not describe
on account of the lack of available material for study, he recognizes
as valid all the species previously published. He intimates, how-
ever, that P. pierospermum is probably a synonym of P. fissi-
squamum. Besides transferring to Plagiochasma the various
species described under Aitonia he restores to P. Attonia and P.
Colsmannianum their original specific names and describes them
as P. rupestre and P. nepalense respectively. He likewise proposes
the following seven species as new: P. dschallanum, P. tenue, P.
262 EvANsS: THE GENUS PLAGIOCHASMA
algericum, P. Beccarianum, and P. Schimperi, all from Africa; P.
coerulescens Nadeaud, from Tahiti;‘and P. brasiliense, from
Brazil. Since 1898 the following new species of Plagiochasma (or
Aytonia) have been published: Ayionia Evansii Haynes and A.
jamaicensis Haynes, both from Jamaica (13); P.. abyssinicum
Gola, from Abyssinia (9, p. 62); and P. articulatum Kashyap,
from India (15, p. 320, f. 8). It should be noted further that
P. italicum was reduced to a synonym of P. rupestre by K. Miiller
in 1907 and that this reduction was sanctioned by Schiffner (25,
pp. 252, 254). In the review of the literature just given no allusion
is made to papers which simply record extensions of range of
individual species.
If, now, the four species of Griffith are ignored and the reduc-
tion of P. italicum to synonymy is accepted, the seven species
added by Stephani in 1898 and the four subsequently published
increase the total number of species of Plagiochasma to thirty-four.
Of these species P. rupestre has been recorded from various stations
in Europe, Asia, and Africa, and P. elongatum from both North
America and Asia, while the others are distributed as follows:
Asia, seven species; Africa, ten; North America, six; South Amer-
ica, six; Australia and the islands of the Pacific, three.
The genus is most at home in arid portions of the tropics,
many of the species growing on calcareous rocks. P. rupestre,
however, extends as far north as the Mediterranean portions of
Europe and P. australe (as already noted) was based on New
Zealand material. On account of the dry habitats the sperms
often fail to reach the archegonia, and even where fertilization
takes place the normal development of carpocephala and sporo-
phytes may be prevented by periods of drought. Immature and
abnormal carpocephala due to such causes have produced much of
the confusion in the interpretation of the genus and its various |
species.
MORPHOLOGY OF THE GENUS
Our knowledge of the morphology of Plagiochasma is largely
based on the work of Leitgeb (18, pp. 63-68, fl. 1), although later
writers have added important details. For a resumé of the work
done on the genus the Inter-Relationships of the Bryophyta by
Cavers may be consulted (4). Recent writers agree in placing
EvaANs: THE GENUS PLAGIOCHASMA 263
Plagiochasma in Leitgeb’s group Operculatae (or its equivalent) of
the Marchantiaceae. It represents the most simply organized
member of the group, but there is a strong tendency at the
present time to consider it reduced rather than primitive.
The species of Plagiochasma grow in depressed and intricate
mats. The thallus is remarkably uniform in structure throughout
the genus and shows clearly the usual differentiation into an
epidermis, a photosynthetic layer with intercellular spaces, and a
compact ventral layer, from the lower surface of which arise the
ventral scales and the rhizoids. The thallus is sometimes con-
tinuous in its growth, maintaining the same width, and sometimes
seems to be divided into a succession of joints, each joint springing
from the apex of an older joint and rapidly increasing in width.
_ According to Leitgeb this condition is not due to apical innovations
but simply to the fact that the growth of the thallus is subject to
interruptions, the new portions being narrower when the growth
is first resumed. According to Kashyap, however, true apical
innovations occur in P. articulatum. In addition to the usual
branching by dichotomy, in which both branches are alike, the
thallus often bears ventral adventive branches, which broaden out
rapidly from a stalk-like base. The authors of the Synopsis
Hepaticarum (11, pp. 511-521) supposed that these branches
were never produced by species in which the thallus was jointed
and they therefore divided the genus into two sections, as follows:
§ 1. Frons ex apice cordato articulatim innovans; § 2. Frons e
ventre innovans, apice saepe subcontinua. Leitgeb showed, how-
ever, that P. intermedium of § 1 often developed ventral branches,
that P. appendiculatum of § 2 often appeared jointed, and that
the distinction relied upon in the Synopsis was therefore inconstant
and to be used with caution.
The epidermis, as in most genera of the Marchantiaceae,
consists of a single layer of pale or colorless cells, although the cell-
walls are sometimes more or less pigmented with purple. The
walls are usually firm but thin and this is true even of the cuticle.
The latter, however, sometimes appears thickened, roughened, and
slightly opaque, owing to the deposition of a waxy substance on
the surface. When this is present the thallus is bluish or glaucous
green instead of bright green. Trigones can always be demon-
264 EVANS: THE GENUS PLAGIOCHASMA
strated in the epidermis and are often large and conspicuous, but
in certain species their size is very variable and depends on external
conditions.
The epidermal pores of the thallus as noted by Leitgeb, are
of two distinct types. In the first (Fic. 1) the opening is small
and often difficult to demonstrate, especially when more or less
stopped up by the waxy secretion on the cuticle. Around this small
opening there are usually from four to six bounding cells, the radial
walls between which are normally more or less thickened. Some-
times one or more of the bounding cells will be divided by a
FIG. I. PLAGIOCHASMA RUPESTRE (Forst.) Steph.
A series of epidermal pores from various specimens, X 300. A. Italy, D. Lanza,
in V. Schiffner’s Hep. Europ. Exsic. No. 10. B, C. Canary Islands, C. J. Pitard, in
Plant. Canar. No. 76 D. Baja California, T. S. Brandegee. E. Puebla, F. Lieb-
mann, type of P. mexicanum. F. Jalisco, Barnes & Land 123. G. Morelos, Barnes
& Land 480. H. Oaxaca, F. Liebmann, type of P. elongatum. 1. Jalisco, Barnes &
Land 134. J-L. Arizona, G. E. Nichols, the last figure drawn from material culti-
vated in the laboratory.
Evans: THE GENUS PLAGIOCHASMA 265
periclinal wall (with respect to the pore), and the opening under
such circumstances may be described as surrounded by two more
or less complete concentric series of cells. The cells surrounding
the openings are not elevated and are usually much like the other
epidermal cells. When the radial walls between them are slightly
or not at all thickened, as is sometimes the case in certain species,
the entire pore-structure is very inconspicuous.
In pores of the second type (Fics. 5-8, A), although the
opening itself is not necessarily larger than in pores of the first
type, the whole structure is much more evident because it is more
or less raised above the general level of the epidermis and because
the cells surrounding the opening are in two or more concentric
series with six to eight or even more cells in each series. It is
convenient to describe these cells, which differ considerably in
form from the other epidermal cells, as being arranged in radiating
series with two or more cells in each series. In many cases the
walls separating these are distinctly thickened. Immediately
bounding the opening a circular ridge is present which appears
to be formed by the thickened and coalescent inner walls of the
surrounding cells. Deutsch (6, p. 494, f. 9), however, has recently
shown that in the very similar pores of Targionia hypophylla L.
the ridge consists of a circular row of dead and collapsed cells with
thickened and highly cutinized walls. Apparently his explanation
would apply equally well to the pores of the second type in
Plagiochasma, and it is sometimes possible to demonstrate the
radial walls between the collapsed cells. The presence of this
circular ridge in the second type of pore and its absence from the
first type is perhaps the most important difference between them.
It will be noted when the pores of Plagiochasma are compared
with those of other genera of the Marchantiaceae that those of the
first type are essentially like the pores found in Oxymitra and in
the various genera of the Astroporae, while those of the second
type find their counterparts in the other genera of the Operculatae,
as well as in certain genera of other groups. Both types stand in
marked contrast to the complex pores of Marchantia, Preissia,
and Bucegia, where the cells surrounding the opening are in
several layers. Even in Plagiochasma, however, complex pores
of the marchantiaceous type are present on the carpocephala.
266 EvaANs: THE GENUS PLAGIOCHASMA
The photosynthetic tissue contains several layers of inter-
cellular spaces, separated from one another by plates of green cells
a single cell in thickness. These spaces, which vary greatly in
size in different species, are connected with one another by means
of holes through the green partitions, and some of them com-
municate directly with the outside air by means of the epidermal
pores. According to Leitgeb (18, p. 64) the thallus first forms
primary intercellular spaces or air chambers, each of which has its
pore, and these chambers afterwards become divided up by
secondary plates of cells which grow out from the primary parti-
tions. Barnes and Land (2, pp. 210-213, f. 77-22) admit that
such a partitioning of chambers in Plagiochasma may perhaps
occur to a limited extent, but they show clearly that some of the
air-spaces without epidermal pores are formed directly, in very
young segments, by the splitting apart of cells originally united.
The compact ventral tissue is not highly developed. In the
median portion of the thallus it attains a thickness of perhaps ten
to twenty cells, but it thins out rapidly on each side until it
becomes only one cell thick and forms the ventral walls of the
air-chambers. It is composed of thin-walled cells, a few of which
contain oil-bodies, but it shows no signs of the slime cavities or
the fibrous cells which occur in some of the more complicated
Marchantiaceae. Sometimes, but not always, mycorrhiza is
present in the ventral tissue, occupying cells in the median portion
just below the green tissue. The cells containing the mycorrhiza
sometimes have their walls pigmented with purple, but this is not
invariably the case.
The rhizoids are of the two types usually found in the Mar-
chantiales and require no especial comment. The ventral scales,
however, which are in two regular rows, are unusually well
developed and exhibit considerable variety in form and structure, .
sometimes even in a single species (see Fics. 2,3). They show
the characteristic division into basal portion and appendage or
appendages. The basal portion is ovate to lunulate in outline and
is attached by an oblique line extending forward from the median -
region half way to the margin or beyond. In the apical part it
narrows gradually or abruptly into the appendage or appendages,
the number of these structures often varying from one to three on
EvANS: THE GENUS PLAGIOCHASMA 267
a single thallus. The appendages vary in form from narrowly
lanceolate to orbicular, when the genus as a whole is taken into
account, and they likewise vary markedly in size. As in other
genera of the Marchantiaceae the appendages are the first part of
Bs
Fic. 2. PLAGIOCHASMA RUPESTRE (Forst.) Steph.
A series of ventral scales (or their lag
parts of Europe and the Atlantic Islands. A, B. er Lanza, in V. Schiffner’s s
Hep. Europ. Exsic. No. 10. C, D. Madeira oo W. Trelease, X 40. E-G.
anary Islands, O. Kuntze, X27. H. From the same specimen, X 17. I-L.
upg te Islands, C. J. Pitard, in Plant. Canar. No. 762, X 40. M. Ascension Island,
er 22, type of P. limbatum, x 27. N. From the same specimen, ‘
the scale to develop, covering over the embryonic region at the
growing point and being gradually pushed into a ventral position
by the apical elongation of the thallus. The basal portion and
the appendages are usually deeply pigmented with purple, but the
color of the appendages in particular often becomes bleached
268 EvANs: THE GENUS PLAGIOCHASMA
Fic. 3. PLAGIOCHASMA RUPESTRE (Forst.) Steph.
A series of ventral — (or their appendages) from various North American
specim A-F. Arizona, G. E. Nichols, X 40, the last three figures drawn from
ake aed in rs tines G, H. Arizona, W. A. Cannon, X 40. I, J.
Baja California, T. S. Brandegee, X 40. K,L. Morelos, C. G. Pringle 15310, X 40.
M-O. Jalisco, Barnes & Land 123, X17. P. Morelos, Barnes & Land 480, X 40.
Q. From the same specimen, X17. R. Oaxaca, F. Liebmann, type of P. elongatum,
X 40.
EvaANsS: THE GENUS PLAGIOCHASMA 269
out with age. The scales are a single cell in thickness except near
the base. The cell-structure is sometimes uniform throughout,
but the marginal cells are sometimes irregular and project as
indistinct teeth bearing slime papillae or their vestiges (Fic.
5, D). The cells containing oil-bodies, on account of their small
size and lack of pigmentation, form a striking feature of the
scales. They are sometimes confined to the basal portion and
sometimes occur in the appendages as well.
With regard to the inflorescence certain species of Plagio-
chasma are described as dioicous and others as monoicous, while
some are said to be either dioicous or monoicous. Unless the
male and female plants grew in separate mats, a dioicous inflores-
cence would be difficult to demonstrate beyond doubt on account
of the fact that the androecia are often borne on ventral branches,
which easily become separated and present the appearance of
being distinct plants. It is therefore probable that some of the
so-called dioicous species are really monoicous. Stephani (37,
Pp. 777) mentions an interesting case which shows how variable
the inflorescence can be and how easily a hasty observation could
lead to error. In the thallus he described he found five joints,
the first, second, and fourth of which were male, the third and
fifth female. Each joint bore ventral branches, the male joints
in most cases bearing female branches and the female joints male
branches. Among the ventral branches he found further certain
ones which bore both male and female inflorescences in succession
and others which forked, one fork being male and the other female.
Schiffner (28, p. 15) has described material of P. italicum in
which the inflorescence was equally variable. In both these cases
the inflorescence might properly be described as heteroicous, and
this term would probably apply to most monoicous species and to
many which are supposed to be dioicous.
The antheridia, as in most of the other Marchantiaceae, are
grouped in definite discoid inflorescences or androecia, slightly
raised above the dorsal surface of the thallus and apparently
not affecting its subsequent growth. The androecia vary in out-
line from circular to more or less crescentic or cordate, with the
convex side turned toward the base of the thallus. These cres-
centic androecia, as Leitgeb notes, should not be confused with the
270 EvANS: THE GENUS PLAGIOCHASMA
dichotomous androecia occasionally found in Lunularia, where
two growing points are involved (18, p. 26). They represent,
according to his ideas, a persistence of the two-lobed condition
at the apex of an ordinary thallus, where the single growing point
lies in the indentation between the lobes. Kashyap (15, pp. 318,
321), however, says that in both P. appendiculatum and P. articu-
latum the androecium usually has two growing points and implies
that the horseshoe shape is therefore due to a forking. Un-
fortunately, he presents no conclusive evidence to support his
position. The antheridia are of the usual marchantiaceous type.
Each one lies at the bottom of a deep depression, the roof of which
forms a distinct bluntly conical ostiole with a minute opening at
the tip. Between the ostioles the epidermis of the androecium
shows numerous pores and these, as recently demonstrated by
Kashyap, are of the same type as in the vegetative thallus. In
cases where the normal pores are complex and show many radiating
series of cells with several cells in each series, the androecial pores
tend to be more simple and to show fewer series of cells and fewer
cells in each series. A row of delicate and slender scales, usually
pigmented with purple, surrounds the androecium.
The carpocephala arise singly or in short median rows from
the dorsal surface of the thallus and apparently in most cases
neither stop nor interrupt its further elongation. Each carpo-
cephalu mcommonly bears three or four archegonia evenly dis-
tributed around its margin (Leitgeb, 18, p. 29) although in P.
appendiculatum, according to Kashyap, the number is frequently
five or six and may be as high as nine. Through the extensive
growth of the upper median portion and of the parts adjacent to
the archegonia, the latter are displaced to the lower part of the
carpocephalum and each one eventually lies at the bottom of a
groove which is bounded by two folds of tissue. In case fertiliza-
tion has taken place the grooves become much deeper through the
continued growth of the folds, the edges of which come together
and grow inwards toward the developing sporophyte, as originally
described and figured by De Notaris in the case of P. rupestre
(5, p. 480, pl. 1, f. 20-26); where the folds meet at their upper
ends they form a more or less distinct apiculum or horn. No
pseudoperianths of any sort are developed. Around the margin
EvANS: THE GENUS PLAGIOCHASMA o7t
of the carpocephalum a circle of slender scales (see Fics. 4, E;
5, E-G; etc.) makes its appearance very early; these scales,
according to Leitgeb (18, p. 67), are in two series and they show
the same purple pigmentation as the androecial scales, becoming
bleached out with age. When the capsules have reached maturity
the region between the carpocephalum and the thallus undergoes
intercalary growth and elongates into a stalk, some of the scales
remaining at the base and some being carried up with the carpo-
cephalum. The length of the stalk varies greatly in different
species and even in the same species under different conditions.
Sometimes it is so short that it scarcely raises the carpocephalum
above the thallus. The stalk is remarkable on account of the
fact that it possesses no rhizoid-furrow, although it may be ir-
regularly grooved.
With regard to the siocpholaey of the carpocephalum in
Plagiochasma and in Clevea, where similar conditions obtain, the
opinions of writers vary. Leitgeb (18, p. 29) considered the whole
structure a highly developed dorsal outgrowth of the thallus,
comparable with the low outgrowths found in Corsinia. He based
his con@eption on the continued growth of the thallus after the
initiation of the carpocephalum and on the lack of a rhizoid-
furrow in the stalk, both indicating that the growing point of the
thallus was not involved. He therefore considered that the carpo-
cephalum in these two genera was morphologically different from
the very similar carpocephalum found in Reboulia, Sauteria, and
more complex forms, where the initiation of the carpocephalum
brought the growth of the thallus to an end and where the stalk
showed one or two rhizoid-furrows. In these cases he considered
that the carpocephalum represented a modified shoot (or shoot-
system) in which the growing point of the thallus was directly
involved. Leitgeb’s views regarding the androecium were very
similar. Not only in Plagiochasma but also in Reboulia and other
genera, where the development of the androecium failed to stop
the growth of the thallus, he considered the structure a mere
dorsal outgrowth and therefore morphologically different from
the androecium in Preissia and related genera, where it represented
a modified shoot.
Goebel (8, p. 46) has recently pointed out that these ideas
Zi2 Evans: THE’ GENUS PLAGIOCHASMA
involve certain difficulties. He emphasizes the improbability of
having the male and female inflorescences morphologically dif-
ferent in a single genus, such as Reboulia, or in a group of related
genera, and he brings out the fact that if the carpocephalum of
Plagiochasma is really a dorsal outgrowth, as Leitgeb claimed, then
its scales would have to have a different significance from the
very similar scales found on a carpocephalum which represents a
branch system. In the latter case the scales are modified ventral
scales, homologous with those found on an ordinary thallus. In
Plagiochasma, however, the scales if considered homologous with
the ventral scales would imply an inversion of the dorsi-ventrality
of the thallus to account for their dorsal position; if considered
morphologically different from the ventral scales, they would have
to be regarded as developed from structures normally present on
the dorsal surface, such as the hairs found in certain species of
Riccia, or the slime-papillae which occur in the cupules of Mar-
chantia. Neither of these interpretations seems plausible.
Goebel shows further (8, pp. 85-88) that these difficulties of
interpretation are avoided if such genera as Reboulia and Plagio-
chasma are considered as reduced from the more complex Mar-
chantiaceae rather than as primitive forms from which the more
complex genera have been evolved. In Marchantia, for example,
where the complexity perhaps reaches its highest expression, both
the androecium and the carpocephalum represent stalked branch-
systems, the sexual organs arising in acropetal succession on the
evident branches. A first indication of reduction appears when
the branches lose their distinctness although the acropetal succes-
sion of the sexual organs remains apparent. This is seen in the
androecium of Preissia. A second indication is a reduced develop-
ment of the stalk, leading to its final disappearance, as seen in the
androecia of Conocephalum and its allies. A third step in the
reduction, according to Goebel’s account, is a change in the posi-
tion of the inflorescence, which is perhaps caused by a very early
appearance of an apical innovation. This may occur even in
inflorescences which retain their stalks. In many cases such an
innovation is not started until the inflorescence is already estab-
lished, when it shows its character clearly as a new shoot broaden-
ing out from a narrow base. Such a condition is evident in the
EVANS: THE GENUS PLAGIOCHASMA 213
carpocephalum of Reboulia and also in the curious Plagiochasma
articulatum, according to Kashyap’s account (15, p. 320). If,
however, the innovation starts much earlier it might seem to
continue the growth of the thallus without an interruption and
thus to displace the inflorescence to an apparently dorsal position.
The latter would still represent a modified shoot, the scales would
still be homologous with ventral scales, and a thallus which showed
a median dorsal row of such inflorescences would really be a
sympodium. The change in position just described is clearly seen
in the androecium of Reboulia and in both the male and female
inflorescences of most species of Plagiochasma. In the case of the
androecium of Bucegia, recently investigated by Schiffner (30),
all gradations in the process have been demonstrated.
It is perhaps not necessary to assume that the carpocephalum
in Plagiochasma actually involves the growing point of the thallus
and that a new growing point must be differentiated before the
growth can be continued. If the important part played by
intercalary growth in the development of the carpocephalum is
taken into account such an assumption becomes superfluous.
Even in such complex carpocephala as those of Marchantia the
growth of the stalk and of the upper part leading to the displace-
ment of the archegonia to the lower surface is intercalary in
character. In Reboulia, after the earliest rudiment of the carpo-
cephalum is laid down in the young segments derived from the
growing point, the later development is entirely intercalary, as
Leitgeb himself admits (18, p. 31), and the growing point, carried
upward by the elongating stalk, plays no part whatever in the
process. . In Plagiochasma the first beginning of the carpocephalum
is developed from young segments just as in Reboulia, and from
this rudiment the mature structure develops by intercalary growth
in the same way. The mere fact that the growing point is not
carried upward by the elongating stalk but continues directly
the growth of the thallus does not invalidate the homology of this
carpocephalum with that of Reboulia. In both cases the structure
owes its initial stages to the activity of the growing point of the
thallus and its later stages to intercalary growth, and in neither
case does the growing point play any active part in these later
stages. Of course what is here said about the carpocephalum
would apply in most respects to the androecium as well.
274 Evans: THE GENUS PLAGIOCHASMA
In case the growing point of the thallus is not used up in the
development of the inflorescence the latter becomes a lateral or
dorsal branch, and the shoot which bears a series of such in-
florescences would be a monopodium and not a sympodium.
Such a change from a sympodium to a monopodium would not
be unique. Goebel (8, p. 91) quotes similar examples from the
inflorescences of the Boraginaceae and from the vegetative shoots
of the Vitaceae. And the further reduction from such a mono-
podium through Corsinia to Riccia, where the sexual organs are
not in distinct inflorescences and where the intercalary growth
so prominent in the carpocephala of Reboulia and Plagiochasma
has become largely or wholly eliminated, would be easily intel-
ligible.
In classifying the species of Plagiochasma Stephani (37) lays
a great deal of stress upon the shape of the carpocephalum. He
divides the genus into two groups, “a’’ and ‘‘d.”” In “a’’ the
carpocephalum is said to be more or less convex; in ‘‘d, ’? more or
less excavated at the vertex. From the writer’s observations this
distinction is not constant. In at least one species which Stephani
places in division ‘‘a,’’ namely P. japonicum, the carpocephala
examined were distinctly concave, and even if this condition
should be proved exceptional its occasional presence would in-
validate Stephani’s classification. It is quite possible that the
form of the carpocephalum is influenced by external condition,
although no experimental evidence can be brought forward to
support this idea.
The sporophyte of Plagiochasma is described in its essential
features by Leitgeb (18, p. 67). He calls attention to its well-
developed foot and scarcely evident stalk and to the fact that the
wall of the capsule consists of a single layer of cells except in the
apical portion, which shows a circular region three cells thick,
This he regards as a rudimentary lid although it does not separate
at maturity as a typical lid would do. The capsule, in fact, de-
hisces irregularly and leaves behind a shallow urn with indefinite
teeth or lobes around the rim. The cells of the capsule wall are
destitute of the annular or half-annular thickenings found in so
many of the Hepaticae. Some of them show more or less distinct
trigones, but many of them are thin-walled throughout. The
Evans: THE GENUS PLAGIOCHASMA 275
very rudimentary lid was apparently considered by Leitgeb a
sufficient reason for placing Plagiochasma among his Operculatae.
Solms-Laubach (29, p. 12) has shown, however, that in Leitgeb’s
group Astroporae an equally definite lid is developed but that it is
easily overlooked on account of the thickenings in the cells of the
capsule wall. In‘spite of this fact the lack of annular thickenings
in the capsule wall and the structure of the photosynthetic layer
of the thallus would seem sufficient to exclude Plagiochasma from
the Astroporae and to place it in the same group as Reboulia,
Grimaldia, and Neesiella, whatever this group may be called.
Stephani (37, p. 776) describes the spores as large, yellowish,
tetrahedral, and loosely reticulate on the convex face with high
and rough lamellae. The tetrahedral form is of course due to the
original arrangement of the spores in the tetrad resulting from the
division of the spore mother cell. One face of each spore, the
spherical or convex face, represents a fourth of the original surface
of the tetrad (see Fics. 4, F; 5,H; 7,H; 8,G). The other three
faces, the plane faces, represent the regions of contact with the
three other spores of the tetrad (see Fics. 4,G; 8,H). Through-
out the genus a broad border is formed where the three plane faces
meet the spherical face, and three narrow ridges are developed
along the edges of the pyramid formed by the three plane faces.
In order to gain a clear idea of the reticulum described by Stephani
it is advisable to pass in review a large series of spores and to select
for study those which show the spherical face plainly and those in
which the apex of the pyramid is turned toward the eye of the
spectator. If this is done the meshes of the reticulum will be ob-
served in their most diagrammatic position, and it will be seen
that the reticulum extends over both spherical and plane faces.
A regular reticulum has been observed by the writer in all the
the species examined. Sometimes, however, in some or all of the
spores in a capsule, the reticulum will be incomplete owing to the
partial development of the bounding ridges. Since this condition
occurs in species where a normal reticulum is usually developed
it can not be used as a distinguishing peculiarity. The spores of
Plagiochasma, in fact, in contrast to many genera of the Hepaticae,
yield no very helpful differential characters in separating species.
The elaters in the majority of cases are like those found in other
276 Evans: THE GENUS PLAGIOCHASMA
Hepaticae and consist of long tapering cells with thin walls strength-
ened by from two to four spiral bands of thickening. Two in-
teresting modifications, however, should be noted. In the first,
the spiral bands, which are entirely free from one another in most
elaters, are coalescent by means of one or more longitudinal bands
(Fic. 5, I). In the second, no spirals at all are present but
the wall of the elater is uniformly thickened and the cavity re-
duced to a narrow median canal (Fic. 8,1). Coalescence of the
type found in the first modification seems to be unusual. It was
first detected by Miyake (22, p. 223, pl. 3, f. 3) in the remarkable
Makinoa crispata (Steph.) Miyake, of Japan, but it has since been
observed by Schiffner (29, p. 24) in Neesiella rupestris (Nees)
Schiffn., a widely distributed species of Europe and North Ameri-
ca, and probably occurs in other genera. The second modifi-
cation is apparently confined to the genus Plagiochasma. It was
first noted by Gottsche in P. intermedium (10, p. 264) and is
mentioned by Stephani as one of the characteristic features of
P. japonicum (36, p. 84). Its occurrence has likewise been re-
corded by Kashyap (15, p. 320) in the case of P. appendiculatum
and P. articulatum, and the writer has observed it in P. eximium.
Although the uniform thickening of the elater-wall has been con-
sidered a good specific character, it is not a constant feature and
must be used with caution. Even in P. intermedium, where it is
certainly the usual condition, Leitgeb (18, p. 68) failed to find
it in the material which he examined. In the case of P. japonicum
Massalongo (21, p. 48, pl. 13, f. 19) has described and figured a
var. chinense, based on Chinese specimens collected by Father
Giraldi, in which some of the elaters showed rudimentary spiral
bands, although most were of the more solid type. Miss Haynes,
also, in a series of unpublished drawings based on Japanese
material of the same species, collected by T. Yoshinaga, has
represented elaters in which poorly developed and irregular bands
were present. In P. eximium some of the elaters show spiral
bands and others uniform thickenings and there are various
gradations between them. In P. appendiculatum the solid type
is said to be less common than in P. articulatum, and Kashyap
associates it with exposure to dryness during the development of
the capsule. In spite of these variations there are certain species
EvANS: THE GENUS PLAGIOCHASMA OTF
in which a uniform thickening has never been observed, so that
it is allowable to associate this peculiarity with certain species,
even if they may sometimes develop elaters of the usual type.
DESCRIPTION OF THE NORTH AMERICAN SPECIES
The seven North American species with their known ranges,
according to published records, are the following: P. crenulatum,
Mexico; P. elongatum, Mexico and China; Aytonia Evansit,
Jamaica; P. intermedium, Mexico and Guatemala; Aytonia
jamaicensis, Jamaica; P. mexicanum, Mexico; and P. Wrightw,
Texas. In the writer’s opinion, however, P. elongatum, A ytonia
Evansii, and P. mexicanum represent synonyms of the widely
distributed P. rupestre. This reduction to synonymy would
diminish the total number of North American species to five, if
it were not for a Mexican species proposed as new, which raises
the number to six. In certain cases it has proved possible to
extend the known ranges of individual species to a considerable
extent. The six species recognized in the present paper may be
distinguished as follows:
Key to the North American species
Epidermal pores of the first type (see p. 264), incon-
spicuous, not elevated, usually surrounded by from
four to six cells. 1. P. rupesire (p. 277).
Epidermal pores of the second type (see p. 265), con-
spicuous, at usually surrounded by six o
more is
Elaters a spiral bands of thickening.
Appendages of ventral scales almost as long as
the scales, ovate to orbicular, abruptly acute
or apiculate. 2. P. crenulatum (p. 288).
Appendages of es ah ee wee
the scales, ovate, apiculate. 3. P. jamaicense (p. 292).
Appendages of ventral scales lanceolate to
ovate, acute or obtuse. 4. P. Wrightii (p. 294).
Sapo to ri of ventral scales subulate, acu-
5. P. Landii (p. 298).
toe pone e with uniformly thickened walls
showing no signs of spiral bands of thickening. 6. P. intermedium (p. 301).
1. PLAGIOCHASMA RUPESTRE (Forst.) Steph.
A ytonia rupestris Forst. Char. Gen. Plant. 147. pl. 74. 1776.
Rupinia lichenoides Linn. f. Suppl. Plant. oe Veg. Ed. 13, 69.
1781.
278 Evans: THE GENUS PLAGIOCHASMA
Rupinia rupestris Swartz, Meth. Musc. 39. 1781.
Reboulia maderensis,Raddi, Gior. Sci. Lett. Arti. 1821.
Corsinia lamellosa Nees & Bisch. Flora 13: 401. 1830.
Sedgwickia hemisphaerica Bisch. Nova Acta Acad. Caes.-Leop.
Carol. 17: 1079. pl. 70, f. 4. 1835. Not Bowdich.
Plagiochasma Aitonia Lindenb. & Nees; Nees von Esenbeck,
Naturg. Europ. Leberm. 4: 41. 1838.
' Anthrocephalus italicus Sassi, Atti Prim. Riun. Sci. Pisa 160. 1840.
Jungermannia (Fegatella) australis Tayl.; Hook. f. & Tayl. Jour.
Bot. 3: 572. 1844.
PRN (Fegatella) limbata Tayl.; Hook. f. & Tayl. 1. c. 4:
8
45.
Plagiochasma australe Nees; G. L. & N. Syn. Hep. 515. 1846.
Plagiochasma limbatum Nees; G. L. & N. 1. ¢. 516. 1846.
Plagiochasma elongatum Lindenb. & Gottsche; G. L. & N. 1. c. 519.
1846.
184
Plagiochasma mexicanum Lindenb. & Gottsche; G. L. & N. lL. c.
519. 1846.
Plagiochasma italicum De Not. Mem. Real. Accad. Sci. Torino IT.
18: 476. pl. r. 1859.
Aitonia ttalica Lindb. Not. F. et Fl. Fenn. g: 291 (footnote).
1868.
Otionia rupestris Dumort. Hep. Europ. 148. 1874.
Otionia ttalica Dumort. |. c. 149. 1874.
Rupinia italica Trevis. Rend. Ist. Lomb. II. 7: 785. 1874.
Rupinia rupestris Trevis. Mem. R. Ist. Lomb. III. 4: 437. 1877.
Rupinia mexicana Trevis. l. ¢. 437. 1877.
Rupinia elongata Trevis. 1. c. 437. 1877.
Rupinia limbata Trevis. 1. c. 437. 1877.
Rupinia australis Trevis. 1. ¢. 437. 1877.
Aitonia lanigera Spreee Trans. Bot. Soc. Edinburgh 15: 568.
1885.
Aitonia australis Steph. Hedwigia 28: 129. 1889.
Aytonia elongata Underw. Bot. Gaz. 20: 66. 1895.
Aytonia mexicana Underw. l. c. 66. 1895.
Plagiochasma rupestre Steph. Bull. Herb. Boissier 6: 783. 1808.
Plagiochasma lanigerum Steph. 1. c. 188. 1898.
Aytonia Evansit Haynes, Bull. Torrey Club. 34: 57. pl. 5. 1907.
EvANs: THE GENUS PLAGIOCHASMA 279
Thallus pale green and glaucous above with a narrow purple
border, plane or somewhat canaliculate, strap-shaped, sometimes
forking, sometimes innovating at the apex or with ventral branches,
margin undulate, scarcely or not at all crispate, mostly 1-2 cm.
long, and 5—7 mm. wide, thickness of thallus about one tenth the
width; epidermis composed of thin-walled cells with small trigones
and a thin minutely roughened cuticle with a waxy deposit, the cells
mostly 18-30 w in diameter; pores not elevated, small, surrounded
by four to six cells in a single series or in an incomplete double
series, radial walls usually distinctly thickened; aeriferous layer
about one half the thickness of the thallus in the median portion,
fairly compact with the air-spaces a little wider than the green
cells, tending to extend at right angles to the epidermis; ventral
scales deep purple, obliquely lunate to ovate, entire, with a few
scattered cells containing oil-bodies, gradually narrowed into one
or two (very rarely three) ovate-lanceolate to lanceolate or subu-
late appendages, acute to acuminate at the apex, slightly or not
at all constricted at the base, entire: inflorescence monoicous:
o inflorescence on a small ventral branch or on an ordinary
branch: 9 receptacles borne singly or in a median longitudinal
series; peduncle short, mostly 3 mm. long or less; carpocephalum
concave at the apex, with two or three apiculate lobes, maturing
two or three sporophytes; scales of carpocephalum abundant,
narrowly subulate with elongated filiform points, entire: spores
400 » in length and about 10 uw in diameter, with two to four spirals,
sometimes more or less coalescent. [F1Gs. 1-4.]
The following specimens have been examined:
NEw Mexico: Organ Mountains, Filmore County, altitude
1800 m., July, 1897, E. O. Wooton.
ARIZONA: without definite locality or date, H. H. Rusby; Santa
Catalina Mountains, April, 1913, W. A. Cannon; Soldier’s Canyon,
Santa Catalina Mountains, September, 1913, G. E. Nichols.
BayA CALIFORNIA: without definite locality or date, T. S.
Brandegee 25. :
DuRANGO: Tejamén, August, 1906, E. Palmer 473, 474; city
of Durango and vicinity, 1896, E. Palmer 709 (mixed with Tar-
gionia hypophylla L.).
JALIsco: wet banks and rocks, Barranca de Oblatos, Guadala-
jara, September, 1908, Barnes & Land 122, 123, 134.
280 - EvANs: THE GENUS PLAGIOCHASMA
MoreELos: San Antonio Falls, Cuernavaca, October, 1908,
Barnes & Land 480; near Cuernavaca, July, 1908, C. G. Pringle
15310.
Oaxaca: Comaltepec, Chuapam, F. Liebmann (type of P.
elongatum); Barranca San Luis, April, 1906, C. Conzatti 1619.
PUEBLA: San Baltanzan, near Puebla, May, 1909, J. Nicolas
2; San Lorenzo Tehuacan, F. Liebmann (type of P. mexicanum).
SAN Luts Potos!: near San Luis, 1878, Parry & Palmer ro16.
ZACATECAS: near Conception del Oro, August, 1904, E. Palmer
307.
JAMAICA: on banks near Portland Gap, Blue Mountains,
July, 1903, A. W. Evans 213 (type of Aytonia Evansii); on banks
and rocks near Content Gap, September, 1906, A. W. Evans 613.
BraziL: Rio Janeiro, April, 1875, Glaziou 7402 (listed by
Spruce as ‘ Fimbriaria sp.”’ in Rev. Bryol. 15: 34. 1888); Rio
Grande de Sul, August, 1905, A. Bornmiiller 4994 (distributed
under a manuscript name of Stephani).
Ecuapor: near Bajios, Andes, altitude 2,000 m., R. Spruce
(distributed in Hepaticae Spruceanae as ‘‘ Aitonia lanigera’’ and
probably representing a part of the original material).
GALAPAGOS IsLaNDs: Abingdon Island, September, 1906, A.
Stewart 8676; Charles Island, October, 1906, A. Stewart 940.
Peru: Ollantaytambo, July, 1911, H. W. Foote (listed by the
writer as “P. chlorocarpum”’ in Trans. Conn. Acad. 18: 298.
1914).
Bo.tvia: near Mapiri, January, 1893, M. Bang 1748 (listed
by Rusby as ‘“Aitonia valida” in Mem. Torrey Club 6: 129.
1896); Pelichuco, altitude 3350 m., May, 1902, R. S. Williams
2728; lower Pelichuco Rio, altitude 900 m., April, 1902, R. 5S.
Williams 2738.
ARGENTINA: Sierra de Ventana, January, 1892, R. Hanthal 67.
CuILE: Desert of Atacama, September—October, 1890, 7.
Morong.
ITaLy: on walls near Voltri, Liguria, J. Baglietto (distributed in
Rabenhorst’s Hep. Europ. No. 85, as P. ttalicum); near Palermo,
Sicily, D. Lanza (distributed from specimens cultivated at Prague,
Austria, in Schiffner’s Hep. Europ. Exsic. No. 10, as P. italicum).
MaApvEIRA ISLANDS: without definite localities or dates, G.
EvaANs: THE GENUS PLAGIOCHASMA ae
Raddi (type of Reboulia maderensis), G. Don; near Funchal,
1858, Johnson; Como do Lobos, August, 1896, W. Trelease.
Canary IsLANDs: Palma, 1815, C. Schmidt; February, 1888,
O. Kuntze; Bocco de Bufadero, Teneriffe, January, 1906, C. J.
Pitard (distributed in Plantae canarienses, No. 762).
Care VERDE IsLANDs: Green Mountains, St. Vincent, date
and collector unknown.
ASCENSION IsLAND: without definite locality or date, J. D.
Hooker 22 (type of Jungermannia | Fegatella| limbata).
St. HELENA: without definite locality or date, Lieut. Houghton
33:
ABYSSINIA: locality, date, and collector unknown.
TRANSVAAL: Miiller’s Farm, MacLea (distributed in Reh-
mann’s Hepaticae austro-africanae, No. 32, as “ Plagiochasma
muricatum Steph.,” but probably representing a part of the type
material of P. tenue).
AUSTRALIA: Swan River, West Australia, 1846, 7. Drummond.
NEw ZEALAND: without definite locality or date, J. D. Hooker
(type of Jungermannia | Fegatella| australis).
Aside from several other ‘localities in Italy and the At-
lantic Islands, P. rupestre has been recorded from Greece, from
Dalmatia, from southern France and Corsica, from Portugal,
from Cape Colony, and from Asia Minor. Whether the plant
from Schen-si, China, collected by Father Giraldi in 1896 and
described by Massalongo (21, p. 49) under the name P. elongatum
8 ambiguum, belongs here or not cannot be decided from lack of
specimens. According to the description, which tells nothing
about the epidermal pores, the spores are subrugose (or rugulose)
and not reticulated. Such a peculiarity would exclude the plant
not only from P. rupestre but probably from the genus Plagio-
chasma altogether. The author intimates, however, that the
spores may not have been mature, so that no definite conclusions
can be drawn from them.
Although P. rupestre has not been considered an especially
variable member of the Marchantiaceae, it nevertheless exhibits
considerable diversity in size, in color, in the distribution of its
male and female inflorescences, and in the structure of certain of
its organs and tissues. Some of this diversity at least is due to
282 Evans: THE GENUS PLAGIOCHASMA
external conditions. The variations in size and in inflorescence
have already been considered. With regard to color the thallus
is sometimes green throughout but is usually more or less pig-
mented with purple. The pigmentation may be confined to the
ventral scales, but it usually affects also the ventral surface from
the scales to the edge and a narrow border on the dorsal surface;
sometimes, in fact, the entire dorsal surface may be more or less
colored, although this condition is rare.
The structure of the epidermis and its pores is also worthy of
comment (Fic. 1). Not only do the epidermal cells vary
markedly in size, but their walls differ in thickness and the trigones
are sometimes conspicuous, with straight or convex sides, and
sometimes hardly to be made out at all. The waxy layer on the
cuticle, moreover, is sometimes so thick and opaque that it is
difficult to see the epidermal cells beneath. The epidermal pores
escaped the observation of the early authors, and even Nees
von Esenbeck (26, p. 45) and DeNotaris (5, p. 477) said that .
no pores were present. Apparently Voigt, in 1879, was the
first to detect them (40, p. 747, pl. 9, f. 24, 25). In P. Aitonia
he says that the pores are the smallest known to him, that
the cells bounding the opening differ from the other epidermal
cells only in their thickened radial walls, that the inner angles
of these cells are often cut off by walls thus forming two circles
(of four cells each), and that the inner cells of these circles are
often so small that they can easily be overlooked. In P. italicum
he says that the pores are larger, that the radial walls are scarcely
thickened, and that the inner cells of the two concentric circles
are almost always distinct. His figure of the surface view, which
is said to represent P. Aitonia, does not show the radial thickenings
very clearly, but his account of the two species which he dis-
tinguishes brings out the amount of variation to be expected in»
P. rupestre. In the vast majority of cases the number of cells
around the openings varies from four to six. In very rare cases
three cells or more than six may be demonstrated. In the accom-
panying figures the great variations in the size of the opening, in
the thickness of the radial walls, etc., are shown. Apparently the
degree of thickness which the radial walls show varies with the
size of the trigones in the neighboring epidermal cells. In culti-
Evans; THE GENUS PLAGIOCHASMA 283
vated material (Fic. 1, A, L) the cells tend to be large, the
trigones to be small, and the radial thickenings to be very incon-
spicuous, but similar conditions are sometimes present in plants
growing naturally, so that no very definite conclusions can be
drawn with respect to the causes of such variation.
The older writers laid but little stress on the peculiarities of
the ventral scales in distinguishing the species of the Marchan-
tiaceae. Perhaps some of the recent writers have over-emphasized
the importance of these characters, but in the genus Plagiochasma
the scales are often very helpful in the determination of species.
They are, however, subject to considerable variation, and this is
seen especially well in the case of P. rupestre. In their most
characteristic form the basal portion of the scale varies from
Fic. 4. PLAGIOCHASMA RUPESTRE (Forst.) Steph.
A-C. Epidermal pores, cross-sections, X 300. D. Cells from the margin of a
i F.
Figs. A and F-H were drawn from a specimen collected in Jalisco, _—— & Land
134; Fig. B, from a specimen collected in Arizona, G. E. Nichols; and Figs. C-E,
from a specimen collected in Morelos, Barnes & Land 480.
284 Evans: THE GENUS PLAGIOCHASMA
lunulate to ovate, while the appendages, one or two in number,
are subulate and acute, without constriction at the base (see
Fic. 2, A, B). Sometimes, when only one appendage is present
there is no sharp boundary where it passes into the basal portion.
The deviations from this characteristic condition are shown most
clearly in the form of the appendages, which are sometimes ovate
and more or less plicate at the base, and in their apices, which
may be acuminate or, on the other hand, obtuse or even rounded,
several of these variations being sometimes present on a single
thallus or in a single clump (Fics. 2, 3). The. pigmentation
of the scales is usually pronounced, although it may be absent in
very moist environments and may disappear to a greater or less
extent with age. The cells of the scales are fairly uniform through-
out (see Fic. 4, D). Some of the marginal cells occasionally
project as blunt or sharp denticulations, and in rare cases a distinct
marginal tooth is present. It is much more usual, however, for
the margin to be quite entire, and the peculiar slime papillae found
in some of the other species are so evanescent that they leave no
vestiges. The basal portion of the scale shows scattered cells
with oil-bodies, and sometimes one or two such cells are to be
seen in the appendages. One of the figures by DeNotaris shows
a ventral view of a thallus with several scales in position. One
of these scales shows four appendages. The writer has never
observed so high a number, although three appendages were found
in a very few cases.
In reducing P. italicum to a synonym of P. rupestre Miiller
calls attention to the fact that Stephani recognizes both species
in his Species Hepaticarum. He adds, however, that he has
sought in vain to find differential characters separating them,
although he has examined specimens under both names in the
Stephani herbarium. He found these to be almost exactly alike
even in their details. The differences brought out by Stephani
are the following: in P. rupestre, the dioicous inflorescence, the
pore surrounded by five cells, the single narrowly lanceolate and
acute appendage of the ventral scales: in P. italicum, the monoi-
cous inflorescence, the pore surrounded by four cells, the one or
two narrowly triangular appendages with broad bases. It will
be seen at once that these differences are based on exceedingly
EVANS: THE GENUS PLAGIOCHASMA 285
variable characters, and: it is not probable that many writers on
the Hepaticae will dissent from Miiller’s reduction.
In the present paper several other species are considered
synonyms of P. rupestre, and it might appear at first as if the
reduction had been carried on in a somewhat wholesale way. In
each case, however, the writer has based his opinion on the careful
comparison of type specimens with authentic material of P.
rupestre from Europe and the Atlantic Islands. It should perhaps
be added that most of the species here reduced were proposed at
a time when a plant was assumed to represent a new species if it
came from a distant or isolated locality. The various species
may be discussed in order.
In P. australe, which Stephani places just before P. elongatum,
the following characters are emphasized: the monoicous inflores-
cence, the androecium being near the carpocephalum, the pore
bounded by five cells, the lack of thickenings in the epidermal
cells, the two appendages of the ventral scales, abruptly ligulate
in outline, shortly acuminate, as long as the scales. In the
portion of the type specimen examined some of these characters
were not materialized. The cells of the epidermis, for example,
although difficult to see on account of the thick and waxy cuticle,
had distinct trigones, and the cells bounding the pores were four
in number about as often as five. The radial walls, moreover,
separating these cells were strongly thickened. With regard to
the scales many of them showed a single appendage, and the form
of this structure varied from lanceolate to ovate, the apex being
acute. Of course these are only slight deviations from Stephani’s
account, which was based on Australian material collected by F.
von Miiller, and they are no more than would be expected in a
variable species. Nothing was found in the type specimen which
would warrant a separation from P. rupestre.
In the case of P. limbatum, which Stephani places in his section
“@’’ (see p. 274), although he had not himself examined carpo-
cephala, a portion of the original material, likewise in the Mitten
herbarium, was examined. Here again nothing was found to
warrant a separation from P. rupestre. The distinguishing
characters, according to Stephani’s account, are the following:
the pronounced thickenings in the radial walls of the cells bounding
286 EvANs: THE GENUS PLAGIOCHASMA
the pores, the appendages of the ventral scales, often double,
long-acuminate from a broad base, papulose on the margin.
The papulose margin is unfortunately no more pronounced than
it sometimes is in specimens of P. rupestre from other localities,
and the other differences, even if they were constant, would not
be of much significance (see Fic. 2, M, N).
The characteristics of P. elongatum, according to Stephani,
are the following: the monoicous (paroicous) inflorescence, the
‘very minute pores with only two or three cells around the opening,
the trigones in the epidermal cells, the semilunate scales abruptly
contracted into one or, rarely, two long, narrowly lanceolate and
attenuate appendages, and the shortly emarginate carpocephala
bearded below by lanceolate or filiform scales. A study of the
type-specimen, however, shows that the pores are almost in-
variably surrounded by four cells (Fic. 1, H), and although the
appendages for the most part agree with the description (Fic.
3, R), some of them are subulate rather than lanceolate, the con-
traction is more gradual than is implied, and even the narrowest
appendages are occasionally duplicated in European material of
P.rupestre. The other characters, even if constant, would scarcely
be differential.
In proposing Aytonia Evansti as a new species Miss Haynes
compared it with P. elongatum and naturally laid a good deal of
emphasis on Stephani’s description. She points out differences
between the two species in size, in the structure of the epidermal
pores, and in the form of the ventral scales. She separates her
species because the pores are bounded by four or five cells instead
of by two or three, and because the scales taper gradually instead
of abruptly into the appendages. It has just been shown that
the characters upon which she relied are subject to considerable
variation in P. elongatum and that this species cannot be kept
apart from P. rupestre. It therefore becomes necessary to reduce
Aytonia Evansit to a synonym of the same species. At the same
time it is a pleasure to recommend the careful and accurate
description and figures in Miss Haynes’s paper to students of the
Hepaticae.
In P. mexicanum the original authors apparently had only
slight confidence. They say that it is closely allied to P. rupestre
Evans: THE GENUS PLAGIOCHASMA 287
but that it differs in the more crispate margin of the thallus,
in the fact that it is wholly convolute when dry, and in its less
conspicuous median canal when moist. They likewise quote a
specimen collected by Ecklon and Pappe at the Cape of Good
Hope, and although they doubtfully refer this specimen to P.
rupestre, Lindenberg expresses the opinion that it may be P.
mexicanum. Stephani adds no differential characters of impor-
tance, stating merely that the epidermal cells are surrounded by,
five or six cells with strongly thickened radial walls, that the
epidermal cells have trigones, that the ventral scales have a
single appendage, narrowly lanceolate and long-attenuate, and
that the carpocephalum is excised at the apex with obtuse,
recurved-connivent horns. A portion of the type specimen from
the Gottsche herbarium showed most of these characters, although
some of the epidermal pores were bounded by only four cells
(Fic. 1, E). The specimen showed nothing, however, to justify
the maintenance of P. mexicanum as a valid species.
Of P. lanigerum authentic material was distributed in Spruce’s
Hepaticae Spruceanae. In the original description the ventral
scales were said to be obliquely triangular and acuminate and the
elaters to have four spirals. Stephani adds an account of the
epidermal cells, which are said to have trigones, and of the pores,
which are said to be surrounded by five cells. He states further
that the single appendage of the scales is as long as the basal part,
lanceolate in form, and acute, and that the elaters have from two
to four spirals. It will be seen at once that there is nothing in
either description which would not apply to the variable P.
rupestre, and Spruce’s specimens amply support this position.
In addition to the four species just discussed there are several
others recognized by Stephani which are evidently very closely
related to P. rupestre. Among these are the following: P.
dschallanum, P. tenue, P. algericum, P. nepalense, P. chlorocarpum,
P. validum, P. caerulescens, P. brasiliense, P. peruvianum, and P.
subplanum. It is probable that some of these will prove to be
synonymous with P. rupestre, but unfortunately authentic
material has not been available for study. A possible exception
may be made in the case of P. tenue, which Stephani quotes from
three South African regions: the Transvaal, MacLea, Wilms;
288 EvANs: THE GENUS PLAGIOCHASMA
Usambara, Holst; and Angola, Welwitsch. In the Underwood
herbarium there is a specimen from A. Rehmann’s Hepaticae
austro-africanae, No. 32, collected by MacLea at Miiller’s Farm
in the Transvaal and labeled ‘‘ Plagiochasma muricata Steph.”
This species was never published but the specimen probably
represents P. tenue, because no other species of MacLea’s Trans-
vaal collection is quoted. Since MacLea’s specimen is cited first
_ by Stephani it is even probable that the plant in the Underwood
herbarium is part of the type material. The distinguishing
characters of P. tenue are said to be the following: the dioicous
inflorescence (androecia unknown), the very minute epidermal
pores surrounded by four cells, the triangular ventral scales bifid
two-thirds with parallel, contiguous, narrowly triangular divisions
(the latter of course being really the appendages), and the strongly
convex carpocephala. MacLea’s specimen shows most of these
characters except that the carpocephala are distinctly concave.
The other characters, moreover, are subject to variation, the pores
being sometimes surrounded by five cells and the ventral scales
sometimes having a single appendage and sometimes two more
or less divergent appendages. In other words the specimen
agrees in all essential respects with P. rupestre. Whether the
other specimens quoted by Stephani represent P. rupestre cannot
of course be decided, but the probability is that they do.
If P. rupestre is accepted in the broad sense of the present
paper its geographical distribution becomes very extensive. It is
not, however, unique in this respect among the Marchantiaceae.
Both Targionia hypophylla and Reboulia hemisphaerica have a
similar and even widet range. Probably if P. rupestre extended
as far north in Europe as these other two spécies, it would long ago
have been thoroughly known to European writers, and the con-
fusion regarding its synonymy would already have been dispelled.
2. PLAGIOCHASMA CRENULATUM Gottsche
Plagiochasma crenulatum Gottsche, Mex. Leverm. 266. 1863.
Aytonia crenulata Underw. Bot. Gaz. 20: 66. 1895.
Thallus pale green above but not glaucous, with a narrow purple
border, plane or broadly canaliculate, strap-shaped, sparingly fork-
ing, sometimes with adventive branches, rarely (if ever) innovat-
EvANS: THE GENUS PLAGIOCHASMA 289
ing at the apex, margins undulate-crenate, more or less crispate,
mostly 1-3 cm. long and 5-10 mm. broad, thickness of thallus
about one sixth the ih: epidermis composed of thin-walled cells
with distinct trigones and a thin smooth cuticle, the cells about
30 w in diameter; pores slightly elevated, rather small, surrounded
by about six (four to seven) radiating series of cells with two or
three cells in each series, radial walls slightly thickened; aeriferous
layer about three fourths the thickness of the thallus in the median
portion, fairly compact with the air spaces about as wide as the
green cells or even narrower close to the epidermis; ventral scales
lunulate or ovate, purple, slightly overlapping, irregularly sinuous-
dentate along the margin with contorted cells, bearing evanescent
slime-papillae, cells containing oil-bodies few and scattered, the
scales abruptly contracted into one or (more rarely) two large,
broadly ovate appendages, short-apiculate or abruptly acute at
the apex, irregularly sinuate-dentate along the margin or bearing
one or several sharp teeth or even one or two sharp basal lobes:
inflorescence monoicous (autoicous): co inflorescence borne on a
short ventral branch: @ receptacles borne singly or in tandem
pairs; peduncle mostly 1-1.5 cm. long; carpocephalum concave
at the apex, or obtusely indented, and two- or three-lobed with
apiculate lobes, maturing two or three sporophytes; scales of
carpocephalum lanceolate to narrowly ovate, irregularly sinuate
on the margin or with an occasional tooth: spores about 80 uw in
diameter, yellowish brown, minutely verruculose, regularly reticu-
late, spherical face usually with eighteen to twenty meshes (ten
to twelve marginal), plane faces with about seven (five to eight)
meshes, those near apex of pyramid less distinct; elaters gradually
attenuate, mostly 270-325 u long and 12m wide in maximum
diameter, spirals usually three, often coalescing along a longi-
tudinal line. [Fic. 5.]
The following specimens have been examined:
Curapas: San Cristobal, altitude 2,100 m., November, 1907,
G. Miinch 5607 (distributed by Levier).
Sonora: Huchuerache, December, 1890, C. V. Hartman 307
(Lumholtz Expedition).
VERA Cruz: Orizaba, F. Miiller (type); February, 1885, W. G.
Farlow; January, 1892, J. G. Smith; December, 1900, Conzattt &
Gonzales; November, 1908, Barnes & Land 685; on wet ground,
Cordova, February, 1885, W. G. Farlow; on wall along roadside,
Cordova, February, 1907, Miss L. MacIntyre; along sides of Metlac
Gorge, October, 1908, Barnes & Land 640, 659.
Gottsche likewise reports the species from Chinantla, PUEBLA,
F. Liebmann. No other localities are at present known.
290 Evans: THE GENUS PLAGIOCHASMA
Stephani’s treatment of this large and distinct species is
difficult to understand. He states that the thallus always inno-
vates at the apex, thus implying that dichotomy never occurs, and
he adds among other details that the epidermal pores are large
and surrounded by eight or nine radiating series of cells with four
cells in each series, that the appendages of the ventral scales are
ovate-oblong or broadly linear with shortly acuminate apices, and
that the apex of the carpocephalum is plano-convex. It would
almost seem as if he had not seen the true P. crenulatum at all
but had drawn his description from some other species. Gottsche’s
original account is much more accurate, and Leitgeb (16, p. 64,
pl. 1, f. 6) has given an excellent description, with a figure, of the
epidermal pores. He calls attention to the fact that the con-
centric rings of cells surrounding the pore appear as if partially
shoved under one another. The pore-opening thus acquires the
form of a canal, although the cells all belong to the same epidermal
cell-layer. In the material studied by the writer the pores are
surrounded by from four to seven radiating series of cells, six being
the usual number (Fic. 5, A), and each series is composed of two or
three cells. The radial walls are thickened but not very strikingly
so. The circular ridge immediately surrounding the opening is
distinct in P. crenulatum and in all the following species.
The ventral scales develop large and characteristic appendages,
usually singly but occasionally in pairs (Fic. 5, C, D). The
scales themselves, as in the other members of the genus, are
lunulate or ovate in form and overlap slightly. Their cells di-
minish in size toward the margin and become exceedingly irregular
in form and in the direction of their long axes. Some of the
marginal cells project and bear vestiges of slime papillae. The
appendages are strongly contracted at their junction with the
scales and vary considerably in size. Their form, in general, is
broadly ovate to orbicular and they narrow abruptly into an
apiculate or shortly acute apex, which is tipped with a single cell
or with a row of two or three cells. The cell-structure is very
similar to that in the scale itself and the border, composed of
irregular and contorted cells, is very different from what is found
in the narrower scale-appendages of P. rupestre, where the cell-
structure of the appendages is practically uniform throughout.
Evans: THE GENUS PLAGIOCHASMA 291
A species closely allied to P. crenulatum is the East Indian P.
appendiculatum Lehm. & Lindenb., one of the two species upon
which the genus Plagiochasma was based. This interesting plant
is known to the writer from a series of specimens, including a part
.
VIO
Or
Fic. 5. PLAGIOCHASMA CRENULATUM Gottsche
A. Epidermal pore, surface view, X 300. B. Epidermal pore, cross section,
300. C, D. Appendages of ventral scales, X 40. E-G,. Scales from a carpo-
cephalum, X 40. H. Spore, spherical face, X 225. I. Elater, X 225. The figures
were all drawn from specimens collected by Barnes & Land at Orizaba (685).
of Wallich’s original collection in the Mitten herbarium. It is
characterized, as Stephani notes, by the large appendages of the
ventral scales, cordate or broadly ovate in form, entire, obtuse,
and strongly contracted at the base. This description brings out
their peculiarities clearly. It may be added that the cell-struc-
ture of the appendages is fairly uniform except for a very narrow
border, where the cells are somewhat irregular and contorted.
292 Evans: THE GENUS PLAGIOCHASMA
The epidermal cells and pores are much the same as in P. crenula-
ium, although the number of radiating series of cells around a pore
is sometimes as high as eight. The spores are distinctly smaller
measuring 50-70 uw in diameter (51 » according to Stephani), but
they are reticulate in much the same way, and the elaters usually
show more or less coalescence of their spiral bands.
The specimens collected by Hartman in the state of Sonora
come particularly close to P. appendiculatum, and their reference
to P. crenulatum must be considered provisional. The appendages
of the ventral scales are distinctly orbicular in most cases with
rounded apices and entire margins, so that they simulate those
of the East Indian species very closely. If it were not for the
occasional presence of appendages distinctly narrowed toward
the apex or even acute it would be quite justifiable to consider
the Sonora plant distinct from P. crenulatum. Unfortunately the
material is both scanty and sterile, and the appendages are often
so thoroughly bleached that their true structure cannot be well
made out. It is possible, therefore, that more complete or
fertile material would show that the striking peculiarities of the
plants were of an accidental character, and for this reason they
are included under P. crenulatum.
Another specimen which requires a few words of comment is
the one collected by Miinch at San Cristobal. This specimen was
distributed by E. Levier in his Bryotheca under a manuscript name
of Stephani. The writer finds no sufficient grounds, however, for
separating it from P. crenulatum although the appendages of the
ventral scales are somewhat smaller and narrower than in typical
forms of the species and taper more gradually toward the apex.
Combined with this slight difference the epidermal pores exhibit
thicker radial walls than is usual and tend to show more than six
radiating series of cells around the opening. In other respects the
specimen is like P. crenulatum.
3. Plagiochasma jamaicense (Haynes) comb. nov.
Aytonia jamaicensis Haynes, Bull. Torrey Club 34: 58. pl. 6.
1907.
Thallus pale green above but not glaucous, with a reddish
purple border, plane or broadly canaliculate, strap-shaped, some-
EvaANs: THE GENUS PLAGIOCHASMA 293
times with adventive branches, sometimes innovating at the apex,
rarely (if ever) dichotomous, margins crenulate, more or less
crispate, mostly about 1 cm. long and 5-7 mm. wide, thickness of
thallus about one eighth the width; epidermis composed of thin-
walled cells with small but distinct trigones and a smooth cuticle,
the cells about 30 uw in diameter; pores slightly elevated, rather large,
surrounded by about eight (five to ten) radiating series of cells
with two to four cells in each series, radial walls slightly thickened;
oo layer about three-fourths the thickness of the thallus
the median region, fairly compact, with the air-spaces a little
viider than the green cells; ventral scales lunulate to ovate, purple,
slightly overlapping, irregularly sinuate along the margin with
vestiges of slime papillae, cells containing oil-bodies very few
and scattered, the scales abruptly contracted into one or (rarely)
two ovate appendages, constricted and plicate at the base, entire
or vaguely and irregularly dentate or crenate along the margin,
often showing vestiges of slime papillae, gradually narrowed
toward the apex into an apiculum usually two to three cells long;
inflorescence monoicous: co’ inflorescence (in the few observed
cases) near the apex of a thallus, the Q receptacles being borne
(usually in a series) on the following joint or apical innovation;
peduncle of carpocephalum 1~-1.5 cm. long; carpocephalum mostly
plano-concave at the apex and bluntly two- to four-lobed, usually
maturing two sporophytes; scales of carpocephalum lanceolate
to linear-lanceolate, acuminate, entire or sparingly and irregularly
dentate: spores 70-85 » in diameter, yellowish brown, minutely
verruculose, reticulate but with the meshes often indistinct,
spherical face with about twelve meshes (eight or nine marginal),
plane faces with about four meshes; elaters gradually attenuate,
mostly 250-375 wu long and 8» in maximum diameter, with two
or three spirals, sometimes slightly coalescent.
The following specimens have been examined:
Jamaica: Chestervale, Blue Mountains, altitude 900 m.,
February, 1903, L. M. Underwood 1173 (type), 1177 (both mixed
with Reboulia hemisphaerica [L.] Raddi).
No other localities for the species are known.
The above description is largely compiled from the original
paper of Miss Haynes, only a few details having been added. The
specimens studied bear a very strong resemblance to the Reboulia
with which they are growing, the size, general habit, and pigmented
margin being almost exactly the same. The color, however, is a trifle
redder in the Plagiochasma and of course the dorsal position of the
carpocephala will at once separate it from the Reboulia, where the
294 Evans: THE GENUS PLAGIOCHASMA
carpocephala are terminal. The appendages of the ventral scales
are likewise very different, those of the Reboulia being narrowly
linear or lanceolate.
A close relative of P. jamaicense is P. crenulatum, a somewhat
larger and firmer species, the aeriferous layer being a little more
compact. In distinguishing the species the best differential
characters are drawn from the epidermal pores and ventral scales.
Although the pores in the two plants are built up on the same
plan, those of P. jamaicense tend to be more complex and usually
show a greater number of radiating series of cells around the
opening with more cells in each series. The ventral scales are
much the same in their basal portions except that those of P.
crenulatum are larger and show a broader border of contorted
cells. It is in their appendages that the scales are most distinct.
In P. crenulatum these are broadly ovate to orbicular, the largest
reaching a length of I-1.4 mm. and a width of about I mm.; in
P.. jamaicense they are more narrowly ovate, the largest reaching
a length of 0.45 mm. and a width of 0.3 mm. In P. crenulatum the
apical contraction is usually abrupt, in P. jamaicense it is more
gradual, and the apex seems to be invariably tipped with a row of
two or three cells. Another difference to be noted is in the position
of the androecium. In P. crenulatum it is borne on a short
_ ventral branch and is apparently never succeeded by female
receptacles as is the case in P. jamaicense. Unfortunately, as
already pointed out, it is unwise to place too much confidence in
a difference of this sort. The spores of P. crenulatum, finally,
are more finely reticulated than in P. jamaicense, the spherical
face showing eighteen to twenty meshes instead of only twelve.
4. PLAGIOcCHASMA WriGutit Sulliv.
Plagiochasma Wright Sulliv.; A. Gray, Manual, Ed. 2, 688.
pl. 6. 1856.
Attonia Wrightit Underw. Bull. Illinois State Lab. Nat. Hist. 2:
43. 1884.
Thallus pale green and glaucous above, with a narrow purple
border, plane or broadly canaliculate, strap-shaped, sometimes
forking, sometimes innovating at the apex, sometimes with ventral
adventive branches, margins undulate-crenate, more or less cris-
pate, mostly 1.5-2 cm. long and 3-5 mm. broad, thickness of
Evans: THE GENUS PLAGIOCHASMA 295
thallus about one fourth the width; epidermis composed of thin-
walled cells with distinct trigones often having bulging sides and
a thin cuticle with a waxy covering, the cells averaging about 22 u
in diameter; pores slightly elevated, rather small, surrounded by
about six (five to eight) radiating series of cells with two or three
cells in each series, radial walls distinctly (and sometimes strongly)
thickened; aeriferous layer about three fourths the thickness of
the thallus, of a rather loose structure, the air spaces about twice
as wide as the green cells; ventral scales lunulate, purple, slightly
overlapping, irregularly sinuous-dentate along the margin with
somewhat contorted cells, bearing evanescent slime-papillae, cells
containing oil bodies few and scattered, the scales abruptly
contracted into one or two ovate-lanceolate appendages, obtuse
to acute at the apex, entire, more or less constricted and plicate at
the base: inflorescence Sabena (autoicous): o inflorescence
borne behind the 9 receptacle o on a short ventral branch:
2 receptacles borne singly or in a ea row of two or three;
peduncle mostly 1.5-2 mm. long (3-4 mm. long, according to
Stephani); carpocephalum narrowly concave at the apex, two-
or three-lobed with apiculate lobes, maturing two or three sporo-
phytes; scales of carpocephalum linear-lanceolate, acuminate,
entire; spores brownish yellow, 75-85 in diameter, minutely
rugulose, regularly reticulate, spherical face usually with twelve or
thirteen meshes, plane faces each with four meshes, reticulum often
incomplete; elaters gradually attenuate, mostly 225-250 win length
and 7-Qy in maximum seu aie spirals two or three, often
somewhat coalescent. ([Fic. 6.]
The following specimens have ete examined:
TEXAs: under overhanging rocks along steams, San Marco,
1848, C. Wright 9 (type).
ARIZONA: Nogales, February, 1902, D. T. MacDougal.
CoanuiLa: San Lorenzo Canyon, about 3.5 km. southeast of
Saltillo, September, 1904, E. Palmer 423.
DuRANGO: near the city of Durango, 1896, EZ. Palmer 862.
VeRA Cruz: Cordova, without date, C. Sallé; Orizaba, Feb-
ruary, 1885, W. G. Farlow 17 (in Farlow herbarium).
No other localities have been reported.
In studying the present species the writer has had the privilege
of examining a portion of the type material. Although carpo-
cephala and sporophytes are present, the spores and elaters are
poorly or abnormally developed so that it is impossible to gain from
them an accurate idea of the typical structure of these organs.
296 Evans: THE GENUS PLAGIOCHASMA
This is fortunately supplied by the specimens collected by Sallé
at Cordova.* The specimens from Arizona and the other Mexican
specimens listed above lack carpocephala but show clearly the
vegetative characters of the species.
Sullivant’s original account, which is reproduced with but
slight alterations by Underwood, describes the more striking
characters of the plant but does not call attention to the structure
of the epidermis and of the ventral scales. These deficiencies
are partly supplied by Stephani, who states that the epidermal
pores are slightly convex and surrounded by two to three concen-
tric series of cells with six cells in each series and that the append-
ages of the scales are constricted at the base, ovate, acute, and
entire. In her discussion of Aytonia jamaicensis, Miss Haynes
notes that the concentric series of cells around the pores of P.
Wrighttt are composed of from five to eight cells each, instead of
being uniformly composed of six cells, as Stephani’s, description
states.
The epidermal pores in P. Wrightti agree in all essential
respects with those of P. crenulatum, except that the radial walls
tend to be more distinctly thickened. The epidermis is further
distinguished by its somewhat smaller cells and better developed
trigones and by the presence of a waxy cuticular covering, these
features being apparently associated with a more xerophytic
habitat.
Although Stephani’s description implies that the ventral
scales bear a single appendage it is not unusual to have two such
appendages present. They are characterized by their ovate-
lanceolate form and acute or obtuse apices, the margins being
usually entire and the cell structure fairly uniform. The appen-
dages are somewhat constricted at the base, but the constriction
often appears more marked than it really is on account of the
folds in the basal portion. The marginal cells of the basal por-
tion (Fic. 6, D) are more or less irregular and contorted and show
vestiges of slime-papillae.
As pointed out by Miss Haynes, P. Wrightii is related to P.
jamaicense. She finds differences, however, in the epidermal
*It may be noted that Berkeley in his Introduction to Cryptogamic Botany
(p. 443, f. 93 a~c), published in 1857, has figured a carpocephalum and a spore from
Sallé’s material, under the name P. mexicanum.
Evans: THE GENUS PLAGIOCHASMA 297
pores, in the ventral scales, in the length of the peduncle, in the
scales of the carpocephalum, and in the spores. Those drawn from
the length of the peduncle, from the
scales of the carpocephalum and from
the spores are perhaps a little untrust-
worthy and difficult to apply, but the
others will serve to distinguish the two
species. In P. Wrightii the pores are
much the same as in P. crenulatum,
although perhaps a little less elevated;
the opening is usually surrounded by
six radiating series of cells with two or
three cells in each series. In P. jamai-
cense there is a tendency at least to-
ward a greater complexity, many of the
pores showing eight radiating series of
cells with three or four cells in each
series. Of course here, as in so many
other cases, it is unsafe to base conclu-
sions on the study of only a few pores,
because those of P. Wrightii sometimes
show as many as eight radiating rows
while those of P. jamaicense may show
as few as six or even five. In most cases
the radial wall-thickenings and the
trigones are better developed in P.
Wrightt than in P. jamaicense. Ac-
cording to Miss Haynes the append-
ages in P. Wrightii are less constricted
and narrower than in P. jamaicense
with their margins always entire. A
study of a considerable amount of
material shows that these differences
are usually, but not always, present;
there are rare cases, for example, where
the appendages in P. Wrightii are
broad enough to be called ovate, where
the basal constriction is marked, and
Fic. 6. PLAGIOCHASMA WRIGHTIL
Sulliv.
A. Epidermal pore; surface view,
X 300. B. Ventral scale with two
appendages, X 40. C. Append-
age from another ventral scale,
x 40. D. Cells from the margin
of a ventral scale, basal portion,
X 225. The figures were all drawn
from the specimenscollected in the
state of Coahuila, E. Palmer 423.
298 EvANs: THE GENUS PLAGIOCHASMA
where the margin is vaguely denticulate. These broad appendages,
however, seem to be always associated with narrower appendages
of the usual type and the apices show a much greater degree of
variation than in P. jamaicense, being sometimes acute and some-
times obtuse.
Although P. Wright is doubtless a close relative of P. crenula-
tum the two species differ strikingly in appearance. P. crenulatum
is not only a larger plant but it looks much firmer and more com-
pact owing to the smaller intercellular spaces in the aeriferous layer.
It also lacks the waxy covering on the cuticle, which is sometimes
conspicuous in P. Wrightwi and gives the thallus a glaucous aspect.
The margin is about as crenulate in P. Wrightii as in P. crenulatum
but the whole marginal region is thinner and more markedly
crispate. With respect to branching, P. crenulatum rarely, if ever,
develops a jointed thallus, while such a condition is usual in P.
Wrightu. This difference, however, as already emphasized, is
one which must be used with caution.
5. Plagiochasma Landii sp. nov.
Thallus pale green above but not glaucous, with a narrow
purple border, plane or broadly canaliculate, strap-shaped, rarely
forking, usually innovating at the apex or with ventral branches.
margins undulate, slightly crispate, 1 cm. or less in length, 4-7
mm. in width, thickness of thallus about one sixth the width;
epidermis composed of thin-walled cells with small but distinct tri-
gones and a thin smooth cuticle, the cells averaging about 28 yu in
diameter; pores slightly elevated, rather small, surrounded by
five to seven radiating series of cells with two or three cells in each
series, radial walls slightly thickened; aeriferous layer about three
fourths the thickness of the thallus in the middle, fairly compact
with the air spaces about as wide as the green cells or a little wider,
tending to extend at right angles to the epidermis; ventral scales
broadly lunulate, purple, entire, cells along the margin smaller,
more or less elongated parallel with the margin and with scattered
evanescent papillae, cells containing oil-bodies few and scattered,
the scales gradually narrowed into a single subulate, acuminate
appendage, varying greatly in length, entire or rarely with a sharp
tooth, scales sometimes with two such appendages: inflorescence
monoicous (paroicous): Q receptacles usually borne singly, pe-
duncles very short, usually about 1 mm. long, scales of the carpo-
cephalum lanceolate, acuminate, somewhat contracted at the
Evans: THE GENUS PLAGIOCHASMA 299
base, variable in size, the largest about 1.2 X 0.3 mm., about ten
cells wide, entire or sometimes with one or two short and irregular
teeth, oil-containing cells few and scattered; carpocephalum con-
cave at the apex, usually with two short-apiculate and sometimes
connivent lobes and maturing two sporophytes: spores yellowish
brown, mostly 70-80 uw in diameter, minutely verruculose, regularly
areolate, the spherical face mostly with twelve to fifteen meshes
(nine to twelve marginal), plane faces each with four (or five)
meshes; elaters about 225 wu long and 12 w in maximum diameter,
usually with two distinct spirals, rarely with three. [Fic. 7.]
The following specimens have been examined:
MorELos: near Cuernavaca, October, 1908, Barnes & Land
466; same locality and date, C. G. Pringle 10669 (distributed in
Plant. Mex. as P. elongatum). No. 466 may be designated the
type.
A remarkable specimen collected by W. G. Farlow at Orizaba,
state of Vera Cruz, and preserved in the Underwood Herbarium,
should be noted in this connection. It bears the number 17 and
is referred to P. crenulatum (see Underwood, Bot. Gaz. 20: 66.
1895). The long and slender appendages of the ventral scales,
however, with acuminate apices, would at once exclude it from
P. crenulatum and indicate an affinity with P. Landit. Possibly
it should be referred to this species, but the epidermal pores are
not quite characteristic, and it is impossible to decide from the
material examined whether the slight differences which the pores
show come within the range of variation to be expected. Appar-
ently No. 17 was made up of more than one species; as already
noted the specimen in the Farlow herbarium preserved under this
number is P. Wrightii.
In P. Landii the group of species to which P. crenulatum, P.
jamaicense and P. Wrightii belong has another member, the rela-
tionship being perhaps closest to P. Wrightit. In size and texture
the species approaches P. crenulatum but it differs in habit on
account of the many apical joints or innovations which it produces.
Compared with P. Wrightii the thallus is a trifle larger and firmer.
Here again the epidermal pores and the ventral scales yield
some of the most trustworthy differential characters. The epi-
dermal pores tend to be somewhat less complex than in the other
members of the group, each radiating series bounding the opening
300 Evans: THE GENUS PLAGIOCHASMA
being often composed of only two cells. In this respect it of
course recedes most widely from P. jamaicense. The cells forming
the pore show in a marked degree the peculiarity described ‘by
Leitgeb in the case of P. crenulatum. They appear as if shoved
under one another, and this condition is often so marked that the
Fic. 7. PLaGiocHasma Lanpi Evans
A. Epidermal pore, surface view, X 300.
X 300. C. Ventral scale with appendage, X 4o.
scale, X 40. E-G. Scales from a carpocephalum, x 40. H. Spore, spherical face,
X 225. I. Elater, X 225. The figures were all drawn from the type specimen.
B. Epidermal pore, cross-section,
second and third cells in each radiating series lie almost directly
under the cell next the opening. The cells in each series are
separated by very thin walls, and the lines of junction between
these walls and the cuticle are more or less convex when seen from
above, thus giving the pore a somewhat distinctive, flower-like
appearance.
The ventral scales resemble those of P. rupestre, although the
basal portion shows a very narrow border of contorted cells and
Evans: THE GENUS PLAGIOCHASMA 301
often bears vestiges of slime-papillae. The basal portion is un-
usually deeply pigmented, and the pale scattered cells contain-
ing oil-bodies stand in sharp contrast to the others. The appen-
dages of the scales, sometimes borne singly and sometimes in
pairs, are narrowly triangular or subulate in outline and gradually
taper to acuminate points, although the extreme apex is often
tipped with two cells bearing the vestiges of a slime-papilla
between them. There is no sharp line of demarcation between
the basal portion and the appendages and the latter are not
constricted at the base. For the most part the margins are entire
but occasionally an indistinct tooth is present. The appendages
are very different from the broad and strongly constricted append-
ages of P. crenulatum and P. jamaicense. They resemble more
closely the narrow appendages exceptionally found in P. Wrightit.
The latter species, however, is sure to show appendages of the
usual type on some or most of the scales.
6. PLAGIOCHASMA INTERMEDIUM Lindenb. & Gottsche
Plagiochasma intermedium Lindenb. & Gottsche; G. L. & N. Syn.
Hep. 513. 1846.
Rupinia intermedia Trevis. Mem. R. Ist. Lomb. III. 4: 437.
1877.
Aytonia intermedia Underw. Bot. Gaz. 20: 66. 1895.
Thallus pale green above but not glaucous, with a narrow
urple border, plane or broadly canaliculate, strap-shaped,
thallus about one seventh the width; seideriale composed of thin-
walled cells with small but usually distinct trigones and a thin,
smooth cuticle, the cells averaging about 25 » in diameter; pores
distinctly elevated, large, surrounded by about eight (six to ten)
radiating series of cells, with two to four cells in each series, radial
walls strongly thickened; aeriferous layer about two thirds the
thickness of the thallus, of a loose structure, the air spaces often
three or four times as wide as the green cells; ventral scales
lunulate, purple, sinuous to crenulate along the margin, some of
the marginal cells often contorted, cells containing oil-bodies few
and scattered, the scales gradually contracted into two appendages,
less often into only one, the appendages slightly or not at all
constricted at the base, lanceolate to ovate, entire, usually acute
302 EvANS: THE GENUS PLAGIOCHASMA
and tipped with a single cell, mostly seven to ten cells wide:
inflorescence monoicous: o inflorescence borne near the apex,
sometimes on a ventral branch: @ receptacles usually borne
singly near the apex of the thallus, but sometimes in a median
series of two or more, the stalk mostly only 1-2 mm. long; carpo-
cephalum concave with apiculate lobes, commonly maturing two
or three sporophytes; scales of carpocephalum lanceolate, acute
to acuminate, entire, or rarely with a tooth; spores mostly 60-70 u
n diameter, minutely verruculose, sometimes reticulated, some-
de not, the spherical face then showing a periclinal ridge and
one or two irregular supplementary ridges, and each plane face
a periclinal ridge; elaters 200-250 uw long and 9-12 uw in maximum
diameter, tapering gradually to blunt extremities, usually with
uniformly thickened walls and a ae narrow cell cavity, rarely
with rudimentary spiral bands. [Fia. 8.]
The following specimens have been examined:
JaLisco: near Guadalajara, 1889, C. G. Pragie: zoo (see
Underwood, Bot. Gaz. 20: 66. 1895); wet banks and rocks,
Barranca de Oblatos, Guadalajara, September, 1908, Barnes &
Land 136; moist rocks, Barranca Ibarra o Portella, below Exper-
iencia, September, 1908, Barnes & Land 146; banks of streams,
road to San Domingo Mine and adjacent gullies and hills, Etzatlan,
October, 1908, Barnes & Land 261.
PurEBLA: banks along Avenida Hidalgo and path to Barranca
Tezuitlan, October, 1908, Barnes & Land 538.
VeRA Cruz: San Antonio Huatusco, mixed with Targionia
hypophylia L., 1857, C. Mohr 146.
GUATEMALA: Guachipilin, Dept. Santa Rosa, September, 1893,
Heyde & Lux; Cuajiniquilapa, Dept. Santa Rosa, August, 1894,
Heyde & Lux (see Underwood, Bot. Gaz. 20: 66. 1895). Both
specimens were distributed by John Donnell Smith under No. 6292.
JAPAN: Ogawa-mura, Tosa, July, 1900, T. Yoshinaga 6.
The species has likewise been reported from several other
localities in Mexico and Japan and from the province of Shen-si in
China. Type Locariry: Hacienda de Jovo, VERA Cruz, F.
Liebmann.
The remarkable elaters of P. intermedium with their uniformly
thickened walls will usually serve to distinguish the species from
its American allies, even if thickening of this type is not absolutely
constant. In the case of sterile specimens the epidermal pores
EvANs: THE GENUS PLAGIOCHASMA 303
and the ventral scales afford differential characters of importance.
The pores are much like those of P. jamaicense and tend, if any-
thing, to be even more complex, frequently showing nine radiating
series of cells around the opening with four cells in each series.
The radial walls separating the series are, moreover, strongly
thickened. The ventral scales bear considerable resemblance to
those of P. Landii and P. Wrightii.. The basal portion shows a
vague border of contorted cells, the margin itself being indistinctly
Fic. 8. PLAGIOCHASMA INTERMEDIUM Lindenb. & Gottsche
A. Epidermal pore, surface view, X 225. B. Epidermal pore, cross-section,
X 225. C. Ventral scale with two appendages, X 40. D-—F. Scales from a car-
. G. Spore, spherical face, X 225. H. Spore, plane faces,
X 225. I. Elater, X 225. The figures were all drawn from the specimens collected
by Barnes and Land at Etzatlan (261).
sinuous or crenulate with vestiges of slime papillae. The appen-
dages, usually borne in pairs, are lanceolate to ovate and commonly
acute. According to Stephani they are strongly constricted at
the base, but this condition is apparent rather than actual and
seems to be due to the presence of basal folds. When these are
not present the appendages show no appearance of constriction.
304 EvaANs: THE GENUS PLAGIOCHASMA
Although the appendages are much like those of P. Landii and
P. Wrightti they are distinguished from the first by their blunter
apices and from the second, usually at least, by their narrower
form and lack of basal constrictions.
In proposing Aitonia japonica as a new species Stephani
emphasized the solid elaters and spoke of them as a unique
feature, apparently overlooking the fact that Gottsche had as-
signed similar elaters to P. intermedium many years earlier. In his
Species Hepaticarum, however, Stephani recognizes both P.
japonicum and P. intermedium and ascribes solid elaters to both.
A comparison of his descriptions brings out the following dif-
ferences. In P. japonicum the thallus is said to be dichotomous
and to innovate from the apex and laterally from the midrib; the
epidermal cells to have trigones, and the pores to be surrounded
by three or four concentric series of cells with six to eight cells
in each series; the appendages of the ventral scales to be ovate-
oblong or elliptical; the carpocephalum to be plano-convex with
rounded lobes; and the spores to be 50u in diameter. In P.
intermedium the thallus is said to be rarely dichotomous but to
innovate from the apex; the epidermal cells to have thick walls
but no trigones; the pores to be surrounded by three concentric
series with eight cells in each series; the appendages of the ventral
scales to be strongly constricted, ovate-oblong, acute, and entire;
the carpocephalum to be concave, the lobes showing erect-
recurved and connivent horns; and the spores to be 60 uw in di-
ameter.
It will at once be seen that most of these differences are slight
or relate to characters which may be expected to be variable.
Those drawn from the epidermal cells and from the carpocephala
seem at first sight to be moreimportant. Unfortunately they prove
to be either inconstant or based on hasty observations. The
epidermal cells in P. intermedium, for example, have distinct
trigones even if they are sometimes small; and the carpocephala
in Yoshinaga’s specimens of P. japonicum are distinctly concave
at the apex and show apiculate lobes, agreeing with the usual
condition in P. intermedium. The writer, in fact, has been able
to find no valid characters distinguishing the North American
from the Asiatic species and therefore feels compelled to reduce
the latter to synonymy, in spite of its wide geographical separation.
>
Evans: THE GENUS PLAGIOCHASMA 305
The spores reproduced (Fic. 8, G, H) were drawn from the
specimens collected at Etzatlan, 261, and show very few ridges
except those along the edges of the tetrahedron. These ridges
do not form a reticulum. On the spherical face a continuous wavy
ridge is present about half way between the periphery and the
center, and from one to three additional ridges extend partly or
wholly across the enclosed space. On each plane face a single
ridge extends across about half way between the base and apex
of the triangles. Unfortunately these spore characters, which at
first sight appear so striking, are subject to variation. In the
spores of the Guadalajara material, 136, a regular reticulum is
present in most cases, and the two extreme types of spore-structure
are connected by a series of intergradations. The Japanese
specimens of P. japonicum show spores with a diameter of about
70 and a well-developed reticulum. In Massalongo’s figures,
drawn from his variety chinense, the reticulum is clearly shown
but the number of meshes represented is greater than in the
Japanese material. In the Synopsis Hepaticarum a specimen
from Nepal is doubtfully included under P. appendiculatum as
‘““? 8 depauperata.”’ Gottsche states that this plant is close to
P. intermedium and implies that the elaters are of the solid type,
while Massalongo cites it as a possible synonym of P. japonicum
8 chinense. Unfortunately the lack of specimens has made the
study of this form impossible to the writer. It should be re-
membered, however, that Kashyap occasionally finds solid elaters
in P. appendiculatum, so that their presence in this doubtful form
would not necessarily exclude it from that species. The Synopsis
tells us nothing about the epidermis or ventral scales, and Stephani
does not mention the plant at all.
The present paper is based on the study of a large series of
specimens, including type material of the following previously
published species: P. appendiculatum, P. australe, P. cordatum,
P. crenulatum, P. elongatum, Aytonia Evansti, P. eximium, P.
jamaicense, P. lanigerum, P. limbatum, Reboulia maderensis, and
P. mexicanum. For the privilege of studying this rich material
the writer is largely indebted to Dr. M. A. Howe, of the New York
Botanical Garden, Dr. W. G. Farlow, of Harvard University,
306 Evans: THE GENUS PLAGIOCHASMA
Dr. G. Lindau, of the University of Berlin, Professor C. R. Barnes
and Dr. W. J. G. Land, of the University of Chicago, and Miss
Caroline C. Haynes, of New York City. The collection belonging
to the New York Botanical Garden, which includes specimens from
the Mitten and Underwood herbaria, has proved particularly help-
ful. A series of unpublished drawings by Miss Haynes has like-
wise been of much assistance.
SHEFFIELD SCIENTIFIC SCHOOL,
ALE UNIVERSITY
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. Cavers, F. The Inter-relationships of the Bryophyta. New
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. Haynes, C. C. Two new species of Aytonia from Jamaica. Bull.
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to
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Evans: THE GENUS PLAGIOCHASMA 307
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6-8. New York. 1856.
Trevisan, V. Schema di una nuova classificazione delle epatiche.
Mem. R. Ist. Lomb. II. 4: 383-451. 1877
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chantiaceen. Bot. Zeit. 37: 729-743, 745-759. pl. 9. 1879.
INDEX TO AMERICAN BOTANICAL LITERATURE
1913-1915
e aim of this Index is to include all current botanical literature written by
Americans, published in America, or based upon American material ; the word Amer-
ica being used in the broadest sense.
Reviews, and papers that relate exclusively to forestry, agriculture, horticulture,
manufactured products of vegetable origin, or laboratory methods are no and
no attempt is made to index the literature of bacteriology. An occasional exception is
made in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not mentioned unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
*o errors or omissions, their since will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card. Selections of cards are not permitted ; each
subscriber must take all cards published during the term of his subscription, Corre-
spondence relating to the card issue should be addressed to the Treasurer of the Torrey
Botanical Club,
Ames, A. The temperature relations of some fungi causing storage
rots. Phytopathology 5: 11-19. F 1915
Arthur, J. C., & Fromme, F. D. A new North American Endophyllum.
Bull. Torrey Club 42: 55-61. pl. 2+f.1, 2. 2 Mr 1915.
Endophyllum tuberculatum sp. nov.
Ayres, J. A. Flower of Adenocaulon bicolor. Bot. Gaz. 59: 154-157.
Pott; 22. 17 F018,
Bailey, F. D. Notes on miscellaneous potato diseases. Oregon Agr.
Exp. Sta. Bien. Crop Pest & Hort. Rep. 2: 245-256. f. 20-28.
15 Ja 1915.
Baquedano, R. E. Nuestros bosques de Araucarias. Bol. Bosques.
Pesca i Caza. 2: 509-524. Ap 1914.
Barss, H. P. Bacterial gummosis or bacterial canker of cherries.
Oregon Agr. Exp. Sta. Bien. Crop Pest & Hort. Rep. 2: Bere
f. 13-19. 15 Ja 1915.
Barss, H. P. A new filbert disease in Oregon. Oregon Agr. Exp. Sta.
Bien. Crop Pest & Hort. Rep. 2: 213-223. f. 4-12. 15 Ja 1915.
Bartlett, H. H. Additional evidence of mutation in Oenothera. Bot.
Gaz. 59: 81-123. f. 1-17. 17 F 1915.
Bartlett, H. H. Mutation en masse. Am. Nat. 49: 129-139. f. I-9.
Mr 1915.
309
310 INDEX TO AMERICAN BOTANICAL LITERATURE
Benedict, R. C. Fern hats and fern cigar cases. Am. Fern Jour. 5:
7, 8. $1.2. Mr 191s.
Berry, E.W. The Mississippi river bluffs at Columbus and Hickman,
Kentucky, and their fossil flora. Proc. U. S. Nat. Museum 48:
293-303. pl. 12, 13. 28 Ja 1915.
Includes descriptions of Salix viminalifolia and Tecoma preradicans, spp. no
Boérgesen, F. The marine algae of the Danish West Indies. Part 2.
Phaeophyceae. Dansk. Bot. Arkiv 2: 1-66. f. I-44. 1914.
Bovie, W. T. The visible effects of the Schumann rays on protoplasm.
Bot. Gaz.-59: 149-153. 17 F 1915.
Britton, E. G. Mosses of Bermuda. Bull. Torrey Club 42: 71-76.
pl. 6,7. 2 Mr 1015.
Burd, J. S. The economic value of Pacific Coast kelps. Calif. Agr.
Exp. Sta. Bull. 248: 183-215. F 1915. [Illust.]
Chamberlain, C. J. A phylogenetic study of cycads. Proc. Nat.
Acad. Sci. 1: 86-90. 15 F 1915.
Conklin, E. G. Phenomena of inheritance. Pop. Sci. Mo. 85: 314-
837s te gO-50.. OO rotg.
Cook, O. F. Brachysm, a hereditary deformity of cotton and other
plants. Jour. Agr. Research 3: 387-400. pl. 53-62. 15 F 1915.
Copeland, E. B. Hawaiian ferns collected by M. l’Abbé U. Fauree.
Philip. Jour. Sci. g: ay 435-441. S 1914.
Includes seven new spec
Crocker, W., & Geieoks 4. F. A method of prophesying the life
duration of seeds. Proc. Nat. Acad. Sci. 1: 152-155. 15 Mr 1915.
ister A. Two new Mariposas. Bull. S. Calif. Acad. Sci. 14: 11,
2.3. B20 a Fore.
aes discolor and C. campestris, spp. nov.
Dendy, A. Progressive evolution and the origin of species. Am. Nat.
49: 149-182. Mr Iog15.
Derby, O. A. Illustrations of the stem structure of Tietea singularts.
Am. Jour. Sci. IV. 39: 251-260. f. I-zo ME ONS,
Dixon, H. N. New and rare Australasian mosses, mostly from Mitten’s
herbarium. Bull. Torrey Club 42: 93-110. pl. 9. 16 Mr 1915.
Includes descriptions of 10 new species.
Drayton, F. L. The Rhizoctonia lesions on potato stems. Phyto-
pathology §: 59-63. pl. 6 +f.5. F 1915.
East, E. M. An interpretation of self- -sterility. Proc. ‘Nat. Acad.
Sci. I: 95-100. 15 F 1915
Ehlers, J. H. The eapeeauase of leaves of Pinus in winter. Am.
Jour. Bot. 2: 32-70. f. 1-4. 18 F 19 gI5.
Farr, C. H. The origin of the inflorescences of Xanthium: Bot. Gaz.
59: 136-148. pl. ro. 17 F 1915. 2
INDEX TO AMERICAN BOTANICAL LITERATURE aa k
Fawcett, G. L. Fungus diseases of coffee in Porto Rico. Porto Rico
Agr. Exp. Sta. Bull. 17: 1-29. pl. 1-8. 9 F 1915.
Fawcett, H. S. The known distribution of Pythiacystis citrophthora
and its probable relation to mal di gomma of Citrus. Phyto-
pathology 5: 66,67. F 1915.
Frye, T. C., & Zeller, S. M. Hormiscia tetraciliata sp. nov. Puget
Sound Marine Sta. Publ. 1: 9-13. pl. 2. - 15 Ja 1915.
Gates, F. C. Notes from the tropical strand: Ipomoea pes-caprae and
Canavalia lineata. Torreya 15: 27, 28. f. 1, 2. 11 Mr 1915.
Gates, F. C. The pioneer vegetation of Taal volcano. Philip. Jour.
Sci. 9: (Bot.) 391-434. pl. 3-10. S 1914.
Hall, H. M. Notes on Baeria and Lasthenia. Bull. Torrey Club 42:
IlI-116. 16 Mr 1915.
Hall, H. M., & Yates, H. S. Stock poisoning plants of California.
Calif. Agr. Exp. Sta. Bull. 249: 219-247. f. 1-7. Mr 1915.
Hall, J.G. Notes upon Washington fungi. Phytopathology 5: 55-58.
l.
ApS Neottiospora yuccaeafolia and Tureenia juncoidea, spp. nov.
Hance, R.T. Pollen development and degeneration in Zebrina pendula,
with special reference to the chromosomes. Bull. Torrey Club 42:
63-70. pl. 3-5. 2 Mr 1915.
Hartley, C. Injury by disinfectants to seeds and roots in sandy soils.
U.S. Dept. Agr. Bull. 169: 1-35. pl. 1 +f. 1, 2. 20 F 1915.
Hartley, C., & Brunner, S.C. Notes on Rhizoctonia. Phytopathology
5: 73, 74. F 1915.
Hartley, C., & Merrill, T. C. Storm and drouth injury to foliage of
ornamental trees. Phytopathology 5: 20-29. f. 1-3. F 1915.
Hartwell, B. L., & Damon, S.C. The comparative effect on different
kinds of plants of liming an acid soil. Rhode Island Agr. Exp. Sta.
Bull. 160: 407-446. f. I-10. O 1914.
Heald, F. D., & Studhalter, R. A. Longevity of pycnospores and
ascospores of Endothia parasitica under artificial conditions. Phyto-
ieee 35-44. $1: 2) F -19TS.
«Je a otes on plants of the Chicago region. Torreya 15: 21-26.
11 Mri
Hoffer, G. a The more important fungi attacking forest trees in
Indiana. Indiana State Board Forestry Rep. 1914: 84-97. f. I-5-.
1915.
Hopkins, L. S. The ferns of Allegheny County, Pennsylvania, their
haunts and habits and something of their folklore. Bot. Soc. W.
Pennsylvania Publ. 3: 1-129. Au 1914. [Illust.]
312 INDEX TO AMERICAN BOTANICAL LITERATURE
Jackson, H. S. Notes, observations and minor investigations on
plant diseases. Oregon Agr. Exp. Sta. Bien. Crop Pest & Hort.
Rep. 2: 261-283. f. 30-44. 15 Ja 1915.
Jackson, H. S. A Pacific coast rust attacking pear, quince, etc.
Oregon Agr. Exp. Sta. Bien. Crop ba & Hort. Rep. 2: 204-212.
f. 1-3. 15 Ja 1915.
Johnson, J. Black rot, shed burn, and stem rot of tobacco. Wisconsin
Agr. Exp. Sta. Research Bull. 32: 63-86. f. 1-7. Je 1914.
Jones, L. R. Control of potato diseases in Wisconsin. Wisconsin
Agr. Exp. Sta. Circ. 52: 1-19. f. 1-4. N 1914.
Linnell, M. B. Wild and cultivated clovers of Ohio. Ohio Nat. 15:
443-448. F 1915.
Loeb, J., & Wasteneys, H. The identity of heliotropism in animals
and plants. Second note. Science II. 41: 328-330. f. z. 26 F
1915.
Longyear, B. O. Some Colorado mushrooms. Colorado Agr. Exp.
Sta. Bull. 201: 1-34. f. 1-25. N 1914.
Lutman, B. F., & Johnson, H. F. Some observations on ordinary beet
scab. Phytopathology 5: 30-34. f. 1-4. F 1915.
MacDougal, D. T. The Salton Sea. Am. Jour. Sci. IV. 39: 231-250.
f. 1-6. Mr 1915.
Mansfield, W. The structural variation of allspice. Practical
Druggist 33: 28, 29. f. 1-3. Mr 1915
Maxon, W. R. The North American species of Psilogramme. Bull.
Torrey Club 42: 79-86. 2 Mr 1915
Includes Psilogramme chiapensis, P. glaberrima, and P. villosula, spp. nov.
Maxon, W.R. Notes on American ferns: IX. Am. Fern Jour. 5: 1-4.
Mr 1915.
Maxon, W.R. Notholaena Aschenborniana and a related new species.
Am. Fern Jour. 5: 4-7. Mr 1915.
Includes Notholaena hyalina Maxon, sp. no
Maza, M. G. de la, & Roig, J. T. Flom de Cuba.
Sta. Bull. 22: 1-182. pl. 1-33. D 1914.
McClelland, C.K. Grasses and forage plants of Hawaii.
Exp. Sta. Bull. 36: 1-43. pl. 1-9. 20 F 10915.
Meinecke, E. P. Robert Hartig (1839-1901).
I-3. $1. 7... F-34915.
Merrill, E. D. Sertulum bontocense: new or interesting plants col-
lected in Bontoc subprovince, Luzon, by Father Morice Vanover-
bergh, II. Philip. Jour. Sci. 9: (Bot.) 443-459. S 1914.
Includes many new species and new combinations
Cuba Agron. Exp.
Hawaii Agr.
Phytopathology 5:
Meyer, R. Echinocactus macrodiscus Mart. var. multiflorus R. Mey.
var. nov. Monats. Kakteenk. 24: 150-154. 15 N 1914. [Illust.]
INDEX TO AMERICAN BOTANICAL LITERATURE 313
Meyer, R. Weiteres tiber Echinocactus horizonthalonius Lem. Monats.
Kakteenk. 24: 157. 15 N 1914.
Miles, F. C. A genetic and cytological study of certain types of
albinism in maize. Jour. Genetics 4: 193-214. pl. 8 +f. 1-9. 3F
I9I5. :
Mitchell, M. R. The embryo sac and embryo of Striga lutea. Bot.
Gaz. 59: 124-135. pl. 8, 9. 17 D 1915.
Moxley, G. L. Fern notes. Am. Fern Jour. 5: 9, 10. Mr 1915.
Newcomb, E. L. Belladonna and Hyoscyamus. Am. Jour. Pharmacy
86: 531-542. f. 1-5. D 1914.
O’Gara, P. J. The Pacific coast cedar rust of the apple, pear, quince
and related pome fruits caused by Gymnosporangium Blasdaleanum.
Office Path. U. S. Weather Bureau Sta. Rogue River Valley Bull.
Tech. Ser. 2: [1-7.] pl. r-4+f. 7. 1S 1913.
Parish, S. B. Notes on some southern California plants. Bull. S.
Calif. Acad. Sci. 14: 12-16. Ja 1915.
Parish, S. B. The Whitewater sands. Muhlenbergia 9: 133-139.
pl. TI-13.° 15 F 1915.
Peirce, G. J. The physiological aspects of California for the botanist.
Pop. Sci. Mo. 86: 270-273. Mr 1915.
Pierce, R. G. Chestnut blight in Nebraska. Phytopathology 5: 74.
F 1915.
Quehl, L. Mamillaria cephalophora Quehl spec. nov. Monats,
Kakteenk. 24: 158. 15 N 1914. [Illust.]
Rock, J. F. A new Hawaiian Cyanea. Bull. Torrey Club 42: 77, 78.
pl. 8. 2 Mr 1915.
Cyanea Larrisonii sp. nov.
Rogers, J. T., & Gravatt, G. F. Notes on the chestnut bark disease.
Phytopathology 5: 45-47. F 1915.
Rumbold,; C. Notes on chestnut fruits infected with the chestnut
blight fungus. ®Phytopathology 5: 64, 65. F 1915.
Shafer, J. A. Collecting in the mountain region of eastern Porto
Rico. Jour. N. Y. Bot. Gard. 16: 33-35. 28 F 1915.
Shaw, J. K. The technical description of apples. Massachusetts
Agr. Exp. Sta. Bull. 159: 73-90. f. 3-14. D 1914.
Sinnott, E. W., & Bailey, I. W. Investigations on the phylogeny of
the Angiosperms. 5. Foliar evidence as to the ancestry and early
climatic environment of the Angiosperms. Am. Jour. Bot. 2: 1-22.
pl. 1-4. 18 F igts.
Steil, W. N. Some new cases of apogamy in ferns. Preliminary note.
Science II. 41: 293, 294. 19 F 1915.
314 INDEX TO AMERICAN BOTANICAL LITERATURE
Stone, R. E. The life history of Ascochyta on some leguminous plants,
—II. Phytopathology 5: 4-10. f. 7. F 1915.
: Includes description. of Mycosphaerella ontarioensis sp. nov.
Taubenhaus, J. J. The problem of plant diseases which confronts the
gardener. Gard. Chron. Am. 17: 301-304. Ja 1913. [Illust.]
Taylor, N: The growth-forms of the flora of New York and vicinity.
Am. Jour. Bot. 2: 23-31. Ja 1915
Tolaas, A. G. A bacterial disease of cultivated mushrooms. Phyto-
pathology 5: 51-54. pl. 3. F 1915.
Tracy, H.H. Notes on Jsoétes. Am. Fern Jour. 5: 12, 13. Mr 1915.
Tufts, W. P. An inquiry into the nature of a somatic segregation of
characters in the Le Conte pear. Oregon Agr. Exp. Sta. Bull. 123:
$16. p1.. £,.:21 . D 1914.
Van Horne, A. Some rare fungi found at St. Andrews. Canadian
Record Sci. 9: 328-338. Ap 1914. |
Victorin, M. Random botanical notes from Portneuf County, Que.
Ottawa Nat. 28: 155-160. f. 1-5. 27 F 1915.
Weingart, W. Phyllocactus ‘‘ Carl Rettig’? Weing. hybr: nov. Monats.
Kakteenk. 24: 145, 146. 15 N 1914.
Weir, J. R. A new host for a ‘species of Razoumofskya. Phyto-
. > pathology 5: 73. F 1915.
Weir, J. R. New hosts for some forest tree se ak Phytopathology 5:
Fie 92. 1 SOtR,
Weir, J. R. Some observations on abortive sporophores of wood-
destroying fungi. Phytopathology 5: 48-50. F 1915.
Wilcox, E. V. Report of the Hawaii agricultural experimental station
1914; 1-73. $1.43: 20 F 10s.
Includes considerable information of a botanical nature
Winslow, E. J. Notes on Maine ferns. Am. Fern Jour. 5: 13, 14.
Mr 1915.
VOLUME 42, PLATE I2
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POLYGONATUM CANALICULATUM (MunL.) PursH B. POLYGONATUM C oe ee AMERICANUM
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POLYGONATUM CANALICULATUM GIGANTEUM (Dretr.) FARWELL
Vols AE gon No. 6
BULLETIN
OF THE
TORREY BOTANICAL CLUB
ee re ee
JUNE, 1915
Some new species and varieties of Bihai*
ROBERT F. GRIGGS
(WITH PLATE I9 AND SIX TEXT FIGURES)
For a number of years the writer has contemplated publishing
a revision of the musaceous genus Bihai. Inasmuch as this has
been much delayed, it has been thought wise not to withhold the
publication of the several new species which have come to light
in the course of the work. In order to enable the student rightly
to place the new species among the old a key to all the species
is also given. The preparation of this in turn has involved the
revision of the subgenera and minor groups which is herewith
presented.
Intrageneric Relationships.—In previous treatments the species
have been grouped under two subgenera, which under somewhat
varied characterizations included respectively the large species
and the small species. The first of these to receive a name,
Taentostrobus, was founded by Kuntze on his B. imbricata. Baker,
however, ignored Kuntze’s name and included this species under
his own subgenus Platychlamys, with B. Bihai as type, which was
taken as synonymous with TJaeniostrobus by Schumann and
replaced by it. The narrow bracted species were thrown together
by Baker under the name Stenochlamys (type B. psittacorum) and
this was adopted by Schumann. While one recognizes a more or
less natural subdivision of the genus around these types there
are no characters by which it can be so divided without tearing
* Contribution from the Botanical Laboratory of the Ohio State University,
oO. .
[The BULLETIN for May (42: 247-314. pl. 12-18) was issued May 22, 1915.]
315
316 Griccs: SOME NEW SPECIES OF BIHAI
asunder closely related species, such for example as B. longa, B.
pendula, and B. marginata, which though very closely related,
differ in size and width of the bracts so much that they are neces-
sarily thrown into different subgenera by the key characters.
As one goes through the genus he finds so many difficulties of this
sort that it becomes clear that some other method of subdivision
is necessary. A new classification is therefore proposed which
while retaining the old subgenera as recognizable primary sub-
divisions, even if they are somewhat indefinitely characterized,
is mainly built around smaller groups of rather closely similar
species. The names of these groups have been taken from the
specific names of characteristic species, indicating therefore cate-
gories of lower rank than the subgenera with their substantive
designations. Before proceeding to the key it may be well to
outline this proposed classification.
TAENIOSTROBUS
EPIscoPALES, including only B. episcopalis.
IMBRICATAE, including B. conferta, B. Wagneriana, B. imbricata,
B. densa, B. reticulata, B. punicea, B. Mariae, and B. curtispatha
(intermediate). |
PENDULAE, including B. rostrata, B. longa, B. revoluta, B. pendula,
.B. Collinsiana, B. villosa, B. platystachys, and B. marginata.
CHAMPNEIANAE, including B. Bihai, B. Champneiana, B. barqueta,
B. borinquena, B. rutila, B. purpurea, B. stricta, and B. humilis.
STENOCHLAMYS
DISTANTEs, including B. velligera, B. metallica, B. spissa, B. ee
_antha, B. tortuosa, B. geniculata, B. librata, B. adflexa, B.
- lingulata, B. distans, B. latispatha, B. brasiliensis, B. acuminata,
and B. Burchellw.
CANNOIDEAE, including B. glauca, B. densiflora, B. angusta, B.
crassa, B. aurantiaca, B. straminea, B. Harrisiana, B. hirsuta,
and B. psittacorum.
Key to the species of Bihai
I. Branch-bracts closely imbricated forming a sere — OG yet Ue gee ae
ba ea008 standing apart, distant or gseake flow :
2. Bracts persistent (IMBRICATAE)
3t..
ee ee eee as Pre 3°
GrRIGGS: SOME NEW SPECIES OF BIHAI Olt
ty
. Bracts deciduous, strictly ascending, scarlet with yellow margins.
B. episcopalis (Vellozo) Kuntze.
Rueseiprescence 25 cml, broad at basesois Gi ke ee es
ee PerCOMUADOUL £5: Cio MTOR eS ee a ae 5.
4. Rachis straight, leaves rounded to the base...B. conferta (Peters.) Kuntze.
4. Rachis somewhat flexuose, leaves cuneate, very ¥ sedialeth ral.
B. Wagneriana (Peters.) Kuntze.
Noe meoreacciice fect. Or: suberect. 2 eo oc, eae 6.
5. Inflorescence gsonauied ate hu Ne ane Shi hy a EL SNe te ee ea | 8.
eaves Sreen, iniorescence peduncled.. 5. x A en Se a
6. Veins and veinlets 3 uously marked with red, inflorescence sessile, close to
fue wound, bem and: Yellow Se ee B. reticulata Griggs.
- 7. Inflorescence 15 cm. wide, bracts 10 cm. long............ B. imbricata Kuntze
g
7. Inflorescence scarcely exceeding 10 cm. in width, middle bracts 4—5 cm. lon
‘ . densa Goa.
ay Bracts suborbicular, widest at the middle, rose red, plant very large.........
~>8, Bracts ovate, widest below the middle, medium sized plants................
9. orescence deltoid, somewhat loose, red purple
a aa bl de le
Sa Se aE B, punicea pe
9. ne. Cense, Scarlet: iis 6 oo ee oy ee B. densa Griggs.
‘To. ely imbricated, concealing the rachis, flowers rose colored where
Rieke ins en Pee cca Me pce B. Mariae pores .) Kuntze.
Io, sivaey eachicig only at the middle, not concealing the rachis, flowers yellow.
B. sc RNR (Peters.) Kuntze.
Tx: Inlorescence pendent, otis
at Inflorescenc Bi lene vn et ee eS ek oh) ame 20.
12. Bracts feat. pra as wide as ne hee When tattered: OG. 2 oo eee ae
i ba. bracts Harrow, mot. more than half.as wide ag long 3. oe Se ek 15.
13. Bracts rhomboid, widest at the pe abe close and slightly im
B. ciao (Peers ) Kuntze.
“B. . rostrata (Ruiz & Pavon) Griggs.
e, Scarlet :, bright rufous pubescent within,
nt very large...... paren B. longa oe
Cer ee a oe ah ae eee Nera eee et ee a ae ee
a re ee ee ee ed
ry: Glabrous throughout, Hachis aise straight, leaves nevi glaucous
stiasteiie Griggs. —
; aor een and Sinick tein i villous with aie nae rufous hair, flowers
: ee a ey ee B. villosa (Klotzsch) Kuntze.
18. Inflo orescence mescty. puberulent i 7 ah pm A he . Baa os ehh cig ie =o.
19. Plant larce ’ 4041 L rel i yw x am &
SB. Platystachys (Baker) Griggs.
B. + marginata See
318 GRrIGGS: SOME NEW SPECIES OF BIHAI
. Leaves green or glaucous, bracts colored
. Leaves purple beneath, bracts green... B. metallica (Planch. & Linden) Kuntz
. Inflorescence persistently hairy
. Inflorescence ones unless when yo
20. Rachis about 2 cm. in diameter, bracts glabrous, thick and fleshy, shriveling much
ta diteiam UmARIAMARY | Ses os ok ee ee ee 21
eae BO Rachie seldom exceeding 1 cm. in diameter, bracts mostly chartaceous, SS
ai acs long-peduncled, scarlet... 2. 2.62. bees B. Bihai (L.) cis
PD. RREGIEDONO® SRRGHO 5.50580) eis. oe oc eA a ca Meme 2G ee ae RE 2.
22. Bracts of o OE oe a Eee Ue fe als Way ee a hae ee ey 23.
ar raed Gase Oat 6 See Cee at ie ag or a eh eee ea gw 6.
23. Flower with a subgibbous expansion on the anterior side, filaments strongly
ene petals of nearly the same texture asthe sepals..............-+-+-. a5,
23. Flower merely arcuate, petals very delicate, shorter than the sepals.......-.. 24.
24 irifiorescence oranie. 0 ae ee B. Cham pneiana Griggs.
94. Inflorescence Tose colored ooo a eek, Ae B. barqueta (Loes) Griggs.
Os. InGoredrente Grange. os Sn a. ee ae ee ee B. boringuena rete
as. Inflovescente ecatleti. i 6c oie B. borinquena coccinea Grig
26. Bracts margined with seuiny sets eR Ok rede ea 5 i ae dona ig oso ah eae es yw Wane “g
BG. Brocts red and sPPeni sin oa Se a eh es eas ss dae nee wae 28.
27. Bracts red and yellow, narrowly ovate, about 6 cm. wide, barely touching each
other, perianth 5 cm. long, posterior sepal broadly lanceolate. . B. rutila Griggs.
. Bracts purple and — 8-10 em. wide, close together, perianth segments 6
JONES nearsobions: 3 ya ee a Pe B. purpurea Griggs.
é pon scarlet with green pastes, few, plant rather small (2 m.), leaves scarcely
exceeding I m
be a Rie gE: ee aie ew le dle Ae NCR R mw Weel bo eee eS lelce wee ele BLS) eye eo oO ene re
. Bracts green with a patch of red on the cheeks, more numerous, plant up to 4 m
ves elongate, cuneate-oblong............... B. elongata ESAS
tall, lea
. Bracts very broad, 6 cm. deep, plant about 1.5 m. tall, flowers tipped with white.
stricta (Huber) Griggs.
. Bracts about 3 cm. deep, larger, flowers colorless baton: green where exposed.
B. humilis (Jacq.) Griggs.
; Mediqat sized plants, leaves commonly surpassing 50 cm., habit musoid or si
DISTANTES)
PSE SSeS Re Pie ee eke ee pm iy Oe wee ee lee wee ele hoa ie! Bee (8 ke I.
S at ae commonly about a meter tall, if taller very slender, leaves scat
we cm. long, habit isis (CANNOIDEAE)
villous
: setts thickly covered with very long hair.
B. velligera (Poepp.) ies
ihe oe ede ee AC TES a Ge ae ey at ar eee Cet le apres
ate a et ek tet Wien te ee tag i ee Ree ee tala gre mae Uae Berra are
PRR RR ee ALOT erase Se eae ee lie Atay a wet ee
fe ee ae ee 6 eee ete le eee ae Ew ee et el eye ee mee We we miay preity (pe cates he ale Ok Pe
Bracts solid
red
. Bracts green with red cheeks, flowers yellow.
B. dasyantha (Koch. & Bouché) Kuntze.
exserted
. Bracts ovate-lanceolate, flowers green, only their tips
GRIGGS: SOME NEW SPECIES OF BIHAI 319
36. Bracts narrowly lanceolate, flowers yellow, geniculate, standing out of the
bracts at SUMBs en Re he eat ae B. geniculata Griggs.
39: sa slightly concave, i CURET TREVOR nt pe ae eee 38.
Pome Oe s boat-shaped, not EEO ae Pa eg rte or ee oa ee 30.
38
: pee) bracts a. 15 cm. long, yellow with peratet
Bit
daaiéla (Ruiz & Pavon) Griggs.
. Middle bracts about 7 cm. long, scarlet, bach is long eeignee
ad ae Griggs.
39. Perianth 15-20 mm. long, inflorescence deltoid, compact.. librata Griggs.
~ 39. Perianth 30-60 mm. long, inflorescence oblong, rather lax................. 40
aR TROT CRCETCE OPENED a rt aay ch es Su eS Dee eae AI.
40. srvinbbt ees Re ea el AW 3 ha ele od Ce eS Co Cee 42.
41: F. ted, rachis very flexuose, bracts, at least the sat strongly falcate.
B. distans Griggs.
- 41. Flowers and their bracts entirely included, rachis and bracts nearly straight.
B. latispatha (Benth.) Griggs.
Wor Bias seaciine Fs i IONth ai eRe es hoe © a es 43
42. Bracts 5 cm. long, all deflexed, flowers golden yellow.
B. pus ed eacoeel! Griggs.
1
. Flowers atoie>t or slightly arcuat tcl erected . 44.
. Perianth geniculate at anthesis, flowers numerous but opening one or two at a
time, appressed to the channel of the bracts un til near anthesis, then quickly
raised by a bend in the pedicel, bracts scarlet, horizontal, three ranked, of
WGP aiveiee aes ae Soe ee B. geniculata Griggs.
44. Margins of bracts not involute, flowers reddish (or yellow), habit cannoid.
B. acuminata (Rich.) Kuntze.
44. Margins of the bracts involute, all arcuat di owers greenish white. . 45.
45. Leaves ese beneath 6 Gee oS B. bekd tliensis (Hook.) Kuntze
45. Leaves glaucous.) .0 2.04 .. ps B. brasiliensis pulverulenta (Hook.) trices,
46. Bracts as signi colored as the rachis, leaves green on both sides..........
47
. Bracts and flowers yellowish green, rachis and pedicels red, leaves glaucous.
46
B. glauca (Poit.) Kuntze.
ays Bracts exceeding 10cm. in length... 25. a a es Ce 48.
—~47- ad pers vers ie: pen saraenaiel teachinie 10 (mio eo Se 5 50.
48 P g ot involute t ti 1 with black . 49.
ae: y ling at fi ing , involute and hevonek pointed, flowers
tipped with) black. : 60.06 fe Reo B. densiflora (Verlot) Kuntze.
. Leave 8 narrowly oblong, flowers white, pedicels orange.
B. angusta (Vellozo) Griggs.
49. Leaves éiliptical: flowers reddish (or yellowish), ee red.
B. acuminata (Rich.) Kuntze.
-~ ‘go. Flowers not darkened at the tip, dances Do a ee as 51.
50. Flowers maculate.............. Ga Ld ae ek eer ees =
woe BPS SPRATT) eee kt i ee ay ee Os A
-51. Bracts orange, leaves mostly etic’ Bes Sh Oy Gere e cake ye B. crassa Pi
~ 52. Bracts somewhat ascending, — green, leaves about 30 cm. long, sessile,
52.
subcordate, peduncle short, flowers pale yellow.
te dasaalincs (Ghiesb.) Kuntze.
Bracts all deflexed, ie reaching 60 cm., Sees at the base, peduncle elon-
ones, flowers golden - yellow...... aoe aes B. Burchellii (Baker) Griggs.
320 GRIGGS: SOME NEW SPECIES OF BIHAI
53- Inflorescence brightly colored.............6.-- sees tees e etter rere 54.
53. Inflorescence greenish yellow, pubescent............... B. straminea Griggs.
54. Bracta not patticolored. ...- .-. . 2. a ewe eee eee eee ere es 55
54. Bracts scarlet above, greenish yellow below........... B. Harrisiana Griggs.
55. Bracts scarlet, concave, acute
55. Bracts orange, revolute and scarcely concave, obtusish.
B. psittacorum (L.f.) Kuntze.
56, Inflorescence pubescent... 22003. 0. ee eee ee se B. hirsuta (L. f.) Griggs.
56. Inflorescence glabrous............-..- ....B. hirsuta cannoidea (Rich.) Griggs.
Fic. 1. Bihai densa Griggs; three-eighths natural size.
Bihai densa sp. nov.
Plant of medium size, petiole 70 cm. long, leaf 80 cm. long,
30 cm. wide, acute, subcordate at the base, thin and tough, main
veins 5-10 mm. apart, green and glabrous except the midrib and
petiole which are pilose with projecting ferruginous hairs about
1 mm. in length; inflorescence erect (?), 18 cm. high, half as wide,
oblong, nearly 1 cm. thick (dry), nearly glabrous, peduncle 20 cm.
long in the type, somewhat shriveled but 1 cm. thick, rachis nearly
straight, concealed, except at the very base, stiff; bracts ascending,
fifteen in the type, scarlet, thick and firm, shriveling little, broadly
ovate, rostrate, inclined to be involute at th i ,
Le . e tips, the lowest
standing somewhat apart, the rest closely imbricated, middle
GRIGGS: SOME NEW SPECIES OF BIHAI 321
bracts 4-5 cm. long, 5-6 cm. wide, widest near the base; flowers
about a dozen to a bract, only their tips visible, bracteoles delicate,
o X12 mm., flower very small, perianth 20-25 mm., spa
pubescent, thick, indurated at the tips, the posterior 2-3 m
longer than the other floral parts, petals 3, pedicels finally Aas
12 mm., berries 9 mm. in diameter (dry) concealed within the.
bracts. [TEXT FIG
PANAMA: shied Dok Bocas, Rio Faté Valley, province of
Colon, altitude 40-80 m., in dark shady forest, August 16, 1911,
H. Pitter. Type in National Herbarium, sheet 678445.
Bihai densa is nearest B. punicea from which it differs especially
in the shape and color of the bracts, and the flower characters.
The only one of the previously known species which could be
mistaken for either is B. imbricata.
Fic. 2. Bihai punicea Griggs; three-eighths natural size.
Bihai punicea sp. nov.
Plant of medium size, leaf about 1 m. long, 25 cm. wide, green
and glabrous, acute; inflorescence pendent, deltoid, about 12 cm.
wide, a little longer, peduncle 30 cm. long, 7 mm. thick (dry),
322 Griccs: SOME NEW SPECIES. OF BIHAI
glabrous, rachis nearly straight, stiff, glabrescent, concealed above
but visible below, bracts twenty-three in the type, mostly hori-
zontal, red-purple, covered with tawny hair, boat-shaped, ovate
when opened out, widest near the base, slightly arcuate, very
obtuse, except the lower, those from the middle of the inflorescence
about 5 cm. long, of the same breadth, the lowest 14 cm., the high-
est only about 1.5 cm.; flowers a little shorter than the bracts, less
numerous than in allied species, bracteoles ample, 40 X 25 mm.,
obtuse and frayed at the tip, persistent, flowers curved up out of
the bracts toward the large sepal, perianth about 3 cm. long, lat-
eral sepals free, corolla limb tridentate, the middle tooth smaller
than the other two. [TEXT FIG. 2.]
PANAMA: between Gorgona and Gatun, altitude 10-50 m.,
Smithsonian Biological Survey, January 7, 1911, H. Pittier 2290.
In National Herbarium, sheets 677161, 676544, 676545.
Bihai revoluta sp. nov.
Plant of medium size, leaf (known to me only from a sucker)
elliptical, 50 X 15 cm., acuminate, rounded to the slightly de-
current base, green and glabrous; inflorescence pendent, red,
rachis straight, flexible, internodes mostly about 3 cm. long,
covered with abundant evanescent ferruginous tomentum, bracts
numerous, horizontally divaricate, more or less tomentose at the
base, linear-oblong, obtuse, slightly concave, their margins con-
spicuously revolute soon after opening, middle ones about 12 cm.
long, 2 cm. wide: bracteoles persistent, about I X 5 cm., glabrous,
flowers rather numerous, pedicel and ovulary red, perianth yellow,
glabrous, about 5 cm. long, sepals thick, free, petals two (in the
type), shorter by 2-3 mm. and much more delicate than the sepals.
[TEXT FIG. 3.]
Fic. 3. Bihai revoluta Griggs; about half natural size.
Cotomp1a: Santa Marta, 1898-1901, H. H. Smith 2315, with
the following field note: ‘‘Thickets and forest near streams, 3,500-
5,000 feet. Bract and pedicel red, ayers! yellow. Specimen is
a from nn eae 3,500 feet, March 1." Type in the herbarium
GRIGGS: SOME NEW SPECIES OF BIHAI 323
of the New York Botanical Garden; specimens seen also from the
Missouri Botanical Garden and from the National Herbarium.
The following station may likewise be recorded: Orizaba, MExIco,
Botteri 523.
The narrow revolute bracts of this species at once differentiate
it from all other known species except B. lingulata and B. adflexa.
There is, however, no close resemblance between the stiff erect
inflorescence of B. adflexa and the lax panicle of the present species.
The resemblance to B. lingulata is much closer, but that species
has yellow bracts and a compact inflorescence with internodes only
one third as long.
Fic. 4. Bihai marginata Griggs; about three-sevenths natural size.
Bihai marginata sp. nov.
Plant about 4 m. tall, trunk 6.cm. in diameter at base, petiole
50 cm. long, leaf blade 60 X 15 cm., oblong, acute, rounded at the
324 GRrIGGS: SOME NEW SPECIES OF BIHAI
base, green and glabrous on both sides; inflorescence pendent, less
than 20 cm. long in the type, 10-12 cm. wide, exposed portion of
the peduncle less than 10 cm. long, slender, about 5 mm. in diam-
eter, softly puberulent with light brown hair, rachis strongly flex-
uose in smooth curves, stiff, minutely puberulent, internodes about
25 mm. long, bracts eight in the type, all reflexed, scarlet with yel-
low margins, somewhat puberulent, shallowly boat-shaped, ovate-
lanceolate when flattened out, gradually narrowed to an obtuse tip,
those in the middle of the inflorescence 8-9 cm. long, 4 cm. wide;
flowers rather numerous, the younger pushing the ripening fruit
out beyond the edges of the branch bracts, bracteoles 60 X 15 mm.,
triangular-lanceolate, very acute, pubescent on the back, persistent
in fruit, perianth 5 cm. long, curved toward the larger sepal, lipped,
sepals all free, corolla limb three-toothed, middle tooth somewhat
smaller than the others, pedicel about 15 mm. long in fruit, berry
(apparently ripe) oblong, 15 mm. long, half as wide. [TEXT
FIG. 4.]
PANAMA: Marraganti and vicinity, 3-60 m. altitude, April 3-9,
1908, R. S. Williams 696. Type in the herbarium of the New
York Botanical Garden.
Bihai barqueta (Loes) comb. nov.
Heliconia barqueta Loes, Verh. Bot. Verein. Brandenburg 51:
18. 1909. ‘‘Mexico in prov. Vera Cruz in distr. Jalapa ad
fluvii ripam. in coffeeto Sel. n. 3609 Flor. Dec.”
Closest to B. Champneiana from which it differs in the rose-
colored inflorescence whose bracts are more distant than in that
species. While this plant would appear to be safely distinct from
B. Champneiana the relations found to exist between the orange
and red bracted forms in Jamaica (see below) makes it desirable
to have information concerning the flower characters not furnished
by the description since recent studies have shown that these
characters have great diagnostic value.
Bihai borinquena coccinea Griggs & Harris, var. nov.
Closely similar to B. borinquena in the floral parts but differing
in the brilliant scarlet-crimson bracts which are numerous and
somewhat narrower than in the typical form of the species.
Jamaica: cultivated at Hope Gardens, W. Harris. Pressed
specimens in the herbarium of the New York Botanical Garden.
It is of interest to add in this connection that the orange-
GRIGGS: SOME NEW SPECIES OF BIHAI oan
bracted Bihai borinquena is common in Jamaica along with the
red-fruited variety. This Jamaican plant differs but slightly
from the Porto Rican type. From it, thanks to formalin specimens
and very beautiful colored drawings made by Mr. Harris’s
assistant, Miss H. A. Wood, I am able to describe the flower which,
by a typographical error, was stated to be 11 cm. long in the
original description:
Flowers about fifteen, bracteoles yellow, firm when fresh, ovate-
triangular, enveloping the flower; ovulary yellow, perianth 6 cm.
long, white below, green on the exposed tips, bent somewhat
toward the posterior sepal, with a conspicuous subgibbous expan-
sion on the anterior side, filaments conspicuously bent into the
expansion, perianth segments free above but partially fused below,
staminode about 3 cm. long, reaching to the bend in the filaments;
berry dark blue, ro X 13 mm., pedicel stout, fleshy, ringed, about
25 mm. long at maturity, seed 10 X 4 mm., tubercled, horny.
Bihai stricta (Huber) comb. nov.
Heliconia stricta Huber, Bol. Mus. Goeldi 4: 543. 1906.
Type LOcALItTy: “Esta especie é bastante frequente nas fraldas
do Cerro de Couchahuya; ella é cultivado no horto Fs gece do
Museu Goeldi.”
Bihai stricta is evidently closest to B. humilis but differs in the
much wider branch-bracts.
Bihai adflexa sp. nov.
Plant of,medium size, leaves 60-80 cm. long, 15-20 cm. wide,
oblong, narrowed to the base, tip not seen, green and glabrous on
both sides, main veins prominent, 5-7 mm. apart: inflorescence
deltoid, stiffly erect, peduncle apparently elongated, about 8 mm
in diameter, glabrous, rachis stiffly zigzag, puberulent, middle in-
ternodes about 15 mm. long, bracts twelve in the type, all ascend-
ing, scarlet, somewhat puberulent on the back, narrow, linear-
oblong, obtuse, revolute, little concave, strongly bent upward
near the base, whence the name, middle bracts about 7 cm. long,
scarcely 2 cm. wide, the upper much smaller, the lower much
larger: flowers numerous, bracteoles persistent, 4 X I cm., with a
thin evanescent pubescence both within and without; perianth 4
cm. long, sparingly pubescent like the bracteoles, sepals free,
corolla limb shige toothed. [TEXT FIG. 5.]
GUATEMALA: Coban, altitude 1,600 m., in deep forest, June
326 GRIGGS: SOME NEW SPECIES OF BIHAI
1908, H. von Tuerckheim II 2356. Type in the herbarium of the
New York Botanical Garden.
The very narrow revolute bracts at once distinguish B. adflexa
from all other species of the genus except B. lingulata and B.
revoluta.
. Uy
ANS Ni
\ tig A
Fic. 5. Bihai adfexe Griggs; about half natural size.
There is small likelihood of its being confused with B. revoluta
with its elongated, lax, pendent inflorescence. The resemblance
to B. lingulata is closer but that species has larger, yellow inflo-
rescence with wider bracts. It is, moreover, Peruvian.
Bihai brasiliensis pulverulenta (Lindl.) var. nov.
Heliconia pulverulenta Lindl. Bot. Reg. 19: ~ 1648. 1833.
ook. Bot. Mag. 78: pl. 4685. 1852.
Bthaia pulverulenta Kuntze, Rev. Gen. Pl. 2: 685. 1891.
GRIGGS: SOME NEW SPECIES OF BIBAI 327
In every detail except its slightly inferior size and its heavily
farinose leaves B. pulverulenta corresponds precisely with B.
brasiliensis. Although recognizing that glaucescence was some-
times evanescent and variable in amount, I have heretofore sup-
posed that marked differences in this matter were sufficient
criteria of specific‘distinctness. But the correspondence between
these two species is so exact in every other respect that one has
no other recourse than to throw them together. A specimen
brought back from Rio de Janeiro by the United States Exploring
expedition under Captain Wilkes is very interesting in this connec-
tion. It has the typical inflorescence of the two species; some of
the leaves are cordate but others are narrowed to the base; some
of them were clearly green and glabrous but others show evident
signs of a heavy pulverulence which has been rubbed off except
in a few protected spots! The reason that the similarity of the
two plants was not recognized long ago is that glaucescence was
made a primary character in the key so that they were never
put close enough together to invite comparison.
Bihai angusta (Vellozo) comb. nov.
Heliconia angusta Vellozo, FL Flum. Text 107. 1825. Plates 3:
20. 1827.
Heliconia bicolor Benth. Maund’s Botanist 3: pi. ror. No date,
1837-1840? Regel, Gartenflora 5: 289. pl. 172. 1856.
Heliconia angustifolia Hook. Bot. Mag. 75: pl. 4475. 1849.
Bihai straminea sp. nov.
Plant rather small, erect, habit cannoid, leaves reaching 35 cm.
in length and 10 cm. in breadth, smaller in the type, the upper, at
least, sessile, elliptical, acuminate, round at the base, glabrous,
green on both sides; inflorescence erect, pale greenish yellow,
eure glabrous, about 18 cm. long in rs material seen, 2-3
thick, rachis somewhat zigzag, minutely puberulent, inter-
ais 10-15 mm.; bracts half a dozen or less, ascending, glabrous
unless at the hake. narrowly triangular, concave and boat-shaped,
straight sided, witlest at the base, acute, 3-10 cm. long, the largest
about 12 mm. broad, tips of the lowest exceeding the uppermost:
‘flowers few, bracteoles about 15 mm. long, less than 5 mm. wide,
caducous after anthesis, glabrescent; pedicel, ovulary and calyx
thickly and softly pubescent, perianth about 3 cm. long, with a
328 GRIGGS: SOME NEW SPECIES OF BIHAI
dark spot near the tip, strongly bent toward the large sepal some-
what as in B. geniculata, large sepal rolled back in flower, lateral
sepals almost completely fused with the corolla or free, pedicel
about 15 mm. (not mature). [TEXT FIG. 6.]
Fic. 6. Bihai straminea Griggs; about half natural size.
PANAMA: dry wooded hills around Alhajuila, prov. Panama,
altitude 30-100 m. (Smithsonian Survey), January II, 1911, #7.
Pittier 2328 (type in the National Herbarium, sheet 677138);
between Miraflores and Corazal, Canal Zone, altitude 20-30 m.
(Smithsonian Survey), January 3, 1911, H. Pittier 2200.
Bihai Harrisiana sp. nov.
Erect, up to 4 m. tall, habit cannoid, leaves sessile, up to
30 cm. long and 11 cm. broad, broadly elliptical, acuminate, round
GRIGGS: SOME NEW SPECIES OF BIHAI 329
at the base, green and glabrous: inflorescence erect, glabrous,
peduncle of variable length, rachis somewhat flexuose, internodes
about 15 mm. long, bracts few, the lowest fertile one asp ing
10 cm. long, the upper about 6 cm., all about 15 mm. wi
narrowly lanceolate, obtuse or some of them acute, be ienibied.
upper half bright crimson fading into greenish yellow below;
flowers few, bracteoles about 20 X 7 mm., early deciduous, peri-
anth 25-30 mm. long, posterior sepal red, fading into orange on
the edges and tip, conspicuously spotted with black at the tip,
lateral sepals free, red, petals two, sometimes with the rudiment
of a third, red, fading into green on the tip, pedicels 10-15 mm. long.
[PLATE 19.] :
Jamaica: Leicesterfield, Upper Clarendon, W. Harris 10841,
February 28, 1910; with the field note, ‘1,800 ft. alt. An erect
growing plant with reed like stems up to 12 feet high, basal third
of stems leafless, lower half of bracts orange yellow, upper half
bright crimson, rhizome stout, creeping, flowers scarlet, tipped
with green and white.” (Type in the herbaria of the Ohio State
University and the New York Botanical Garden; also cultivated
in Hope Gardens, Jamaica.) ‘‘Jamaica, Wilson’’! in Herb. Gray.
In addition to the information contained on the label Mr.
Harris has favored me with a letter describing the habitat of the
plant in greater detail as follows: ‘With regard to the new species
from the Upper Clarendon I might say that it is fairly plentiful—
usually in light shade but sometimes in the open. | When growing
in shade or partial shade the stems are usually twice as high as
those growing in open situations. The most luxuriant clumps
I saw were growing in partial shade near the edges of small streams,
whilst plants in the open were yellowish in appearance and did
not look happy. It is evidently a shade and moisture-loving plant.
‘“‘T saw some old stems this year that were fully 2 or 3 feet
higher than the specimens measured by me last year, and it would
be correct to give the height as ‘up to 12 feet or over’ and the
habitat might be given as the ‘upper reaches of the Rio Minho
Valley, Upper Clarendon, 1,200 to 2,000 feet altitude.’ ”’
It is a great pleasure to be able to associate with this beautiful
species the name of so able and indefatigable a botanist as William
Harris.
A dried specimen of B. Partiden would not ordinarily be
distinguished from B. hirsuta cannoidea but in well dried material
330 GRIGGS: SOME NEW SPECIES OF BIHAI
it is evident that the scarlet of the ends of the bracts does not
extend below the middle. The leaves, too, are larger and more
broadly oval than in any of the specimens of the variety cannoidea
(or related species) which have come under my observation while
the stature far exceeds anything else recorded in the Cannoideae.
Bihai hirsuta (L.f.) comb. nov.
Heliconia hirsuta L.£. Suppl. 158. 1781.
Type LocaLity: “ Habitat in America Meridionali.”’
IMPERFECTLY KNOWN SPECIES
The following species have also been described recently, but the
descriptions given are so lacking in diagnostic characters as to
make it inadvisable to assign them places in the key:
HELICONIA ROBUSTA Pax; Fedde, Rep. Nov. Sp. 7: 107. 1909
TyPE LocALity: “ Bolivien Walder um San Carlos bei Mapiri
700 m. (Buctien n. 1267 flowered July 1907).”
This plant is evidently one of the Pendulae but the description
is inadequate either to differentiate it from previously described
species or to identify it with them.
HELICONIA PEARCE! Rusby, Bull. N. Y. Bot. Garden 6: 494. 1910
“Probably at Santa Ana’’ (Bolivia).
Said to be “about intermediate between H. psittacornu
[psittacorum|] and H. brasiliensis.’ From the description, however,
it would appear almost identical with B. acuminata.
Explanation of plate 19
Briar HARRISIANA Griggs
The figures, natural size, were drawn from specimens flowering in the Hope
Gardens, Kingston, Jamaica, by Mr. Harris’s assistant, Miss H. A. Wood.
The ferns and flowering plants of Nantucket—XV
EUGENE P. BICKNELL
SOLANACEAE
PHYSALODES PHYSALODES (L.) Britton.
Nicandra physalodes (L.) Pers.
Waste lots and yards on the eastern side of the town. Still in
full flower late in September, 1899. At Edgartown, Martha’s
Vineyard, I have seen it in bloom as early as July 1 and as late as
October 12.
*PHYSALIS HETEROPHYLLA Nees.
Siasconset, a few sterile plants in a mowed field, September
16, 1899.
*PHYSALIS PRUINOSA L.
In waste ground, scarce, and undoubtedly introduced. Mrs.
Flynn has sent me for examination specimens collected by her
August 25, 1904, in early bloom, and October 2, 1902, in flower and
fruit. On Martha’s Vineyard and on Chappaquiddick Island, it
is frequent as a garden weed and in soil that has been under
cultivation, but I saw it nowhere in surroundings that at all
conveyed the suggestion that it might be indigenous. Identifica-
tion of this and the foregoing species confirmed by Dr. Rydberg.
Note.—Mr. Floyd has informed me that Mr. Dame, in a letter
to Mrs. Owen, reported that Physalis peruviana L. had appeared
at Gibb’s swamp in several flourishing colonies in 1895, after an
extensive brush fire. No specimens are known to have been
preserved.
SOLANUM NIGRUM L.
Frequent by street sides and in waste places in and near the
- town and occasional in other parts of the island; sometimes in the
rubbish back of pond shores. Plants very small June 17, 1910,
first flowers June 27; remains in flower through September.
Common on Martha’s Vineyard.
332 BICKNELL: FERNS AND
_ The more usual form of this plant on Nantucket is only slightly
pubescent, and has more or less zig-zag and wing-angled stem and
branches; the leaves are ovate to lanceolate, often rhombic and
attenuate, cuneately narrowed into rather short petioles, and
entire to openly low sinuate or irregularly dentate, usually with
acutish teeth, the lower surface minutely appressed pubsecent;
inflorescence subumbelliform, the flowers 2-6, on strigillose pedi-
cels; corolla white, yellowish-green at the center; calyx lobes ovate-
oblong to oblong, becoming 2-3 mm. leng, loosely spreading or
recurved in fruit; anthers 1.5-2 mm. long, the filaments slightly
pubescent; ovary pubescent; style not protruding; berry be-
coming over I cm. in diameter; seeds 1.5 mm. broad. On
Martha’s Vineyard I have collected a much smaller variety,
reduced and more delicate in all its parts; also a larger form, with
purplish-tinged flowers.
*Solanum peregrinum sp. nov.
Similar to Solanum nigrum, duller green, rather low, branched
from the base, the stem and branches slightly if at all zig-zag,
somewhat angled but not winged, roughish with scattered in-
curving hairs or, above, like the younger parts of the plant,
clothed with a whitened strigose pubescence; leaves rather strongly
venose, thinly roughish pubescent on both surfaces with loosely
appressed hairs, the veins strigose; blades broadly ovate, obtuse,
abruptly contracted to slender petioles, sometimes slightly cordate
or subhastate, prevailingly deeply dentate with prominent, often
regular, obtuse or rounded teeth; inflorescence corymbiform, the
flowers six to nine, on finally reflexed strigose pedicels; corolla white,
3-4-5 mm. long, the lobes ovate-oblong, obtuse, densely pubescent:
on the outer surface, finally recurved; calyx lobes short, I-1.5
mm., broadly ovate or triangular, usually only obscurely veined,
loosely spreading in fruit; anthers pale yellow, 2-2.5 mm. long,
the filaments bearded, I mm. or more in length; ovary densely
white pubescent; style slightly exserted; berry black, becoming 8
mm. in diameter, the seeds 2 mm. hea
Street sides and waste places in Nantucket where it is rather
more common than S. nigrum, at least in the town; Surfside;
Siasconset. In full flower July 14, 1912; continues in bloom *
through September. Not seen on Martha’s Vineyard. Type:
Nantucket street, September 12, 1899, deposited in the herbarium
of the New York Botanical Garden.
FLOWERING PLANTS OF NANTUCKET pao
This plant is evidently introduced on Nantucket and its
native habitat is unknown. Whatever may be its history there
seems little reason to doubt that it has been included in the species
Solanum nigrum. Nevertheless I have not found that its char-.
acters properly correspond with those of any one of the numerous
varieties that have been described of this wide-ranging and poly-
morphic species. Among the extensive series at the New York
Botanical Garden the following are to be referred to S. peregrinum:
Montana: Columbia Falls, September 17, 1894, R.S. Williams.
FLORIDA: Pensacola, August 6, 1901, A. H. Curtiss, ‘ No. 6863,
Second distribution of plants of the Southern States,” labeled
“S. nigrum var. Dillenwi,” a very different plant. SWITZERLAND:
August 28, 1897, S. L. Clarke. :
Solanum nigrum, as now generally accepted, is unmistakably a
-composite species, and in the subdivision which it must sometime
receive the precise application of the collective name will be a
matter for very critical determination. The ordinary Nantucket
form agrees closely with the description of ‘‘true S. nigrum”’ as
understood by Dr. Gray.* From this S. peregrinum is clearly
distinct showing obvious differences throughout—duller green
color, coarser pubescence, straighter and less angled stem and
branches, shorter and broader, deeply. dentate-lobed leaves, the
blades abruptly contracted into longer petioles, corymbiform
instead of umbelliform inflorescence, broader divisions of the
corolla, larger anthers, bearded filaments, shorter calyx lobes,
larger seeds.
Comparison of S. peregrinum with the type of S. interior
Rydberg shows it to be quite distinct from that western species.
SoLANUM DULCAMARA L.,
Neglected places in the town; cedar thicket at Monomoy;
among pines near Miacomet Pond; the Thorn lot and at several
stations on the western side of the island. First flowers June 28,
1912; in full bloom June 15, 1910; late flowers and well fruited
September 10, 1907. :
LYCIUM VULGARE (Ait. f.) Dunal.
Occasional i in waste ground and along fences i in and near the
* Syn. Fl. 2!: 227-228.
+ Bull. Torr. Club 31: 641, 1905. |
304 BICKNELL: FERNS AND
town, and at one place on the harbor shore. In full flower May
30, 1909, June 15, 1911, mid-September, 1899.
DATURA STRAMONIUM L.
Infrequent and apparently never of very strong growth on
Nantucket. A few small plants at Siasconset, below the bluff,
in 1899, 1904, 1912; small plants by the railroad in Nantucket,
1904; waste ground south of the town September 16, 1907, in full
flower.
*DaTuRA TATULA L.
One small plant in a town barnyard September, 1899; a
solitary plant by a streetside in full flower August 10, 1906.
Note.—Petunia axillaris (Lam.) B.S. P. is occasionally to be
seen as a transient waif in the town streets.
SCROPHULARIACEAE
VERBASCUM THAPsSUs L.
Nowhere abundant, but scattered over the island in dry open
places. Spikes beginning to show June 18, 1908; first flowers
June 26, 1910, June 28, 1912.
*VERBASCUM -PHLOMOIDEsS L. ;
Two stout plants together on a bank at Siasconset, June 13,
1908, in full flower, thus earlier blooming by over two weeks than
the closely related Verbascum Thapsus.
VERBASCUM BLATTARIA L.
Seen only in a grassy yard on North Water Street, September
10, 1907, a number of fruiting plants, and two still in bloom, their
flowers white. Mrs. Owen has recorded it from a single station in
Nantucket and from Siasconset. It is thus not less scarce today
than it was thirty years ago, and is one of a number of common and
widely distributed weeds that on Nantucket do not seem to find
conditions favorable to their spread.
*CYMBALARIA CYMBALARIA (L.) Wettst.
Linarta Cymbalaria Mill.
Established along old walls at several places in the town.
Just in bloom June 16, 1911; in full flower June 5, 1912.
FLOWERING PLANTS OF NANTUCKET 335
LinarRia LinariaA (L.) Karst.
Linaria vulgaris Hill.
Another one of our common weeds that does not make head-
way on Nantucket. I have met with it there on only three
occasions: in 1899, at Siasconset, a few plants together; in 1904,
a single patch on the south shore at the site of the old Surfside
hotel; and in 1912, by the Wauwinet road in Squam, a small colony
in full flower July 12. As far back as 1888, Mrs. Owen reported
it as “not infrequent on the edge of the town,”’ where today, if it
has not died out entirely, it must be extremely scarce. It occurs
on Tuckernuck and is frequent on Martha’s Vineyard, but seems
to be absent from Chappaquiddick Island.
LINARIA CANADENSIS (L.) Dumort.
- One of the common plants of sandy and sterile soils. First
flowers May 30, 1909, June 4, 1911; in full flower June 7, 1908;
continues to bloom well into September.
*SCROPHULARIA LEPORELLA Bicknell.
Scarce. In September, 1899, a colony of some twenty plants,
the tallest nearly five feet high, grew at the border of a thicket
in Squam. Five years later, and many times since then, this
station was searched for in vain; probably none of the plants had
survived the encroachment of the woody growth that had at first
given them protection. Not until 1910 was the species seen
again when two spindling flowerless plants were found at Watt’s
Run. The next year much of it, in full flower on June 9, raised
its tall stems along a low shore thicket in Quaise, and on July 5,
1912, a group of some fifteen plants were in full flower in low open
ground near Miacomet Pond. Collected in a yard on West
Silver street July 1, 1911, by Miss Grace B. Gardner. It occurs
on Martha’s Vineyard, there being two stations on Chappa-
quiddick Island.
It is interesting that the wide variation in leaf form to which
this species is subject may be seen in its extremes in the small and
isolated colonies found on Nantucket. In plants rising side by
side the leaves of one may be evenly close-serrate, of the other
coarsely and irregularly laciniate, or even doubly laciniate, with
salient triangular to lanceolate-attenuate teeth. Even this ragged-
336 BICKNELL: FERNS AND
ness of outline may be sharply accentuated by a more prominent
tooth here and there standing forth quite out of line with its
fellows either at right angles to the axis of the leaf or actually
recurved.
The plants of these coastal islands compared with those of more
hilly country inland show a marked tendency toward a different
form of corolla, the tube wider, even campanulate, with its upper
lip abbreviated and with short rounded lobes, much like that of
the western S. occidentalis, to which this species is evidently very
closely related.
GRATIOLA AUREA Muhl.
Abundant on wet shores and in low penal. as well as in
many fresh water ponds, where it develops after the manner of a
true aquatic producing lengthened and attenuate semi-pellucid
leaves. Young plants beginning to appear May 30, 1909, and
June 8, 1910; mostly less than one inch high June 17, 1910; in
full flower August 7, 1906, and until after the middle of September.
On Martha’s Vineyard I have seen it in flower as early as June 26
(1913) and as late as October 8 (1909).
ILYSANTHES DUBIA (L.) Barnhart.
Ilysanthes gratioloides Benth.
Infrequent. Shores of Sachacha Pond; at several stations
near Siasconset; Maxcy’s Pond; pool on Madequet road. Still
in flower September 12, 1907, September 16, 1899.
LIMOSELLA TENUIFOLIA Hoffm.
Abundant on the wet sandy shores of ponds on the south side
of the island, especially Hummock Pond; also at Sachacha Pond.
Here, and at Miacomet Pond, it may be seen growing beneath the
water with elongated attenuate leaves, some of them Io cm. or
more in length. Ordinarily the leaves are mostly 2-4 cm. long
and very narrowly linear, either obscurely spatulate or tapering
to an obtuse tip. Not yet in flower May 31, 1909; in full flower
June 17, 1908, continuing in bloom through September.
*VERONICA OFFICINALIS L.
__ Asmall patch in full flower on a lawn on Main Street, June 16,
1911, doubtless introduced.
FLOWERING PLANTS OF NANTUCKET 337
*VERONICA SERPYLLIFOLIA L.’
Infrequent and possibly introduced. Damp fields westward
from the town at several stations; Grove Lane; Crooked Lane;
Millbrook swamp. In full flower May 31, 1909, June 11, rg1t.
*VERONICA PEREGRINA L.
Apparently of very recent introduction. It grew in profusion
in a neglected yard at the Springfield House, June 11, 1911, in
flower and fruit, but has not been seen anywhere else.
VERONICA ARVENSIS L.
Frequent by streetsides in the town; abundant on a lawn on
North Water street, 1910; dry open thickets in Shawkemo, Polpis,
and Millbrook swamp. In full flower May 30, 1909, June 7, 1908,
June 27, IgIo.
*VERONICA TOURNEFORTIL Gmelin.
Near Sea Cliff Inn, June, 1893, Mrs. Mabel P. Robinson;
Brant Point road “abundant and seemingly established’’ 1896,
Mrs. Maria L. Owen, fide F. G. Floyd.
*VERONICA TEuCRIUM L.
Grove Lane, one plant, 1897, Miss Elizabeth S. Kite, fide
F. G. Floyd. An estray from farm house gardens at several
places on Chappaquiddick Island. In full flower in September.
*AUREOLARIA PEDICULARIA (L.) Raf.
Gerardia pedicularia L.
A colony of scattered plants in full flower September 16, 1899,
in an open growth of scrub oak a mile or more northeast of Siascon-
set. I have not seen it since on Nantucket, although it is locally
common on Chappaquiddick Island, where it continues in bloom
sometimes to the middle of October. Doubtless referable to Dr.
Pennell’s variety caesariensis, although no specimens are now at
hand for verification.
AGALINIS PURPUREA (L.) Britton.
Gerardia purpurea L.
Common in low grounds, either in ee soil, fresh or brackish,
or on dry sandy levels, blooming in August and September and
later. The corolla varies much in size and is largest and most
pubescent in stout and somewhat fleshy horizontally wide branched
338 BICKNELL: FERNS AND
plants of brackish soils which also develop relatively large broad-
ened capsules; the largest corollas seen were 3.8 cm. long and of
equal width.
*AGALINIS PAUPERCULA (A. Gray) Britton.
Gerardia paupercula Britton.
Flowering specimens collected August 16, 1906, in damp places
in the hills south of Polpis, have been determined by Dr. F. W.
Pennell. I met with it nowhere else on Nantucket where it can
scarcely be other than a scarce and local plant.
AGALINIS MARITIMA Raf.
Gerardia maritima Raf.
Locally common in many salt marshes along the north shore
of the island; the Creeks; Swain’s Neck and Bache’s Harbor;
Eatfire; Pocomo; Coskaty; Little Neck. Blooms in August and
September.
J * Agalinis acuta Pennell, sp. nov.
“Annual. Plant light green, not tending to blackenin drying.
Stem 1-4 dm. tall, simple or moderately branched, rather con-
spicuously striate four-angled. Leaves opposite, linear, slightly to
moderately scabrous above; those of the stem I-2.5 cm. long,
0.6-I.3 mm. wide. Pedicels slender, in flower 5-15 mm. long, in
fruit 12-20 mm. long, 1-2(-3) times the length of the bracts.
Calyx-tube 3 mm. long, evidently reticulate-veiny (firmer in tex-
ture than in A. decemloba), 24-34 the length of the capsule, its lobes
0.5-I1 mm. long, triangular-acuminate, not or scarcely callose.
Corolla 10-13 mm. long, membranous, its tube 7-9 mm. long, its
lobes 3-4 mm. long, retuse to emarginate, all spreading; within
pubescent below sinus and over entire width of basal portions of
posterior lobes; posterior lobes ciliate, anterior finely ciliolate;
‘‘rose-pink”’ (not seen fresh). Filaments lanose toward apex,
anterior more densely so; anther-sacs lanose with white hairs on
the valvular surface, glabrous on the sides. Style glabrous.
Stigma I-1.2 mm. long. Capsule 3.8-4.2 mm. long, ovoid, yel-
lowish-brown. Seeds 0.4-0.6 mm. long, triangular-rounded, tur-
gid, yellowish-brown; testa finely reticulated.
“Type: dry sandy downs, Edgartown, Martha’s Vineyard,
Massachusetts, collected in flower September 12, 1901, M. L.
Fernald, 45 in United States National Herbarium; cotypes in
Gray Herbarium and Herbarium New York Botanical Garden.
FLOWERING PLANTS OF NANTUCKET 339
“ Flowers, late-August to October. Fruit, late-September to
October. Dry sandy soil, rare or locally frequent in the Coastal
Plain, Cape Cod region of Massachusetts, Nantucket, Martha’s
Vineyard and Long Island, westward as far as the Hempstead
Plains; very occasional inland (above the Fall-Line) in eastern
Massachusetts, Rhode Island and Connecticut.
‘* Differs from A galinis decemloba (Greene) Pennell, which occurs
from Lancaster County, Pennsylvania, to northern Alabama, by
its lower habit, pedicels mostly 1-2 (not 2-3) times the length of
the bracts, calyx-tube campanulate (not hemispheric), firmer in
texture, 3-? (not 3-3) the length of the capsule, calyx-lobes
0.5-I mm. (not 0.05-0.2 mm.) long, not or scarcely callose, and its
seeds smaller, 0.4-0.6 mm. (not 0.6—-0.8 mm.) long, strongly (not
obscurely) reticulated.
“Differs from Agalinis Skinnertana (Wood) Britton, which
occurs from the St. Clair River, southwestern Ontario to eastern
Kansas, by its less conspicuously striate-angled stem, angles gla-
brous or nearly so (not scabrellous-roughened), its corolla-lobes
more or less emarginate (not truncate), stigma I-1.2 mm. (not
1.5-2 mm.) long, capsule 3.8-4.2 mm. (not 4-5 mm.) long, and seeds
smaller, 0.4—-0.6 mm. (not 0.7—0.9 mm.) long.”” Francis W. Pennell.
A characteristic autumn-flowering plant of the dry plains in
the southern half of the island; on the north side observed only
in Pocomo. In full flower September 2, 1904, continuing in
bloom through the month. Common also on the south side of
Martha’s Vineyard and on Chappaquiddick Island. Well grown
plants become much branched and 3 dm. in height; diminutive ex-
amples may be not over 2 cm. high with only a terminal flower.
Comparison of Nantucket specimens collected in 1904 with
Wood’s type of Gerardia Skinneriana in the herbarium of Columbia
University, to which species the New England plant had long
been referred, showed that they were not at all the same; and
when, through the kindness of Professor Greene, I received speci-
mens of his then recently described Gerardia decemloba, from
Washington, that more nearly related plant was readily seen not
to be identical. Dr. Pennell’s studies in this group have brought
him quite independently to the same conclusion, and have shown
further that our eastern plant, and A. decemloba as well, are
340 BICKNELL: FERNS AND ©
distinct from the more southern Agalinus parvifolia (Chapm.)
Small, to which both had been recently referred.
SCHWALBEA AMERICANA L.
Mrs. Owen’s catalogue contains the record of all that is known
of this plant on Nantucket: “‘ Plains opposite Bloomingdale; rare,
L. L. D. Plants of luxuriant growth but few in number.” It
has never been reported by any other collector than Mr. Dame.
PEDICULARIS CANADENSIS L.
Frequent in dry thickets and open Sean throughout the
eastern side of the island from Pocomo and Squam to Siasconset;
Quaise; near Madequecham Pond. Observed in full flower from
June 4 to 15.
MELAMPYRUM LINEARE Lam.
An inhabitant of dry thickets and copses on the eastern side
of the island from Shimmo and Shawkemo to Squam and Siascon-
set. Just in flower June 2, 1909, June 7, 1908; still in bloom
July 11, 1912.
LENTIBULARIACEAE
UTRICULARIA MACRORHIZA LeConte.
U. vulgaris var. americana A. Gray.
Reed Pond; Maxcy’s Pond; pool west of the town; reported
from Polpis by Mrs. Owen. In flower at Reed Pond June 8, 1908,
June 18, 1910, July 3, 1912, September 10, 1907.
UTRICULARIA GEMINISCAPA Benj.
U. clandestina Nutt.
In a ditch on Swain’s Neck, also in Polpis; not seen in flower.
‘Ditches in Polpis choked with it.” M. L. O. catalogue.
UTRICULARIA INTERMEDIA Hayne.
‘““A single non-flowering specimen in Reed Pond. Morong,
1887.”" M.L. O. catalogue.
SETISCAPELLA SUBULATA (L.) Barnhart.
Utricularia subulata L.
Taupawshas swamp in open places among masses of cranberry,
both in wet spots and in merely damp soil, July 6, 1912, just in
flower. Reported by Mrs. Owen in her catalogue from Tom
FLOWERING PLANTS OF NANTUCKET 341
Never’s and Gibb’s Ponds, where it was collected by Mr. Walter
Deane (see Rhodora, 6: 160).
SETISCAPELLA CLEISTOGAMA (A. Gray) Barnhart.
Utricularia cleistogama Britton.
Mrs. Owen, in her catalogue, has reported her discovery of
this diminutive plant on Nantucket, where she had found it in
abundance in a part of Tom Never’s swamp and also at Almanac
Pond. Flowering specimens, sent by Mrs. Owen, as I am informed
by Professor Fernald, are preserved in the Gray herbarium together
with a letter from their collector dated August 28, 1881. The
citation of the plant from Nantucket in the sixth edition of Gray’s
manual evidently rests on this material.
Although now held to be a species distinct from S. subulata it
cannot be said of this plant that it is yet conclusively known to be
anything more than a reduced state of that species. Whatever
the truth may be the two plants do not always show that wide
diversity in their corollas that, by comparison of typical examples,
does undoubtedly seem to attest them as distinct. Near Edgar-
town, on Martha’s Vineyard, on September 30, 1912, there fell
to me a most favorable opportunity of observing the extent of
variation natural to the flowers of S. cleistogama among the
plants of a single colony. The situation was a few square feet of
damp sandy soil in open ground. In the weakest examples,
some of them not over I cm. high, the corollas, ‘‘not larger than
a pinhead,” were subglobose or saccate, and white or faintly
bluish in color, precisely as descriptions require them to be. But
in stronger plants the corollas increased doubly in size and came
also, by an exact gradation, to a distinctly two-lipped form, the
blunt lower lip dusky or purplish lineate and with an evident
white spur, the most open flowers showing an unmistakable
yellowish tinge. The spur, obsolete in the smallest ‘corollas, —
varied in the larger ones from rounded to oblong and acutish; in
one instance it was bifid.
In very small examples of S. subulata, unmistakable as to
identity because components of colonies of the typical plant, the
corolla, perhaps from arrested development, may be somewhat
abortive and reduced to a fraction of its normal size, and is some-
times palest yellow, or even whitish with a faint bluish tinge.
342 BICKNELL: FERNS AND
Such plants were collected on Martha’s Vineyard, near Lambert’s
Cove, July I, 1913.
OROBANCHACEAE
*THALESIA UNIFLORA (L.) Britton.
Orobanche uniflora L.
Probably not uncommon, although not sila reported
from Nantucket. On June 4, 1909, when it was in full flower and
therefore easily noticeable it was found along the foot of Rattle-
snake bank, and also by a low thicket in Shawkemo and on an
open sandy hillside eastward towards Polpis.
Note.—The beech-drops, Leptamnium virginianum (L.) Raf.,
grows in association with almost all the groups of beech trees
on Chappaquiddick Island, and is abundant at several stations
and of strong growth, some plants attaining a height of 5 dm.
On Nantucket I have never missed an opportunity of seeking it
wherever beech trees were seen but always without success. It
seems altogether probable, however, that some day it will be
discovered there.
BIGNONIACEAE
*CATALPA CATALPA (L.) Karst.
Catalpa bignonioides Walt.
Small trees not over three feet in height, were observed, in
1904 and 1906, scattered among an extensive growth of pines east
of the town; a single small tree among introduced pines at
Wauwinet, 1912.
PLANTAGINACEAE
PLANTAGO RUGELII Dene.
Reported in Mrs. Owen’s catalogue as ‘“‘very scarce’? and
found, by Judge Churchill and Mr. Dame, only at two stations at
the southern end of the town. I did not succeed in finding it
until 1907 when, in September, a few flowering plants were
observed in a shaded yard on Broad Street. Two years later it
had become established by nearby streetsides at several places,
and was also seen in a farmyard south of the town and at as
far distant a point as a thicket in Shawkemo.
FLOWERING PLANTS OF NANTUCKET 343
PLANTAGO MAJOR L.
_A weed of roadsides, old fields and waste places. July to
September.
*PLANTAGO HALOPHILA Bicknell.
A characteristic plant of salt marshes and brackish shores,
sometimes extending along roadsides. Flowering during August
and September.
PLANTAGO LANCEOLATA L.
Excessively abundant in fields and grassy places everywhere.
Dark green glabrate plants and paler and narrower leaved very
hairy forms, mainly of poorer soils, constitute markedly divergent
varieties of this species. Forms with dark, ovate, compound
heads are frequent, as well as sharply contrasting forms having
the spikes linear and elongated. Blooms from May until October.
PLANTAGO DECIPIENS Barneoud.
Frequent or, locally, rather common in salt marshes and along
shores, blooming from August through September.
*PLANTAGO ARISTATA Michx.
A few plants on Little Neck, September 10, 1904, in flower and
fruit; near Brant Point, September 13, 1907; one plant in a waste
yard June 27, 1910; a scattered growth in a dry field in Shawkemo,
in full flower July 1, 1912. Abundant and evidently long estab-
lished on Martha’s Vineyard, and also extending all along the sandy
roadway that crosses Chappaquiddick Island, but on Nantucket
apparently of only recent introduction.
RUBIACEAE
HouUsTONIA COERULEA L.
Abundant throughout, not only in low moist places but also
over the dry plains and commons, and to the tops of the exposed
rolling hills. In the southwest quarter of the island it even
overspreads the sandy wastes near the shore, in association with
Sisyrinchium arenicola, these being the most noticeable plants
flowering there in early June. In full flower May 30, 1909; it
continues quite generally in bloom into July, and produces belated
flowers in August and even in September.
344 BICKNELL: FERNS AND
Large flowered forms having white, brightly yellow-eyed
corollas, with relatively short, evenly graduated tube, seem to
answer quite exactly to the description of var. Faxonorum Pease
& Moore.
HousTONIA LONGIFOLIA Gaertn.
Recorded by Mrs. Owen from ‘‘near Long Pond. L. L.
D[ame].”’ No other collector has reported it. About Long Pond
are conditions of soil and surroundings suggestive of localities on
Long Island where this is a characteristic plant, but I have not
succeeded in rediscovering it on Nantucket where, if it occurs
today, it must be of extreme rarity. |
CEPHALANTHUS OCCIDENTALIS L. ;
A common shrub of wet swamps and the borders of muddy
ponds. Leaves appearing June 7, 1911, June 8, 1908; flower
heads less than half size July 4, 1912; still in bloom September 10,
1907.
MITCHELLA REPENS L.
A scarce plant on Nantucket where I have met with it only
twice. In 1910 a small patch was discovered in a dense thicket
near the head of Tom Never’s swamp, and a few plants in a
secluded spot at Beechwood, in Polpis. It had not flowered at
either station. Mrs. Owen has reported it from Trot’s swamp and
from Polpis.
*GALIUM ERECTUM Huds.
On Grove Lane, June 28, 1912, a large mass in full flower.
*GaLium APARINE L.
Everywhere in shaded thickets on banks and in low grounds.
In full flower May 30, 1908, May 31, 1909; well fruited June 28,
1912; by midsummer only the withered stems are to be found.
Varies remarkably in proportions and in the size of the leaves and
fruit.
GALIUM PILOsuM Ait.
Frequent, especially in the northwestern quarter of the island.
Not yet in flower June 26, 1910; past flowering and with mature
fruit September 2, 1904. Varies from densely pubescent to
—— glabrous, the more pubescent forms occurring in exposed
FLOWERING PLANTS OF NANTUCKET 345
sandy places and in pure sand. The glabrate plants are not
common and are found in more protected situations among open
growths of scrub pines. Their stems are shining and sometimes
quite glabrous, their leaves hispidulous ciliate on the margins
and on the midrib beneath, var. puncticulosum (Michx.) T. & G.
GALIUM TRIFLORUM Michx.
Frequent in shaded thickets on the eastern side of the island,
especially in Squam, Shimmo and Polpis. In full flower and with
young fruit August 13, 1906. As elsewhere two forms may be
distinguished, the leaves of one narrow and tapering, of the other
broadened and rounded or abruptly narrowed to the cuspidate
apex.
GALIUM CLAYTONI Michx.
Common in low grounds flowering from the middle of June
through September.
The ordinary form of this bedstraw is a small plant of damp
or wet open places often where the soil loses much of its moisture
or becomes quite dry in summer. In partial shade, or in very
wet situations, which encourage a vigorous growth throughout
the season, it comes to an enhanced phase of development that
gives it quite the aspect of being a different species. Nowhere
have I seen this larger state of the plant of more pronounced
character than on Nantucket.
The more common smaller form of the plant is very scabrous
and of a pale green color, not readily discoloring when pressed;
the leaves are often firm with revolute margins, those of the stems
sometimes reduced to fours and only 8-10 mm. long and 1-2 mm.
wide; the minute flowers of three- or, not rarely, four-parted corol-
las, are in pedunculate clusters of three on short finally divergent
pedicels 2-5 mm. long, and the fruit is 1 mm. or less in diameter.
The larger form is only slightly scabrellous and is thin leaved and
deeper green in color, turning dark or black on the herbarium sheet.
In its luxuriance of growth along ditches and in very wet spots
it sometimes forms close masses 6 dm. or more in height, from
which it is possible to disentangle individual plants as much as
13 dm. in length. The leaves, in whorls of six on the stems and
four on the branches, are narrowed into evident petioles, and be-
346 BICKNELL: FERNS AND
come as large as 3 cm. long and 6 mm. wide; the corolla, 1.5—2 mm.
in expanse, is four-parted or, in smaller flowers, three-parted, the
lobes either acute or obtuse; the fruit I-1.5 mm. in diameter, is
borne on slender spreading pedicels sometimes fully 12 mm. in
length which are disposed to be arcuate and slightly scabrous.
Smaller and narrower leaved plants having pedicels of this char-
acter so closely simulate Galium trifidum L. that they might easily
be misidentified. It would be hard to believe of these divergent
phases of Galium Claytoni that they were other than well-defined
species, were it not that the gradation from one to the other is
gradual and complete and did not as many examples lie midway
in the range of variation as at either extreme. Almost the same
extent of variation in the species may be observed on Long Island
and about New York. If it be thought expedient to recognize
the larger form by a name we have for it the varietal designation
Galium Claytoni var. subbiflorum Wiegand (Rhodora 12: 228-230.
1910).
It could be wished that the name Galium Claytoni were better
accredited in its application to this plant for which we have come
to use it. A careful reading of Michaux’s description leaves little
doubt in my own mind that his G. Claytoni was no other plant
than the G. trifidum of Linnaeus. Michaux was particular to
emphasize of his species that its leaves were in fours or, rarely, in
fives, never in sixes, thus meeting exactly the case of G. trifidum;
and it is further suggestive in his description that the rather
ambiguous phrase “fasciculus ramorum terminalibus’’ at once
becomes clear and at the same time diagnostic of G. trifidum, if we
may understand it to mean that, except at the ends of the branches,
the flowers were solitary. Upon these indications, and others not
necessary now to bring forward, there would seem to be little
reason to doubt that the species we have been calling Galium
Claytont has never received a name proper to its own identity,
unless, indeed, it should bear the name Galium tinctorium L.,
now otherwise appropriated. Bigelow and some others of our
earlier botanists took this view, that is to say, their descriptions
make it plain that this name was understood by them as applying
to the larger phase of the plant now called G. Claytoni; and, as a
matter of fact, the description of G. tinctorium in Species Plantarum
FLOWERING PLANTS OF NANTUCKET 347
undoubtedly applies better to this plant than to the species that
has come to bear it, and which Bigelow described as Galium
obtusum. It is to be hoped that there may be still in existence
original specimens by which the true application of these names
may be made clear.
On Nantucket the smaller form of G. Claytoni comes into
bloom rather earlier than the larger one—June 15, 1908, June 18,
1910; the latter showed its earliest open flowers June 26, I9gI0.
Galium tinctorium L. (G. obtusum Bigelow) occurs on Martha’s
Vineyard and is to be looked for on Nantucket.
*GALIUM PALUSTRE L.
Abundant in the meadows about Shawaukemmo Spring,
where, in June and early July, it is conspicuous from its clustered
masses of white bloom about springy places and along runs and
_ rills; also found near Abram’s Point. Just in flower June 8, 1911;
in full bloom June 25, 1910, July 1, 1912. Flowers cymulose
clustered, the corollas spreading 2—-5-4.5 mm., four-parted, the
lobes acute; flower buds often purplish-pink or rose color.
Note.—Galium circaezans Michx. is included, by name only,
in Mrs. Owen’s list. It is not known that it has ever been collected
on Nantucket. .
*SHERARDIA ARVENSIS L.
In full flower all through the short grass of a lawn on Chester
street, June 15, 1910.
CAPRIFOLIACEAE
SAMBUCUS CANADENSIS L.
_ Very common both in moist and in dry thickets and among
the scattered patches of bayberry and mixed growth on the open
plains. Not yet in bloom June 27, 1910; cymes whitening June
30, 1912, the first open flowers July 4, in showy bloom July 10;
still some flowers August 4, 1906; fruiting well September, 1907.
VIBURNUM DENTATUM L. ;
In swampy thickets, as well as in dry places, and even in pine
barrens; often growing with V. venosum, but much less abundant
and blooming somewhat earlier. Cymes close and green June 7,
1908; just in flower June 22, 1910; bearing green fruit, no flowers
348 BICKNELL: FERNS AND
remaining, July 12, 1912. In the autumns when I have observed
it it has fruited much less freely than V. venosum.
‘VIBURNUM VENOSUM Britton.
A noteworthy shrub of Nantucket abounding throughout in
damp thickets, on dry banks and in open ground; also in pine
barrens. Cymes small and green June 7, 1908; forward bushes
just in flower June 30, 1912, everywhere in showy bloom July 4
to 13, and later; fruiting abundantly in September. :
The leaves show great diversity of size and form, even in
shrubs growing side by side, and the fruit is no less variable,
sometimes globose, again oval and somewhat pointed. The
largest shrubs seen measured 22 cm. around the trunk near
the base and were over 3.5 m. in height. The type of this fine
shrub was collected on Nantucket, September 19, 1899. It
occurs also on Tuckernuck, and is much more common on
Martha’s Vineyard than V. dentatum which it replaces completely
on Chappaquiddick Island.
Reported in Mrs. Owen’s list under the name V. molle Michx.
*TRIOSTEUM PERFOLIATUM L.
Another native plant that has been found only at a single
station on Nantucket—at the foot of a steep thicketed bank by
a small pond near Acquidness Point. Here, on September 11,
1907, were two strong clusters not two feet apart consisting to-
gether of about twenty stems. Two years later it was seen that
these plants were succumbing to the closer pressure of the investing
woody growth, and in 1911, when I last visited them, one group
had been reduced to three, the other to six weakened stems. In
1910 the plants had still been able to bloom, but bore flowers with
light yellow corollas instead of the normal red ones they had
produced in their days of vigor. Flower buds just visible June 2,
1909; nearly open June 7, 1908; in full bloom June 19, ee
LINNAEA AMERICANA Forbes.
Mrs. Owen says of this plant: ‘‘A patch 15~20 feet in diameter
discovered in the pines in 1868 or 1869 by Mr. L. L. Dame. A
patch in the pines on the South Shore road discovered in 1872
by Miss C. L. Tallant.’’ At the latter locality the plants were
seen not more than two years, and Mr. Dame looked in vain for
FLOWERING PLANTS OF NANTUCKET 349
his plants in 1886. Mr. Floyd writes me that “Mrs. Owen told
me not very long before her death that the plant still persisted
at one station on Nantucket.”
*SYMPHORICARPOS SYMPHORICARPOS (L.) MacM.
Symphoricarpos vulgaris Michx.
“Road opposite Barzilliai Coffin farm September 1, 1912,
in full ower. Miss Grace Brown Gardner.”
*LONICERA JAPONICA Thunb.
Occasional by fence rows and roadsides in the general town
region, less often by fieldsides and the borders of meadows. In
full flower June 27, 1912.
CUCURBITACEAE
MICRAMPELIS LOBATA (Michx.) Greene.
Occasional in waste places in and near the town. In full
flower late in August.
SICyos ANGULATUS L.
At a few places in damp lots and by fences in the town; by’ the
railroad in Washington street.
CAM PANULACEAE
*CAMPANULA RAPUNCULOIDES L.
Occasional by street sides and in waste ground in the town,
more rarely by roadsides in the suburbs. First flowers June 27,
1910, June 29, 1912; in full bloom at the end of August, 1904.
SPECULARIA PERFOLIATA L.
Rare: Acquidness Point, knoll and hillside thicket east of
Shimmo farm, June 7, 1911; Rattlesnake bank, July 1, 1912.
Only a few plants at each station and no petaliferous flowers.
LOBELIACEAE
LOBELIA CARDINALIS L.
A few plants along Watt’s Run September 6, 1904, and much
of it in flaming bloom at the same locality September 17, 1907;
Quaise and Polpis, M. L. O.
Note.—Lobelia inflata L. is included in Mrs. Owen’s catalogue.
Nothing is known of it as a Nantucket plant at the present day,
nor have I seen it on Martha’s Vineyard.
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Notes on Michigan Liliaceae*
OLIVER ATKINS FARWELL
(WITH PLATE 20)
NEW VARIETIES OF ALLIUM CANADENSE
In Michigan there are three well-defined races or varieties of
the common wild garlic, Alum canadense L. The type, with
narrow leaves and numerous, small, ovoid, whitish, obtuse or
acutish bulblets in the simple umbel; a form with noticeably
broader leaves but with fewer, larger, long-acuminate bulblets .
_which are red at maturity; and a larger, glaucous, form with more
fleshy leaves, often with two or three sessile, contiguous umbels,
which bear a larger number of obovoid or nearly spherical bulblets
of a yellowish white color. There is such a marked difference in
the appearance of these forms, at least in the living state, that it
seems advisable to name them and place them on record.
“ Allium canadense ovoideum var. nov.
Bulb ovoid, about 19 mm. long, outer coats fibrous-reticulated,
5-8 cm. below the surface; plant 3-4 dm. high, green, with four
or five leaves near the base, these equalling the scape or a little
shorter; umbel terminal, mostly of a few, twelve or less, large,
ovoid-acuminate, sessile bulblets with or without a few long-pedi-
celled flowers and some secondary umbels of one to three bulb-
lets, on long rays 5-10 cm. long; involucre of one to three ovate,
acuminate, scarious bracts, 2.5—4 cm. long; bulblets of the umbel
gradually tapering from a broad base, the larger about 22 mm.
in length, mostly rose colored at maturity; generally there are no
flowers but some umbels may have from one to four, and occa-
sionally there may be one in one of the umbellets; the flowers are
pale purple, on pedicels 5 cm. in length, 8 mm. across when open,
5-6 mm. high; segments acute, the outer narrowly ovate, the
inner oblong or linear.
On wet banks in open fields. MICHIGAN: near Rochester, June
11, 1914, Farwell 3667.
* Contributions to the Botany of Michigan, No. 12.
OL
352 FARWELL: NOTES ON MICHIGAN LILIACE4
Differs from the typical form of the species in its large long-
acuminate, rose-colored bulblets, in the narrower perianth-seg-
ments and longer pedicels.
” Allium canadense robustum var. nov.
Bulb spherical or nearly so, 2.5 cm. in diameter or less, the
outer coats fibrous-reticulated, deep-seated, 15-18 cm. below the
surface; scapes, often two, 3-4.5 dm. in height, 6 mm. or less
in diameter, and, with the two to six leaves near the base, glaucous;
leaves rather fleshy, 6 mm. wide, channelled above, convex below,
two thirds to three fourths the length of the scape; scape bearing
one to three sessile, contiguous umbels, mostly of bulblets, at its
apex with one or more rays, 12.5 cm. or less in length, each bearing
an umbellet of from one to three bulblets; bulblets often foliaceous,
from spherical or nearly so to obovoid, tipped with a minute
curved point, the larger about 12 mm. in longest diameter, very
numerous, often more than thirty, yellowish white; involucre one
to four ovate, acuminate scarious bracts, the acumination often
prolonged to a length of 7.5 cm.; flowers, when present, few, on
pedicels 19-38 mm. long, white or pale rose, 7 mm. high, 10 mm.
across when open, segments obtuse, the outer ovate, the inner oval
or obovate.
In rather moist meadow lands. MuicnHiGANn: near Rochester,
June 9, 1912, Farwell 2629; June 13, 1912, Farwell 2729; June 11,
1914, Farwell 3606.
Differs from the typical form of the species in its more robust
habit, compound umbel with differently shaped bulblets, obtuse
petals and general glaucousness. Listed as Allium rubrum Oster-
haut? in Rpt. Michigan Acad. Sci. 15: 169. Nov. 1913.
NEW SPECIES OF LILIUM
I have long considered that the wild red lily of the middle west
is specifically distinct from the yellow-flowered Lilium canadense L.
of the Atlantic seaboard. _
Our plant is much taller and coarser; the flowers generally are
strongly revolute and red instead of yellow; if the two plants
were specifically the same, it would be reasonable to expect to see
a yellow-flowered individual at times in the middle west but such
has never come under my observation during the twenty-five years
I have been observing these plants. I had become so thoroughly
FARWELL: NoTES ON MICHIGAN LILIACEZ 353
imbued with the idea that our red lily of the middle west was the
true Lilium canadense that when I first saw the smaller, yellow-
flowered lily east of the Appalachians, from the window of a
slowly moving train, I was unable to recognize it as of that species,
it appeared to be so radically different from my conception of
what Lilium canadense should be. The red-flowered plant is
unquestionably a distinct species.
‘ Lilium michiganense sp. nov.
Leaves remotely whorled, six to ten in a whorl, three- to
seven-nerved and rough on the nerves beneath as well as on the
margins, lanceolate (12 mm. wide by 9 cm. long) to ovate-lanceo-
late (19 mm. wide by 4.5 cm. long); flowers in a pyramidal
cluster (two to four axial from the uppermost whorl and about
four racemosely disposed on the terminal portion of the stem), the
peduncles 10-12 cm. long, often bearing a foliaceous bract near
the middle.
MicuHicAN: Wiard’s Crossing, July 9, 1910, Farwell 21621%.
Similar to L. canadense in foliage and to L. superbum in its
flowers, which are of a beautiful orange-red, copiously spotted
inside with purplish brown or crimson spots. The upper leaves
of this plant are often smooth on the veins beneath, a transition to
L. superbum, which, though credited to Michigan, I have not yet
detected here. The new species occurs in three well-defined forms,
and I have taken the most highly developed form, although 1 not
the commonest, as the type.
’ Lilium michiganense umbelliferum var. nov.
Differs in having the stem end abruptly at the uppermost node,
the inflorescence being composed of an umbel only, of from two to_
five flowers, from the axils of the leaves of the uppermost node.
MICHIGAN: Wiard’s Crossing, July 9, 1910, Farwell 216214;
Birmingham, September 7, 1903, Farwell 1261c; Belle Isle, July
16, 1892, Farwell 1261; Palmer Park, July 16, 1902, Farwell 1261b;
Rochester, July 4, 1896, Farwell r261a.
v Lilium michiganense uniflorum var. nov.
The simplest form, in which the stem is continued beyond the
uppermost whorl of leaves and bears a single, nodding flower.
MIcHIGAN: Wiard’s Crossing, July 9, 1910, Farwell 216214;
304 FARWELL: NOTES ON MICHIGAN LILIACEZ
Rochester, July 14, 1912, Farwell 2848; Parkedale Farm, October
5, 1913, Farwell 3523; Rochester, July 14, 1913, Farwell 2848.
It is a misnomer to call L. michiganense, as is usually done here,
“the wild yellow lily’’ as the flowers are never yellow; the buds
are at first green, then yellow (some, however, never show any
sign of yellow externally), and finally red; the flowers when opened
are orange-red externally and on the blade internally, the midvein
being orange-yellow, the claw pale yellow or whitish and copiously
blotched with crimson spots which extend upward on the blade
for one half to two thirds its length. I have included in this
species certain forms in which the buds show no yellow color, and
the flower has brownish purple instead of crimson spots with orange-
red extending down the claw, but in which the leaves are charac-
teristic. Further study may prove these forms to be distinct.
The segments are recurved to below the middle, often spirally
coiled, so that the apex is again ascending.
’ Lilium peramoenum sp. nov.
Similar to L. michiganense, but the leayes are more numerous,
eight to sixteen in a whorl, with a corresponding increase in the
number of flowers in the inflorescence; the flowers are orange-red
throughout, dotted with copious but smaller spots; the leaves are
narrower, linear-lanceolate, 10 mm. wide by 9 cm. long to oblong-
lanceolate 15 mm. wide by 7 cm. long.
MICHIGAN: Parkedale Farm, July 19; 1914, Farwell 3726.
Elder thickets in rich muck lands. L. michiganense is found
usually in poorer soil though it may be in dry fields and on banks
or in wet swamps and meadows. Both species range from 1-2
m. in height.
NOTES ON UNIFOLIUM BIFOLIUM
In Rhodora for December, 1914, Mr. Fernald describes a
pubescent form of Maianthemum canadense as a new variety (var.
interius) and gives its range from Illinois north and northwestward
into Canada. He had not seen this pubescent form east of
Illinois or Wisconsin. The late Mr. G. H. Hicks collected it at
Owosso, Michigan, in 1889, and I found it at Stoney Creek,
Michigan, in 1913. Both stations are in the southeastern part
of the state, about ay miles apart.
FARWELL: NOTES ON MICHIGAN LILIACEZ 355
A careful study of a series of specimens, ranging from Oregon
to Hungary, allows of but one conclusion—that there is but a single
species and that the plant recently described by Mr. Fernald as
Maianthemum canadense var. interius, is, in reality, identical with
the Linnean Convallaria bifolia and is, consequently, Unifolium
bifolium (L.) Greene. The plant from the Black Hills, Fics. 7,
8, it will be observed, corresponds with that from Denmark,
Fic. 6; the collections of Mr. Hicks, Fics. 9, 10, are nearly
identical with those from Hungary and Bavaria, Fics. 1, 2; and
my own collection from Stoney Creek, Fic. I1, is very close to
another plant, Fic. 3, from Bavaria; the outlines of the leaves of
the var. canadense, FIGs. 12-19, are the same as those of the
type, no other difference, save that of pubescence, being detectable;
the apex in both is acute or obtuse, the base cordate or sagittate;
the western variety kamtschaticum, FiGs. 22, 23, shows the broad-
est sinus; the European type, the deepest; the var. canadense, the
shallowest; and the var. ovale, FIGs. 20, 21, the narrowest; but
there is a regular graduation from one extreme to another. The
var. kamtschaticum has the longest petiole with the blade prom-
inently decurrent on it; the plants of Loher, Fic. 3, and Cavas-
sius, Fic. 4, show the blades slightly decurrent on the petioles,
thus in a measure paralleling the western form. The pedicels
vary from 2-6 mm. in length in all the forms; the petioles of the
upper leaf measure I-6 mm., and of the lower leaf, 2-16 mm., in
all the forms except the western, which has much longer petioles.
There is in Michigan a plant with broadly ovate or oval leaves,
Fics. 20 (stem-leaf), 21 (root-leaf) ; Pursh traversed the region of
“the Lesser and Great Lakes” and I have no doubt that this form
is the basis of his Smilacina canadensis var. ovalis; the base of the
leaf is cordate or rather sagittate but the sinus is closed or nearly
so, which renders it inconspicuous. There is a form which has
three leaves to the stem and in Europe forms are found with but
one leaf. The differences in length of petioles, pedicels, and the size
of the leaf-sinus cannot be correlated with other characters with
a sufficient degree of accuracy to warrant separating the different
forms as species; such differences as exist are of degree and not of
kind and therefore are not of specific value. The species and its
forms may be listed as follows:
356 FARWELL: NOTES ON MICHIGAN LILIACEZ
UNIFOLIUM BIFOLIUM (L.) Greene
Maianthemum canadense var. interius Fernald, Rhodora 16:
ati. “ila:
MICHIGAN: Owosso, May 21, 1889, G. H. Hicks; Stoney Creek,
May 25,1913, Farwell 3412. Illinois westward to the Black Hills
and northward into Canada; also in Europe.
Unifolium bifolium monophyllum forma nov.
Differs from the species only in having but one leaf on the stem.
SWEDEN: Moheda, C.G. H. Cavassius, in the herbarium of Parke,
Davis & Company.
Unifolium bifolium canadense (Desf.) comb. nov.
Maianthemum canadense Desf. Ann. Mus. Paris 9: 54. 1807.
Convallaria bifolia var. canadensis Poir. Enc. 4: 371. 1816.
MIcHIGAN: Detroit, May 9, 1895, Farwell 379a; Algonac,
September 13, 1914, Farwell 3898. From Michigan eastward,
ranging north to Newfoundland and south to the Carolinas.
Unifolium bifolium trifolium (Pursh) comb. nov.
Smilacina canadensis ¥ trifolia Pursh, Fl. Am. Sept. 233. 1814.
The stem has three leaves, otherwise like the variety canadense.
MICHIGAN: Detroit, May 9, 1895, Farwell 379).
Unifolium bifolium ovale (Pursh) comb. nov.
Smilacina canadensis B ovalis Pursh, Fl. Am. Sept. 233... 1814.
I here adopt Pursh’s varietal name for the form in the Lake
Superior district, with broadly ovate or oval leaves.
MICHIGAN: Keweenaw Peninsula, June, 1886, Farwell 370;
October, 1914, Farwell.
UNIFOLIUM BIFOLIUM KAMTSCHATICUM (Gmel.) Piper
California to Idaho and Alaska; also Siberia.
FARWELL: NOTES ON MICHIGAN LILIACE ood.
NOTES ON VAGNERA
VAGNERA STELLATA (L.) Morong
The common woodland form, 3-4.5 dm. in height, bearing seven
to twelve leaves with a short, harsh pubescence on the veins on the
lower surface, becoming, with age, glabrate or smooth, at least to
the touch.
MIcHIGAN: Ypsilanti, May 29, 1891, Farwell r121; Parkedale
Farm, May 19, 1912, Farwell 2567; Rochester, June 28, 1914,
Farwell 3600.
This I take to be the typical form of the species. There isa
wide range in the size of the plant and leaves, in the pubescence,
and in the habitat of this species; the rhizome seems to vary in
size correspondingly; the fruit, while young, is green; but as it
approaches maturity it becomes purple on the angles; then inter-
mediate lines of a paler color appear; later it is finely lined or
spotted and ultimately becomes a ruby red as in V. racemosa
and the other species of the genus. Fruit, however, in this
species is a rarity; most of the flowers are sterile and drop from
their pedicels at an early stage as though the plant were of hybrid
origin; if such is the case the parents probably are V. racemosa
and V. trifolia.
Vagnera stellata mollis var. nov.
Differs from the typical form of the species in having the leaves
densely and permanently velvety pubescent all over the lower
surface; in the more numerous leaves, six to fourteen, I-5 cm.
wide and 3.5-20 cm. long; plant often larger, attaining a height
of nearly 1 m., but sometimes smaller and not exceeding 18 cm.
MicuiGANn: Parkedale Farm, May 30, June 11, and July 4,
1914, Farwell 3651, 366014, 372014; growing in muck lands, on
barren hilltops, and in Sphagnum, in the order named. SOUTH
Dakota: Black Hills, W. S. Rusby.
The habitat of the variety is as variable as that of the typical
form. The plants growing in Sphagnum are the smallest but seem
to perfect fruit more readily than those in other situations; those
growing in rich muck in elder thickets are the largest and do not
perfect fruit at all; the plant does not disdain the exposed tops
and sides of more or less barren hills but fruit is seldom seen in
such localities. With all its variableness, however, the soft vel-
vety pubescence of the plant remains constant.
358 FARWELL: NOTES ON MICHIGAN LILIACE
VAGNERA TRIFOLIA (L.) Morong
This species and Unifolium bifolium are locally known as wild
lily-of-the-valley. In the Lake Superior region of Michigan both
are quite common. Besides the ordinary three-leaved form of
V. trifolia there are two others that are frequently met with; one
has two leaves to the stem, and the other, one leaf. These may
be recorded as follows:
Vagnera trifolia bifolia forma nov.
A form with two leaves on the stem.
MICHIGAN: Keweenaw Peninsula, June 16, 1886, Farwell 378a.
Vagnera trifolia unifolia forma nov.
A form with only one leaf on the stem.
MICHIGAN: Keweenaw Peninsula, June 16, 1886, Farwell 378).
DEPARTMENT OF BOTANY,
ParKE, Davis & COMPANY,
ETROIT, MICHIGAN.
Explanation of plate 20
Fics, I-11. UNIFOLIUM BIFOLIUM (L.) Greene
1. Pressburg, Hungary, Richter. 2. Woods near ager Germany, May, 1891,
C. J. Mayer. 3. Woods near Munich, Germany, May, her. eda,
Sweden, C. G. H. Cavassius. 5. Same locality and ae (forma monophyllum).
6. pe Hs Denmark, June, 1865, J. Lange. 7, 8. age Hills, South Dakota, W. S.
Rusby, 10. Owosso, Michigan, May 21, 1889, G. Hicks. 11. Stoney Creek,
aie c Farwell 3412
Fics. 12-19. UNIFOLIUM BIFOLIUM CANADENSE (Desf.) Farwell
. Westfield, oe ites 29, 1873, H. H. Rus 15. East Green-
a phe York, 1867, A 9. Algonac, Michigan, September 13, 1914,
Farwell 3808. 17, 18. feos ule. May 9, 1893, Farwell 3709a
FIGs. 20, 21. UNIFOLIUM BIFOLIUM OVALE (Pursh) Farwell
. Lower leaf; Kewenaw Peninsula, Michigan, os 16, 1886, Farwell 379.
ai. deat sak: same ater a collector, October, 1
FIGs. 22, 23. UNIFOLIUM BIFOLIUM KAMTSCHATICUM (Gmel.) Piper
22. Lower leaf; Cascades of the Columbia River, pirates June 3, 1882,
W.N. Suksdorf 23. Upper leaf; same locality, collector, and dat
GS. 1-8, 12-15, 22, and 23 were drawn from specimens in the herbarium
of pees Davis & Company; the remaining figures, from specimens in the herbarium
of the writer. All are reduced one half
INDEX TO AMERICAN BOTANICAL LITERATURE
1908-1915
The aim of this Index is to include all current botanical literature written by
Americans, published in a or based upon American material ; the word seni
ica being used in the broadest s
Reviews, and papers that eer exclusively to forestry, agriculture, hhovlicalice:
ee products oi vegetable origin, or laboratory methods are not included, and
no attempt is made to index the literature of Lopate An occasional exception is
made in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not sasidtonel unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is mimes monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
subscriber must ‘ale all cards published during the term of his subscription, Corre-
spondence esis to the card issue should be addressed to the Treasurer of the Torrey
Botanical Clu
Arthur, J. C. Cultures of Uredineae in 1912, 1913 and 1914. Myco-
logia 7: 61-89. Mr 1915.
Balthis, F.K. An Illinois school campus. Am. Bot. 21: 1-8. F 1914.
[Illust.]
Barbour, E. H. Plant tissue in the carboniferous shales of Nebraska.
Nebraska Geol. Surv. 4: 231, 232. f. z. 30 N 1914.
Bartlett, H. H. The mutations of Oenothera stenomeres. Am. Jour.
Bot. 2: 100-109. f. r-g. F 1915.
Bartlett, H. H. The source of the drug Dioscorea, with a consideration
of the Dioscoreae found in the United States. U.S. Dept. Agr.
Plant. Ind. Bull. 189: 1-25. f. 7-8. 11 N1910.
Bartlett, H. H. Systematic studies on Oenothera,—V. Oc. Robinsonii
and Oe. cleistantha, spp. novv. Rhodora 17: 41-44. pl. I11I.
17 Mr 1915.
Beal, W. J. Weeds growing in Amherst. Rhodora 17: 29, 30. 6 F
1915.
Blake, S. F. Moneses uniflora var. reticulata. Rhodora ‘17: 28, 29.
6 F 1915.
Bovie, W. T. A simplified precision auxanometer. Am. Jour. Bot. 2:
95-99. f. I-3. F 1915.
359
360 INDEX TO AMERICAN BOTANICAL LITERATURE
Brown, H. P. Growth studies in forest trees. 2. Pinus Strobus.
Bot. Gaz. 59: 197-241. pl. 13, 14+f. 1, 2. 17 Mr 1915.
Buchtien, O. Contribuciones a la flora de Bolivia. Parte I. 1-197.
La Paz. 1910.
Burnham, 8. H. A visit to the Saratoga battlefield. Am. Bot. 21:
19-21. F 1915.
Buswell, W. H. A winter flora of a mountain top. Am. Bot. 21:
Q-I!. F 1915.
Castillo, L., & Dey, J. Jeografia vejetal del Rio Valdivia i sus in-
mediaciones. 1-120. f. I-64. 1908.
Clute, W. N. Insects and the color of flowers. Am. Bot. 21: 15-17.
F 1915. [Illust.]
Collins, F.S. November flowers. Rhodora 17: 33-38. 17 Mr 1915.
Conard, H.S. The structure and development of Secotium agaricoides.
Mycologia 7: 94-104. pl. 157+f. 1. Mr 1915.
Connolly, C. J. Beitrage zur Kenntnis einiger Florideen. Flora 103:
125-170... 2, 24+-f--2-24: 1% Ji ort... >
Cook, O. F. Date palm allies in America. Jour. Heredity 6: 117-122.
J. SH10. 26 F lors:
Curtis, O. F., & Colley, R. H. Picro-nigrosin, a combination fixative
and stain for algae. Am. Jour. Bot. 2: 89-92. F IQI5.
Davidson, J. Second annual report of the botanical office. Second
Ann. Rep. Bot. Office British Columbia 1: O 35-0 76. f. 38-61.
IQI5.
Fernald, M. L. The North American representatives of Dryopteris
spinulosa var. dilatata. Rhodora 17: 44-48. 17 Mr 1o15.
Fernald, M. L. Some new or unrecorded Compositae chiefly of north-
eastern America. Rhodora 17: 1-20. 6 F 10915,
Includes seven new species in Solidago (4), Aster (2) and Arnica (x).
Fernald, M. L., & St. John, H. Some anomalous species and varieties
of Bidens in eastern North America. Rhodora BT: 20-25. 6 F
1915.
Describes Bidens colpophila sp. nov.
Fink, B. William Wirt Calkins, amateur mycologist.
57-61. pl. 154. Mr 1915.
Fletcher, E. F. Two more foreign plants found on wool
Westford. Rhodora 17: 32. 6 F 1915.
Mycologia 7:
-waste at
Fromme, F. D. Negative heliotropism of urediniospore germ-tubes.
Am. Jour. Bot. 2: 82-85. f. 1, 2. F 1915.
INDEX TO AMERICAN BOTANICAL LITERATURE 361
Frothingham, E. H. The eastern hemlock (Tsuga canadensis (Linn.)
Carr). U.S. Dept. Agr. Bull. 152: 1-43. pl. 1-5 +f. 1-3. 3 F 1915.
Gates, F.C. A woody stem in Merremia gemella induced by high warm
water. Am. Jour. Bot. 2: 86—88. f. 7, 2.. F 1915.
Gates, R. R. On the nature of mutations. Jour. Heredity 6: 99-108.
i... 25 F ASTS:
Gormley, R. Onagraceae of Ohio. Ohio Nat. 15: 463-468. 12 Mr
IQI5.
Greenman, J. M. Descriptions of North American Senecioneae. Ann.
Missouri Bot. Gard. 1: 263-290. pl. 10-14. 30S 1914.
Griffiths, D. Hardier spineless cactus. Jour. Heredity 6: 182-191.
f. 15-19. 25 Mr 1915.
Harter, L. L. Notes on the distribution and prevalence of three im-
portant sweet potato diseases. Phytopathology 5: 124-126. Ap
IQI5.
Hasselbring, H. Effectos de la sombra sobre la transpiracién y la
asimilacién de la planta del tabaco en Cuba. Cuba Estac. Exp.
Agron. Bull. 24: 1-38. F 1915.
Hasselbring, H. Tipos de tabaco cubano. Cuba Estac. Exp. Agron.
Bull. 23: 1-15. pl. 1-7. Ja 1915.
Hasse, H. E. Additions to the lichen-flora of southern California.
No. 10. Bryologist 18: 22, 23. Mr 1915.
Hawkins, L. A. Some effects of the brown-rot fungus upon the com-
position of the peach. Am. Jour. Bot. 2: 71-81. F 1915.
Haynes, C. C. Hepaticae collected in Florida by Severin Rapp.
Bryologist 18: 19-22. Mr 1915.
Heald, F. D., Gardner, M. W., & Studhalter, R. A. Air and wind
dissemination of asco[s]pores of the chestnut-blight fungus. Jour.
Agr. Research 3: 493-526. pl. 63-65+f. 1, 2. 25 Mr 1915.
Holden, R. On the relation between Cycadites and Pseudocycas.
New Phytologist 13: 334-340. pl. 3+f. z. D 1914.
Jensen, J. Dendrologische Beobachtungen in dem Gebiete am Kopfe
des Michigansees. Mitteil. Deuts. Dend. Gesells. 1914: 184-188.
23 N 1914.
Jones, L. R., & Gilbert, W. W. Lightning injury to potato and cotton
plants. Phytopathology 5: 94-102. pl. 8, g. Ap 1915.
Lipman, C. B. A suggestion of a new phase of the problem of physio-
logical diseases of plants. Phytopathology 5: 111-116. Ap 1915.
362 INDEX TO AMERICAN BOTANICAL LITERATURE
Lloyd, C. G. Letter No. 55. 1-4. f. 627. Cincinnati. Mr 1915.
Includes a note on Polyporus (Ganodermus) gibbosus.
Lloyd, C. G. Synopsis of the Cordyceps of Australasia. 1-11. f. 611-
626. Cincinnati. Mr 1915.
Lloyd, C. G. Synopsis of the genus Fomes. 211-288. f. 570-610.
Cincinnati. Ja 1915.
Lorenz, A. List of Hepaticae to be expected in New England. Bry-
ologist 18: 24-26. Mr I915. ;
Merrill, G. K. Lecanora atrosanguinea Merrill and Blastenia Herret
Hasse identical. Rhodora 18: 29. 17 Mr 1915.
’ Monnet, P. Contributions a l'étude de la végétation du Grand Bassin
Américain. Rev. Gén. Bot. 26: 342-349. pl. g-r2. 15 Au 1914.
Murrill, W. A. The genus Lepista. Mycologia '7: 105-107. Mr 1915.
Murrill, W. A. Marking types in the mycological herbarium. My-
cologia 7: 108. Mr 1915.
Orton, W. A. Environmental influences in the pathology of Solanum
tuberosum. Jour. Washington Acad. Sci. 3: 180-190. f. I-3. 4 Ap
1913. :
Osterhout, W. J. V. Extreme alterations of permeability without
injury. Bot. Gaz. 59: 242-253. f. 1- 4. 17 Mr 1915.
Qsterhout, W. J. V. Normal and abnormal permeability. Am. Jour.
Bot. 2: 93, 94. F 1915.
Pepperberg, R. V. Preliminary notes on the carboniferous flora of
Nebraska. Nebraska Geol. Surv. 3: 313-329. pl. 1-12. Jl 1910.
Perkins, L.S. The pomerange. Jour. Heredity 6: 192. 25 Mr 1915.
A natural hybrid between the orange and pomelo.
Piper, C. V. Andropogon halepensis and Andropogon sorghum. Proc.
Biol. Soc. Washington 28: 25-44. 12 Mr 1915.
Prescott, A. The spinulose shield fern. Am. Bot. 21:17, 18. F 1915.
Reimer, F.C. Blight resistance in pears and pear stocks. Monthly
Bull. State Com. Hort. Calif. 4: 145-149. f. 23. Mr 1915.
‘Rehder, A. Einige neuere oder kritische Gehélze. Mitteil. Deuts.
Dend. Gesells. 1914: 257-263. 23 N IQI4.
Richardson, A. E. V. Wheat breeding. Jour. Heredity 6: 123-141.
f-IF-19. 25 F 1915,
Riddle, L. W. An undescribed species of Cetraria. Bryologist 18:
27, 28. Mr r9ts.
Cetraria pallidula Tuckerman.
INDEX TO AMERICAN BOTANICAL LITERATURE 363
Rolfe, R. A. Mormodes tigrinum. Curt. Bot. Mag. 11: pl. 8597.
fe 2k fO8S,
Rolfs,P.H. Tomatodiseases. Florida Agr. Exp. Sta. Bull. 117: 37-48.
Fat. Mis, IN SL,
Rorer, J.B. Bud-rot of the cocoanut palm. Dept. Agr. Trinidad and
Tobago Bull. 11: 68, 69. 1912.
Rorer, J. B. Some fruit diseases. Dept. Agr. Trinidad and Tobago
Bull. 11: 75, 76. 1912.
Sargent, C. S. Three of Clayton’s oaks in ‘the British Museum,
Rhodora 17: 39, 40. 17 Mr 1915.
Saunders, C. F. A garden of associations. Am. Bot. 21: 12-14. F
1915.
Schaffner, J. H. Peculiar varieties of Amaranthus retroflexus. Ohio
Nat. 15: 469-471. f. z. 12 Mr 1915.
Seaver, F. J. Photographs and descriptions of cup-fungi—I. Peziza.
Mycologia 7: 90-93. pl. 155, 156. Mr 1015.
Shear, C. L. Mycology in relation to phytopathology. Science II.
41: 479-484. 2 Ap 1915.
Sherman, L., & Schier, W. E. Field peace of the Wisconsin Myco-
logical Society for the seasons of 1912, 1913, and 1914. [Mil-
waukee.] [1-4.] [1915.]
Shreve, F. The direct effects of rainfall on hygrophilous vegetation.
Jour. Ecology 2: 82-98. pl. 14. Je 1914.
Smith, R. E. The investigation of ‘‘ physiological ”’ plant diseases.
Phytopathology 5: 83-93. Ap 1915.
Smith, R. E., & Boncquet, A.. New light on curly top of the sugar
beet. Phytopathology 5: 103-107. f. 1-3. Ap 1915.
Stapf, O. Pinguicula seat Curt. Bot. Mag. 11: pl. 8602. Mr
1915.
A plant from Mexico.
Steil, W. N. Apogamy in Nephrodium hirtipes. Bot. Gaz. 59: 254-
Stevens, F.L. Some problems of plant pathology in reference to trans-
portation. Phytopathology 5: 108-110. Ap I9I5.
Stewart, V. B., & Leonard, M. D. The réle of sucking insects in the
dissemination of fire blight bacteria. Phytopathology 5: 117-123.
Ap 1915.
364 INDEX TO AMERICAN BOTANICAL LITERATURE
St. John, H. An insular variety of Solidago sempervirens. “Rhodora
£772:26, 276-4 F-12915:
Stuart, W. Group classification and varietal descriptions of some
American potatoes. Bull. U.S. Dept. Agr. Professional Paper 176:
1-56. pl. 1-19. 27 Mr 1915. !
Turrill, W. B. Eugenia uniflora. Curt. Bot. Mag. 11: pl. 8590.
F 1915.
A plant from tropical South America.
Vries, H. de. The coefficient of mutation in Oenothera biennis L. Bot.
Gaz. 59: 169-196. 17 Mr 1915.
Webster, H. A rash mycophagist. Rhodora 17: 30-32. 6 F 1915.
A case of poisoning attributed by the author to Lepiota Mongaric [Morganil.
Wight, W. F. The varieties of plums derived from native American
species. Bull. U. S. Dept. Agr. Prof. Paper 172: 1-44. 13 Mr
Willis, J.C. On the lack of adaptation in the Tristichaceae and Podo-
stemaceae. Proc. Royal Soc. 87: 532-550. 1914.
Wooton, E. O. Cacti in New Mexico. New Mexico Agr. Exp. Sta.
Bull. 78: 1-70. My rorr. [Illust.]
Wright, C. H. Tillandsia regina. Curt. Bot. Mag. 11: pl. 8506.
F 1915.
A plant from Brazil.
BuLL_. TORREY CLUB VOLUME 42, PLATE I9
WAWooD
F4en
BIHAI HARRISIANA Griccs
BULL. TORREY CLUB VOLUME 42, PLATE 20
'
ot Ne
UNIFOLIUM BIFOLIUM (L.) GREENE
Vol. 42 No. 7
BULLETIN
OF THE
TORREY BOTANICAL CLUB
JULY, 1675
Studies of West Indian plants—VI
NATHANIEL LorRD BRITTON
35. THE GENUS COCCOLOBIS IN CUBA
A. Leaves 2-13 mm. long; inflorescence one- to three-flowered [Rurcra].
Leaves spinulose-mucronate. 1. C. armata.
Leaves emarginate at both ends. 2. C. microphylla,
. Leaves 2 cm. long or longer; inflorescence many-flowered.
1. Leaves spinulosemucronate.
Racemes loosely flowered; leaves acute. 3. C. pilonis.
Racemes densely flowered; leaves acuminate. 4. C. woodfredensis.
2. Leaves not spinulose-mucronate.
Leaves very large, ca pear deeply cordate; fruit
large; halophytic tree or shrub. 5. C. Uvifera.
Leaves oe rarely cae fruit small; mesophytes.
Pedicels filiform, in fruit 8-12 mm. long, as long as
e fruit or longer.
Leaves flat, their margins not revolute. 6. C. coriacea.
Leaves bullate, their margins strongly revolute. 7. C. Cowellii.
Pedicels epreel than the fruit.
of the raceme glabrous.
te almost veinless above, few-veined be-
n unded, or emarginate. 8. C. nipensis.
Leaves uae on both sides.
Fruiting pedicels 2.5-4 mm. long; fruit
ss)
9. C. laurifolia,
Fruiting pedicels 1.5 mm. long or less;
i er.
Leaves obtuse or emarginate at apex,
obtuse a i 10. C. retusa.
_ Leaves acuminate at both ends. tr. C. cubensis.
Leaves acute at apex, oblique at
base. 2. C. colomensis.
[The BULLETIN for June (42: 315-364. pl. r9, 20) was issued June 16, 1915.)
365
366 BriTTON: STUDIES OF WEsT INDIAN PLANTS
Rachis of the raceme puberulent or pubescent
(or glabrous in C. Wrighiii ?).
Raceme stout, 8-20 cm. long; leaves 8-17 cm.
ong; flowers sessile.
Leaves loosely reticulate-veined; flowers
white. ;
13. C. costata.
d ly reticulate-veined; flowers
bright re 14. C. Shaferi.
Raceme slender, shorter; leaves smaller.
Leaves not reticulate-veined above.
Raceme geniculate. 15. C. geniculata.
Fruiting pedicels less than 0.5 mm.
ong. 16. C. rufescens.
Fruiting pedicels 1-2 mm. long.
Leaf-margins reflexed. 17. C. reflexa.
Leaf-margins not reflexed. 18. C. praecox.
Leaves reticulate-veined above.
rt-acuminate. 19. C. benitensis.
icels 0.8 mm. long or less.
coriaceous, oblong-
lanceolate, 5 cm. long orless. 20. C. pallida,
Leaves chartaceous, ovate or
elliptic, larger.
Racemes as long as the
leaves or longer; petioles
5-10 mm. long.
Racemes shorter than the
leaves; petioles 3 mm.
long or less. ‘ . 22. C. brevipes.
Fruiting pedicels 1.5-3 mm. long. 23. C. Wrightii.
NS
H
. C. diversifolia.
1. C. ARMATA C. Wright; Griseb. Cat. Pl. Cub. 62, 283. 1866
TYPE LocALity: Near San Marco.
DIsTRIBUTION: Dry, rocky hills, Oriente, Santa Clara. En-.
demic.
The plant becomes much larger than the original description
indicates; on the Rio San Juan, Santa Clara, it forms a tree 6 m.
high, and on the United States Naval Station, Guantanamo Bay,
Oriente, trees up to 10 m. high were observed.
2. C, MICROPHYLLA C. Wright; Griseb. Cat. Pl. Cub. 62. 1866
TYPE LOCALITY: Western Cuba.
BRITTON: STUDIES OF West INDIAN PLANTS 367
DistRIBUTION: Coastal thickets, Santa Clara, Pinar del Rio.
Endemic.
At Coloma, Pinar del Rio, forms a virgate tree 5 m. high.
3. C. PILONIS Urban, Repert. 13: 445. 1914
Type LocaLity: Loma Pilon, near Holguin, Oriente.
DIsTRIBUTION: Barren rocky soil, Oriente, Santa Clara (?).
Endemic.
The Santa Clara specimens, collected near the city of Santa
Clara (Britton & Wilson 6189, Britton & Cowell 13325), are in
foliage only and thus not determinable with certainty.
Both this species and the following one are related to C. flaves-
cens Jacq., of Hispaniola. The type is from a shrub 1.6 m. high.
4. Coccolobis woodfredensis sp. nov.
Similar to C. pilonis, and to C. flavescens Jacq. (of Hispaniola),
the leaves spinulose-mucronate. A shrub, 6 m. high, glabrous
throughout, the rather stout branches zig-zag; leaves coriaceous,
ovate to ovate-lanceolate, 3-6 cm. long, shining above, dull be-
neath, inconspicuously veined, the under surface very finely
reticulated, the apex spinulose-mucronate, the base rounded,
obtuse or subcordate, the stout petioles scarcely 2 mm. long;
racemes very narrow, 3-4 cm. long, densely many-flowered; fruit-
ing pedicels about 1 mm. long; old fruit ovoid, acutish, 4 mm.
long, 2 mm. thick.
Dry rocky hillside, between Piedra Gorda and Woodfred,
Sierra Nipe, Oriente, 400-500 m. alt. (Shafer 3180).
5. C. Uvirera (L.) Jacq. Enum. 19. 1760
TYPE LOCALITY: Shores of the Caribbean Sea.
DIsTRIBUTION: Coastal thickets, all provinces and Isle of
Pines; occasionally on hillsides back from the coasts: Florida;
West Indies; continental tropical America.
6. C. cortAcEA A. Rich. in Sagra, Hist. Cub. 11: 184. 1850
C. calobotrys Meisn. in DC. Prodr. 14: 157. 1857.
TYPE LocALITy: Vuelta de Abajo, Cuba.
DISTRIBUTION: Pinar del Rio (?), Havana, mountains of
Oriente. -Endemic.
368 BRITTON: STUDIES OF WEsT INDIAN PLANTS
Meisner cites Sagra’s Nos. 290 and 544, on which he based
C. calobotrys, from the vicinity of Havana, but the plant is not
known to grow in Havana Province. I have not seen the type
Specimens. A specimen from Ensenada de Mora, Oriente
(Britton, Cowell & Shafer 13003), is from a slender tree 7 m. high.
7. Coccolobis Cowellii sp. nov.
A glabrous shrub, 1.5 m. high, with few, stout, ascending
branches. Leaves thick-coriaceous, shining, deeply cordate at the
nearly sessile base, bullate, with strongly revolute margins, diversi-
form, some ovate to ovate-elliptic, 5-9 cm. long, rounded at the
apex, some elongated-lanceolate, attenuate-acuminate, 10-18 cm.
ong; principal venation impressed above, prominent beneath;
ultimate venation obscure on both surfaces; racemes terminal,
nearly sessile, 1-3 dm. long, the rachis and pedicels glabrous;
pedicels filiform, 3-6 mm. long, spreading; bracts scarcely I
m. long, ovate, acute, about as long as the nearly truncate
ochreolae; flowers bright red, 3 mm. long; perianth-lobes oblong,
obtuse; fruit ovoid, acute, 4 mm. long, crowned by the calyx-lobes.
Barren rocky soil, savannas near Camaguey (Britton & Cowell
I3151).
8. C, NIpENsIS Urban, Repert. 13: 445. 1914
TYPE LOCALITY: Pine lands, 500-650 m. alt., Sierra Nipe,
near Woodfred, Oriente.
DISTRIBUTION: Mountains of northern Oriente. Endemic.
This species is noted by Dr. Shafer as forming a tree up to
10 m. high.
g. C. LAURIFOLIA Jacq. Hort. Schoen. 3: 9. pl. 267. 1798
C. floridana Meisn. in DC. Prodr. 14: 165. 1857.
TYPE LOCALITY: Caracas.
DISTRIBUTION: Thickets and woodlands at lower elevations
near the coasts, Oriente, Camaguey, Matanzas, Isle of Pines:
Florida; Bahamas; Jamaica; Hispaniola to St. Croix; Venezuela.
10. C. RETUSA Griseb. Cat. Pl. Cub. 61. 1866
C. leoganensis parvifolia Griseb. Cat. Pl. Cub. 61. 1866.
Hyponym.
BRITTON: STUDIES OF WEsT INDIAN PLANTS 369
TYPE LOCALITY: Eastern Cuba.
DIsTRIBUTION: Thickets and hillsides, Oriente, Santa Clara,
Pinar del Rio: Hispaniola (?).
11. C. CUBENSIS Meisn. in DC. Prodr. 14: 162. 1857
TYPE LOCALITY: Cuba [near Santiago].
DIsTRIBUTION: Oriente. Endemic.
Dr. Shafer’s 4229, from rich woods in the alluvial valley of
- Rio Yamaniguey, northern Oriente, referred to this species from
description, is from a tree 5 m. high.
12. Coccolobis colomensis sp. nov.
A glabrous shrub, 2 m. high, with slender twigs. Leaves
chartaceous, ovate to elliptic, 4-10 cm. long, bluntly acute at the
apex, obliquely obtuse at the base, strongly reticulate-veined and
shining on both sides, the rather stout petioles 6-10 mm. long;
racemes glabrous, terminal, nodding, much shorter than the leaves,
5 cm. long or less; ochreolae less than 0.5 mm. long; pedicels
about 1.5 mm. long, spreading; flowers about 1 mm. long, the
ovate perianth-lobes as long as the tube; anthers not exserted;
young fruit ovoid, about 3.5 mm. long, short-coronate.
Marsh near Coloma, Pinar del Rio (Britton & Gager 7037).
13. C. costaTa C. Wright; Sauvalle, Anales Acad. Habana 7: 343.
1870
C. leoganensis cordata Griseb. Cat. Pl. Cub. 61. 1866. Hyponym.,
TYPE LocALity: Cuba [C. Wright 1393, eastern Cuba].
DisTRIBUTION: Mountains of Oriente; Porto Rico.
Dr. Shafer’s 3084, which satisfactorily matches Wright
1393, is from a tree 8 m. high, at 400-500 m. alt. on the Sierra
Nipe, in dry rocky thickets; he noted the flowers as white and the
fruit red-black.
14. Coccolobis Shaferi sp. nov.
A shrub or small tree up to 4 m. high, the twigs and leaves
glabrous. Leaves coriaceous, ovate or ovate-elliptic to lanceolate,
12 cm. long or less, strongly and rather densely reticulate-veined
on both sides, obtuse, acute or acuminate at the apex, cordate at
the base, the stout petioles about 1 cm. long; spikes dense,
370 Britton: StupIEs OF WEsT INDIAN PLANTS
slender, 6-12 cm. long, the rachis, bracts and ochreolae puberulent;
flowers bright red, sessile, about 5 long, the short perianth-lobes
rounded.
Hillsides and thickets, northern Oriente.
Type, Shafer 4165, from between Camp Toa and Camp La
Barga, 400-450 m. alt.
15. C. GENICULATA Lindau, Bot. Jahrb. 13: 141. 1891
Type LocALity: Near Puerto Principe, Cuba.
DisTRIBUTION: Known only from the type locality.
Referred by Grisebach to C. punctata parvifolia Griseb.
16. C. RUFESCENS C. Wright; Sauvalle, Anales Acad. Habana 7:
343. 1870
C. punctata parvifolia Griseb. Mem. Am. Acad. II. 8: 175. 1860.
C. rufescens longifolia Lindau, Bot. Jahrb. 13: 143. 1891.
TYPE LOCALITY: Cuba.
DISTRIBUTION: Mountains of Oriente. Endemic.
17. C. REFLEXA Lindau, Bot. Jahrb. 13: 141. 1891
TypPE LocaLity: Cuba [Wright 2256].
DistTRIBUTION: Britton & Cowell 13115, from a tree 6 m. high,
in savannas near Camaguey, appears to be the same as Wright 2256.
18. C. pRAECOX C. Wright; Lindau, Bot. Jahrb. 13: 142. 1891
TYPE LOCALITY: Cuba.
DiIsTRIBUTION: Known only from the type specimens (Wright
2253).
19. Coccolobis benitensis sp. nov.
A much-branched shrub, up to 2 m. high, the twigs more or
less tortuous, glabrous. Leaves coriaceous, ovate, or elliptic-
ovate, 1.5-7 cm. long, glabrous, punctate, short-acuminate at the
apex, rounded or obtuse at the base, densely prominently reticu-
late-veined on both sides, the primary veins prominent beneath,
the petioles 1.5-4.5 mm. long; spike-like racemes very slender,
short-stalked, 4 cm. long or less, the ochreae and rachis puberulent;
flowering pedicels scarcely longer than the ochreolae, 0.5 mm. lon
or less; fruiting pedicels 1-1.5 mm. long; perianth (young) about
,
BRITTON: STUDIES OF WEST INDIAN PLANTS S71
0.5 mm. long; fruit ovate-ellipsoid, 5-6 mm. long, about 3 mm.
thick, dark red.
Wet mountains of northern Oriente. Type from vicinity of
Camp San Benito, 900 m. alt. (Shafer 4049).
20. C. PALLIDA C. Wright; Griseb. Cat. Pl. Cub. 61. 1866
TYPE LOCALITY: Western Cuba, near San Marco.
DISTRIBUTION: Known only from the type locality [Wright
2254].
21. C. DIVERSIFOLIA Jacq. Enum. 19. 1760
TYPE LOCALITY: Caribbean Islands.
DISTRIBUTION: Cuba, collected by Read, according to Lindau:
Bahamas; Jamaica; Hispaniola; Porto Rico; Vieques; St. Thomas;
St. Jan; St. Croix; Saba; Anguilla (?); Antigua; Montserrat ; Guade-
loupe; Dominica; Martinique; St. Lucia; Barbadoes; Bonaire;
Curacao; Aruba. Planted in Bermuda.
22. Coccolobis brevipes sp. nov.
Twigs slender, densely puberulent. Leaves chartaceous, ellip-
tic, 3-7 cm. long, bluntly acute at the apex, obliquely narrowed or
rounded at the base, shining above, glaucescent beneath, glabrous
and strongly reticulate-veined on both sides, the principal veins
prominent beneath; petioles stout, 2-3 mm. long; racemes terminal,
about 5 cm. long, densely puberulent; bracts puberulent, acute,
scarcely 1 mm. long; ochreolae about as long as the bracts;
flowering pedicels 0.5-0.8 mm. long; flowers about 1.5 mm. long,
the perianth-lobes ovate.
Cuba (Wright 2257 in herbarium of the Missouri Botanical
Garden). Lindau, in Bot. Jahrb. 13: 152, cites this number as
part of C. Wrightii Lindau, but the specimen here described is
different from Wright 1395, the type of C. Wrightit.
C. brevipes is similar to C. rufescens C. Wright in leaf-form and
pubescence, but the venation is quite different.
23. C. Wricutu Lindau, Bot. Jahrb. 13: 151. 1891 |
Type LocaLity: Near Monte Verde, Oriente.
DISTRIBUTION: Mountains of northern Oriente.
Referred by Grisebach to C. tenwifolia L.
372 Britton: STUDIES OF West INDIAN PLANTS
36. ADDITIONAL SPECIES OF TABEBUIA GoMEz
The type species is Bignonia Tabebuya Vell., of Brazil, a
species with simple leaves. It is not possible to separate the
group into two genera, the one with simple, the other with com-
pound leaves, as has been proposed by several authors,* calling
the compound-leaved species Tecomas, and the true type of
Tecoma Juss. is Bignonia stans L. (Fecoma stans HBK., Steno-
lobium stans Seem.). The two-lipped calyx of the type species of
Tabebuia is not constant through the group.
Tabebuia Brooksiana sp. nov.
A tree, up to 30 m. high. Leaves 4-7-foliolate; petioles 12 cm.
long or less; petioles slender, I-4 cm. long; leaflets oblong, oblong-
lanceolate or oblong-oblanceolate, chartaceous, 5—12 cm. long, 4
cm. wide or less, faintly shining above with the veins impressed,
dull, mostly finely reticulate-veined when mature, and lepidote
beneath, with the midvein and lateral veins prominent, obtuse,
acute or retuse at the apex, the base obtuse or acutish; flowers
clustered; calyx 2-lipped, 10-15 mm. long; corolla pink, 5-6 cm.
long, the tube about 5 mm. long, the narrowly campanulate
throat about 3.5 cm. long, the limb about 1.5 cm. long, its un-
dulate lobes ciliolate.
. Woodlands, provinces of Oriente and Camaguey, Cuba, from
sea-level up to 350 m. altitude. Type from Ensenada de Mora,
Oriente (Britton, Cowell & Shafer 12985). Referred by Grisebach
to Tecoma Leucoxylon Mart. and to T. Leucoxylon reticularis
Griseb. The species is named in honor of Mr. Theodore Brooks
_of Guantanamo, who has rendered important aid to scientific
exploration. Tecoma heptaphylla A. Rich. in Sagra, Hist. Cub.
Ir: 106 (not Martius), described as with 7-foliolate leaves from
Vuelta de Abajo (Pinar del Rio), Cuba, is not known to me; I
have seen no species with 7-foliolate leaves from western Cuba.
>
Tabebuia Shaferi sp. nov.
A tree, up to 10 m. high. Larger leaves 6-8-foliolate; passes
3-4 cm. long; petiolules 2 cm. long or less; leaflets ovate-oblong to
suborbicular, chartaceous, the larger 6 cm. long, 3-5 cm. wide,
dark green above, paler beneath, minutely and densely reticulated
* See Urban, Symb. Ant. 7: 377.
BRITTON: STUDIES OF WeEstT INDIAN PLANTS 373
on both sides, the midvein impressed above, prominent beneath,
the apex rounded or obtuse, the base subcordate; ‘flowers
pinkish’’; capsule 12 cm. long; seeds 5 mm. long, 7 mm. wide, the
wings 5 mm. wide.
Along a small stream in the Pinales southeast of Paso Estancia,
Oriente, Cuba (Shafer 1710a). Apparently the same as C. Wright
3043, referred by Grisebach to Tecoma Leucoxylon Mart., forma
foliolis latioribus.
Tabebuia pachyphylla sp. nov.
A tree, upto12m.high. Petioles stout, 6-8 cm. long; petiololes
stout, 4 cm. long or less; leaflets 5-7, oblong to ovate-oblong,
coriaceous, 5-12 cm. long, 2.5-6 cm. wide, lustrous above, dull
and densely minutely reticulate-veined beneath, the midvein
prominent, the lateral veins slender, the apex bluntly acute, the
base rounded or subcordate; flowers pink, clustered; calyx 15 mm.
long, bluntly lobed; corolla pink, glabrous, about 5 cm. long, its
narrow throat about twice as long as the limb, its lobes rounded.
Mountains of northeastern Cuba. Type specimen collected
at Arroyo del Medio, Sierra Nipe, Oriente, Cuba, 450-550 m.
alt. (Shafer 3645).
Tabebuia trinitensis sp. nov.
tree up to 10 m. high. Leaves 4-foliolate or 5-foliolate;
petioles slender, 8-11 cm. long; petiolules slender, 4 cm. long or less;
leaflets thin-chartaceous, oblong or oblong-lanceolate, dark green
above, paler and lepidote beneath, the larger ones 11-15 cm. long,
3-4 cm. wide, bluntly acute or acuminate at the apex, obtuse at
the base, finely skier oan and the primary venation rather
prominent on both s
Hanabanilla Falls, ‘Trinidad Mountains, province of Santa
Clara, Cuba (Britton, Earle & Wilson 4866)
The description is from leaf-specimens only but these indicate
that the tree is a species distinct from all others of Cuba. The
ultimate venation and texture of the leaflets differentiate it from
T. Brooksiana Britton.
Tabebuia calcicola sp. nov.
A slender tree, up to 6 m. high. Leaves 2-5-foliolate; petioles
stout, lepidote, 4-5 cm. long; petiolules lepidote, 1.5 cm. long or
less: leaflets coriaceous, light green, ovate to ovate-oblong, 3 5-8
te
wR
et
374 Britton: STUDIES OF West INDIAN PLANTS
cm. long, 5 cm. wide or less, acute or bluntish at the apex, rounded
at the base, finely reticulate veined and densely lepidote on both
sides, the primary venation impressed above, prominent beneath;
flowers clustered; pedicels stout; calyx 12-14 mm. long, shortly
2-lipped; corolla about 5 cm. long, the tube about 5 mm. long,
the campanulate white throat 3 cm. long, the pink limb about
1.5 cm. long, the undulate lobes ciliolate; capsule 10-14 cm. long.
Limestone rocks and cliffs, province of Pinar del Rio, Cuba.
Type collected near Guane (Britton & Cowell 9772).
Tabebuia moaensis sp. nov.
Ashrub, about m. high. Leaves 3-5-foliolate; petioles stout,
2 cm. long or less; petiolules rather stout, 4-15 mm. long; leaflets
elliptic to obovate, coriaceous, 3-5 cm. long, 1-2 cm. wide, lustrous
above, dull and minutely reticulate-veined beneath, the primary
venation not very prominent; calyx 10 mm. long, 2-lipped;
corolla pink, about 4 cm. long, its rounded lobes ciliolate.
Camp La Gloria, south of Sierra Moa, Oriente, Cuba (Shafer
6264). This is, apparently, the same as C. Wright 3047, referred
by Grisebach to Tecoma haemantha.
Tabebuia pinetorum sp. nov.
A shrub, about 2.6 m. high. Leaves 3-foliolate or 4-foliolate;
petioles very stout, only 3 cm. long or less, petiolules stout; 3-8
mm. long; leaflets elliptic or ovate-elliptic, coriaceous, 5-14 cm.
long, 2.5-8 cm. wide, shining above, obtuse, rounded or bluntly
short-pointed at the apex, obliquely cordate at the base, dull,
lepidote and finely reticulate-veined beneath, the primary venation
impressed above, prominent beneath; peduncles stout; calyx 12
mm. long; somewhat 2-lipped; ‘flowers lilac’; pod 11 cm. long,
about I cm. thick.
Pine woods, Baracoa, Oriente, Cuba (Underwood & Earle 1362).
Tabebuia arimaoensis sp. nov.
A small tree with slender, whitish twigs. Leaves 3-foliolate;
petioles slender, 1-2.5 cm. long; leaflets subcoriaceous, shining
above, with the midvein impressed dull beneath with the mid-
vein prominent, lepidote on both sides, acute, the terminal one
with a petiolule 2-5 mm. long, cuneate-oblanceolate, 4-5 cm. long,
8-12 mm. wide, the lateral ones sessile, narrowly oblong, inequi-
lateral, obliquely narrowed at the base. Flowers and fruit
unknown.
‘cat
hare
gg
BRITTON: STUDIES OF WEsT INDIAN PLANTS 375
Rocky hillside, Rio Arimao, province of Santa Clara, Cuba
(Britton & Wilson 5797).
Tabebuia arenicola sp. nov.
A tree 7 m. high. Leaves 3-foliolate; petioles slender, 1-2.5
cm. long; leaflets subcoriaceous, shining, sparingly lepidote, and
obscurely veined above, dull, closely lepidote and with prominent
midvein and slender lateral veins beneath, inconspicuously
reticulate-veined, obtuse or emarginate at the apex, the terminal
one oblanceolate, 6-7 cm. long, 1.2-2 cm. wide, acute at the base,
with a petiolule 4-6 mm. long, the lateral ones oblong, 4-6 cm.
long, sessile by a very oblique base.
Sandy plain, Conde Beach, Guantanamo Bay, Oriente, Cuba
(Britton 2142).
Tabebuia geronensis sp. nov.
Twigs stout. Leaves 1-3-foliolate; petioles slender, 1 cm.
long or less; in 3- foliglate leayes,. the lateral leaflets sessile, the
terminal one shorfstalked = leaflets oblong to ovate-elliptic or
ovate-oblong, subcoriaceous, 3.5-7 cm. long, 3 cm. wide or less,
obtuse and rounded or apiculate at the apex, rounded, subcordate,
or lateral ones obliquely narrowed at the base, dull on both sides,
lepidote above, finely and strongly reticulate-veined and densely
lepidote beneath; fruiting calyx densely lepidote, 1 cm. long;
capsule 8-9 cm. long, about 8 mm. thick, with a slender tip 6 mm.
long.
Nueva Gerona, Isle of Pines, Cuba (A. H. Curtiss, May, 1904).
Tabebuia Curtissii sp. nov.
Young twigs, leaves, pedicels and calyx densely lepidote-
scurfy. Leaves 3-foliolate, or the upper and lower 1-foliolate;
leaflets coriaceous, those of 1-foliolate leaves elliptic, 3-6 cm. long,
obtuse at both ends, those of 3-foliolate leaves obovate or oblong-
obovate, 8 cm. long or less, abruptly acute at the apex, narrowed
at the base, the terminal one short-stalked, the lateral ones sessile,
all smooth and shining above, dull, reticulate-veined and densely
lepidote beneath; flowers in terminal clusters; pedicels 2 cm.
long or less; calyx narrowly campanulate, 1.5 cm. long, its teeth
triangular, acute; corolla 5 cm. long, its lobes broad and rounded.
Nueva Gerona, Isle of Pines, Cuba (A. H. Curtiss, May, 1904).
376 BRITTON: STUDIES OF WEsT INDIAN PLANTS
Tabebuia crassifolia sp. nov.
A tree, 5 m. high. Leaves simple, oblong-elliptic, coriaceous,
6-12 cm. long, 4.5 cm. wide or less, obtuse or emarginate at the
apex, somewhat narrowed at the base, when young densely
lepidote and dark green above, pale and more densely lepidote
beneath; when old strongly shining, elepidote and with impressed
midvein above, remaining densely lustrous-lepidote and with very
prominent midvein beneath, the secondary venation slender,
neither surface reticulate-veined, the stout lepidote petioles 10-14
mm. long; flower solitary in an upper axil, its slender peduncle 3
cm. long, bearing a linear bractlet 2 mm. long below the middle;
calyx densely lepidote, 1 cm. long, lobed but scarcely two-lipped;
corolla light purple, about 5 cm. long.
By a spring, barren savannas southeast of Holguin, Oriente,
Cuba (Shafer 1285).
Tabebuia angustata sp. nov.
A tree up to12m. high. Leaflets 3-7-foliolate; petioles slender,
5-13 cm. long; petiolules slender, 5 cm. long or less; leaflets
chartaceous, oblong or oblong-lanceolate, 5-18 cm. long, 9 cm.
wide or less; dull on both sides, more or less lepidote, at least
when young, densely and finely reticulate-veined beneath with a
prominent midvein and slender lateral veins, the apex bluntly
acuminate or acute, the base obtuse or acute: flowers clustered ;
calyx 2-lipped, 12-16 mm. long; corolla rosy-white, 5-6 cm. long,
its narrow throat somewhat longer than the limb, its lobes ciliate;
' capsule 10-25 cm. long.
Woodlands and along streams at lower and middle elevations
_in wet or moist parts of Jamaica. Type collected on the south-
_ western slope of Dolphin Head (Harris 9253). This tree has
been referred to Tecoma platyantha Griseb.
Tabebuia jamaicensis sp. nov.
A tree up to 12 m. high. Leaves 5-foliolate; petioles stout,
15 cm. long; petiolules stout, 2-9 cm. long; leaflets chartaceous,
the three upper obovate, 18-23 cm. long, 9-12 em. wide, somewhat
narrowed at the base, the two lower ovate-elliptic, about 10 cm.
long and 6 cm. wide, rounded or subtruncate at the base, all
abruptly acuminate at the apex, finely reticulate-veined and loosely
lepidote on both sides, the midvein and lateral veins prominent
beneath; corolla whitish, glabrous, 5 cm. long, its cylindric tube
about 8 mm. long, its campanulate throat 2.5 cm. long, its spread- -
ing limb about 1.5 cm. long, its lobes erose. _
BRITTON: STUDIES OF WEST JNDIAN PLANTS 377
Hillside, Negril, Jamaica (Britton 2020). The description is
based on one leaf and one flower.
Tabebuia actinophylla (Griseb.)
Tecoma actinophylla Griseb. Cat. Pl. Cub. 194. 1866.
TYPE LOCALITY: Western Cuba.
DISTRIBUTION: Limestone cliffs, province of Pinar del Rio,
Cuba. .
Tabebuia Sauvallei nom. nov.
Tecoma sanguinea C. Wright; Sauvalle, Anales Acad. Habana
6: 320. 1870. Not Tabebuia sanguinea DC..
TYPE LocALity: Border of Rio Carbuni, Manacal, near
Trinidad, Cuba.
DistRIBUTION: Hillsides and river banks, near Trinidad,
province of Santa Clara, Cuba.
Tabebuia Buchii (Urban)
Tecoma Buchiti Urban, Symb. Ant. 3: 375. 1903.
Known only from the type locality, dry calcareous mountains,
Morne La Pierre, Haiti.
Tabebuia lepidota (HBK.)
Bignonia lepidota HBK. Nov. Gen. 3: 139. 1818.
Tecoma lepidota DC. Prodr. 9: 220. 1844.
_ ‘TYPE LocaLity: Havana, Cuba.
DISTRIBUTION: Barren soil, provinces of Camaguey, Santa
Clara, Matanzas and Havana, Cuba; Bahamas.
Referred by Grisebach to Tecoma lepidophylla and to Tecoma
Leucoxylon and by Combs to Tabebuia lepidophylia.
’ The species consists, apparently, of many races, differing in
size, number and form of leaflets and size of flowers.
Tabebuia Berterii (DC.)
Tecoma Berterit DC. Prodr. 9: 220. 1845.
_ TYPE LocaLity: Santo Domingo.
DistRIBUTION: Hispaniola, ascending to 1,200 m. altitude.
i
378 BRITTON: STUDIES OF. WEST INDIAN PLANTS
Tabebuia domingensis (Urban)
Tecoma domingensis Urban, Symb. Ant. 7: 376. 1912.
TyPE LocALITY: Near Barahona, Santo Domingo.
DISTRIBUTION: Known only from the type locality.
Tabebuia revoluta (Urban)
Tecoma revoluta Urban, Symb. Ant. 7: 539. 1913.
TYPE LOCALITY: Between La Vega and Jarabacoa, Santo
Domingo, at 700 m. alt.
DistTRIBUTION: Known only from the type locality.
Tabebuia acrophylla (Urban)
Tecoma acrophylla Urban, Symb. Ant. 3: 374. 1903.
TYPE LOCALITY: Mountains near Bilboro, Haiti, at 600 m. alt.
DISTRIBUTION: Known only from the type locality.
Tabebuia bibracteolata (Griseb.)
Tecoma bibracteolata Griseb. Cat. Pl. Cub. 193. 1866.
TYPE LOCALITY: Eastern Cuba.
DISTRIBUTION: Known only from the type specimens.
TABEBUIA HETEROPHYLLA (DC.) Britton, Ann. Mo. Bot. Gard. 2:
48. 1915
Rapuntia (?) heterophylla DC. Mém. Mus. Paris 9: 153. 1822.
Tabebuia triphylla DC. Prodr. 9: 214. 1845. Not Bignonia
triphylla L.
Sandy soil and rocky hillsides, Mona, Porto Rico, Vieques,
Culebra, St. Thomas, St. Croix, St. Jan, Virgin Gorda, Anagada;
recorded from St. Barts and Barbadoes.
Referred by Grisebach to Tecoma Berterii DC., and by Urban
to Tecoma Leucoxylon (L.) Mart.; this is not Tabebuia leucoxyla
DC. Prodr.:9= 232:
Tabebuia myrtifolia (Griseb.)
Tecoma myrtifolia Griseb. Mem. Am, Acad. II. 8: 524. 1862.
Tecoma lepidota myrtifolia Maza, Anal. Hist. Nat. Madrid 19:
265. shee
BRITTON: STUDIES OF WEsT INDIAN PLANTS 379
? Bignonia microphylla A. Rich. in Sagra, Hist. Cub. 11: 104.
1850. Not Lam.
TYPE LOCALITY: Matanzas, Cuba.
DISTRIBUTION: Coastal thickets, vicinity of Matanzas, Cuba.
Tabebuia platyantha (Griseb.)
Tecoma platyantha Griseb. Fl. Br. W. I. 447. 1864.
Tecoma Brittonit Urban, Symb. Ant. 5: 496. 1908.
Tecoma Brittonii decussata Urban, loc. cit. 497. 1908.
TYPE LOCALITY: On rocks, St. James, Jamaica.
DISTRIBUTION: Rocky woodlands, central parishes of Jamaica.
Tabebuia bahamensis (Northrop)
Bignonia pentaphylla L. Sp. Pl., ed. 2, 870. 1763 (as to the
Bahama plant of Catesby). Not Tabebuia pentaphylla (Juss.)
Hemsl.
Tecoma bahamensis Northrop, Mem. Torr. Club 12: 65. pl. 15.
1902.
TYPE LOCALITY: Near Nassau, New Providence, Bahamas.
DISTRIBUTION: Bahamas; Cuba. _
37- THE GENUS ANASTRAPHIA D. Don
Type species: Anastraphia ilicifolia D. Don.
Founded on a specimen in the Lambertian Herbarium said
to have come from South America; the plant was correctly
attributed to Cuba by De Candolle (Prodr. 7: 26. 1838)
1. Involucral bracts straight, erect or nearly so.
Involucre 2.5—3 cm. long
At least some of the leaves spinulose-dentate.
Involucre campanulate.
All the involucral bracts lanate. 1. A. ilicifolia.
Only the gies ncehied lanate, the inner glabrous.
Leaves distinctly reticulate-veined above.
All the invoices bracts linear-lanceo-
acuminate and erect. 2. A. mantuensis.
merely acute, all somewhat ing. 3. A. montana.
Leav
above, but wrinkled; outer bracts ovate. 4. A. Cowellii.
Inyo’ long-attenuate at base, all its many ;
bracts lanate 5. A. aitenuata.
380 Britton: STUDIES OF WEST INDIAN PLANTS
None of the leaves spinulose-dentate.
Involucral bracts tomentose. 6. A. Picardae.
Inner involucral bracts glabrous. 4. A. crassifolia.
Involucre 2 cm. long or less.
Involucre 1.5—2 cm. long.
Leaves or some of them 1.5~2 cm. lon
Inner involucral bracts obtusish or short-
pointed. A. oligantha.
Inner involucral bracts acuminate. 9. A. bahamensis.
Leaves 12 mm. long or less.
Leaves strongly spinulose-dentate. 10. A. calcicola.
Leaves repand-denticulate. 11. A. parvifolia.
Involucre 1.5 cm. long or less
Leaves 2-3 times as long as wide
Leaves pee atE denticulat
Lea rongly Rae beneath. 12. A. Wilsoni.
Leaves ah reticulated beneath, or faintly
reticulated.
Achenes short-tomentose; leaves reticu-
late-veined above. 13. A. intertexta,
Achenes villous; leaves smooth above. 14. A. Shaferi.
Leaves entire-margined. - 15. A. obtusifolia.
Leaves not longer than wide, or sete little longer.
eaves obtuse or truncate a
Involucre 6 mm. long. 16. A. microcephala,
Involucre 10 mm. long. 17. A. Rosei
Leaves cuneate at base. 18. A. Buchii.
z: pease oi pe recurved, or with recurved tips.
nvolucral bracts arachnoid. 19. A. Northropiana,
volucral bewehe villous. 20. A. recurva.
3. Involucre known only in a young condition; leaves oblong,
4 cm. long, 1.8 cm. wide, entire-margined or with
tooth near the apex, 21. A. lomensis.
ANASTRAPHIA ILICIFOLIA D. Don, Trans. Linn. Soc. 16: 296.
1830
Gochnatia ilicifolia Less. Linnaea 5: 261. 1830 (attributed to
Guiana doubtfully).
TYPE LOCALITY: South America [in error].
DIsTRIBUTION: Limestone cliffs and river-gorges, provinces of
Matanzas and Havana, Cuba. A shrub 2-3 m. high, the leaves
shining above.
A specimen in Kew Herbarium from rocks on the Rio Canimar,
Cuba, 1823, bears the label Staehelina ilicifolia Mutis., but that
species is a South American Gynoxys.
BRITTON: STUDIES OF WEsT INDIAN PLANTS 381
2. ANASTRAPHIA MANTUENSIS C. Wright; Griseb. Cat. Pl. Cub.
158. 1866
TYPE LOCALITY: Pine lands near Mantua, Cuba.
DIsTRIBUTION: Pine lands and rocky hillsides, near Mantua,
-Proyince of Pinar del Rio, Cuba.
3. Anastraphia montana sp. nov.
A depressed, spreading or prostrate shrub, the young twigs
white-floccose. Leaves oblong, or oblong-oblanceolate, 2-4 cm.
long, 8-16 mm. wide, coriaceous, sharply spinulose-dentate all
around, dark green, shining and densely reticulate-veined above,
white-floccose beneath, obtuse or acutish at the apex, the floccose
petioles 1-2 mm. long; heads solitary at the ends of branches;
involucre campanulate about 2.5 cm. high, its bracts rigid,
somewhat spreading, in about 8 series, the outer ovate to lanceo-
late, slightly lanate, acute, the inner linear-lanceolate, acuminate
glabrous achenes linear, narrowed at base, densely appressed-
pubescent, 6 mm. long; pappus light brown, spreading, about
twice as long as the achene
Top of Sierra Caliente, south of Sumidero, Pinar del Rio,
Cuba, August, 1912 (Shafer 13781).
4. Anastraphia Cowellii sp. nov.
A shrub, 2 m. high, the young twigs lanate- puberulent.
Leaves oblong, 1.5-3.5 cm. long, 6-15 mm. wide, coriaceous,
sharply spinulose-dentate, green, glabrous, shining and indistinctly
veined, but wrinkled above, brownish-floccose beneath, mostly
obtuse at the apex, obtuse or narrowed at the base, the petioles
1-4 mm. long; heads solitary or rarely 2 together; involucre
narrowly campanulate, about 3 cm_ high, its bracts in 7 or 8
series, the outermost ovate, lanate, the middle ones lanceolate, the
inner linear-lanceolate, acuminate, glabrous; corollas orange, 2
cm. long; achenes linear, 6~7 mm long, densely pubescent; pappus
brownish, spreading, about 3 times as long as the achene
Rocky hills, palm barren, city of Santa Clara, Cuba (Britton
& Cowell, 10183, type); same locality (Britton & Wilson 6073);
rocky places in savanna, Queen City to Minas, Camaguey, Cuba
(Shafer 2927); savanna near Camaguey (Britton & Cowell 13248).
5. Anastraphia attenuata sp. nov.
A shrub about 2.5 cm. high, the young twigs densely lanate.
Leaves oblong to oblanceolate, 1.5-5 cm. long, 8-17 mm. wide,
382 Britton: Stupies oF West INDIAN PLANTS
coriaceous, entire or some of them remotely spinulose-dentate,
green, glabrous and reticulate-veined above, whitish-lanate be-
neath, obtuse or acute at the apex, narrowed at the base, the
petioles 1-2.5 mm. long; heads solitary at the ends of branches;
involucre obconic, long-attenuate at the base, 3-3.5 cm. long, its
bracts lanate, imbricated in about 14 series, the lower ovate, the
upper lanceolate; corolla yellow, about 1 cm. long; achenes linear,
densely pubescent, 6 mm. long; pappus tawny, a little longer
than the achene
Moist thickets, Sierra Nipe, Oriente, Cuba, between Piedra
Gorda and Woodfred, at 400-500 meters elevation (Shafer 3113).
6. ANASTRAPHIA PICARDAE Urban, Symb. Ant. 3: 414. 1903
TYPE LocaLity: Near Cadets, Haiti, at 1200 m. alt.
DistripuTION: Known only from the type locality.
7. Anastraphia crassifolia sp. nov.
A shrub about 1.6 m. high, the young twigs densely tomentose.
Leaves oblong, or oblong-oblanceolate, coriaceous, 4-7 cm. long,
I-2 cm. wide, entire-margined, green, shining, and with impressed
veins above, lanate-tomentose and with prominent veins beneath;
involucre campanulate, nearly 3 cm. high, its bracts in 7 or 8
series, the outer ovate, lanate, the inner linear-lanceolate, glabrous,
long-attenuate.
Near mangroves, mouth of Rio Yamaniguey, Oriente, Cuba
(Shafer 4261). Only one old involucre was collected.
8. ANASTRAPHIA OLIGANTHA Urban, Symb. Ant. 3: 417. 1903
TYPE LocaLity: Santo Domingo.
DisTRIBUTION: Santo Domingo. Known only from the type
specimens collected by Wright, Parry and Brummell in 1871,
and from plate 123, f. 2, of Plumier, with which they were identified
by Professor Urban. Referred to by Hitchcock (Rep. Mo. Bot.
Gard. 4: 102) as A. pauciflosculosa C. — an unpublished
species.
9. ANASTRAPHIA BAHAMENSIS Urban, Symb. Ant. 3: 415. 1903
Anastraphia pauciflosculosa Hitchc. ae Mo. Bot. Gard. 4: 102.
pl. 12. teoe Hyponym.
BRITTON: STUDIES OF WEsT INDIAN PLANTS 383
Anastrapma cunetfolia Greenm. Bull. N. Y. Bot. Gard. 4: 126.
1905.
TYPE LocALity: Andros Island, Bahamas.
DIstTrIBuTION: Andros, Eleuthera, Cat Island, Fortune Island,
Crooked Island, Acklin’s Island, Great Exuma, Caicos Islands,
Mariguana, Dellis Cay, Inagua.
I cannot distinguish, specifically, from this species, a plant of
the Sierra Nipe, Oriente, Cuba (Shafer 3185).
1o. Anastraphia calcicola sp. nov.
A shrub, I-1.5 m. high, the young twigs lanate, the branches
stiff. Leaves cuneate-obovate or nearly elliptic, small, 5-10 mm.
long, nearly as wide as long, sharply 1—-3-toothed on each margin,
coriaceous, obtuse or subtruncate at the apex, glabrous and densely
reticulate-veined above, white-lanate beneath, the petioles about
I mm. long; heads solitary on short branchlets; involucre 1.5-2
cm. high, its bracts in about 4 series, loosely and sparingly lanate,
the outer ovate, acute, the inner narrowly lanceolate, acuminate;
achenes linear, 4 mm. long, densely pubescent; pappus yellowish-
brown, about 3 times as long as the achene.
Coral rock hillsides and coastal cliffs, United States Naval
Station, Guantanamo Bay, Oriente, Cuba (Britton 2042, type;
2220).
11. Anastraphia parvifolia sp. nov.
An upright shrub, 2 m. high, the young twigs lanate. Leaves
oblong to oblong-obovate, small, 5-10 mm. long, 4-6 mm. wide,
coriaceous, obtuse at the apex, obtuse or narrowed at the base,
dark green, smooth and shining above, white-lanate beneath, the
margin repand-denticulate, the slender petiole 1-3 mm. long;
involucre about 2.4 cm. long, its bracts in 5 or 6 series, the outer
ovate to ovate-lanceolate, loosely lanate, acute, the inner linear-
lanceolate, acuminate, with lanate tips; corolla orange-yellow,
18 mm. long; achenes linear, densely pubescent, about 3 mm.
long; pappus yellowish-brown, about 5 times as long as the achene.
Dense thickets in barren savannas southeast of Holguin,
Oriente, Cuba (Shafer 2938).
12. Anastraphia Wilsoni sp. nov.
An intricately branched shrub 2 m. high, the young twigs
somewhat lanate. Leaves oblong to oblong-obovate, subcoria-
384 Britton: STUDIES OF WeEsT INDIAN PLANTS
ceous, I-2.5 cm. long, 1 cm. wide or less, obtuse or acute at the
apex, narrowed at the base, green, shining, and densely finely
reticulate-veined above, brownish-tomentulose and prominently
veined beneath, the margins strongly spinulose-dentate, the slender
petioles 1.5-3 mm. long; heads solitary on short branchlets;
involucre 1-1.5 cm. high, its bracts in about 8 series, the outer
ovate, acute, densely lanate, the inner lanceolate to linear-lanceo-
late, attenuate, nearly glabrous; corolla 1.5 cm. long; achenes
densely pubescent, 5 mm. long; pappus tawny, 3 times as long
as the achene.
Rocky bank, Rio Toyaba, near Trinidad, province of Santa
Clara, Cuba (Britton & Wilson 5573).
13. ANASTRAPHIA INTERTEXTA C. Wright; Griseb. Cat. Pl. Cub.
158. 1866
Type LocALIty: Near San Marcos, district of Bahia Honda,
Pinar del Rio, Cuba.
DistRIBUTION: Mountain sides, Pinar del Rio, Cuba.
14. Anastraphia Shaferi sp. nov.
An intricately branched shrub 2-2.5 m. high, the young twigs
lanate. Leaves oblong to oblong-oblanceolate, 10-18 mm. long,
8 mm. wide or less, coriaceous, sharply spinulose-dentate, obtuse at
the apex, narrowed at the base, green, glabrous and smooth above,
whitish-lanate and rather prominently veined beneath, the petioles
about 1.5 mm. long; heads solitary or sometimes 2 together at the
ends of branchlets; involucre about 1 cm. long, its villous-lanate
bracts in about 5 series, the outer ovate, the inner lanceolate,
acuminate; corolla “whitish,” about 11 mm. long; achenes vil-
lous-pubescent, 3 mm. long; pappus tawny, 2-3 times as long
as the achene.
Dry cliff, below the falls of Rio Naranja, Oriente, Cuba, 450-
550 m. elevation (Shafer 3865).
15. Anastraphia obtusifolia sp. nov.
An intricately branched shrub, the young twigs lanate-
tomentulose. Leaves oblong to oblanceolate, 2.5 cm. long or less,
4-10 mm. wide, coriaceous, entire-margined, dark green and
finely densely reticulate-veined above, white-lanate and incon-
spicuously veined beneath, obtuse at the apex, narrowed at the
base, the petioles I-1.5 mm. long; head (only one old one seen
BRITTON: STUDIES OF West INDIAN PLANTS 385
attached) sessile, lateral; involucre 9-10 mm. long, its bracts
in 5 or 6 series, the outer ovate, acute; the inner linear-lanceolate,
acuminate.
Camp La Gloria, south of Sierra Moa, Oriente, Cuba (Shafer
8189).
16. ANASTRAPHIA MICROCEPHALA Griseb. Cat. Pl. Cub. 158. 1866
TYPE LOCALITY: Eastern Cuba.
DISTRIBUTION: Known only from the type specimens collected
by Wright.
17. Anastraphia Rosei sp. nov.
A shrub with tortuous branches, the young twigs floccose.
Leaves broadly elliptic, elliptic-obovate or suborbicular, 10-15
mm. long, 8-13 mm. wide, obtuse or nearly truncate at the apex,
obtuse or subtruncate at the base, dark green, loosely floccose or
becoming glabrate, and finely reticulate-veined above, densely
white-tomentulose and with few rather prominent veins beneath,
spiny-toothed, with 2 or 3 teeth on each side, the slender tomentu-
lose petioles 3-5 mm. long; heads solitary and sessile on short
spurs, about 10-flowered; involucre narrowly obconic, 10-11 mm
long, its bracts in about 5 series, all appressed-villous, acute, the
outer ovate, the inner linear-lanceolate; achenes densely silky,
4 mm. long; pappus yellow-brown, 8 mm. long.
Hillside, Azua, Santo Domingo, March 18, 1913 (Rose, Fitch
& Russell 4023). Related to A. Buchii Urban, of Haiti, which
has leaves cuneate at base, smaller heads and less hairy involucre-
bracts.
18. ANASTRAPHIA Bucutt Urban, Symb. Ant. 5: 527. 1908
TYPE LOCALITY: Dry woods, Poste Coudau, Haiti, at 100 m,
elevation.
DIsTRIBUTION: Known only from the type locality.
19. ANASTRAPHIA NORTHROPIANA Greenm.; Combs, Trans. Acad.
Sci. St. Louis 7: 435. pl. 36. 1897
Anastraphia Northropiana Combsii Urban, Symb. Ant. 3: 417.
1903.
TYPE LOCALITY: Fresh Creek, Andros, Bahamas.
BawaMA IsLanps: Andros, New Providence, Cat Island.
386 BRITTON: STUDIES OF WEST INDIAN PLANTS
Cusa: On rocky coastal hills, Punta Piedra, Nipe Bay,
Oriente (Britton & Cowell 12459); harbor of Santiago, Oriente
(Britton 1877); near Nuevitas, Camaguey (Shafer 839); Calicita,
Santa Clara (Combs 521); Rio San Juan, Santa Clara (Britton,
Earle & Wilson 5830, 5864); apparently the same species,
collected in foliage only, on Cayo Ratones, Bay of Mariel, Pinar
del Rio (Britton & Gager 7678). 1 cannot specifically distinguish
the Cuban plant from the Bahamian; on the southern coast of
Santa Clara Province it forms a tree up to 10 m. high with a
trunk 2 dm. in diameter; its bark is very rough.
20. Anastraphia recurva sp. nov.
A much-branched shrub up to about 3 m. high, the young
twigs tomentulose. Leaves oblong to elliptic or oblanceolate,
coriaceous, 4 cm. long or less, 8-15 mm. wide, green and smooth
or somewhat reticulate-veined above, lanate beneath, obtuse or
acute at the apex, cuneate-narrowed, acute or obtuse at the base,
spinulose-dentate or entire, the petioles 1.5-3 mm. long; heads
solitary at the ends of branchlets; involucre broadly campanulate,
1.5-1.8 cm. high, its bracts densely villous, imbricated in 6 or 7
series, recurved-spreading, linear-lanceolate, acuminate; corolla
yellow or yellowish, about 1 cm. long; achenes densely whitish-
pubescent, about 4 mm. long; pappus tawny, 2-3 times as long
as the achene.
Serpentine hills near mouth of Rio Yamaniguey, Oriente, Cuba .
(Shafer 4257; type); between Rio Yamaniguey and Camp Toa,
400 m. alt. (Shafer 4011); vicinity of Camp San Benito, goo m.
alt. (Shafer 4177); Camp La Gloria, south of Sierra Moa (Shafer
8187, 8266); between Camp La Barga and Camp San Benito
(Shafer 4120).
A species which has wide variation in leaf-forms; the involucre
in all specimens seen appears to be the same.
21. Anastraphia (?) lomensis sp. nov.
A shrub about 2.5 m. high, the twigs tomentulose. Leaves
oblong, coriaceous, about 4 cm. long and 1.8 cm. wide, entire-
margined, or rarely with a spinulose tooth near the apex, spinulose-
tipped and acute, or rounded at the apex, narrowed or obtuse at
the base, faintly reticulate-veined, green and glabrous above,
lanate and rather prominently veined beneath, the petioles 2-3.5
BRITTON: STUDIES OF WEsT INDIAN PLANTS 387
mm. long; heads solitary or sometimes 2 together at the ends of
twigs; young involucre I cm. long, its floccose bracts in about 5
series, acute, the outer ovate, the inner lanceolate.
Rocky mountain side, Loma Mensura, about 800 m. alt.,
Oriente, Cuba (Shafer 3811).
The leaves are similar to those of A. Picardae Urban, of Haiti.
38. UNDESCRIBED WEST INDIAN SPECIES
Cyperus calcicola sp. nov.
Perennial by short rootstocks; culms slender, tufted, smooth,
1-2.5 dm. high. Leaves shorter than the culm, 0.5-1 mm. wide;
bracts of the involucre 2 or 3, the longer sometimes 4 cm. long,
similar to the leaves; head globose, solitary, 5-7 mm. in diameter;
spikelets about 3-flowered, 3.5—4.5 mm. long; scales dark brown,
the lowest empty ones persistent on the axis after the fall of the
rest of the spikelet, lanceolate, the others ovate to ovate-lanceolate,
acute, faintly striate-nerved.
Limestone rocks, Cayo Muertos, Porto Rico (Britton, Cowell
& Brown 4976).
Related to C. fuligineus Chapm., in which the scales of the
spikelet are more numerous, strongly striate-nerved and mucro-
nate.
Psilocarya portoricensis sp. nov.
Annual, with fibrous roots; culm rather slender, about 4 dm.
high. Leaves shorter than the culm, 2-3 mm. wide; umbels several,
slender-stalked; spikelets 3-5 in the umbels, ovoid, acutish, 4-6
mm. long, 2 mm. thick; scales ovate, dark brown, sharply acute;
style rather deeply 2-cleft; achene oblong, I mm. long, trans-
versely wrinkled; tubercle triangular, acute, one third to one half
as long as the achene, and distinctly decurrent upon it.
Shore of Laguna Tortuguero, Porto Rico (Britton, Cowell &
Brown 3850).
Nearest to P. nitens (Vahl) A. Wood, of eastern continental
North America, the achene of which is smaller, suborbicular and
the tubercle scarcely decurrent.
Rynchospora borinquensis sp. nov.
Perennial by short rootstocks; culms slender, trigonous, about
6dm. high. Leaves flat, about 2 mm. wide, shorter than the culm;
388 BRITTON: STUDIES OF WEsT INDIAN PLANTS
corymbs 2 or 3, distant, filiform-stalked, compound, narrow, 2-7
cm. long; spikelets clustered, fusiform, 5 mm. long, narrowed at
both ends, bearing only 1 achene; scales lanceolate or oblong-
lanceolate, dark brown, acute; bristles about 6, upwardly barbed,
longer than the achene and tubercle; style-branches about one-
third as long as the undivided part; achene oblong-obovate, faintly
transversely wrinkled, 1.5 mm. long, a little longer than the
smooth, conic, acute tubercle.
Open wet places in the forest, Rio Icaco and adjacent hills,
Sierra de Naguabo, Porto Rico, 465-720 m. alt. (Shafer 3515, type);
Sierra Nipe, Oriente, Cuba (Shafer 3453, 3638).
Apparently nearest R. glauca Vahl, which has shorter, ovoid
spikelets, and a shorter tubercle.
Cassia clarendonensis sp. nov.
A finely pubescent shrub about 1.3 m. high. Leaves linear in
outline, 10-15 cm. long, bearing an obtusely conic gland 1 mm. high
between the lowest pair of leaflets, the slender, pubescent rachis
angular; leaflets 10-15 pairs, thin, pubescent on both sides, with
very short pubescent petiolules, apiculate, inequilateral, the upper
2 or 3 pairs oblong, 1.5-2 cm. long, 6-7 mm. wide, the others
elliptic to obovate-elliptic, shorter and slightly broader; flowers
in several short pubescent panicles in the upper axils; pedicels
2-4 mm. long; sepals obliquely elliptic, obtuse, pubescent, 5-6
mm. long; petals yellow, obovate, veiny, about 9 mm. long, rather
abruptly short-clawed; sepals 7; larger anthers curved, 7-8 mm.
long, the smaller nearly straight, about 5 mm. long; style curved,
about 11 mm. long; pods short-stalked in the calyx, linear, many-
seeded, densely puberulent, 7-10 cm. long, 6 mm. wide, flat,
impressed between the seeds, the margins scarcely thickened;
seeds oblong, transverse, pubescent, about 4 mm. long.
Inverness, Clarendon, Jamaica (Harris 11693).
‘Purdiaea velutina Britton & Wilson, sp. nov.
A shrub 3 m. tall, or a small tree; twigs villous. Leaves obovate,
4-10 cm. long, 2-3.3 cm. broad, rounded and emarginate or
mucronate at the apex, gradually tapering to a rather broad
sessile base, 5-7-ribbed, reticulate-veined, glabrous; bracts oblong-
obovate or obovate, 15-24 mm. long, 6-9 mm. broad, velutinous on
the back, glabrous within above the middle, below clothed with
rather short, appressed silky hairs; peduncles and pedicels villous:
one ciliate, the three outer ones unequal, ovate, 9-12 cm. nae:
BRITTON: STUDIES OF WEsT INDIAN PLANTS 389
4-6 mm. broad, rounded or acutish and apiculate at the apex,
papery, several nerved, the two inner sepals lanceolate, 5-6 mm.
long, acuminate; petals lanceolate to narrowly ovate-lanceolate,
5-5-6 mm. long, 2-2.5 mm. broad, acute, glabrous; filaments
subulate, glabrous; anthers oblong; ovary subglobose, villous.
Type collected along trail, Rio Yamaniguey to Camp Toa,
Oriente, Cuba (Shafer 4474); also collected in the vicinity of Camp
San Benito, Oriente, Cuba (Shafer 4092).
Purdiaea Shaferi Britton & Wilson, sp. nov.
A shrub 2 or 3 m. tall; twigs glabrous or nearly so. Leaves
broadly elliptic-obovate, 4.5-9.5 cm. long, 2.5-4.5 cm. broad,
rounded and emarginate or mucronate at the apex, sessile, 7—9-
ribbed, rather faintly and coarsely reticulate-veined, glabrous;
bracts obovate, 12-14 mm. long, 7-8 mm. broad, papillose;
peduncles and pedicels villous; sepals 5, ciliate, the three outer
sepals very unequal, broadly ovate to ovate-oval, 8-12 mm. long,
4-8 mm. broad, rounded or acute and apiculate at the apex,
long, 2 mm. broad, acuminate; petals elliptic, 5 mm. long, 2.5—
3.2 mm. broad, mucronate, glabrous; filaments subulate, glabrous;
anthers oblong; ovary globose-ovoid, hirsute; style subulate.
Type collected in pinelands, vicinity of Baracoa, Oriente,
Cuba (Shafer 4285).
Purdiaea microphylla Britton & Wilson, sp. nov.
shrub I or 2 m. tall; young twigs and branches of the
inflorescence more or less hirsute. Leaves oblanceolate to obovate,
10-15 mm. long, 5-7 mm. broad, acute at the apex, cuneate at the
base, sessile, rigid, faintly 3-ribbed, glabrous; sepals unequal,
rigid, the three outer ones ovate to oval, 4 mm. long, 3 mm. broad,
acute, the two inner sepals ovate, 3 mm. long, 1.2-1.5 mm. broad,
apiculate, ciliate; fruit 5-lobed, the angles keeled, 5-celled, glab-
rous; style subulate, persistent.
Type collected at Camp La Gloria, south of Sierra Moa,
Oriente, Cuba (Shafer 8265).
Piriqueta cubensis Britton & Wilson, sp. nov.
shrub 1-3 m. tall, the young twigs grooved, velvety-ferrugi-
nous with stellate hairs. Leaves oblanceolate or obovate, 4-8 cm.
390 Britton: STUDIES OF WeEsT INDIAN PLANTS
long, I-2.2 cm. broad, obtuse or acutish at the apex, cuneate at the
base, petioled, above clothed with scattered stellate hairs, beneath
velvety ferruginous with stellate hairs and reticulate-veined, the
midvein and lateral nerves obscure or impressed above, prominent
beneath; calyx-lobes oblong-lanceolate to ovate-lanceolate, 1.2—-1.6
cm. long, 6-8 mm. broad, velvety-ferruginous; petals elliptic to
somewhat oblanceolate, 1.6—-1.8 cm. long, 5-6 mm. broad, glandu-
lar-ciliate; filaments filiform, glabrous; anthers oblong-lanceo-
late; ovary narrowly ovoid, tomentose; styles filiform, 2.5-2.7
cm. long, glabrous; capsule velvety-ferruginous, the valves ovate;
seeds pyriform.
Type collected along trail, Rio Yamaniguey to Camp Toa
(400 m. alt.), Oriente, Cuba (Shafer 4190); also collected on the
Sierra Nipe, Oriente (Shafer 3109).
Rheedia Hessii sp. nov.
Young branches angled, slender, the older ones terete. Leaves
opposite, coriaceous, clustered on short lateral twigs, narrowly
lanceolate, 1.5-2.5 cm. long, 6 mm. wide or less, narrowed at the
base into stout petioles 2-3 mm. long, spinulose-acuminate at the
apex, the midvein prominent beneath, very indistinct above, the
lateral venation wholly obscure, the margins thickened; staminate
flowers solitary in upper axils on pedicels about 4 mm. long;
sepals suborbicular, about 3 mm. broad; petals obovate-elliptic,
rounded at the apex, 5 mm. long; stamens numerous, the stout
filaments 2—2.5 mm. long; anthers less than 0.5 mm. broad.
Indiera Fria, near Maricao, Porto Rico (F. L. Stevens & W.E.
Hess 3333). In habit and in leaf-form this somewhat resembles
R. fruticosa C. Wright, of Cuba.
Mayepea cubensis P. Wilson, sp. nov.
A shrub 6 m. tall, the young twigs glabrous or nearly so. Leaves
oblong-oblanceolate, 6-9 c
inconspicuous; panicles terminal, shorter than the leaves, the
branches glabrous; bracts oblong-lanceolate or narrowly lanceo-
late, hirsutulous; pedicels about 2 mm. long; calyx-lobes tri-
angular, obtuse or acutish, sparingly hirsutulous on the outside
hirsutulous within mostly at or near the apex; petals oblong or
somewhat oblong-lanceolate, 5.5-6.5 mm. long, 1.2-2 mm. broad,
BRITTON: STUDIES OF West INDIAN PLANTS 391
glabrous, obtuse, several-nerved; stamens shorter than the petals;
filaments 1 mm. long; anthers elliptic-ovate to broadly elliptic;
ovary ovoid, glabrous; stigma subsessile, globose-ovoid or sub-
globose, sometimes slightly emarginate.
Type collected at mouth of the Rio Yamaniguey, Oriente, Cuba
(Shafer 4253).
Agalinis albida Britton & Pennell, sp. nov.
Annual; plant green, tending to blacken in drying. Stem
4—6 dm. tall, slender, with virgately ascending branches, obscurely
striate-4-angled, glabrous; leaves opposite, or somewhat sub-
opposite above, ascending or recurved-spreading, linear-subulate
below to linear and longer above, entire, acute, those of the stem
I-2.5 cm. long, 0.5-I mm. wide; bracts gradually much reduced;
leaves thickened, minutely scabrous to scabro-roughened above.
axillary fascicles none; racemes elongated, 2-16-flowered; pedi-
cels ascending, slender, clavate, glabrous, in flower 0.5—-1 mm. long,
in fruit 2-3 mm. long, much shorter than the bracts; calyx-tube
2-2.5 mm. long, depressed-hemispheric, obscurely veined, 1/3-2/5
the length of the capsule, somewhat truncate, its lobes 0.7—1 mm.
long, broadly triangular to ovate-triangular, acute to acuminate;
apex of tube and lobes within sparingly puberulent; corolla 10-
15 mm. long, spreading, 7-10 mm. wide, membranous, its tube
8-12 mm. long, straight to slightly upcurved, its lobes 2-3 mm.
long, rounded to truncate, all somewhat spreading, without
minutely pubescent, within slightly pubescent about the bases of
the filaments, pubescent below sinus and over most of basal
portions of posterior lobes; lobes all ciliate, white, at times shaded
with violet; posterior filaments 2.5-3.5 mm. long, anterior
5-6.5 mm. long, all somewhat lanose; anther-sacs I-1.5 mm. long,
oblong-lanceolate, acute to mucronate at base, lanose-pubescent
with white hairs on the valvular surface, glabrous on the :sides;
style 4-5 mm. long, filiform, glabrous; stigma 2.5 mm. long;
capsule 4 mm. long, depressed-globose, dark-brown; seeds 0.6-
0.8 mm. long, lunate-triangular to narrowly quadrangular, slightly
less than one-half as broad as long, irregular; testa pale, with reticu-
lations delicate, brown, enclosing elongated angular spaces; intra-
reticular lines very fine, forming a network.
Wet, grassy pineland, western Cuba, Isle of Pines, and in
Jamaica.
Type, Colpothrinax savanna, vicinity of Herradura, province
of Pinar del Rio, Cuba, collected in flower and fruit August 26-
30, 1910 (Britton, Earle & Gager 6475).
392 BRITTON: STUDIES OF WeEsT INDIAN PLANTS
Cuspa. Pinar del Rio: Guane (Shafer 10662); Herradura
(Hermann 291, 565); Laguna Jovero (Shafer 10937); Laguna los
Indios (Shafer ro8o0r); Pinar del Rio (Wright 2991 p.p.) ; San Cristo-
bal (Wright 2001 p.p.), Santa Clara: Cieneguita (Combs 443),
Isle of Pines: Managua (Palmer & Riley 1102).
Jamaica. Shooter’s Savanna (Harris 11160) ; Upper Clarendon
(Harris 11100).
Differs from other species of A galinis bearing short-pedicelled
flowers, spreading corolla-lobes and dark brown seeds (A. purpurea
and near allies) by its leaves strongly ascending, but I-2.5 cm.
long, its corolla much paler, nearly or quite white, and relatively
small, and by its seeds which are less than one-half as broad as long.
Jacaranda Cowellii Britton & Wilson, sp. nov.
A shrub 1.5-2 m. tall, the young twigs, pedicels, calyx and
corolla more or less glandular-pubescent; leaves once-pinnate,
2.5-6 cm. long, occasionally longer; petioles and rachis narrowly
grooved; leaflets 8-15 pairs or more, opposite or subopposite,
elliptic to oval or orbicular, 2-5 mm. long, 1.5-3 mm. broad, sessile,
rigid, green and lustrous above, paler beneath, the margin strongly
involute, the midvein impressed above, prominent beneath; bracts
of the inflorescence oblong; calyx-ldbes triangular-ovate or ovate,
acutish, 2 mm. long; corolla-tube 2-2.5 cm. long, the lobes
pia filaments of the fertile stamens subulate, glabrous, the
filament of the sterile stamen flattened, bearded below the middle,
glabrous above, densely bearded at the apex; anthers. oblong-
elliptic; ovary ovoid, glabrous; style subulate; fruit elliptic or
oblong-elliptic, the apex rounded, or abruptly short-acuminate
with the tip acutish or obtuse.
Type collected in palm barrens in the vicinity of the city of
Santa Clara, Cuba (Britton & Cowell 13316); also collected at the
same locality (Britton & Wilson 6071; Britton & Cowell 10174);
dry hill, Holguin, Oriente (Shafer 12434); barren savanna south-—
east of Holguin, Oriente (Shafer 2940). :
*
Mosses from the west coast of South America
R. S. WILLIAMS
(WITH PLATES 21-25)
A list of mosses collected by Dr. and Mrs. J. N. Rose in Peru,
Bolivia and Chile, July to October, 1914, while on a trip under-
taken chiefly for the purpose of obtaining Cactaceae.
This collection of 41 mosses, coming as it does largely from the
dry cactus region of the west coast of South America, contains for
its size an unusually large number of interesting species.
CAMPYLOPUS INCRASSATUS (Kze.) C. M.
Perv: near Mollendo, 18996.
Campylopus peruvianus sp. nov.
Sterile: plants in compact cushions with more or less branching
stems, radiculose below, and 10-15 mm. high; leaves ovate-lanceo-
late, entire except at the apex, closely imbricate, erect-appressed
‘when dry, somewhat spreading when moist, the upper rather
larger, 3-4 mm. long, mostly with a short, rough, hyaline point
rarely more than 0.5 mm. long; costa about one half the width of
the leaf in the lower part, in cross-section showing stereid bands
above and below the guide cells and bearing on the back about 24,
not serrate, lamellae up to 4 cells in height; alar cells enlarged,
mostly pale, extending to the costa; marginal cells of lower leaf
narrow and elongate, forming a narrow, hyaline border sometimes
extending half wavy up the leaf; the median cells mostly
short-rhomboidal, those just above the alar, short rectangular, all
with slightly thickened not pitted walls.
Peru: near Mollendo, August, 78997.
Astomum chilense sp. nov.
oicous, the inconspicuous male flowers containing 5 or 6
antheridia with few paraphyses, enclosed by 2 or 3 very small —
inner perigonial leaves; the outer leaves being scarcely differen-
tiated: plants in rather compact mats with simple stems about I
mm. high, bearing radieles at base; stem-leaves up to 1.5 mm.,
rarely 2 mm. long, ovate-lanceolate, more or less folded and
crispate when dry, rather widely spreading when moist, with flat,
entire margins; costa excurrent into a short apiculus, in cross-
393
394 WrutLiaMs: MossEs FROM WEST COAST OF SOUTH AMERICA
section showing 2 large guide cells, a rather small stereid band
between them and the row of large cells next the upper surface, and
below the guide cells a much larger stereid band with outer cells
somewhat enlarged; cells in lower part of leaf pale, smooth,
mostly short-rectangular, about 8 u wide by 12 » long, in upper
part densely papillose, rather distinct, nearly square, about 6 X
6 uw; perichaetial leaves scarcely differentiated; seta often somewhat
bent and twisted when dry, erect when moist, about 2.5 mm. long;
capsule oblong, slightly wrinkled when dry, about I mm. long,
including the short, oblique beak; spores rough, about 164 in
diameter; calyptra cucullate, smooth, descending about half way
down the capsule.
CHILE: vicinity of La Serena, October, 79290.
GYROWEISIA BOLIVIANA R. S. Williams, Bull. N. Y. Bot. Gard. 3:
104. 1903.
PERU: Cuzco, September, 19047.
LEPTODONTIUM GRACILE C. M.
PERU: vicinity of Lima, July, 18594.
BARBULA FUSCINERVIA Mitt.
Peru: Juliaca, September, rg092.
BARBULA FUSCO-VIRIDIS Broth.
CHILE: Palos Quemnados, October, 19184; La Ligua, October,
19532.
BARBULA REPLICATA Tayl.
Peru: near Oroya, July, 18717; Arequipa, July, 19531;
Posco, July, 18813; Cuzco, August, 19055.
Pterogoneurum Roseae sp. nov.
Autoicous, the male flower terminal, the outer perigonial
leaves not differentiated, the inner with a very small, pale-golden,
roundish, scarcely costate blade bearing a rough, hyaline hair-
point 3 or 4 times as long, enclosing 5 or 6 antheridia about 50 u
long and numerous, somewhat club-shaped paraphyses twice as
long as the antheridia: plants appearing somewhat gregarious, only
2-3 mm. high, with very short stems and crowded, bud-like
branches; stem leaves when dry, erect, appressed-imbricate, when
moist, widely spreading, broadly ovate, serrulate nearly to the
base, the blade about 1 mm. long with a rough, hyaline hair-point
often exceeding the blade in length, the margins of the upper half of
the blade broadly inflexed over the costa; costa expanded in the
upper half, in cross-section showing a broad stereid band below
WILLIAMS: MossEs FROM WEST COAST OF SoUTH AMERICA 399
the single row of large cells bearing about 20 rows of filaments
3-5 cells high with the terminal cell often contracted into a
mamillate apex; basal cells of leaf pale, short-rectangular to
square, up to 20 u wide and about 25 u long, with walls slightly
thickened at the angles, the cells about one half up the leaf becom-
ing more or less transversely elongate, especially toward the
margin, with unequally thickened, not pitted, walls; outer peri-
chaetial leaves not differentiated, the 1 or 2 inner leaves very
small, pale, narrowly lanceolate, nearly ecostate; seta reddish, 8-9
mm. high; capsule ovate-oblong, nearly symmetric and erect,
2 mm. long with the lid; lid conic, about as high as its basal
diameter; peristome papillose, very irregular, from a low basal
membrane, the teeth varying from long-lanceolate to little more
than lobes scarcely projecting above the annulus; annulus per-
sistent, of a single row of large cells; calyptra cucullate, smooth,
extending about half way down the capsule; spores slightly
rough, up to 24 w in diameter.
PERU: vicinity of Lima, July, 78505 (type); Posco, August,
18813a.
Crossidium Rosei sp. nov.
Autoicous, the male flowers at the apex of the branches, the
inner perigonial leaves smaller than the outer, otherwise scarcely
differentiated, the 6 to 8 antheridia about 0.25 mm. long, with rather
numerous, often longer, filiform paraphyses: plants in rather
compact mats, the short stems, with radicles at base, bearing
several short branches crowded together bud-like, and 2-3 mm.
high;leaves when dry appressed-incurved, often somewhat crispate,
when moist, erect-spreading, ovate-oblong to oblong-spatulate,
2-2.5 mm. long, in the upper third finely papillose on both sides,
the margins entire and flat or more or less recurved; costa slightly
excurrent, enlarged in the upper half, in cross-section showing a
large stereid band on the under side and on the upper side some 6-8
large cells, mostly in one row, bearing numerous, short, somewhat
club-shaped filaments 2 or 3 cells high with the terminal cell
papillose; leaf cells distinct throughout, the basal pale green,
nearly square to short-rectangular, up to 20 wu wide by about 35 u
long, the median cells scarcely elongate, about 16 X 16», with
walls scarcely thickened; perichaetial leaves hardly differentiated,
the inner usually smaller; seta rather flexuous, about 8 mm. long;
capsule erect, nearly symmetric, oblong, about 1.5 mm. long with
the lid; lid conic, acute, a little higher than its basal diameter, the
cells, except at the base, elongate in oblique rows; peristome a
papillose basal membrane extending 3 or 4 rows of cells above the
persistent annulus, its margin more or less incised and bearing
396 WILLIAMS: MOSSES FROM WEST COAST OF SOUTH AMERICA
here and there short lobes; calyptra cucullate, smooth, descending
well below the lid; spores rough, up to 25 uw in diameter.
PERU: vicinity of Lima, July, 78774.
PSEUDOCROSSIDIUM gen. nov.
Dioicous. Small plants with simple or slightly branched
stems having a distinct central strand and leaves, when dry, erect-
appressed or sometimes twisted about the stem. Stem-leaves
with margins in the upper part once to twice revolute, the upper
side of the revolute part with cells often much inflated or with
greatly enlarged papillae. Costa broader in the upper part than
below, nearly percurrent or excurrent; in cross-section about two
thirds up showing 5 to 8 guide cells, mostly in one row, beneath
them a large stereid band and on the upper side numerous lax,
thin-walled cells in 1 to 3 layers forming a continuous, densely
papillose surface or sometimes more or less broken up into filaments
2 or 3 cells high with the terminal cells papillose (occasionally, also,
a thin band of stereid cells just above the guide-cells in P. apicu-
latum). Cells of the upper part of leaf not or slightly elongate,
mostly obscure with crowded papillae on one or both sides, in the
lower part, smooth, pale, rectangular or short, more or less trans-
versely elongate. Perichaetial leaves greatly differentiated, the
2 or 3 inner much larger than those of the stem, convolute, pale
throughout, with narrow costa mostly vanishing below the obtuse
or truncate, nearly or quite entire apex. Capsule nearly cylindric
and erect with a slightly oblique, subulate lid. Peristome of 32
slender, papillose teeth twisted to the left, from a low basal mem-
brane. Annulus of 2 or 3 rows of persistent cells. Calyptra
cucullate.
Type species, P. chilense.
Pseudocrossidium chilense sp. nov.
Evidently dioicous, the male flowers not found: plants in
brownish green cushions with somewhat branching stems, 2-4 mm.
high, with a very distinct central strand and walls of outer cells
scarcely thickened; stem-leaves erect-appressed when dry, more
or less broadly ovate, obtusely or acutely pointed, the upper about
1 mm. long, the lower shorter, rather deeply keeled above with
the margins strongly revolute half way down or more; costa
nearly percurrent, enlarged in the upper half, 120-140 w wide, in
cross-section above the middle showing 6 to 9 guide-cells mostly in
one row, below them a large stereid band and on the upper side
2 or 3 layers of lax, thin-walled cells, those at the surface densely
papillose and sometimes more or less separated; cells of upper
WILLIAMs: MOossEs FROM WEST COAST OF SoUTH AMERICA 397
part of leaf scarcely elongate, about 6 » in diameter, more or less
papillose on both sides, in lower part, paler, smooth, mostly nearly
square to transversely elongate, often 12 X 12 yu; perichaetial
leaves projecting well above those of the stem, the 2 or 3 inner
convolute, about 2 mm. long, mostly broadly rounded or truncate
at the entire or nearly entire apex, the leaf-cells mostly slightly
elongate throughout with lax, thin walls, smooth or finely peniliose
on the back in the upper part; seta erect, about 1 cm. lon
capsule cylindric, about 2.25 mm. long without lid, the stoutly
beaked lid nearly erect, I mm. long, with the elongate cells in
oblique rows; peristome of 32 slender papillose teeth, twisted
about one half around to the left, from a rather low basal mem-
brane; annulus of 2 or 3 rows of small cells; calyptra cucullate,
smooth; spores smooth, about 10 uw in diameter.
CHILE: near Valparaiso, September 19132).
Pseudocrossidium apiculatum sp. nov.
Dioicous, the outer perigonal leaves like those of the upper
stem, the inner much shorter, broadly ovate, with narrow costa,
enclosing numerous antheridia, 0.4 mm. long, and abundant,
slightly club-shaped paraphyses: plants in compact, greenish
brown mats with mostly simple stems 1 cm. high, with large,
very distinct central stand and outer walls of 1 or 2 rows of rather
small cells with scarcely thickened cell-walls; stem-leaves ovate-
lanceolate, acute, the upper about 1.5 mm. long, appressed and
mostly twisted about the stem when dry, the margins in the upper
half often twice revolute; costa excurrent, apiculate, in the widest
part 140-160 yw across, in cross-section showing 6 to 8 guide cells,
below them a large stereid band and on the upper side 3 or 4 layers
of lax, thin-walled cells forming a compact densely papillose surface
or broken up into distinct filaments 2 or 3 cells high with the
terminal cell papillose (sometimes also a thin band of stereid cells
occurs just above the guide-cells); cells of upper leaf mostly not
elongate or transversely elongate, often 12 X 12 u, papillose on
both sides, those of the revolute part often much inflated or with
greatly enlarged papillae on the upper face; lower cells of leaf
paler, mostly rectangular, 12-16 uw wide and 20-40 pw long; peri-
chaetial leaves pale, convolute, the inner 2.5 mm. long or more,
projecting well above the stem-leaves, somewhat obovate to
oblong, the apex obtusely rounded and more or less erose, the
costa narrow and vanishing a little below the apex; fruit unknown.
PERuU: above Arequipa, August, 78977.
[Pseudocrossidium excavatum (Mitt.) comb. nov.
To this genus I would also refer Tortula excavata Mitt. Jour.
398 WutLt1amMs: MOossES FROM WEST COAST OF SOUTH AMERICA
Linn. Soc. 12: 154. 1869. It has a cross section of leaf and
perichaetial leaves very similar to the preceding but is a much
smaller species with stem leaves only about 0.5 mm. long and
lacking the sharp apiculate point. From P. chilense it may be at
once distinguished by the basal cells which are rectangular, not
mostly transversely elongate or square. The costa is also nar-
rower, about 60 uw in the widest part. It seems to be known only
from the Andes near Quito.]
Desmatodon subtophaceus (R.S. Williams) comb. nov.
Didymodon subtophaceus R. S. Williams, Bull. N. Y. Bot. Gard. 3:
550; TOs: :
Peru: above Arequipa, August, 19533; Juliaca, September,
19099.
CHILE: Las Vacas, near Chiapa, October, 19230.
The stereid band is nearly or quite lacking in the upper part
of the costa but present in the lower part.
ToORTULA CONFUSA Card.
Peru: Cuzco, September, 19519.
TorTULA KuNZEANA (C. M.) Mitt.
CuILE: La Ligua, 19383; Paloma, October, 19354.
Tortula limensis sp. nov.
Apparently mostly autoicous but occasionally synoicous, the
inner perigonial leaves much smaller and paler than the outer
and enclosing abundant antheridia, 0.3-0.4 mm. long, with numer-
ous slightly longer, club-shaped paraphyses: plants in compact
mats with more or less branching; stems about 1 cm. high having
a very small central strand or none; stem-leaves mostly oblong, the
larger about 3 mm. long by 1.5 mm. wide, appressed-contorted
when dry, somewhat spreading, recurved when moist, with flat
or slightly recurved crenulate and papillose margins; costa terete,
slightly rough on the back, about 40 wide in the lower part,
slightly tapering upward, excurrent into a reddish not quite
smooth hair-point sometimes over one half the length of the
blade, in cross-section showing 2 large guide-cells, below them a
stereid band and on the upper side a few thin-walled cells in two
layers, the upper surface finely papillose; cells of upper leaf not
elongate, densely papillose on both sides, 16-20 u in diameter, those
of basal part pale, smooth, up to 25 p wide and 60-80 uw long; inner
perichaetial leaves very similar to those of the stem but a little
larger; seta about 12 mm. long; capsule oblong, slightly curved
WILLIAMS: MOSSES FROM WEST COAST OF SOUTH AMERICA 399
and nodding, 2.5 mm. long without the lid, the lid conic, in height
often but little exceeding its basal diameter; peristome of 32 pale,
nearly erect, very papillose teeth more or less contracted at the
joints, from a basal membrane about one-tenth the height of the
teeth; annulus of I or 2 rows of small cells adherent to the rim
of the capsule; calyptra cucullate, descending well below the base
of the lid; spores slightly rough, up to 25 » in diameter.
PERU: Lima, July, 18603c¢ (type), 18603a; north of Lima,
July, 19523.
A species much like some forms of T. montana in general
appearance.
Tortula minuscula sp. nov.
Dioicous, the terminal bud-like male flower at length appearing
lateral by the growth of a subapical branch; the antheridia about
0.3 mm. long, with abundant slightly club-shaped paraphyses:
plants in compact, dusky green mats with often branching stems
5-8 mm. high, having a distinct central strand; stem-leaves about
m. long, appressed or somewhat erect- spreading when dry,
rather broadly ovate-lanceolate, entire, the margins somewhat
recurved along the middle and thickened with 1 row of additional
cells in the narrow-grooved point; costa percurrent, often slightly
broadened and flattened near the middle of the leaf with a cross-
section showing 10 to 12 guide cells, a single row of cells nearly as
large on the ventral side and on the dorsal side a row of slightly
smaller cells with a few stereid cells between them and the guide-
cells; the cross-section near apex shows no stereid cells and only 2
guide-cells; cells of upper part of leaf very papillose on both sides,
the median often slightly transversely elongate, about 5 X 8 u,
the basal short-rectangular to nearly square; perichaetial leaves
very similar to those of upper stem but with slightly larger and
paler base; seta 6-7 mm. long; capsule erect, ovate-oblong,
about 1 mm. long without the lid; the lid slightly oblique, conic-
subulate, scarcely 0.5 mm. long; peristome from a papillose basal
membrane projecting well above the annulus, of 32 erect or nearly
erect, very papillose teeth of irregular length; annulus of 2 or 3
rows of persistent cells; calyptra cucullate, smooth, descending
well below the lid; spores smooth, about 10 yw in diameter.
PERU: Cuzco, September, 19520.
TORTULA MURALIs (L.) Hedw.
CHILE: Vifia del Mar, near Valparaiso, September, 79105; Las
Vacas, near Chiapa, 19230a; vicinity of La Serena, 19289; vicinity
of Illapel, 19462, 19462d; La Paloma, October, 19522.
400 WrtL1aMs: MossEs FROM WEST COAST OF SOUTH AMERICA
ENCALYPTA EMERSA C. M.?
Botrvia: Comanche, August, 18884.
This plant is perhaps distinct from E. emersa, of which I have
seen no specimens. The Bolivian plant is autoicous with stems
rather stout, about 2 cm. high; stem-leaves oblong, about 4 mm.
long by 2 mm. broad, the apex rounded, the margins somewhat
crenate and papillose; costa vanishing a little below the apex,
rough on the back with pale spines, simple toward the apex,
branching at the middle and lower part of the costa; capsule
cylindric, nearly smooth, emergent, without peristome; calyptra
entire at base, somewhat scabrous at apex.
ALIGRIMMIA PERUVIANA R. S. Williams, Bull. N. Y. Bot. Gard. 3:
124. 1903.
PERU: vicinity of Arequipa, August, 18823, the locality
from which the type specimens were described.
GRIMMIA MICRO-OVATA C. M.
Bottvia: Comanche, August, 18882a; vicinity of Comanche,
August, 18882; vicinity of La Paz, August, 18870.
GRIMMIA SAXATILIS Mitt.
Peru: Juliaca, September, rgogr.
ZYGODON CIRCINATUS Mitt.
Peru: near Mollendo, August, 19526. Type specimens col-
lected on the island of Chiloe, Chile, and apparently unknown
elsewhere, excepting from this Peruvian collection.
Physcomitrium Roseae sp. noy.
Antheridia not found: plants in thin mats or somewhat gre-
garious, with stems about 2 mm. high, scarcely projecting above
the earth over which they grow with the rosette-like cluster of
leaves resting on the surface; stems with a few radicles at the
base and 5 to 7 leaves clustered at the apex; leaves mostly ovate,
acute, with flat, entire margins, about 2.5 mm. long by 1.5 mm.
wide, with sometimes I or 2 much smaller inner leaves; costa
percurrent, weak, in cross-section showing 5 or 6 cells of somewhat
variable size without thickened walls; cells of upper part of leaf
more or less hexagonal, 20-25 u in diameter, toward the base
square to rectangular, about 25 4 wide and 30-50 x long; seta 2
mm. long; capsule subglobose, about 1.5 mm. high, without
annulus and peristome, the cells about the rim elongate in 5 or 6
WILLIAMS: MOSSES FROM WEST COAST OF SoUTH AMERICA 401
rows, forming a rather ill-defined border, those below to near the
base very irregular, becoming at the base smaller, quite regular
and with small stomata in several rows; lid nearly flat, with an
ill-defined border of transversely elongate cells that become toward
the center of lid much broader and larger; spores mostly roundish,
smooth, 28-30 in diameter; calyptra (perhaps not normal
remaining attached to the seta half way down the capsule, terete-
cylindric, smooth.
CHILE: near Valparaiso, September, 19176.
FUNARIA CALVESCENS Schwaegr.
BoLivia: vicinity of La Paz, August, 18858.
FUNARIA HYGROMETRICA (L.) Sibth.
PERU: vicinity of Chosica, June, 18549; Arequipa and vicinity,
August, 18822, rg9001; Juliaca, September, 19098.
CHILE: near Valapraiso, September, 19106, 19107, 10132a;
Santiago, September, 19781.
FUNARIA MACROSPORA R. S. Williams, Bull. N. Y. Bot. Gard. 3:
133- 1903.
Bouivia: La Paz, August, 19463; Araranca, August, 10527.
PERU: Arequipa, August, 19528. The spores in all these speci-
mens are rough and the larger measure 25-30 u. Possibly the
species is not distinct from F. hygrometrica. The plants are all
much larger than any specimens of F. hygrometrica arctica |
have seen.
FUNARIA SUBERECTA Mitt.
CHILE: vicinity of Illapel, October, 19462a.
MIELICHHOFERIA CAMPYLOTHECA C, M.
Botiv1a: Comanche, August, 18884a.
HAPLODONTIUM JAMESONI (Tayl.) Hpe.
PERU: Juliaca, September, 19097.
HAPLODONTIUM SERIOLUM C, M.
PERU: Juliaca, September, 19518.
LEPTOBRYUM PYRIFORME (L.) Wils. _
Bo.iviA: Araranca, August, 19464.
LEPTOBRYUM WILSONI (Mitt.) Broth.
Peru: Arequipa and vicinity, August, 19514, 19517.
402 WuLLtAMs: MossEs FROM WEST COAST OF SOUTH AMERICA
ANOMOBRYUM FILIFORME (Dicks.) Husn.
PERU: above Toyatoya, August 18941; Juliaca, September,
TQIO!.
BRYUM CONCAVUM Mitt.
Boxivia: La Paz, August 18895, 18806, 19463a.
Peru: Arequipa, August, 79529; Cuzco, September, 19056,
19058.
BARTRAMIA FRAGILIFOLIA C. M.
BOLIviA: vicinity of Comanche, August, 19524.
Philonotis fragilicaulis sp. nov.
Flowers and fruit not known; plants in compact cushions with
slender, branching stems, mostly denuded of leaves in the lower
part and without radicles, 3-4 dm. long and 0.25 mm. in diameter,
having a large well-defined central strand and often rough surface
in the older parts from the large, mamillose-inflated outer cells;
leaves nearly erect and loosely imbricate-incurved when dry,
scarcely spreading when moist, ovate-pointed, about 1.5 mm.
long and 0.75 mm. wide, the margins flat, serrulate in the upper
part, sometimes crenulate nearly to the base; costa stout, rough
on the back in the upper half, excurrent into a serrulate, short or
somewhat elongate and subulate point; cells of leaf on the under
surface, more especially the median cells, mamillose-inflated at
the base, those of the upper leaf mostly somewhat elongate, more
or less quadratic, the median 8-12 w wide and 20-30 p long, the
basal rather larger and more rectangular.
Perv: Araranca, 4260 m. alt., August, 19513.
POGONATUM POLYCARPUM (Schimp.) Broth.
Peru: Juliaca, September, rozoo.
HEDWIGIDIUM IMBERBE (Sw.) Bry. Eur.
Bottvia: vicinity of Comanche, 18883.
FABRONIA ANDINA Mitt. ;
BOLiviA: vicinity of La Paz, August, 18857, 18871; Comanche,
August, 19515.
LESKEA GRACILLIMA Tayl.
PERU: Cuzco, September, 19059, 19530.
One specimen (No, 18603b) is not determined. It is possibly a
small Pohlia, but lacks fruit and may belong elsewhere.
WiLuiaMs: MOSSES FROM WEST COAST OF SOUTH AMERICA 403
Explanation of plates 21-25
PLATE 21
Campylopus peruvianus. 1. Plant about natural size. 2. Stem-leaf, X 18. 3.
Hyaline point of leaf, X 103. 4. Basal cells on one side of costa, X 124. 5. Me-
dian cells of leaf, X 124. 6. Half a cross-section near the middle of leaf, x 124.
Astomum chilense. 7. Plant about natural size. 8. Plant, X 10. 9. Apex of
leaf, X 183. 10 and 11. Upper stem leaves, X 20. . Lower stem leaf, X 20.
13. Median cells of leaf, X 183. . Cross-section ai bak about half way down,
124. 15. Basal cells on one sinc a costa, X 124.
PLATE 22
Pterogoneurum Roseae. 1. Plant about natural size. 2. Upper leaf, X 10. 3.
Lower leaf, X 10. 4. Inner perichaetial leaf, X 10. 5. Calyptra, X 10. 6. Cells
at the shoulder of leaf about half way up blade, X 124. 7. Apex of epee x 46.
8. Capsule, X 10. 9. Half a cross-section near the middle of leaf, X 1 10. Part
of peristome, annulus and rim of capsule, XK 124. 11. Basal cells ek margin
extending only part of the way to eosta, X 124
Crossidium Rosei. 12. Plant about natural size. 13. Calyptra, X II. 14 and
15. Upper and middle stem aaa xX 18. 16. Basal cells on one side of costa,
124. 17. Capsule, X II. . Median leaf cells, X 12 19. Cross-section about
one third down leaf, X-103. 20. Part of peristome, meeutca and rim of capsule,
wad. :
&
PLATE 23
Pseudocrossidium chilense. 1. Plant about natural size. 2. Plant, X 9. 3 and
4. Perichaetial leaves, X 20. 5. Basal cells on one side of costa, X 124. 6 and 7.
Upper and middle stem leaves, X 20. 8. Median cells of leaf, X 183. 9. C
section of leaf below the middle, X 124. 10. Cross-section above the middle, X 183.
Pseudocrossidium gory a 11. Plant about natural size. 12. Plant, X 6.
13. Stem-leaves, X 13. . Perichaetial leaf, X 1 15. Cells in upper leaf about
midway between margin on costa, X' 183. © 16. ik just above the rectangular
basal cells, X 183. 17. Cross-section of leaf about one third down, X 124. 18.
Basal cells of leaf from margin to costa, X 124. 19. Cross-section of a second leaf,
X 183
PLATE 24
Tortula minuscula. 1. Plant about natural size. 2. Calyptra, X 14. 3. Cap-
sule, X 14. 4. Median exothecal cells, X 124. 5. Median cells of leaf, X 183. 6.
art of peristome, annulus and rim of capsule, X 183. 7- Perichaetial leaf, X 20.
8 and 9. Stem leaves, XK 20. 10. Basal cells of leaf from margin to costa, X 183.
11. Cross-sections of costa in upper part and near the middle, X 183.
Tortula limensis. 12. Plant about natural size. 13 and 14. Stem-leaves, X 6.
15. Capsule, X 6. 16. Calyptra, X 6. 17. Cross-section of leaf, X 124. 18. Apex
of leaf-blade and costa on under side, X 124. 19. Median cells of leaf, X 124. 20.
Basal cells of leaf from margin to costa, X 124. 21. Part of peristome, annulus an
rim of capsule, X 124.
404 Wuiiiams: MossEs FROM WEST COAST OF SOUTH AMERICA
PLATB 25
Physcomitrium Roseae. 1. Plant, X 5.
2. Leaf, X 14. 3. Apex of leaf, X 100.
4. Cells po rim of cap
sule, Ioo. 5. Cells at base of capsule surrounding a
stoma, X 10 6. icine cells of leaf, X 100. 7. Cells of lid from margin to center,
a 8. tai cells of leaf from margin to near the costa, X 100. 9. Part of
cross-section of leaf near the middle, K 100
Philonotis fragilicaulis. 10. Plant, about natural size.
Ir and 12. Stem leaves,
X 23. 13. Median cells of leaf peat mamilla:
I
at the lower and under surface,
asal cells of leaf from margin to costa,
cross-section a leaf near 2H middle, X 183. 17. Part of
cross-section of leaf a little below the middle,
X 183. 14. Leaf at apex, X 18
16. Part o
x
H
w
Notes on Carex—VIll
KENNETH KENT MACKENZIE
CAREX STRICTA LAM. AND ITS ALLIES
Probably the only species of Carex growing in the vicinity of
New York, which is abundant enough to make it the distinctive
feature over large areas, is the tussock sedge, Carex stricta Lam.
Every swamp is full of the great clumps of this plant, usually
very largely to the exclusion of everything else. It is with diffi-
culty that proper herbarium specimens can be made, as the root-
stocks are very heavy and stout. In fact an axe or heavy spade
is needed to dig out characteristic specimens. The result is that
herbarium specimens are mostly very poor and usually consist
of the culms broken off as close to the ground as possible.
The distinguishing features of this common plant are its
habit of forming dense tussocks (or bogs in the vernacular sense),
its not being loosely and strongly stoloniferous, its intensely deep
green leaves, triangular or channelled at the base, usually light
brown culm bases, and but slightly hispidulous sheaths. It is a
very rough plant, the perigynia are strongly granular roughened,
and the scales are usually light reddish brown tinged. _
Another species which is widely distributed throughout the
northeastern United States can be readily told in the field, but
much less easily from the usual run of herbarium specimens.
This plant grows in great beds and does not form tussocks. It
produces many long horizontal stolons; the leaves, at first glau-
cous green, in age become light green or blue green, and contrast
strongly with the leaves of Carex stricta, when growing near it;
the culms at base are brown-purplish tinged, the sheaths are
strongly hispidulous, and the scales vary from blackish to dark
reddish brown in age, while the leaves are flat or nearly so to the
base. The perigynia are slightly more beaked than in Carex
stricta, and the lowest bract is noticeably larger. This species is
Carex strictior Dewey. It has often been mistaken for Carex
406 MACKENZIE: NOTES ON CAREX
aquatilis Waht: by discriminating collectors; who have noticed
that it was distinct from Carex stricta, and who could find no other
name apparently available for it.
By far the most widely distributed species of this group in the
eastern part of the United States, is a plant to which I am applying
the name Carex Emoryi Dewey. This isa freely stoloniferous plant
growing in beds like Carex strictior. It, however, is a much stouter
plant, and the basal sheaths are not filamentose or hispidulous.
The perigynia, which are abruptly and prominently short-beaked,
are smooth under an ordinary lens except towards the apex
and much resemble the perigynia of the C. aquatilis group.
This species ranges from New York, western New Jersey and
Maryland to Manitoba, Colorado and New Mexico, and has at
various times been referred to Carex stricta or Carex aquatilis.
A slender species of this group with a northerly range is Carex
Haydeni Dewey. This plant has broadly oval or suborbicular
perigynia which are noticeably inflated and light brownish at
maturity, in these characters being readily separable from other
members of the group here discussed. The scales always exceed
the perigynia and the spikes are small in size. While the culms
are cespitose they are much less densely so than in Carex stricta.
As far as I can make out the habit of growth is much like that of
Carex torta, i. e., the plant has very short ascending stolons. The
basal sheaths are often, but not always, entirely free from the
filamentose fibers so evident in Carex stricta, and the culms are
usually much darkened at the base.
The first three species referred to above all vary to a very
considerable extent in the size of the pistillate spikes. Specimens
with very short spikes, referable to Carex strictior, have been
described as Carex stricta var. curtissima Peck. Such specimens
have the perigynia very crowded, and on this account the peri-
gynia not having a chance to properly develop often present a
different appearance from specimens where they are less crowded.
I regard these plants only as exhibiting individual peculiarities,
and do not believe they should be given varietal recognition.
These same three species also often produce slender elongated
pistillate spikes. Such specimens have of late years been going
under the name Carex stricta var. angustata (Boott) Bailey, and
MACKENZIE: NOTES ON CAREX 407
very narrow spiked forms have been called Carex stricta var.
xerocarpa (S.H. Wright) Britton. However, the original Carex
angustata Boott, on which the variety angustata is founded, is a _
plant from the Oregon-Washington region, and is entirely distinct
from the eastern species. The narrow spiked specimens of the
eastern plants do not seem worthy of recognition.
While it is not my intention in this paper to consider all the
Pacific Coast species which are closely related to Carex stricta, I
have included Carex angustata and certain plants confused with it.
Carex angustata itself is distinguished from its eastern relatives by
the finely many-nerved perigynia. It seems to be exactly the same
plant afterwards described by Professor Bailey as Carex acutina
and is of rare occurrence in collections. It is a slender plant
with rather loosely cespitose culms and short ascending stolons.
The lowest bract does not exceed the inflorescence, the lower
sheaths are nearly if not entirely smooth, and the foliage is much
less rough than in its eastern relatives.
More common in the same region is a much larger plant which
has often been distributed as Carex acutina: Bailey. In it the
lowest bract much exceeds the inflorescence and the lower sheaths
are strongly hispidulous. This species seems not to have been
described and accordingly I am now proposing it as Carex egregia.
The plants from the eastern parts of the Rocky Mountains
called Carex acutina seem to be either forms of Carex aquatilis
Wahl. (C. variabilis Bailey), having sharp pointed scales exceeding
the perigynia, or species closely related to it. They are but dis-
tantly related to the genuine Carex acutina.
The species here treated are separable as follows:
Culms very densely cespitose, forming bogs; short ascending stolons
or long horizontal stolons not conspicuously developed. C. stricta.
Culms loosely cespitose, in clumps or ee short ascending stolons or
long horizontal stolons freely dev
Loosely cespitose; with short Lissisting sto ae
Perigynia broadly oval or suborbicular, Sinise: brownish at
maturity (eastern United States).
Perigynia narrower, aa Sigs not inflated (Oregon and
C. Haydeni.
Washington
Lowest bract eabaeked by inflorescence; lower sheaths
smooth. C. angustata,
Lowest bract exceeding inflorescence; lower sheaths
o
hispidulous. C. egregia.
AO8 MACKENZIE: NOTES ON CAREX
Forming beds; long horizontal stolons numerous.
Lower sheaths filamentose; perigynia granular-roughened,
green; culms slender to base; lower sheaths little septate-
nodulose. C. strictior.
Lower sheaths not filamentose; perigynia granular-roughened
towards apex only, straw-colored; culms stout at base;
lower sheaths strongly septate-nodulose. C. Emoryt.
CAREX STRICTA Lam. Encyel. 3: 387. 1789
“Carex acuta L.’’ Muhl. Descr. Gram. 263, 1817.
Carex virginiana Smith, Rees’ Cycl. 7 sp. 100. 1819.
Carex commutata J. Gay, Ann. Sci. Nat. II. 11: 198. 1839.
Carex Kelvingtoniana Steud. Syn. Pl. Cyp. 215. 1855.
Carex virginica Steud. Syn. Pl. Cyp, 217. 1855.
Carex tenuispica Boeck]. Flora 39: 225. 1856.
Carex tristicha Boeckl. Flora 41: 651. 1858.
Carex xerocarpa S. H. Wright, Am. Journ. Sci. II. 13: 334. 1866.
Carex angustata var. a typica and var. B (in great part) Boott,
Ill. Car. 4: 173. pl. 586, f. 1, 587. 1867. Not C. angustata
Boott; Hooker Fl. Bor. Am. 2: 218. 1867.
Carex angustata vat. xerocarpa Bailey, Carex Cat. 1884.
Carex stricta var. angustata Bailey; A. Gray, Man. Ed. 6, 600.
1890.
Carex stricta xerocarpa Britton, Bull. Torrey Club 22: 222. 1895.
Growing in very dense tussocks, the culms aphyllopodic, 3-10
dm, high, slender, strict, exceeding leaves, very rough on the
sharp angles, the faces concave, strongly filamentose and brownish
or light purplish brown at base, the basal sheaths subcarinate.
Leaves with well-developed blades three to five to a fertile culm, on
lower fourth, often somewhat clustered, the sheaths slightly
hispidulous, tight, hyaline and often brownish tinged opposite
the blades, the blades 1.5-3 mm. wide, usually 1-3 dm. long,
stiffish, deep green, the margins revolute, but blade itself channeled
and keeled towards base, strongly roughened. Principal staminate
spike usually one (with one or two smaller sessile ones near its
base), erect, peduncled, 2-4 cm. long, 2.5 mm. wide, the scales
oblong-obovate, obtuse, light reddish brown with lighter center
and hyaline margins. Pistillate spikes usually two or three,
erect, sessile, or lower slightly peduncled, more or less strongly
separate, the upper often staminate above, the better developed
2-6 cm. long, 3-5 mm. wide, the perigynia closely arranged in
several ranks (or spikes at times somewhat attenuate below);
MACKENZIE: NOTES ON CAREX 409
bracts sheathless, their auricles inconspicuous, slightly dark-
colored, the lower bract 1.5—3 mm. wide, shorter than inflorescence;
the upper reduced. Scales variable, oblong-obovate to lanceolate,
obtuse to acuminate, reddish, with lighter center and narrow
hyaline margins, narrower and usually rather shorter than peri-
gynia, appressed. Perigynia broadly to narrowly ovate, closely
enveloping achene, plano-convex, not inflated, two-edged, granular-
roughened, puncticulate, dark green, obscurely few-nerved
dorsally, nerveless ventrally, 2.5 mm. long, 1.5 mm. wide, rounded
and substipitate at base, short-tapering to the beakless or nearly so
subentire whitish-tipped apex, the style short-exserted; achenes
lenticular, broadly ovate, substipitate, 1.75 mm. long, 1.25 mm.
wide; stigmas two.
SPECIMENS EXAMINED
MAssACHUSETTS: Cambridge, Fernald, June 10, 1891 (C.);
Cambridge, Underwood 2644, May 9, 1891 (C.); Boston, B. D.
Greene (C.); West Roxbury, Forbes, June 8, 1911 (K.M.).
RuHopE IsLAND: Providence, Olney, June 8, 1871 (N. Y., C.);
Providence, Olney, labeled C. strictior Dewey (C.).
New HAMPSHIRE: base of White Mountains, Tuckerman (C.).
New York: Bull’s Head, Staten Island, Britton, June 2, 1889
(C.); Grant City, Staten Island, Britton, May 26, 1889 (C.);
Suffern, Schrank (C.); West Hampton, Delafield, June 13, 1888
(C.); Junius, Sartwell 54 (N. Y.,C.); Prattsburgh, Steuben County,
Wright (N. Y., C.), type of C. xerocarpa S. H. Wright; Philips-
town, Barratt (C.); New York, Torrey (C.).
NEW JERSEY: Plainfield, Tweedy, June, 1884 (C.); Woodbridge,
Lighthipe, May 3, 1890 (C.); Manchester, Torrey (C.); Pitman,
Gloucester County, Long 3425, May 16, 1910 (P.); Farmingdale,
Long & Brown 3644, May 28, 1910 (P.); Glassboro, Gloucester
County, Long 3360, May 14, 1910 (P.); Delair, Camden County,
Long 3353, May 10, 1910 (P.); Cape May Court House, Long 72209,
June 1, 1912 (P.); Delanco, Burlington County, Long 3466, 3467,
May 19, 1910 (P.); Cold Spring, Cape May County, O. H. Brown,
June 26, 1911 (P.); Mickleton, Crawford, May 10, 1892 USE
Kaighn’s Point, Camden County, MacElwee, May 7, 1892 (P.);
Batsto River, Burlington County, Bassett 45, May 25, 1912 (Py;
Quinton, Salem County, Long 3029, April 22, 1910 (P.); Cranberry
Lake, Sussex County, Mackenzie 1294, May 30, 1905 (K. M.);
410 MACKENZIE: NoTES ON CAREX
Netcong, Morris County, Mackenzie 3167, June 21, 1908 (K. M.);
Lakehurst, Ocean County, Mackenzie 4546, May 15, 1910 (K. M.);
South Amboy, Mackenzie 1223, May 21, 1905 (K. M.); Charlotte-
burg, Mackenzie 3083, May 24, 1908 (K. M.); Chatham, Mackenzie
174, May 30, 1903 (K. M.); Greenwood Lake, Mackenzie 2567,
May 19, 1907 (K. M.); Delaware, Warren County, Mackenzte 5354,
May 10, 1913 (K. M.); Landisville, Gross, 1872 (P.); Camden,
Boice, May 16, 1878 (P.).
PENNSYLVANIA: West Chester, Pennell 1333, May 29, 1909
(P.); Frazier’s bog, Montgomery County, Long 3354, 3350, May
12, 1910, and 3773, June 1, 1910 (P.); Tullytown, Crawford (P.);
Corning, Lehigh County, Pretz 5422, May 30, 1913 (P.); Willistown,
Chester County, Stone (P.); East Nantmeal Township, Chester
County, Pennell 2817, June 14, 1911 (P.); Oxford, Crawford, June -
8, 1892 (P.); Williamson, Delaware County, Pennell 3770, June
23, 1912 (P).; Allentown, Pretz 4703, june 28, 1912 (P.); Tully-
town, Keller, June 12, 1897 (P.); Nottingham, Chester County,
Pennell & Long 3700, June 22, 1912 (P.);Chester County, Pennell
3910, June 27, 1912 (P.); Penn Valley, Crawford, May 7, 1899
(P.); Nockamixon, Crawford, June 30, 1893 (P.); Penn Valley,
Crawford, June 1, 1899 (P.); York County, Glatfelter, May 30, 1895
(K. M.); Penn Valley, Pretz, June 30, 1899 (P.); Centre Square,
Brinton, May 20, 1892 (P.); Nockamixon, Crawford, June 30, 1893
(P.); Chester County, Pennell 1398, May 30, 1909 (P.); Lancaster,
Carter, May 24, 1904 (P.); Stacy’s Hollow, Wister 46, June 3,
1862 (P).
DELAWARE: Ashland, Commons, June, 1865 (P.); Greenbank,
Commons, May 26, 1879 (P.).
ILLINoIs: Canton (C. xerocarpa S. H. Wright).
TENNESSEE: Jackson, Bain 489, May 15, 1893 (C.).
Carex Haypent Dewey, Am. Journ. Sci. II. 18: 103. 1845
Carex B erecta Dewey, Am. Journ. Sci. 10: 265. 1826.
“ Carex aperta Boott” Carey; A. Gray, Man. 547. 1848.
Carex aperta var. 6 Boott, Ill. Car. 4: 132. pl. 426. 1867.
Carex aperta var. minor Olney, Exsicc. V. 15. 1871.
Carex stricta var. decora Bailey, Bot. Gaz. 13: 85. 1888.
Carex stricta var. 8 Haydenii Kiikenth.; Engler, Pflanzenreich
a7 A306. 1006. :
MACKENZIE: NOTES ON CAREX 411
Cespitose, the culms slender, 3 mm. thick near base, 5-10 dm.
high, aphyllopodic, sharply triangular and strongly roughened on
the angles above, usually much exceeding the leaves, the fertile
slightly or not at all filamentose at base. Leaves with well-
developed blades two to four to a fertile culm, on the lower fourth
but not bunched, the sheaths papery and yellowish- or whitish-
hyaline ventrally, smooth or nearly so, the blades flat with slightly
revolute margins, about 2.5 mm. wide, usually 1-2 dm. long, very
rough towards the apex; sterile culm leaves longer. Terminal spike
stiminate (usually with an additional sessile smaller one at base),
linear, 2-4 cm. long, 3 mm. wide, the scales oblong spatulate,
obtuse to acutish, reddish brown with lighter center and hyaline
margins. Pistillate spikes two or three, erect, sessile or nearly so,
approximate or a little separate, linear, 1.5-3 cm. long, 4-5 mm.
wide, occasionally staminate at apex, scarcely attenuate at base,
the perigynia very numerous, in few rows, spreading-ascending;
bracts sheathless without dark colored auricles, the lowest 2 mm
wide, normally exceeded by inflorescence; the upper much reduced:
Scales ovate or lanceolate, long acuminate to acute, narrower than
but strongly exceeding perigynia, from dark brown to straw color,
with lighter center and hyaline margins. Perigynia light brownish
at maturity, flattened biconvex, broadly oval to suborbicular,
two-edged nerveless or obscurely few-nerved, 2—2.5 mm. long,
25-1.75 mm. wide, membranaceous and slightly inflated,
puncticulate and resinous dotted, minutely granular towards apex,
round-tapering at base and apex and abruptly very minutely
(0.2 mm. long) beaked, the orifice hyaline, entire or emarginate.
- Achenes lenticular, substipitate, apiculate, yellowish, with orbicu-
lar faces, I mm. long; style exserted; stigmas two.
This species has in times past been confused with Carex
aperta Boott, a phyllopodic species from the Columbia River
region. Although frequently treated as a variety of Carex stricta
Lam., one cannot help feeling that this is due to lack of knowledge
of the two species. They are really thoroughly distinct species,
and I have seen no evidence of any tendency to intergrade.
SPECIMENS EXAMINED
New Brunswick: Bathurst, Fowler, August 4, 1874 (N. Y.).
Maine: Bangor, Knight, June 27, 1905 (K. M.); Bangor,
Mackenzie 3188, July 3, 1908 (K. M.); Orono, Harvey, August,
1891 (N. Y.); Orono, Fernald, June 28, 1890 (C.); St. Francis,
Fernald 135, August 5, 1893 (C.).
New Hampsuire: Keene, Gilbert, June 11, 1877 (N. Y.):
412 MACKENZIE: NOTES ON CAREX
MAssacuuseEtts: Dedham, Forbes, June 15, 1904, and June 9,
1914 (K. M.); Waverley, W. Boott, June 30, 4663 -°(N: X)%
“‘Mass.’”’, Davis (C.).
RHODE IsLAND: Olney (C.).
New York: North Elba, Peck (N. Y.); Amherst, Sartwell
(Ne Y¥).
New Jersey: Lawrence, Mercer County, S. Brown, May 28,
29, 1904 (P.).
PENNSYLVANIA: Sellersville, Bucks County, Pollard, June 2,
1899 (K. M.); Sellersville, Pretz, June, 1883, May 16, 1896, and
June, 1899 (P.); Plumsteadville, ‘‘J.C.M.”, June, 1866 (P.).
Itttnots: Fountaindale, Bebb, 1870 (N. Y.); “Ill.” Vasey
ty Hie Ore, B0) F
Minnesota: Milaca, Sheldon, June, 1892 (C.); Nichols,
Sheldon, June, 1892 (C.).
Iowa: Fayette, Fink, June, 1895 (N. Y.); St. Petersburg,
Parry (C.).
Missourt: Lake City, Jackson County, Mackenzie, May 23, 1897
(K. M.).
NEBRASKA: “Nebraska Ter.”, apparently duplicate of type
(Cs
CAREX ANGUSTATA Boott; Hooker, Pl. Bor. Am. 2: 218. 1840
Carex angustata Boott (in part), Ill. Car. 4: 173. pl. 586, f.2. 1867.
Carex acuta Bailey, Proc. Am. Acad. 22: 86. 1886
Carex acutina Bailey, Mem. Torrey Club 1: aa 1889. Not
“Carex acutina Bailey’’, Holm. Am. Journ. Sci. 16: 34; or
Kiikenthal; Engler, Pflanzenreish 4”: 319.
Loosely cespitose, with short ascending stolons covered with
brown scales, the culms 3-6 dm. high, strongly aphyllopodic,
slender, 3-4 mm. wide at base, 1.5 mm. above, exceeding the
leaves, roughened above on the angles, brownish reddened and
more or less filamentose at base. Leaves with well-developed
blades usually three to five to a fertile culm, on the lower third,
the blades light green ascending, flat to base with slightly revolute
margins, smoothish beneath, 2.5—3 mm. wide, 1-4 dm. long, much
roughened towards the attenuate apex, the sheaths smooth,
membranous, easily breaking and slightly yellowish tinged
ventrally. Staminate spike usually one, from little to strongly
____ rough-peduncled, 2.5—5 cm. long, 3-5 mm. wide, the scales oblong-
MACKENZIE: Notes ON CAREX 413
obovate, obtuse, brownish-black with lighter center and slightly
hyaline margins. Pistillate spikes three to five, erect, more or
less strongly separate, the upper sessile, the lower short-peduncled,
linear-cylindric, 2-4 cm. long, 3.5—-5 mm. wide, the upper especially
often staminate at apex, the lower slightly attenuate at base, the
numerous perigynia ascending, closely packed in several ranks;
lower bract leaflet-like, shorter than or sometimes nearly equalling
inflorescence, sheathless; upper bracts shorter; auricles often
darkened, the upper at least dilated. Scales oblong-ovate to
lanceolate, obtusish to acuminate, narrower and from somewhat
shorter to somewhat longer than perigynia, blackish with lighter
center and slightly hyaline apex. Perigynia straw-colored, obo-
void or oval, sessile or nearly so, two-edged plano-convex, finely
many-nerved or nerveless when young, 3 mm. long, 1.5 mm. wide,
glandular-puncticulate, slightly granular towards apex, not serru-
late, round-tapering at base, abruptly apiculate, beaked, the beak
0.25 mm. long, with entire black-tipped orifice. Achenes lenticu-
lar, sessile, with obovate faces, 1.5 mm. long; stigmas two.
SPECIMENS EXAMINED
OrEGON: Deschutes River, Howell 935, May 9, 1885 (N. Y.,
H.); Sauvie’s Island, Howell, May 15, 1886 (N. Y.); “Oregon”
1883 (H.).
The original specimens of Carex angustata Boott were collected
along the Columbia River in Oregon or Washington by Scouler
and Tolmie. Later Dr. Boott, failing to notice that these speci-
mens represented a different species than the eastern Carex stricta
Lam., applied the name Carex angustata to the eastern plant. His
reason for not taking up the name Carex stricta Lam. was that
there was a common European species known as Carex stricta
Good., and he believed that this name having come into common
use should not be superseded by Carex stricta Lam., even though the
latter was the earlier published name. His views on this point have,
however, not prevailed, but later botanists in taking up the name
Carex stricta Lam. have continued the confusion by tacking on
the name Carex angustata to it as a slender-spiked variety, and
have disregarded the real identity of this species.
Carex acutina Bailey has also been the source of much un-
necessary confusion. The original comparison was with the
European Carex acuta, an aphyllopodic species, and the original
414 MACKENZIE: NOTES ON CAREX
specimens collected by Howell quite justify such a comparison.
Later botanists have created confusion by applying the name to
certain phyllopodic species related to Carex aquatilis Wahl. More
justifiable was the confusion with the species immediately hereafter
described.
v Carex egregia sp. nov.
Loosely cespitose and short stoloniferous from slender creeping
rootstocks, the culms 4.5-9 dm. high, 4.5 mm. wide at base, 1.5
m. wide below spikes, slender but erect, sharply triangular with
concave sides, slightly roughened above, exceeding leaves, strongly
purplish tinged at base, aphyllopodic. Leaves with well-developed
blades three or four to a fertile culm, on lower fourth, somewhat
bunched, the blades flat to base with revolute margins, roughened
towards apex, 2-3 mm. wide, usually 1-3 dm. long, the sheaths his-
pidulous, little carinate and sparingly septate-nodulose dorsally,
and brownish tinged ventrally. Terminal spike staminate, stalked,
4 cm. long, 5 mm. wide, the scales obovate, obtuse, purplish black
with white midvein not extending to apex. Lateral spikes four
or five, the upper one or two small, staminate, the lower two to
four pistillate (often staminate at apex), more or less strongly
separate, sessile or short-peduncled, 2.5-4.5 cm. ong, 5 mm.
ee with very many appressed-ascending perigynia in several
m ows. Lowest bract leaf-like, exceeding inflorescence,
suai dark tinged at base, the upper bracts reduced, not con-
picuously dark-auricled. Scales lanceolate, acute, purplish black
with white midvein, narrower and longer or shorter than perigynia.
Perigynia plano-convex, obovate, 3 mm. long, 1.5 mm. wide, finely
about three- to five-striate on both faces, brownish, puncticulate and
granular-roughened, the margins sharp-nerved, not round, rounded
and substipitate at base, rounded and abruptly minutely apicu-
late at apex, the orifice entire. Achenes lenticular, closely filling
lower part of perigynia, brownish, 1.5 mm. long, substipitate, obo-
vate; stigmas two.
Closely related to Carex eurycarpa Holm, a species with
“‘rounded”’ perigynia, finely many-nerved on both faces ana with
more prominent emarginate, beak. The type specimen was
distributed as that species.
SPECIMENS EXAMINED ©
OREGON: Cascade Mountains, Cusick 29809, August 27, 1902
(N. Y., H.); Cycan Mountains, Cusick 2757, August 14, 1901
MACKENZIE: NOTES ON CAREX 415
(H.); Silver Creek, Lake County, Cusick 2729, August 7, 1901 (N.
Y., H.); Summit of the Cascades, Cusick 2991, August 14, 1902
CN VG, FE):
WASHINGTON: Falcon Valley. Suksdorf 5181, July 15, 1905
(K. M., type).
Ipano: Forks of St. Mary’s River, Leiberg 1174, July 6, 1895
tN. Y.}.
CAREX STRICTIOR Dewey; A. Wood, Class Book 582. 1845
“Carex stricta Good.’ Dewey, Am. Journ. Sci. 10: 269. 1826.
Carex Watsoniana Steud. Syn. Pl. Cyp. 215. 1855.
Carex angustata var. y and var. 6 (in small part) Boott, Ill. Car.
4: 173. pl. 588. 1867.
Carex stricta var. curtissima Peck; Howe, Rep. N. Y. State
Museum 15. 1895.
Cespitose, growing in beds (not in dense tussocks), freely long-
stoloniferous, the culms aphyllopodic, 3.5-9 dm. high, erect,
slender, sharply triangular, 3.5 mm. wide at base, 1.5 mm. above,
very rough above, purplish tinged at base, exceeding leaves, the
basal sheaths hispidulous, strongly filamentose. Leaves with
well-developed blades three or four to a fertile culm, the blades at
first glaucous-green or in age light green, revolute margined,
flat to base, 2.5-3.5 mm. wide, mostly 2-4 dm. long, the margins
very rough, the sheaths strongly yellowish-brown tinged ventrally.
Terminal spike staminate, strongly peduncled, linear, 2—3 cm.
long, 3 mm. wide, the scales oblong-obovate, obtuse, purplish-
black with lighter center and narrowly hyaline margins. Lateral
spikes mostly three, pistillate, approximate and sessile or nearly
so; or the lowest often remote and slender long-peduncled, linear,
4-5 mm. wide, 1.5-3 cm. long, with very many ascending peri-
gynia, closely (or at base more loosely) packed in several ranks.
Lowest bract leaflet-like, about length of culm, sheathless; the
upper much reduced. Scales lanceolate or oblong-lanceolate,
obtuse or acute, narrower and from slightly shorter to slightly
puncticulate, reddish dotted, obscurely few-nerved, 2.25 mm.
jong, 1.5 mm. wide, two-edged, not serrulate, rounded and sessile
at base, abruptly very minutely beaked, the orifice entire. Achenes
lenticular, light brown, oblong-obovate, 1.75 mm. long, 1.25 mm.
wide, es apiculate, the short slender style tardily deciduous;
stigmas tw
416 MACKENZIE: NOTES ON CAREX
The original description of this species by Dewey in Wood’s
Class Book is not satisfactory and it is easy to understand why
subsequent botanists have failed to recognize the plant. It is as
follows:
“Staminate sates 1-2 with oblong and blackish acutish
glumes; pistillate spikes 2-3, cylindric, staminate above, and
hence acutish, lowest short pedunculate; perig. ovate, compressed,
acute, glabrous, entire at the orifice, early falling off, glabrous, a
little longer than the oblong and acute glume; st. a foot and more
high, triquetrous and rough on the angles, with reticulated fila-
ments connecting the leaves towards the base; Ivs. erect, close;
whole plant glaucous, except the spikes.—Wet places, common.”
In 1826, however, in his articles on caricography he had
described two closely related plants, one of which he called C.
acuta, and the other of which he called “Carex stricta Good.”
His description of the latter (Am. Journ. Sci. 10: 269) is an excel-
lent description of the plant I am now treating as Carex strictior
Dewey. After a thorough technical description, in which he
speaks of the nearly black pistillate scales and filamentose sheaths,
he says, ‘‘color of the plants, except the spikes glaucous green,”
and ends up with the following:
“This species, found in England and Sweden, was first recog-
nized in our country, by Mr. Schweinitz. It has probably been
confounded with C. acuta, which it much resembles. There can
be no doubt however that it is a distinct species. Though it
grows in similar situations with C. acuta, it does not form a bog,
but spreads over the surface of the marsh. It differs from that
species too in its color, in the appearance of its spikes, in its more
stiff and erect form and in its fruit being caducous’’ [italics
Dewey’s].
It will be noted that the points emphasized by him are the
same points emphasized in his description of Carex strictior
(supra), namely, color of plant, ‘‘more stiff” form, “strictior,’’
and ‘“‘caducous’’ or ‘‘early falling off’? perigynia. His Carex
strictior grows in wet places and no mention is made of its growing
in bogs (i. e., dense tufts).
Again while he included a description of Carex stricta in Wood's
Class Book he did not emphasize any of the points emphasized
by him in 1826, and he speaks of the plant being found in “wet
- MACKENZIE: NOTES ON CAREX 417
places, as bogs.’’ What then apparently happened is that in the
interval between 1826 and 1843 Dewey found out that his original
“Carex stricta Good.” was different from that European species.
So he made up a new name to apply to his plant, namely C. strictior
Dewey. And at the same time he applied the name Carex stricta
to our common densely tufted species.
As bearing out these views as to the identity of Carex strictior
it is to be noted that in the 1863 edition of Wood’s Class-Book
Dewey added to his description of C. strictior, the words, ‘“‘ Nearer
C. caespitosa than C. stricta,” his Carex caespitosa being the strongly
stoloniferous, bed-forming Carex Goodenovii.
Dewey’s own herbarium does not aid very much. It contains
four specimens kindly lent me by Professor Fernald. Three of
these, collected and distributed by Sartwell, represent the species
I take to be C. strictior. The fourth specimen collected by Olney
is a specimen Of Carex stricta Lam. Similar specimens collected
by Sartwell and Olney are in the Torrey herbarium at the New
York Botanical Garden.
The fact that this species is distinct from Carex stricta has been
noted by a number of botanists. Thus, on a specimen in the
Philadelphia Academy collected in 1892 by Mr. Benjamin Heritage,
we find him noting it as ‘‘a form that does not grow in tufts, but is
stoloniferous.”” Mr. Bayard Long in 1910 collected it at Frazier’s
Bog, near Philadelphia, noting its difference in habit of growth;
and Mr. E. P. Bicknell and Dr. Wm. H. Wiegmann have both
collected it around New York as Carex aquatilis, which in manner of
growth it much resembles. And correspondence with Dr. E. H.
Eames and Mr. F. F. Forbes has resulted in specimens from them
both from plants which they had previously seen were distinct
from Carex stricta,
he specimens listed below are to be referred to this species.
Some are so poor that their identification has been difficult.
SPECIMENS EXAMINED
MatnE: Veazie, Knight, July 15, 1905 (K. M.); Fort Kent,
Mackenzie 3430, 3424, 3427, July 11, 1908 (K. M.); 3663,
July 23, 1908 (K. M.); Somesville, Mt. Desert Island, Rand,
June 24, 1891 (C.); Foxcroft, Fernald 231, June 25, 1895 (C.).
418 MACKENZIE: NOTES ON CAREX
- New Hampsatre: Muik Pond, Lisbon, Graves, June 9, 1893
(C.); base of White Mountains, Oakes (N. Y., C )
Massacuusetts: Chapman, no data (C.); Williamstown, no
data (C.); Tewksbury, Forbes, June 7, 1911 (K. M.).
RuopeE IsLanp: Providence, Olney, June 8, 1871 (K. M.);
Cat Swamp, Providence, Collins, June 19, 1892 (C.).
‘ConNECTICUT: Easton, Eames 8743, June 3, 1914 (K. M.).
OnTARIO: Thedford, Lambton County, Dedge, June 22, 1911
(K. M.). .
~ New York: Penn Yan, Sartwell (one in N. Y. Herb. and three
specimens in Dewey Herb. and one in W. Boott Herb., H.);
Van Cortlandt, Bicknell, June 1, 1890 (C.); Wellesley Island,
Jefferson County, Robinson & Maxon 28, June 28, 1902 (N. Y.);
Torrey, no data (C.); Junius, Sartwell 55 (N. Y.); Junius, Sart-
well, labeled “ Carex aquatilis” (C.); N. Y., Crawe (C.); Litchfield,
Hunt, June 1873 (N. Y.); Herkimer County, Haberer, June 24,
1882 (C.).
New Jersey: Mickleton, Heritage, June 5, 1892 (P.); New
Egypt, J. H. Grove, May 21, 1905 (P.); Repaupo, Crawford,June 15,
1893 (P.); Lindenwold, Githens, June 28, 1902 (P.); Lawrence,
Mackenzie 5868, 5874, May 31, 1914 (K. M.); Oak Ridge, Mac-
kenzie 3185, 3182, 3186, June 28, 1908 (K. M.); Hoboken, A.
Brown, 1877 (C.); “New Jersey’’ Knieskern 57, as Carex salina
Wahl.? (C.); Manchester, Knieskern (C.).
PENNSYLVANIA: Mountainville, Lehigh County, Pretz 319, May
30, 1907, 429, June 22, 1907 (P.); Springfield, Bucks County, Pretz,
June 1886 (P.); “Marshall road” C. E. Smith, May 21 (P.); Penn
Valley, Bucks County, Crawford, June 1, 1899 (P.); Allentown,
Pretz 49, May 18, 1901, 5503, June 17, 1913, 1238, June 11, 1908
(P.); Sellersville, Crawford, June 11, 1899 (P.); Slatington, Bach-
man 2085, May 13, 1911 (P.); Frazier’s Bog, Willow Grove,
Montgomery County, Long 3771, 3767, 3768, 3774, 3760, June
I, 1910, 9434, June 7, 1914, 3355, May 12, 1910; Byberry,
Martindale, June 1865 (P.); Tinicum Meadows, Delaware County,
B. H. Smith, May 30, 1898 (P.); Lowhill, Lehigh County, Prets
4677, June 23, 1912 (P.); Lehigh County, Krout; Easton, Porter,
July 8, 1890 (C.); Long Pond, Monroe County, Britton, June
24, 1895 (C.); Bucks County, Pretz, May 30, 1899 (P.).
MACKENZIE: NOTES ON CAREX 419
DELAWARE: Centreville, Commons, May, 1864, and June 14
1868 (P.); Greenbank, Commons, May 27, 1879, and May 1, 1882
(P.); May 26, 1879 (var. curtissima) 6 a
MARYLAND: Conowingo, Cecil County, Bartram & Long 1250,
June 1, 1913 (P.).
NortH Caroutna: Hendersonville, Biltmore Herb. 1818a, May
27, 1898 (K. M., N. Y.); Highlands, Bilémore Herb. 4990b, June
5, 1897 (N. Y.); Mitchell County, Ashe, May (N. Y.); Macon
County, Buckley (C.).
TENNESSEE: Thompson’s, Ruth 978, May, 1894 (N. Y.).
~ Micuican: Port Huron, Dodge, May 30, 1911 (K. M.); Grand
Rapids, Miss Cole, May 16, 1898 (K. M.); Emerson, Chippewa
County, Dodge, June 16, 1914 (K. M.); Buchanan, Deam 14195, May
29, 1914 (K. M.); Vermilion, Chippewa County, Dodge, June 21,
1914 (K. M.); Olivet, McClatchie, May 24, 1888 (N. Y.); Lake
Superior, Pitcher (C.); Hubbardston, Wheeler, June 25, 1890
C.)
Oxuto: London, Sharp, 1910 (N. Y.); “ad ripas QOhionis,’’
Lesquereux 551 (N. Y.)
Wisconsin: Milwaukee, Hasse, May 27, 1882, and June 3, 1883
(NH. ¥:
~ Ittrvors: Winnebago County, Bebb, 1870 (C.); Chicago,
Moffat 88, June 24, 1893 (N. Y.).
Iowa: Charles City, Arthur (C.).
NEBRASKA: ‘“Nebraska’’ no data (C.).
CAREX Emory Dewey, Bot. Mex. Bound. 230. 1858
Carex virginiana var. elongata Boeckl. Linnaea 40: 432. 1876.
Carex stricta var. Emoryi Bailey, Proc. Am. Acad. 22: 85. 1886.
Carex acuta var. Reverchon’s distr. Texan Pl. 1407; Bailey, |. c.
1886.
“Carex stricta Lam.”’ Mackenzie & Bush, Fl. Jackson Co. Mo. 49.
| 1902:
“Carex stricta var. angustata (Boott) Bailey”, Mackenzie &
Bush, |. c.. 1902.
Carex milligrana Holm, Am. iia Sct 39! 311-1904.
Loosely cespitose, growing in beds, with long slender root-
stocks, the culms SphvOnens, 4-10 dm. ets erect, 6-8 mm
420 MACKENZIE: NOTES ON CAREX
wide at base, slender, sharply triangular and from strongly
roughened to nearly smooth above, exceeding leaves, purplish
tinged at base, the basal sheaths smooth, not flamentose. Leaves
with well-developed blades usually three or four to a fertile culm,
on lower fourth, but not clustered, the blades light green, ascend-
ing, somewhat nodulose, flat to base with slightly revolute margins,
much roughened towards apex, 3-5 mm. wide, 1.5-4 dm. long, the
sheaths very membranaceous ventrally, slightly yellowish tinged.
Terminal spike staminate, strongly peduncled, linear, 2-4.5 cm.
long, 4 mm. wide, the scales oblong-spatulate, obtuse or acutish,
brownish with broad lighter center and hyaline margins. Lateral
spikes four to six, the upper one or two staminate, the lower
pistillate or often staminate at apex, erect, sessile or the lower
short-peduncled, linear, 2-10 cm. long, 4-6 mm. wide, with very
many ascending perigynia closely (or at base more loosely) packed
in several ranks. Lowest bract leaflet-like, from about equalling
to exceeding culm, sheathless; the upper much reduced. Scales
lanceolate, obtusish, short-acute or acuminate, scarcely I mm.
wide, narrower and from somewhat shorter than to strongly
exceeding perigynia, light or purplish brown with broad lighter
center falling with the perigynia. Perigynia plano-convex, ovate
or obovate, closely enveloping achene, at first light green, soon
straw-colored, slightly granular-roughened towards apex, puncticu-
late, somewhat reddish dotted, few-nerved dorsally, 2-2.5 mm.
long, 1.5-1.75 mm. wide, two-edged, not serrulate, rounded and
sessile at base, abruptly apiculate-beaked, the orifice whitish, entire.
Achenes lenticular, oblong-quadrate, substipitate, 1.5 mm. long,
I mm. wide, minutely apiculate, the short slender style tardily
deciduous; stigmas two.
SPECIMENS EXAMINED
New Jersey: Scudders Falls, Mackenzie 5062, June 9, 1912
(K. M.); Red Bank, Gloucester County, Parker, June 6, 1872 (K.
M., P.); Delaware, Mackenzie 4998, May 30, 1912 (K. M.);
Crosswicks Creek, Mackenzie 5809, May 21, 1914 (K. M.):
Coopers Creek, Parker, July 14, 1868 (P.); Camden, Parker
(P., C.); Swedesboro, Lippincott, May 30, 1892 (P).
PENNSYLVANIA: Bethlehem, Schweinitz; York Furnace, Stone,
May 30, 1907 (P.); Penn Valley, Crawford, June 12, 1895 (P.):
Tinicum, Saunders, May 20, 1898 (P.); Tinicum Township,
Saunders, June 19, 1897 (P.); Penn Valley, Bucks County, Craw-
ford, June 1, 1899 (P.); Kintnersville, Van Pelt, June 19, 1905
MACKENZIE: NOTES ON CAREX 421
(P.); Narrowsville, Lippincott, May 30, 1893 (P.); Chester County,
Canby (H.); Fort Washington, iene cca d County, Long, June
II, 1909 (P.).
DELAWARE: Hollyoak, Commons, June 6, 1866 (P.).
MARYLAND: Frederick County, Atkin (C.).
District OF COLUMBIA: Steele, May 18, 1908, and May 12,
1902; Vasey, 188-.
INDIANA: Carmel, Mrs. Deam 10548, May 9, 1912 (K. M.);
Cedarville, Deam 14280, June 14, 1914 (K. M.).
Inuinots: J. Wolf, no data (K. M.).
Manitopa: Brandon, Macoun 13357, June 8, 1896 (N. Y.).
Iowa: Marshalltown, Ball 381, May 15, 1897 (K. M.);
Council Bluffs, Nicollet, 1845 (C.).
Missour!: Greenwood, Bush 6708, May 20, 1912 (K. M.,
N. Y.); Monteer, Bush 2896A, May 17, 1905 (K. M.); Courtney,
Bush 6730, May 28, 1912 (K. M.); Courtney, Bush 6755, 6758,
June 2, 1912 (K. M.); Independence, Bush 703, May 23, 1894
(K. M., C.); Greenwood, Mackenzie, May 9, 1897 (K. M.);
Buckner, Bush 6762, 6776 (N. Y., K. M.), June 5, 1912
(K. M.); Lake City, Mackenzie, May 23, 1897 (K. M.); Sarcoxie,
Palmer 3706, May 19, 1912 (N. Y.); Independence, Bush, June 3,
1888 (N. Y.); St. Louis, Riehl 30, June, 1838 (C.).
NortH DaKota: Towner, McHenry County, Lunell, May 29,
1908, and June 12, 1913 (K. M.); Minot, Ward County, Lunell,
July 7, 1907 (K. M., N. Y.); Ward County, Haigh 224, 1900 (N.
Y.), June 24, 1908 (N. Y.).
Souta Dakota: Rosebud Creek, E. J. Wallace, July 2, 1896,
type of C. millegrana Holm (K. M., N. Y.): Brookings,
Williams & Carter, June 19, 1897 (K. M.); Sylvan Lake, Black
Hills, Rydberg 1085, June, wa (No: 5 | Carter,
June 10, 1897 (N. Y.).
Nesraska: Ashland, Williams, May 1890 (K. M., N. Y.);
Fort Niobrara, Wilcox, May, 1888 (C.).
Kansas: Manhattan, May 16, 1889 (C.); Riley County, Norton
553, May 23, 1895 (N. Y.); Osborne City, Shear 51, May 26, 1894
(C).; Wichita, Carleton, May 9, 1888 (C.).
OKLAHOMA: Sapulpa, Bush 1070, May 2, 1895 (C.).
429 MACKENZIE: Notes on CAREX
Cotorapo: Fort Collins, Marshall 2534, June 12, 1899 (N. Y.);
Fort Collins, State Agr. Col. 2570, June 7, 1898 CO
Texas: Llano County, Reverchon 1407, May 1885 (K. M., N.
Y.); Upper Rio Grande, Bigelow, Mexican Boundary Survey,
duplicate of type (C.).
INDEX TO AMERICAN BOTANICAL LITERATURE
1912-1915
The aim of this Index is to include all current botanical literature written by
Americans, published in America, or based upon American material ; the word Amer-
ica being used in the broadest sense.
eviews, and papers that relate exclusively to forestry, agriculture, mile oe
manufactured products of oe origin, or laboratory methods are not inclu a
attempt is made to index the literature of bacteriology. An occasional trina is
Sins in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not mentioned unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
preciated.
This hi dex is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
nieces must take all cards published during the term of his subscription rre-
ndence relating to the card issue should be addressed to the Treasurer of the Torrey
Bobi cal Club,
Anderson, J. P. Some observations on sycamore blight and accom-
panying fungi. Proc. Iowa Acad. Sci. 21: 109-114. pl. 7, 8. 1914.
Barrus, M. F. Late blight and rot of potatoes. Cornell Agr. Exp.
- Sta. Cire. 19: 77-83. f. 61-67. My 1913.
Bartlett, H. H. The experimental study of genetic relationships. Am.
Jour. Bot. 2: 132-155. 23 Ap 1915.
Belling, J. The evening primrose varieties of De Vries. Am. Nat. 49:
319, 320. My 1915.
Berry, E. W. A species of Copaifera from the Texas Eocene. Torreya
15: 41-44. f. 1-5. Mr 1915.
Copaifera yeguana sp. nov. :
Bessey, E. A., & Byars, L. P. The control of root-knot. U.S. Dept.
Agr. Farm. Bull. 648: 1-19. f. 1-20. 1 Ap 1915.
Blake, S.F. Notes on the genus Sabatia. Rhodora 17: 50-57. pl. 112.
2 Ap I9I5.
Includes Sabatia obtusata sp. nov.
Blake, S.F. Two new Mexican amaranths. Jour. Bot. 53: 103, 104.
Ap 1915 re
Amaranthus annectens and A. lepturus, spp. nov.
Blodgett, F. M. Further studies on the spread and control of hop
mildew. N. Y. Agr. Exp. Sta. Bull. 395: 29-80. At; 2 + f- 21, 2.
F 1915. "493
424 INDEX TO AMERICAN BOTANICAL LITERATURE
Brainerd, E. Viola septentrionalis in British Columbia. Rhodora 17:
70,71. 2 Ap I9gt5.
Bubék, F., & Sydow, H. Einige neue Pilze. Ann. Myc. 13: 7-12.
Includes Helminthosporium obclavatulum sp. nov. from Brazil.
Burlingham, G. S. (Agaricales) Agaricaceae (pars) Lactariae (pars).
N. Am. Fl. 9: 201-236. 30 Ap I9I5.
Includes 18 new species in Russula.
Chase, A. A teretological specimen of Panicum amarulum Hitche. &
Chase. Rhodora 17: 72. 2 Ap 1915.
Cockerell, T. D. A. Notes on orchids. Bot. Gaz. 59: 331-333. f. I-
15 Ap 1915.
Beles poe pediloides (fossil) sp. nov.
Coulter, J. M. Charles E. Bessey. Science II. 41: 599, 600. 23 Ap
1915.
Dachnowski, A. The international phytogeographic excursion of 1913
and its significance to ecology in America. Jour. Ecology 2: 237-
245. D 1914.
Dalbey, N. E. On the anatomy of Grindelia squarrosa. Kansas Univ.
Sci. Bull. 9: 31-41. pl. 2-7. D 1914.
Dearness, J. The myxos of Middlesex. Ontario Nat. Sci. Bull. 7:
3-10. 1912.
Derick, C. M. Dr. David Pearce Penhallow. Canadian Rec. Sci. 9:
387-390. Jai1915. [Portrait]
Elmer, A. D. E. Two hundred twenty six new species—I. Leaflets
Philip. Bot. 7: 2543-2700. 27 Mr 1915;—II. Leaflets Philip.
Bot. 8: 2719-2883. 27 Mr 1915.
Farr, C. H. Notes on a fossil tree-fern of Iowa.
Sci. 21: 59-65. f. 3-5. 1914.
Frazier, Z. R. Notes on the ae of Iowa ihe
Acad. Sciy 21: 67-75. 1914.
Proc. Iowa Acad.
Proc. Iowa
Gates, F. C. Swamp vegetation in hot springs areas at Los Bafios,
Laguna, P. I. Philip. Jour. Sci. 9: (Bot.) 495-516. pl. r1-15. Mr
1915. |
Greenman, J.M. Morphology asa factor in determining relationships.
Am. Jour. Bot. 2: 111-115. 23 Ap 1915.
Grossenbacher, J.G. Medullary spots and their cause.
Bull. Torrey
Club 42: 227-239. pl. 10, rz. Ap 1915.
INDEX TO AMERICAN BOTANICAL LITERATURE 425
Grout, A. J. Some common mosses. Nature-Study Rev. 2: 204-208.
Ap 1915; II. The hair cps. Nature-Study Rev. 2: 229.-232. f.
I-4. My 1915. [Illust.]
Harris, J. A., Lawrence, J. V., & Gortner, R. A. On the osmotic
pressure of the juices of desert plants. Science II. 41: 656-658.
30 Ap I915.
Hasse, C. H. Pseudomonas citri, the cause of Citrus canker. [A pre-
liminary report.] Jour. Agr. Research 4: 97-100. pl. 9, 10. 15
Ap I915.
Headden, W. P. Yellow-berry in wheat—its cause and prevention.
Colorado Agr. Exp. Sta. Bull. 205: 1-38. F 1915. [Illust.]
Hedgcock, G. G. Injury by smelter smoke in southeastern Tennessee.
Jour. Washington Acad. Sci. 4: 70, 71. 19 F 1914.
Herriot, W. The Rosaceae and Leguminosae of Galt, Ont. and
vicinity. Ontario Nat. Sci. Bull. 7: 28-34. 1912.
Hoagland, D. R. Organic constituents of Pacific coast kelps. Jour.
Agr. Research 4: 39-58. 15 Ap 1915.
Hollick, A. A new fossil species of Ficus and its climatic significance.
Jour. N. Y. Bot. Gard. 16: 43-47. pl. §2, 53. 10 Ap 1915.
Ficus interglacialis sp. nov.
Holzinger, J. M. Natural history. History of Winona County.
364-381. Chicago, 1913.
Howe, R. H. Classification de la famille des Usneaceae dans l’Amér-
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Hutchinson, A. H. On the male gametophyte of Picea canadensis.
Bot. Gaz. 59: 287-300. pl. 15-19 + f. a—u. 15 Ap 1915.
Jamieson, C. O. Phoma destructiva, the cause of a fruit rot of the
tomato. Jour. Agr. Research 4: 1-20. pl. a, b, I-6. 15 Ap 1915.
Kaczmarek, R. M. Crocion achlydophyllum (Greene). An ecological
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1915. ‘
Kenoyer, L. A. Notes on variation in Micranthes texana. Proc. lowa
Acad. Sci. 21: 123, 124. pl. 17. 1914.
Kern, F. D. The genetic relationship of parasites. Am. Jour. Bot.
2: 116-131. 23 Ap 1915.
Klugh, A. B. The flora of the sand dunes of Prince Edward County.
Ontario Nat. Sci. Bull. 7: 40-51. f. 1-15. 1912. [lllust.]
Klugh, A. B. The hydrogohytic formations of Lake Opinacon. On-
tario Nat. Sci. Bull. 7: 63. 1912.
426 INDEX TO AMERICAN BOTANICAL LITERATURE
Klugh, A. B. The plant formations of the Bruce Peninsula. Ontario
Nat. Sci. Bull. 7: to-21. f. 1-4. 1912. {Illust.]
Klugh, A. B. A zonal algal habitat. Ontario Nat. Sci. Bull. 7: 64.
1912.
Kock, G. Der nordamerikanische Stachelbeermeltau und _ seine
Bekampfung. Obstziichter. 1-4. f. 1-3. 8 N 10915.
Macbride, T. H. Twenty-five years of botany in Iowa. Iowa Acad.
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Reissued Ja 30, 1914, in corrected form, pp. I-26.
Macdougal, D. T. Department of Botanical Research. Yearbook
Carnegie Inst. Washington 13: 63-104. 1914.
Mackenzie, K. K. Two new sedges from the southwestern United
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x Wootoni and C. Rusbyi, spp. n
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1-32. pl. 1-§ +f. 1-7. . 13 Ap roi.
Maxon, W. R. Polypodium marginellum and its immediate allies.
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McCubbin, B. W. A. The edible toadstools—the smooth Lepiota.
Ontario Nat. Sci. Bull. 7: 53-56. 1912. [I}ust.]
McCubbin, W. A. Experimental results on peach canker. Ann. Rep.
Ontario Fruit Growers’ Assoc. 46: 28-32. 1915.
Merrill, E. D. Dilleniaceae novae. Philip. Jour. Sci. 9: (Bot.) sif-
530. N 1914.
Sixteen new species are described.
Merrill, E.D. Meliaceae novae. Philip. Jour. Sci. 9: (Bot.) 531-541.
Mr 1915.
Eleven new species described.
Merrill, E. D. Notes on Philippine Euphorbiaceae, IT. Philip. Jour.
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Thirty-five new species described.
Miller, C. R. Fresh-water algae occurring in the vicinity of the Island
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Mitchell, F. Weed migration. Ontario Nat. Sci. Bull. 7: 56, $7.
Igi2.
Muller, T. The industrial fiber-plants of the Philippines. Jour. N. Y.
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INDEX TO AMERICAN BOTANICAL LITERATURE 427
Murrill, W. A. Agariceae (pars.). N. Am. Fl. 9: 237-296. 30 Ap
1915.
Includes 70 new species in, Lentinus (2), Crinipellis (4), Polymarasmius (1),
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Scytinotus (1), Pleurotopsis (1), and the new genus Marasmiellus
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Neuman, J. J. The Polyporaceae of Wisconsin. Wis. Geol. & Nat.
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Nichols, G. E. The vegetation of Connecticut, IV. Plant societies
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Pammel, L. H., & King, C. M. Weed survey of Story County, Iowa.
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Pennington, L. H. Temperate species of Marasmius. N. Am. Fl. 9:
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Pool, R. J. The invasion of a planted prairie grove. Proc. Soc. Am.
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Gesells. 21: 183-195. 1912; 22: 254-265. I913-
Includes descriptions of Ampelopsis micans and Parthenocissus lactevivens, tits
pulchra, Rhododendron coreanum and Viburnum Harryanum, spp. nov.
Rehm, H. Ascomycetes novi. Ann. Myce. 13: 1-6. 25 Mr 1915.
; Includes several new species from m North America.
428 INDEX TO AMERICAN BOTANICAL LITERATURE
Rendle, A. B. Three early Jamaican botanists. Jour. Bot. 53: 104-
107. Ap Igr5.
Roberts, J. W. Sources of the early infections of apple biteuabiet
Jour. Agr. Research 4: 59-64. pl. 7. 15 Ap 1915.
Sargent, C. S. Washington and Michaux. Rhodora 17: 49, 50. 2
Ap 1915.
Sheldon, S. M. Notes on the growth of the stipe of Nereocystis
Luetkeana. Puget Sound Marine Sta. Publ. 1: 15-18. 15 Ja 1915.
Sherff, E. E. Studies in the genus Bidens. II. Bot. Gaz. 59: 30I-
316. f. I-3. 15 Ap 1915. -
Five new species are described.
Sinnott, E. W., & Bailey, I. W. Investigations on the phylogeny of
the angiosperms: No. 4. The origin and dispersal of herbaceous
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Smiley, F. J. The alpine and subalpine vegetation of the Lake Tahoe
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Smith, C. P. Carex Tuckermani niagarensis; a neglected sedge.
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Standley, P. C. A new species of Achyranthes from Tobago. Proc.
Biol. Soc. Washington 28: 87, 88. 13 Ap 1915.
Achyranthes ingramiana sp. nov.
Stewart, G. R. Availability of the nitrogen in Some Coast kelps.
Jour. Agr. Research 4: 21-38. 15 Ap 1915.
Surface, F. M., & Pearl, R. Studies on oat breeding. II. Selection
within pure lines. Maine Agr. Exp. Sta. Bull. 235: 1-40. f. 1, 2
Ja 1915.
Sydow, H., & Sydow, P. Novae fungorum species—XIII. Ann. Myc.
13: 35-43. f. 1-3. 25 Mr i9%5.
Includes Phyllachora Anthephorae from Jamaica and Eurytheca trinitensis from
Trinidad.
Truog,E. A new theory regarding the feeding power of plants. Science
I]. 41: 616-618. 23 Ap 1915.
Van Eseltine, G. P. An abnormal specimen of Citrullus vulgaris.
Torreya 15: 44, 45. f. 2, 2. Mr 191s.
Verink, E. D. A preliminary report on the flora of Linn county.
Proc. Iowa Acad. Sci. 21: 77-99. 1914.
Weatherby, C. A. Two extensions of range in Gramineae. Rhodora
a7: 71. 2 Ap 19xs.
Weir, J. R. Observations on Rhizia in flata. Jour. Agr. Research 4:
93-96. pl. 8. 15 Ap 1915.
VOLUME 42, PLATE 21
BuL_. ToRREY CLUB
LES
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1-6. CAMPYLOPUS PERUVIANUS R. S. WILLIAMS
7-15. ASTOMUM CHILENSE R. S. WILLIAMS
BULL. TORREY CLUB
VOLUME 42, PLATE 22
SOOVAGS.
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PNG Nas
I-11. PTEROGONEURUM ROSEAE R. S. WILLIAMS
12-20. CROSSIDIUM ROSEI R. S. WILLIAMS
BULL. TORREY CLUB
VOLUME 42, PLATE 23
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1-10. PSEUDOCROSSIDIUM CHILENSE R. S. WILLIAMS
11-19. PSEUDOCROSSIDIUM APICULATUM R. S. WILLIAMS
BULL. TORREY CLUB VOLUME 42, PLATE 24
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12-21. TORTULA LIMENSIS R. S. WILLIAMS
1-9. PHYSCOMITRIUM ROSEAE R. S. WILLIAMS
10-17. PHILONOTIS FRAGILICAULIS R. S. WILLIAMS
Vol. 42 No. 8
BULLETIN
OF THE
TORREY BOTANICAL CLUB
AUGUST, 1915
The origin of dwarf plants as shown in a sport of Hibiscus
oculiroseus
A. B. STOUT
(WITH PLATES 26 AND 27)
The publication in 1901-1903 of Die Mutationstheorie
aroused a new interest in the well-known sports or spontaneous
variations of earlier writers. While Darwin had recognized the
occurrence of such variations he did not consider that they were
the sole means or even the most frequent means by which species
originated. De Vries sought to separate mutability as a type of
sporadic variability from fluctuating variability, to define quite
different laws for the two, and to ascribe to the former the only
power to give rise to forms of specific rank.
Since 1901 much further evidence has accumulated regarding
the appearance, the behavior, and the heredity of so-called muta-
tions. It is fully established that marked variations do appear
and that these are often hereditary. This phase of the general
doctrine of variation is fully established. Of the various practical
and theoretical considerations presented by these cases none are
more fundamental than those relating to the expression and the
heredity of ‘‘newly born” characteristics.
The general conception of mutation and
havior that indicate the process were combined by
the doctrine that plant characters are to be con
exactly represented by definite hereditary units that they can be
properly called ‘unit characters.” De Vries especially em-
the categories of be-
De Vries with
sidered as so
[The BuLietin for July (42: 365-428. Al. 21-25) was issued July 29, ro15.]
430 Stout: THE ORIGIN OF DWARF PLANTS
phasizes this view in the preface of the English translation of
Die Mutationstheorie (1909).
It has also been shown that numerous cases of hybridization
result in an increased variability in the F, generation over that of
the parents, in the production of intermediate forms, and in unex-
pected ratios. The Mendelian factorial hypothesis, which at-
tempts to explain such phenomena, assumes that the difficulties
of analysis in terms of unit characters indicate that characters
are themselves the results of the combined action or interaction
of hereditary units called factors to which various values can be
attributed as seems necessary. There are some striking cases
of reversion resulting from crosses that seem to support this view;
as, for example, the production in sweet peas of the reversionary
purple bicolor known as “Purple Invincible’”’ from crosses be-
tween certain white flowering strains of ‘‘Emily Henderson”’
(Bateson, 1913, Chap. V). The developments of the Mendelian
theory have led, however, to various and conflicting conceptions
regarding the nature of the hereditary units and of the funda-
mental processes of mutation or discontinuous variation.
De Vries on the one hand considers that mutability is due to
lability of hereditary physiological units; these change from
stable to labile, from active to inactive (or latent) or even to a
semilatent condition, giving irregularities of expression both in
inbred and crossbred lines of progeny. Only in the rare cases of
progressive mutation are new characters added and in none of the
mutations is complete loss of hereditary qualities essential. The
units of De Vries assume different degrees of activity; they are
not uniform and consistent units in their influence.
In marked contrast to this view is the conception of Bateson
that all variation is due to the presence or absence of ‘unit factors’,
that ail true mutation is due to loss of factors, and that cases of
discontinuous variation resulting from crosses are reversions which
are due to recombination of factors. This view minimizes the
occurrence of latency, ignores lability and insists on a rather
rigid unity of assumed hereditary units, not in any sense compar-
parable with the appearance of the visible characters of plants.
These two very different interpretations indicate clearly the
Stout: THE ORIGIN oF DwarF PLANTS 431
difficulties that arise in the attempts to analyze the facts of
heredity and variation in terms of units and raise anew the ques-
tions as to the validity of the conceptions regarding mutation.
It is the purpose of this paper to present the facts as thus
far determined regarding the origin and the behavior of a marked
variation in Hibiscus oculiroseus and to point out some considera-
tions suggested by such cases.
THE DWARF FORM OF HIBISCUS OCULIROSEUS
The two plants shown in PLATE 26 are typical for the appear-
ance of dwarf and robust plants of H. oculiroseus at the end of
the first year of growth as handled in my cultures. Both plants
were from seed planted in January, 1914. The seedlings were
grown in pots in a greenhouse until May, when they were planted
in adjoining beds in the experimental garden. The photograph
was taken September 13, 1914. A glance at the plate will show
the marked differences that exist between the two types. The
dwarf plant is shorter, but is more branched with large well-de-
veloped lateral branches arising close to the base; the internodes
are shorter, making the leaves more crowded; the leaves are smaller
and many of them are somewhat irregular or asymmetrical in
outline and many of them are crinkled. Few of the dwarf plants
have thus far come into flower, but those that have showed flowers
nearly if not fully as large as the robust types.
From the data regarding the ancestry of the dwarf plants, it
appears that a plant of Hibiscus oculiroseus was obtained by the
New York Botanical Garden from the firm of Pitcher and Manda
in the year 1896. ‘This firm obtained their original stock of this
plant from Mr. W. F. Bassett of Hammonton, New Jersey, who
introduced the plant (Britton, 1903) into the trade some years
before. Bassett obtained the first plant of this type about the
year 1880 from a colony of wild plants growing near Absecen,
New Jersey. The plant was propagated by seed, and introduced
into the trade. It was commonly called the ‘‘Crimson Eye.”
Britton (1903) points out that this type differs from H. Mos-
cheutos in several characters. H. Moscheutos in its most abundant
form, at least in the vicinity of New York City, has flowers of a
432 Stout: THE ORIGIN OF DWARF PLANTS
rose color, lighter along the veins and becoming nearly pure white
for about 1 cm. at the base of the corolla lobes. Hibiscus oculi-
roseus has a rose red or Tyrian rose eye about 2 cm. in radius,
beyond which the petal is a sea-foam yellow. The flower pods
are ovoid with a long tapering point, the calyx segments are
triangular-lanceolate and nearly twice as long as broad. Hibiscus
Moscheutos has a nearly globular bluntly pointed pod. On this
account he gives the type specific rank under the name H. ocu-
liroseus.
CULTURES OF H. OCULIROSEUS AT THE NEW YorK BOTANICAL
GARDEN
Open-fed seed was collected from the plant obtained from
Pitcher and Manda and planted for the purpose of increasing
the stock of the species. The progeny (Nash, 1909) was composed
in part of plants conforming to H. oculiroseus and in part of plants
whose flowers suggested that the plants were hybrids between
HI. oculiroseus and the rose-flowered type of H. Moscheutos, an
assumption which the writer has since proved to be true by con-
trolled crosses.
At the time the writer began his investigations with Hibiscus
(1911) there were seven plants of the H. oculiroseus characters
growing in the Garden, all derived from the one parent plant.
All of these were vigorous plants about five feet tall. Five of
these plants have been used as parents and will be referred to as
O No. 1, O No. 2, etc.
One of these plants (O No. 1) produces each year a consider-
able number of leaves, somewhat crinkled and irregular, and the
uppermost internodes of the branches are somewhat shortened.
One would be inclined to attribute this to a fluctuating varia-
tion due perhaps to local soil conditions. The other four plants
show no trace of any of the dwarf characteristics.
PLANTS OF THE FIRST PEDIGREED GENERATION
In 1912, fifteen plants (Series I) were grown from open-fed
seed collected from the plant O No. 1. Fourteen of these were
robust and vigorous in growth, but three of them had some leaves
Stout: THE ORIGIN OF DwARF PLANTS 433
irregular in shape and also exhibited a shortening of internodes at
the tips of stems quite identical to that of the parent plant and
were in general intermediates between the robust type and the
extreme dwarf type. One plant was of the dwarf type quite
identical with the one above described. Ten of the plants, in-
cluding the dwarf and one of the intermediates, were grown in
1913, when all bloomed. The dwarf plant (No. 1 of Series I)
continued small and much branched with many crumpled leaves.
Two capsules of selfed seed were obtained from this plant and one
capsule was obtained from a sister plant (No. 5 of Series I) of the
robust type.
As to flower coloration and pod character, nine of the tén plants
were typical H. oculiroseus. One plant had flowers with faint
pink coloration outside the eye in the blade of the corolla, although
at a short distance or upon casual survey the flowers appeared
to be typical H. oculiroseus. There was outside of this one plant
no indication but what the seed obtained was strictly selfed, and
even in this one case the flowers borne were not such as have been
obtained in the F, hybrids between the parent plant and the only
other type of Hibiscus growing in the vicinity of the seed parent.
The winter of 1913-14 was unusually severe on the Hibiscus
plants growing in the experimental plots, killing about 1,500
plants of various cultures, including the dwarf plant (No. 1) and
the robust plant (No. 5) of Series I, from which selfed seed had
been obtained.
In addition to the plants of Series I, there were also grown
during the summer of 1913 forty-five plants (Series VI) derived
- from selfed seed obtained during the summer of 1912 from the
All of these were dwarf plants.
same parent plant (O No. 1).
No se
Four produced flowers that were typical H. oculiroseus.
ripened and all the plants died during the winter of 1913-14.
All the plants of this series were dwarfs, while of the fifteen plants
of another series (Series I) from a pod of the same parent only
one dwarf appeared as already noted.
The dwarf character of these plants was most conspicuous in
comparison with the selfed progenies of four other plants of
Hibiscus oculiroseus (sister plants of O No. 1) grown during the
434 Strout: THE ORIGIN OF DWARF PLANTS
summer of 1913 from seed of 1912. The data for these can be
briefly summarized as follows: from plant O No. 2, nine plants
were grown, of which four bloomed; from plant O No. 3, forty
plants, of which ten bloomed; and from O No. 5, twenty-seven
plants, of which none came into flower. All of the plants in these
four series, 103 in number, were typical for the robust type with
no trace of any of the dwarf characteristics. The twenty-four
plants that bloomed were true to the H. oculiroseus type of flower.
None of the plants matured seed and every one was killed during
the winter of 1913-14.
PLANTS OF THE SECOND GENERATION
In the summer of 1914 three series of plants were grown,
constituting a second generation in descent from the parent O
No. 1. These may now be described.
Series VII. The parent of this series was a dwarf plant (No. I
of Series I). Thirty-five plants were grown from seed of a single
pod. Twenty-seven were uniformly of the dwarf type; one was
a typical robust plant and seven were intermediates showing in
slight degree the characteristics of the dwarf type. Six plants
bloomed; one had eyed flowers with pale pink in the blade (quite
similar to No. 8 of Series I), three had the eye somewhat diluted,
and on two the eye of the flower was so diluted that the flowers
appeared to be nearly pure white. The one plant of robust habit
had flowers of a type which suggests that it is a stray plant of
hybrid origin.
Series VIII. Of thesame parentage as the Series VII. Forty-
six plants were grown from selfed seed of a single capsule. Forty-
five of these were dwarf quite like the one from this series shown
on PLATE 26. One plant was intermediate but was different from
other intermediates in possessing a considerable development of
lateral branches, the main branch was itself robust and the leaves
were only slightly crinkled. The five plants that bloomed were
all typical for the H. oculiroseus type and were only slightly if any
smaller than the flowers of robust plants. The plants of this
series are shown in the middle of PLATE 27 with the marker
standing in their midst. The plants of Series VII are shown
Stout: THE ORIGIN oF DwarF PLANTS 435
beyond the label stake in the rear. The intermediate plant in
Series VIII just mentioned is shown at point a. To the right in
the foreground are hybrids between O No. 2 and a plant of
Hibiscus Moscheutos, and to the left are hybrids between two
types of H. Moscheutos, all of the same age as the dwarf plants.
Series IX. The thirty-four plants of this series were grown
from the selfed seed of one capsule of plant No. 5 of Series I. The
parent was a robust plant. Thirty-three of the plants were of
the robust type quite uniformly like the one of this series shown
on PLATE 26. One plant was classed as an intermediate; it was
smaller than the others, its leaves were somewhat crinkled, but
it was not branched from the base. Twelve plants produced
flowers; ten of these were typical H. oculiroseus, one had flowers
with slight pink coloration outside the eye, and on one plant the
flowers were quite pink outside the eye, quite like the flowers
produced by hybrids between H. oculiroseus and the pink-flower
type of H. Moscheutos.
Nearly all the plants of these three Series (VII, VIII and IX)
lived through the winter of 1914-15 and were grown during the
summer of 1915 in the same beds as in the previous year. In the
second year of growth several main stems, usually three to five,
are produced by the single cluster of roots belonging to a plant.
As the plants were grown about 30 cm. apart they were much
crowded in the second year, which is a condition that does not
favor the development of secondary branches and hence the
plants were much less bushy than in the first year of growth.
Under these conditions there were fewer differences between
plants previously classed as intermediate and dwarf.
In 1915 the plants of Series VII ranged from 1.25 m. to 1.6 m. in
height with the exception of the plant classed as robust which was
2.08 m. tall but which from the character of its flowers appears to
_ be an accidental hybrid. Series VIII ranged from 1.3 m. to 1.6 m.
in height. The dwarf plant shown to the left in PLATE 26
had four main branches and stood 1.45 m. tall and was typical
for the average plant; the plant shown at point a in PLATE 27,
classed as intermediate in 1914, was of nearly the same height
and appearance. Series IX, described as robust in 1914, varied
436 Stout: THE ORIGIN OF DWARF PLANTS
from 1.75 m. to 2m. in height with the average quite 0.3 m.
taller than that of Series VII and VIII.
During 1915 there was also grown from seed a series of twenty-
eight plants from selfed seed of plant O No. 1, the parent of the
Series I and VI. On August 25, these plants varied from 0.4 m. to
0.75 m. tall and nearly all had the dwarf characteristics. The
series showed greater variation in general vigor than did the plants
of Series VII and VIII in their first year of growth. A series of
twenty-three plants derived from selfed seed of one of the robust
plants of Series I, grown at the same time were in comparison
decidedly robust averaging from 0.8 m. to I m. in height and
with no crumpled leaves or strongly developed lateral branches.
Another series of eleven plants descended from a plant of Series
I classed as intermediate was on the whole intermediate be-
tween the two series just mentioned.
SUMMARY OF THE CULTURES
A total of 288 plants have been grown (at least through a
single season) from seed obtained from five sister plants of
Hibiscus oculiroseus. The dwarf type of plant has appeared only
among the progeny of a single plant (O No. 1); the 103 plants
derived from the other four sister plants were without exception
of the robust type.
There was much diversity among the progeny of the plant O
No. 1, and the two series of plants grown in the first generation
gave quite different proportions. In Series I, there were eleven
robust, three intermediate and one dwarf; in Series VI there
were forty-five plants, all of which were dwarf.
The progeny of a single dwarf plant (the only one that has
thus far matured seed) was composed of one robust (later
appearing to be an accidental hybrid), eight intermediate
and seventy-two dwarf plants. While the dwarf type has not
bred true the greater number of the progeny are of that type.
There is a pronounced tendency to breed true. Robust plants
in this line of descent do not always breed true, as the only one
tested gave one intermediate plant and thirty-three robust plants.
A condensed schematic presentation of these facts regarding
Stout: THE ORIGIN OF DwarF PLANTS 437
the progenies of different lines of descent, not including the series
started from seed in 1915, is given in the accompanying diagram.
1914. Series VII.
27 dwarf
(0t4: Serie eer 1 robust (?).
12 ries f. ie
I dwarf 7 intermediate.
Bion”
og II robust. Eels + IQT4. ae Vill.
a 3 inter- 45 d
yf mediate. \ I risen ea
heave a 1914. Series IX.
O No. 1
ae ™ 33 robust.
I intermediate.
— 1913. Series VI.
dwarf.
ee Serie
Parent4 gaat op lants, su
rake:
O No. ean aug
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27 sen sea
DIscuUSSION
The dwarf type of Hibiscus oculiroseus constitutes a marked
and distinct variation from the usual robust type of the species.
It has smaller leaves, many of the leaves are crumpled, the inter-
nodes are shorter, and there is an increased development of bran-
ches. It is such group-differences as these that constitute the
elementary species of De Vries, for, as he states (1901, p. 176):
‘“‘lementare Arten unterscheiden sich von ihren nachsten Ver-
wandten mehr oder weniger in allen ihren Merkmalen.” All the
changes, however, are considered to be a result of a change in
the condition of a single pangen.
It is difficult or impossible to dence such a mutation in
terms of definite characters that have been gained or lost. The
ordinary type of development of the leaves and internodes have
been modified mainly in the direction of arrested growth, while
the amount of branching has been increased, due chiefly to the
development of basal lateral buds that usually remain dormant.
In comparison with the robust type the characteristics of the
typical dwarfs are readily recognized, the general effect being
438
Stout: THE ORIGIN oF DWarF PLANTS
quite adequately shown in the illustrations on PLATES 26 and 27.
The characteristics of the two types can be listed as follows:
bust form
First year of growth, 0.8—-1.1 m.
Second year of growth, 1.75-2 m
No branching from base of shoots.
Internodes about 30 in number.
Leaves ovate, larger ones lobed, pal-
y
glabrous
a
Flowers and fruit are
oculiroseus.
typical A.
Dwarf form
0.4-0.75 m.
Large branches from base. .
Internodes about 30 in number, but all
proportionally shorter.
Leaves identical except that the largest
are smaller than the average of
robust type; many leaves crinkled.
Flowers and fruit identical with those of
robust form; apparently no smaller.
While typical plants classed as robust and dwarf were very uni-
form in the above characteristics, there were a few plants, as noted,
described as intermediates or semi-dwarfs, which possessed in some
degree of development one or more of the characters of the dwarf
form but which in the second year of growth were less distin-
guishable.
The mutant of Oenothera Lamarckiana that most nearly ap-
proaches in its characters the dwarf Hibiscus is, of course, Oe.
nanella. The principal characteristics of this plant compared
with those of the parent form as given by the description of De
Vries may be given as follows:
Oe. Lamarckiana
Robust: 1.5-1.8 m. tall
Secondary branches abundant and
g
stron
Leaves ovate-lanceolate; long petioled,
much crumpled surfaces
Flowers with petals 3-4 cm. — buds
thin, lt an to a poin
Fruit and seeds n
Oe. nanella
warf: about one-fourth as tall as O.
Lamarckiana, often blooming when
10-20 cm. tall.
gusprme lt branches lacking or nearly
absen:
aes 0 cack shorter — leaves
flowers more compac
Leaves sessile or nearly so; ea at
base, ofte
Fruit and seeds of almost normal size.
The dwarf Oe. nanella differs from Oe. Lamarckiana in nearly
all of its vegetative organs.
De Vries maintains, however, that
such a mutation is brought about by a change in the activity of
Stout: THE ORIGIN OF DWARF PLANTS 439
_a single hereditary ‘‘pangen,” but that the visible result depends
only partly on the direct results of the changed condition in one
pangen. On this point he states (1901, p. 305): ‘Der. dussere,
sichtbare Erfolg hangt also nur zum Theil von der Mutation, zum
Theil aber auch von den alteren Merkmalen ab. Oder mit anderen
Worten, die neue Art kennzeichnet sich in der Regel nicht durch
eine enizige neue Eigenschaft, sondern dadurch, dass veile oder alle
Organe in bestimmter Weise umgestaltet wurden.” This means
that in the case of Oe. nanella when the alta pangen (1913) becomes
inactive it affects not only the height of the plant but also produces
a transformation of many other characters, such as the nature
of the leaves and the branching. De Vries states regarding Oe.
nanella (1904, p. 532): ‘‘The most remarkable feature is the
shape of the leaves. They are broader and shorter, and especially at
the base they are broadened in such a way as to become apparently
sessile. The stalk is very brittle, and any rough treatment may
cause the leaves to break off.” These remarkable differences in
the leaves are brought about, De Vries assumes, by a change in
the ‘‘alta pangen’’, a pangen concerned with the height of the plant.
Such a view is, it would appear, directly opposed to a conception of
independence between heredity units. We get in such cases the
clearest possible conception of the application of De Vries’s idea
of a heredity unit (‘‘pangen’’) as an element of the germ-plasm
which affects and determines the character of a plant as a whole,
constitutionally, and in all or many of its characters. Such a
view is quite different from the units of Weismann, which were
assumed to be individual in effect and sufficiently numerous to
correspond directly to every detailed feature of expression.
The original Mendelian doctrines of unit characters assumed
qualitative germ cell units, independent in action, and each
responsible for the expression of a character. Color and wrinkled-
ness of peas (qualities of the cotyledons) were considered as separ-
able hereditary units quite independent of any preformation of the
cotyledons themselves. Mendelian theories have been concerned
with the phase of development and heredity which involves
qualities, and though evading the difficulties of assumed spacial
relationship in preformation, they assume tiat the qualitative
440 STOUT: THE ORIGIN OF DWARF PLANTS
values of the ‘“‘factors’’ are predetermined. It is an important
fact established especially by Mendelian studies that qualities of
organs may appear and disappear independently of the organ and
to certain degrees independently of each other. This suggests
that the older conceptions of the preformation of organs as such
do not hold and that a hereditary unit for an organ with all its
characteristics does not exist. This is very clearly seen in the
numerous studies of inheritance and variation of color in which
organs like petals, leaves, and stems of plants, eyes of insects, etc.,
re nain quite the same except for color. Such facts emphasize on
the one hand the wide possibilities of latency as most excellently
developed by De Vries, and on the other the most evident fact
that qualities which appear to be localized in organs are more
often general qualities of the entire organism, the development of
which is a matter of intercellular and inter-tissue relationships.
The arguments of De Vries consistently seek to establish the
doctrine of the fewness of the pangens. On this basis dwarfness
is assumed to be the same in all the different cases in which it ap-
pears throughout the plant kingdom.
No consistent uniformity, however, exists among the dwarf
types derived in different species or even in the same species. Some
types of dwarf garden peas appear to differ from tall varieties
chiefly in the character of the internodes. In the ‘“‘brachytic”’
varieties of cotton (Cook, 1915) the shortened internodes a 1d modi-
fied leaves and bracts are usuall confined to the fruiting branches.
Numerous types of dwarf and semi-dwarf types exist in many
species, a notable instance of which has been recently described by
Bartlett (1915), who found that two dwarf types arose in a single
pedigreed generation of Oenothera Reynoldsii. One of these,
semialta, is about half as tall as f. typica and has a very dense
and showy inflorescence, in which the fruits and flowers are very
little smaller than in the parent form. The leaves, however, are
decidedly reduced. As described by Bartlett (1915, p. 130) the
sister dwarf ‘‘debilis is more variable in size than mut. semzalia,
but averages about half as high asthelatter. Its fruits and flowers
are somewhat reduced, but by no means proportionally to the
plant. The leaves, on the contrary, are much more reduced than
those of mut. semzalia’’.
Stout: THE ORIGIN oF DwaRF PLANTS 44]
Bartlett has not discussed these in terms of characters that
have been gained, lost, or modified, but does point out that the
appearance of these two dwarf forms as sister mutants in the rather
large number of individuals “bears a certain degree of resemblance
to Mendelian segregation.” The case is especially interesting in
affording a record of the simultaneous origin of hereditary varia-
tions from a single parentage giving dwarf forms of markedly
different characteristics.
Furthermore, the facts of variation resulting from hybridization
involving dwarf forms indicate a diversity of behavior quite in line
with the above-mentioned facts. Mendel considered that the
dwarf garden pea differed from the tall in a single character pair
and that he obtained neither intermediates nor a greater range of
variation in the F; generation. Darbishire (1911) considers that
this is the case and that the dwarf garden pea differs from the tall
variety only in respect to two types of internodes, long and short.
Punnett (1911, p. 35) applying the conception of presence and
absence very generally to peas states ‘‘all peas are dwarf, the tall
pea is a dwarf plus a unit factor.” This generalization is not borne
out by the data. Lock (1904) in progeny of crosses between a tall
and a semi-dwarf (‘‘Satisfaction”’ and native No. 2) found in the F,
great variation in habit ranging from very robust to very feeble
dwarf. Although he considers that there must be some sort of
Mendelian segregation if the internodes are solely considered, he
admits that ‘‘this cross seems to afford an example of remarkable
intensification of both the allelomorphic characters of the same
pair, namely tallness and dwarfness, the former in the F; and both
in the F, and later generations” (p. 414). Keeble and Pellew
(1910) in crossing two well-known staple semi-dwarfs varieties of
the garden pea obtained in the F; a wide variation, giving both
dwarf and tall as well as intermediates. They conclude that length
of internode and width of stem are both involved and in their
interpretation dwarfs lack both factors, semi-dwarfs lack one and
talls possess both. Height in sweet peas is also of complex charac-
ter, as breeding studies of dwarf, semi-dwarf and tall varieties have
shown. Crosses with bush and ‘‘Cupid” give an F; that is tall and
an F, with variation involving plants that include tall, bush and
442 Stout: THE ORIGIN OF DWARF PLANTS
two types of ‘‘Cupid’’. Bateson attempts to explain this case as he
does all cases of increased variation in terms of presence or absence
of factors, giving different interactions in different combinations.
The point is clear that greater variability does develop and that not
only is height of sweet peas evidently a compound character, but
that the assumed factors take on different values. For example,
the factor T (‘‘tallness’’) when combined with P (erect: branch-
ing) gives the bush variety. The two sets of allelomorphs assumed
in this case are not independent units in expression. It may be
emphasized that the original rigid unit-factor hypothesis is being
given up in favor of the admission, as we see in thiscase, ofa marked
modifiability of the effect of the assumed factors. The integrity
of the assumed units in these cases can only be maintained by
assuming a permanence in them, which is entirely at variance with.
the visible results which they are assumed to produce.
The behavior of the dwarf characteristics of Oenothera nanella
in hybridization is of special interest in this connection. The
group of characters appearing in Oe. nanella go together in crosses
with Oe. Lamarckiana, not asa unit giving alternative expression
but in the splitting that develops in the F, generation giving the
parent forms as twin hybrids. De Vries assumes (1913) that the
alta pangen is inactive in Oe. nanella and labile in Oe. Lamarckiana.
In this case inactive X labile gives both inactive and labile.
When Oe. rubrinervis is crossed with Oe. nanella splitting also
occurs in the F; giving Oe. Lamarckiana and a new type called
Oe. subrobusta. The alta pangen in Oe. rubrinervis is assumed to
be active, so in this case active X inactive = labile and inactive +
active. The Oe. subrobusta form splits in the next generation,
giving dwarf forms which, however, are evidently not Oe. nanella.
None of the types derived are like either of the two parent forms.
Again when Oe. muricata is crossed with Oe. nanella the F; is of
two new types called Oe. laeta and Oe. velutina. Oe. velutina splits
up in the F; giving a dwarf Oe. murinella that is not like Oe. nanella.
None of the types are like the parent forms. The assumed condi-
tion of the Oe. alta pangen in this case is active X inactive = labile,
active and inactive.
The appearance of twin hybrids not like the parents and of sec-
Stout: THE ORIGIN OF DwarF PLANTS 443
ond generation dwarfs not like Oe. nanella is also the rule in crosses
between Oe. nanella and Oe. Hookeri, Oe. Cockerelli, Oe. cruciata
and also Oe. biennis when the latter is the seed parent.
When Oe. nanella is fertilized by pollen from Oe. biennis the F,
is composed largely of dwarf plants. In the earlier crosses De
Vries reports a few tall plants and many dwarfs (1913, p. 241) but
from a cross in 1907 he obtained only dwarfs which, however, he
classed in two types, Oe. semialia and Oe. debilis, each of which
was different from the Oe. nanella parent. In this case the con-
dition of the alta pangen in the hybrids is assumed to be inactive X
labile, which is the same condition assumed for Oe. nanella X Oe.
Lamarckiana.
De Vries finds in such behavior evidence of the production of
new species in groups by hybridization. He assumes that a few
pangens are involved and that those exist in few conditions. The
integrity of such units can only be assumed by calling in such all-
sufficient properties as lability and inactivity which admit of very
sporadic behavior.
It is generally assumed that a new elementary species arises
suddenly without transitional forms. This is not the case with the
dwarf type of Hibiscus here described. The plant O No. 1 from
which all dwarf plants thus far obtained have arisen, possessed a
few of the characteristics of the dwarf. The next generation gave
dwarf, intermediates and robust plants in quite irregular numbers
in the two series grown. Bartlett (1915) describes two dwarf
types arising in a progeny of Oe. Reynoldsii, one of which (semialia)
is intermediate in general stature and has leaves less reduced than
in the extreme dwarf form (dedilis). Variations such as these are
not like ordinary fluctuations in giving a frequency distribution
agreeing with Quetelet’s law and they do not conform to the
Mendelian ratios of segregation, although as Bartlett points out,
there is much in the behavior that suggests the segregation assumed
by Mendelian interpretation. They are irregular and sporadic
variations involving different degrees and intensity of change. The
most marked of these involve changes affecting the character of
several organs,
Further evidence regarding irregular expression of characters is
444 Strout: THE ORIGIN OF DWARF PLANTS
seen in the well-known cases cited by De Vries of non-isolable races,
eversporting varieties, and inconstant species, indicative of irregular
and sporadic inheritance and expression of ever present tendencies
which in these cases rigid selection fails to isolate.
That there are cases of variation that are cumulative is evident.
De Vries especially has given data on two such cases. Ina race of
Chrysanthemum segetum, having an average of twenty-one ray
florets in the terminal inflorescence in a crop of 1,500 plants one
plant was found with four lateral flower heads with twenty-two
ray florets. Seed from this plant gave a progeny of 423 plants
one of which had a terminal inflorescence of thirty-four rays. Seed
from this plant gave a mixed progeny with one plant having sixty-
six ligulate florets, three of which were among the tubular disk
flowers. Seed from this plant gave progeny with florets ranging
from thirty-three to one hundred and one with a few completely
double flowers.
In respect to these results De Vries says (1901, p. 526): ‘‘ Es wird
sich jedem Leser die Frage aufdrangen: ist dieser Uebergang ein
allmahlicher oder ein stossweiser gewesen? Mir scheint das
letztere der Fall zu sein, aber es hangt dabei viel ab von der
Bedeutung, welche man den Wortern giebt. Jedenfalls geschah
die Umwandlung nicht im Laufe der Jahrhunderte, wie es die
Selectionstheorie anzunehmen pflegt, nicht einmal brauchte es
dazu Jahrzehnte. Drei Jahre geniigten, und solches in einer
Cultur von nur wenigen Quadratmetern Umfang.”
De Vries considers that this result is due to the reappearance of
a latent character. It is not clear what the latent character is in
this case. The species already possessed ray flowers in the outer
circle of flowers. The development of a so-called double-flowered
race consisted of a change of tubular flowers to ray flowers. Fur-
thermore, as wholly double flowered plants are sterile, the double
flowered plants are obtained continually by planting seed from
plants not fully double flowered.
A similar case is seen in the development of Linaria vulgaris
beloria. Individuals of certain races produce rarely a single peloric
flower, an evidence De Vries considers of a semi-latent character
which seldom becomes active. Seed from two such plants gave
Stout: THE ORIGIN oF DwarF PLANTS 445
out of a generation of two thousand one hundred plants, twenty
plants having only peloric flowers. Progeny of these peloric plants
gave ninety per cent. true to the type. In both Chrysanthemum
segetum and Linaria vulgaris peloria self sterility made it difficult to
grow pure line progenies. De Vries points out (1901, p. 564) that
while he considers these as cases of mutation they are of a type
quite different from that in Oenothera. In regard to the origin of
such variations that are hereditary the evidence is not at all con-
clusive that slight variations may not be inherited even in a cumul-.
ative manner.
That this is the case is further suggested by the difficulty of
drawing a definite line of distinction between species, varieties and
races. De Vries, who has written most fully on this point from
the standpoint of genetics, states that the best examples of varieties
are those showing latency of a single character which may be just
as constant as species (1901, p. 119); again he treats O. nanella
(1901, p. 256) as a variety simply because somewhat similar dwarf
types recur in a great number of species. He also gives the general
view that varieties usually differ in one character (1901, p. 363);
he states in another connection that the chief difference between
improved races and species, even the smallest of elementary species,
is the instability of the former and the stability of the latter (1901,
p. 84); but we may note that inconstant species do occur (1901,
p. 270).
It is clear that some cases of sporadic variation reproduce more
true to type than others. The so-called law of mutation that
‘‘neue elementare Arten sind meist vdllig constant, vom ersten
Augenblicke ihrer Entstehung an’’ (1900, p. 175) is itself a qualified
statement. The difficulty of assigning a definite heredity in terms
of unit characters or unit factors to “‘newly born’’ characteristics
which appear in mutations has become apparent. This is especi-
ally recognized in part by the view of Gates that mutation is a
phenomenon of variability which is quite distinct from heredity.
Thus far progeny has been grown from only one of the dwarf
plants of Hibiscus oculiroseus. One capsule of seed gave forty-six
plants, all but one of which were dwarf, and another capsule gave
twenty-seven dwarf, one robust and seven intermediates as judged
446 Stout: THE ORIGIN OF DWARF PLANTS
by the growth made the first year. The type appears to be only
slightly inconstant. Further breeding tests will determine if the
type becomes more constant by selection and if differences exist
between the progenies of different dwarf plants.
There has been much discussion recently regarding the possibil-
ity of the association of hybridization with mutation especially in
the Oenotheras. The rather well-known history of Hibiscus
oculiroseus and its dwarf mutant is of significance in this respect.
The wild form of H. oculiroseus was originally found in a region in
which the species H. Moscheutos exhibits much polymorphism,
involving problems which the writer now has under investigation.
Its affiliations with this species are so evident that for some time it
was known as a variety. The several differentiating characters
possessed by H. oculiroseus have already been noted and while
a few variations have been observed in my pedigreed cultures the
species breeds remarkably true totype. In fact the only variations
that have appeared have been among the progeny of the plant
giving dwarf plants, and none of the variations suggest that this
particular plant is a hybrid at least of the usual type. As far as
now known, H. oculiroseus has a limited distribution in nature and
since it is closely associated with H. Moscheutos it may well be
that it has been derived from this species.
Mr. George William Bassett, owner of the William F. Bassett
nurseries, writes in 1915: ‘‘We have never, to my recollection,
observed any dwarf tendency in Hibiscus ‘Crimson Eye.’ Nor
have we had any occasion to throw any out for any cause.” It
does not appear that dwarf forms have appeared in the cultivation
of the species. Mr. Norman Taylor reports to the writer that he
has observed in Long Island colonies of dwarf plants of the pink-
flowered form of H. Moscheutos.
The evidence indicates that the dwarf form is of spontaneous
origin. There is no series of characteristics belonging to either
H. Moscheutos or the parent stock of H. oculiroseus that can be
considered as combining to produce the dwarf, an interpretation
given to the origin of Oe. gigas by Heribert-Nilsson (1912). The
immediate parent (O No. 1) of the dwarf plants possessed in slight
degree the characteristics of crinkled leaves and shortened inter-
Stout: THE ORIGIN OF DwarF PLANTS 447
nodes which later appeared more intensified in the dwarf plants.
This, coupled with the variability of the progeny, might be consid-
ered by Mendelians to indicate that the parent plant was a half-
mutant. While it is no doubt true that such cases do occur (a
most notable case is that of Oe. semigigas), it is evident that the
conception has been applied to many cases of irregular inheritance
and sporadic variation and even to instances of cumulative
variation. |
A most important type of discontinuous variation is that of
simultaneous variation in a group of characters, well illustrated by
the dwarf Hibiscus. Not only is a number of characters modified
but the habit of profuse branching from the base appears as a char-
acter quite new to the parent species. Such phenomena are not
well explained on any conception of continuous unit characters.
Bateson does not admit that such simultaneous variation can
occur as a sporadic variation. He does not believe in the mutation
and inheritance of group-characters as described by De Vries and
already mentioned with respect to characters of Oe. nanella. He
prefers to think of the phenomena as due to a recombination of
factors. His strongest evidence for this view is the apparent
marked reversions that appear in certain crosses. If, however,
recombinations can give new groups or develop characteristics
new to the particular race and species, his analysis is of doubtful
validity. :
To assume that a variation such as the dwarf Hibiscus is due
to a single ciange in a single hereditary unit is to assign to the
units different values and to admit of interaction between units.
On the other hand to assume that there are hereditary factors that
are themselves stable but that can interact upon each other in
various ways is to assign different values to the supplementary or
codrdinating units and to the various interactions between them.
This makes the comprehensive description of the processes of
heredity in terms of units other than characters of doubtful valid-
i
It is quite clear that hereditary variations giving dwarf forms
of various degrees of intensity and extensity do arise, If in all
cases single characters, factors or other hereditary units are con-
448 Stout: THE ORIGIN OF DWARF PLANTS
cerned the evidence is clear that they possess different potencies
and belong to quite different categories, or else that they undergo
quite different sorts of changes not only in different species but in
the same species, variety, strain or even pure line.
The accumulation of evidence from all lines of plant breeding
shows that sporadic and irregular expression and inheritance of
characters are frequent and are widely distributed among plants,
and that ordinary stable characters and combinations become split
up and modified in processes of both mutation and hybridization,
giving variability not conforming to the usual laws of fluctuating
variability.
SUMMARY
A dwarf form of Hibiscus oculiroseus has appeared in a pedi-
greed culture as a sporadic variation. It differs from the robust
form in possessing a smaller stature, shorter internodes, smaller
leaves, many crinkled leaves and in the development of lateral
branches from the base of the main stem.
Plants intermediate between the dwarf and the robust forms
appear. These possess one or more of the characters of the dwarf
type in some degree of development.
All the dwarf plants thus far obtained are the progeny of a
single plant (O No. 1). No dwarf plants appeared among the 103
plants grown as progeny of four sister plants of plant No. 1.
The parent plant of this dwarf (O No. 1) possessed already in
slight degree the characteristics of crinkled leaves and shortened
internodes.
The dwarf plants appeared in varying numbers along with
robust and intermediate types. One series (Series No. I) was
composed of one dwarf, eleven robust and three intermediate plants;
another series (Series VI) was composed of forty-five dwarf plants.
There is a strong tendency for the dwarf form to breed true.
In a total of eighty-one plants grown from seed of a dwarf there
were seventy-two dwarf plants, eight classed as intermediates, and
one that was robust (Series VII and VIII).
It is difficult to describe the dwarf type in terms of characters
that have been lost or gained. The smaller and crinkled leaves and
the shortened internodes are evidences of reduced or arrested
Stout: THE ORIGIN oF Dwarr PLANTS 449
growth. In the marked development of branches from the base
there is increased growth or at least development of buds that
usually remain dormant.
There is no series of characters of either H. oculiroseus or H.
Moscheutos that can be considered as combining in hybridization
to give the dwarf.
The simultaneous appearance of variations involving modifi-
cations of groups of characters and of intermediates of various
kinds exhibit sporadic variations of various degrees of intensity
quite in line with the general evidence of the sporadic nature and
wide range of such variations.
NEW York BoTANICAL GARDEN
BIBLIOGRAPHY
Bartlett, H. H.
1915. Mutation en masse. Am. Nat. 49: 129-139.
Bateson, William.
1913. Mendel’s Principles of Heredity. Third impression. Cam-
bridge.
Britton, N. L. .
1903. The rose mallows. Jour. N. Y. Bot. Garden 4: 219-220.
Cook, O. F.
1915. Brachysm, a hereditary deformity of cotton and other
plants. Jour. Agric. Research 3: 387-400. pl. 53-62.
Darbishire, A. D.
1911. Breeding and the Mendelian discovery. London.
Heribert-Nilsson, N.
1912. Die Variabilitat der Oenothera Lamarckiana und das Prob-
em der Mutation. Zeitschrift fiir Ind. Abst. Vererb. 8:
89-231.
Keeble, F., & Pellew, C.
1910. The mode of inheritance of stature and of time of flowering
in peas (Pisum sativum). Jour. of Genetics 1: 47-56.
Lock, R. H.
1904. Studies in plant breeding in the tropics. Ann. Roy. Bot.
Gardens. Peradenya. 2: 299-414.
Nash, G. V.
1909. The aquatic garden. Jour. N. Y. Bot. Garden 10: 205-208.
Punnett, R. C.
1911. Mendelism. New York.
450 Stout: THE ORIGIN OF DWARF PLANTS
Vries, Hugo de.
1901. Die Mutationstheorie. ' Vol. I. Leipzig.
1904. Species and varieties. Their Origin by Mutation. Chicago.
1909. The mutation theory. Vol. I. Translation by Farmer (J.
B.) and Darbishire (A. D.) Chicago.
1913. Gruppenweise Artbildung unter spezieller Beriicksichti-
gung der Gattung Oenothera. Berlin.
Explanation of plates 26-27
PLATE 26.
To the left, typical plant of the dwarf form of Hibiscus oculiroseus (Plant No.
14 of Series VIII). To right typical plant of the robust form (Plant No. 23 of
Series IX). Photograph taken September 13,1914. Both plants are seven months
old
PLATE 27.
View in garden. In center with marker in the midst is a series (Series VIII)
of dwarf plants. Plant indicated at ais an intermediate. At the right is a series of
hybrids between H. oculiroseus (Plant O No. 2) and the pink-flowered type of H+
pink-flowered forms of H. Moscheutos. The three series are of the same age and
illustrate the difference between the dwarf and the robust forms.
A study of pasture trees and shrubbery
ErRNEsT L. ScoTT
(WITH THIRTEEN TEXT FIGURES)
Some years ago my attention was called to the peculiarly
regular form of the thorn trees in the pastures of northeastern
Ohio. These peculiar shapes are taken by the trees in response to
the constant pruning to which they
are subjected by the cattle. A very
short study revealed the fact that
other species are as characteristic-
ally affected as the thorns; this is
especially true of wild apples, oaks
and elms. Further, the behavior of
each of the different types has feat-
ures peculiar toitself. Many kinds
of trees are not able to withstand
the constant browsing to which
they are subjected when growing in
closely cropped pastures and so
quickly disappear. On the other
hand several species are not mo-
lested at all by the cattle. The
“immunity ’ which is enjoyed by
most nut-bearing trees seems to
depend upon a taste or texture
which is repugnant to the cattle.
Whether or not this explanation fFic.1. A sugar maple found grow-
is of wider application I do not te hes pags a cheat os
know. years. The maximum height was 45
There is of course considerable cm. ‘ This tree appeared to be Senge
variation in the relative abundance ase oe ad cea Lean ae ae
of the different species in the dif- ._ spite af the cattle.
ferent pastures. But considering
452 Scott: A Stupy OF PASTURE TREES AND SHRUBBERY
the region as a whole the haws are the most conspicuous,
not only because their striking contours but because of their
number and size as well. Wild apples are frequently associated
with the haws and resemble them so closely in form that the two
species might easily be confused on superficial observation. Elms
are very common in some pastures, but owing to the small size
which they are able to attain they are inconspicuous, and would
Fic. 2. A haw in the second or spheroidal stage. This particular individual
1.5 m. high and 2 m. in the greatest diameter. Though its age was not ascer-
tained, other trees in the corresponding stage showed annual rings indicating an age
of 10 to 20 years.
often escape the notice of the casual observer. The oaks seem to
have partially solved the problems set for them by the cattle and
by reason of their number and size demand attention. The hick-
ories are, in general, avoided by the cattle and so are usually able to
reach maturity when once started—at least so far as the cattle are
Scott: A Stupy oF PASTURE TREES AND SHRUBBERY 453
concerned. However, when they are attacked they usually
succumb when the browsing is severe, but if the punishment is not
too great they respond in a manner similar to that to be described
for the oaks. Only two or three maples have been found which
bear evidence of close cropping, not enough from which to draw
conclusions (FiG. 1).
Since the thorns, wild apples, oaks and elms are the most
numerous and are able to withstand the severe browsing to which
they are subjected they are the most interesting types found. The
points which I have considered are the forms taken by the several
Fic. 3. The third stage, an erect cone, of the haw. This one was 1.65 m. high
and 1.8 m. in diameter at the base. The age might be anywhere from Io or 20 years
up.
species in response to browsing, and the means by which each
finally gets beyond the reach of the cattle and so becomes indepen-
dent.
The ability to withstand constant browsing is only one factor
in determining the number of individuals of a species found in a
454 Scott: A Stupy OF PASTURE TREES AND SHRUBBERY
given pasture. The quantity of seed, the suitability of seed bed
and soil, together with other conditions of growth, are quite as
important.-
The earliest form assumed by the haws as a direct result of the
browsing is that of an inverted cone. This particular shape seems
to result from the fact that when the tip of a branch is clipped off
three or four buds situated short distances below the wound usually
start to grow. These buds give rise to branches which elongate at
about equal rates, each spreading somewhat from the old axis.
Since the cattle trim all the axes which reach the surface and since
the branches have a general tendency to grow upwards the plant
assumes the form of an inverted cone. Gradually the lower
branches are forced downward by the constantly increasing number
of new axes formed so that a spheroidal shape may be assumed as
shown in Fic. 2. When the lower branches are forced parallel to
the ground the tree takes the form of an erect cone (FIG. 3). In
grazing the cattle rarely bite the new growth back quite to the
dead tips of the older branches, so that there is a gradual increase
in size. Theconical shape is maintained until the treeis of sucha
diameter that the center or apex is no longer reached by the cattle.
The shoots which are now put out at the apex are free to grow
without molestation. The exact duration of the cone stage
depends upon the severity of the punishment to which the trees
are subjected. In some pastures where the grazing is not severe
there may be few or nocones found. In others which are pastured
very closely practically all of the haws and apples present are in
this condition or show distinct evidence of having passed through
it. Under such conditions the plant may be compelled to remain
in the cone stage for many years.
When the cattle are no longer able to reach the apex of the cone
one or more axes quickly spring out so that we find a closely crop-
ped cone with a loose clump of shoots projecting at its apex. This
stage is shown in Fic. 4. In spite of the constant cropping of all
new growth within the reach of the cattle, this protected clump
rapidly increases both in height and diameter. But since above
the reach of the cattle it is free to expand and an hour-glass shape
soon results (FIG. 5). Treesin various stages are shown in FIG. 6.
Scott: A Stupy oF PAstuRE TREES AND SHRUBBERY 455
Gradually the free portion of the tree extends beyond the
lower branches. The lower portions are now shaded and finally
are removed by self pruning until there results the usual rounded
Fic. 4 FIG. 5.
Fic. 4. This haw had become of so great a diameter that the cattle could no
longer reach its apex and so the shoots from this region were able to grow unmolested.
his one was 3 m. high and of the same diameter at the bas
Fic. 5. The hour-glass form which is assumed by he: kiwe after the apex has
been able to grow for a few years without pruning.
head supported by a single trunk or perhaps by a group of two or
three coordinate trunks. However the evidence of the struggle
by which the tree has attained its adult shape is seldom quite lost.
. 6. A common sight in pastures whose owner depends upon the cattle
ae pte the axe.
456 Scorr: A Stupy oF PASTURE TREES AND SHRUBBERY
While the forms taken by the haws are perhaps more striking
they are no more characteristic than those assumed by the oaks.
Here as a rule only a single axis is developed, from the sides of
which short branches are put out. Usually the topmost pair are
longer than any of the others or if the browsing is particularly
severe there may be a swelling at the top from which several
branches arise (Fic. 7). Here the
presence of dead stubs, indicating
the axes which have been killed by
the cattle is a striking feature.
The head shown in Fic. 8 is typical
of what is usually seen.
The reason for the difference
in habit between the oaks and the
haws lies, I believe, in the fact that
the oaks throw almost their whole
strength of growth into a single
bud so that when uninterrupted
growth has been permitted by the
cattle, even for a short time, some
one axis will be seen to be much
longer than any of the others.
While, as has been shown, the
growth of the haws is more diffuse,
the number of axes increasing with
Fic. 7. An oak which represents
the Sorin typically assumed by this
tree when browsed. This one was 12 the removal of each tip and all
years of age and had reached a height elongating at an approximately
of 1.35 m. and a diameter of 0.7 m- equal rate. As amatter of fact it
There were seven annual rings just be- 2
ieee Gah appears that in general the oaks,
when able to reach an independent.
stage at all, do so in fewer years than the haws. On the other
hand the haws are able to stand the more severe punishment.
At times, when the terminal bud is greatly interfered with it
may not be possible to identify the dominant axis. Evidence of
the general tendency is, however, usually furnished by a study of
the growth of successive years. Coupled with the emphasized
growth of a single bud is the habit of putting out but few branches
in response to the removal of the terminal portion of an axis.
Scott: A Stupy or PAstuRE TREES AND SHRUBBERY 457
The gnarled and bent forms which result from severe browsing
may be converted into straight and erect trunks by a stronger
growth over all concave surfaces than is found on the convex
exposures. This together with the tendency of the growing points
to assume the erect position is sufficient to conceal even such a
head as the one shown in Fic. 8. During the process of straighten-
ing many of the dead stubs decay and are broken off so that they
are overgrown. F1G. 9g shows a longitudinal section through a
head similar to that shown in Fic. 7, made after this had long
Fic. 8. Fic. 9.
Fic. 8. When the oaks are severely browsed they are frequently unable to
maintain a successful growing axis until after many trials. The result is a globular
head, from which project many dead and dying spikelike branches. These later
drop off or are overgrown as shown in Fic. 9. About one third natural size.
Fic. 9. A longitudinal section through a head such as is shown in Fic. 8. This
has long since ceased to be the point of conflict and the bases of the stubs are being
overgrown and the stem straightened as described in the text. About one fourth
ural size.
since ceased to be the point of contest and after the main axis
had crowded all the others out of the field. The same means of
458 Scott: A Stupy OF PASTURE TREES AND SHRUBBERY
straightening is used by the other forms studied as well as by those
oaks in which the grazing has been less severe. In FIG. 10 is
shown a longitudinal section through the axis of an oak which
has been subjected to severe browsing, but in which no head had
been formed.
In walking through the pastures in which elms are common
it is noted that almost all of the trees are 30-80 cm. in height.
Fic. to, Another instance of straightening, but in this case the browsing has
not been so severe and there was no head formed. About one fourth natural size.
This is in many cases due to the fact that these trees com-
monly grow in the midst of sedges and reach about the height
of the sedges before they are noticed by the cattle. They usually
reach this height in two or three years and after this are not per-
mitted to increase. Finally the upper portion of the stem dies
from the continued cropping and a new shoot appears lower down
on the axis or even from the roots and, under the protection of the
dead or dying top, soon reaches the height of the latter when it
in turn is subjected to severe browsing. There first results a form
somewhat resembling that assumed by the oaks, a single stem
with a number of side branches, Fic. 11. Finally, if the plant has
sufficient vigor, a form is assumed resembling the first, or inverted
cone stage, of the haws, Fic. 12. This results from branching and
Scott: A Stupy OF PASTURE TREES AND SHRUBBERY 459
development of shoots low down on the axis. An intermediate
stage is shown in Fic. 13. While this shape appears much later
in the elms, it is more accentuated and persistent than in the
haws. This is because the branches of the elm have a more
marked upward tendency and are not so readily induced to
hug the ground as are the branches of the apples and haws.
This upward tendency of the branches, together with the fact
Fig, £1. FiG, 12.
Fic. This elm was in an early gat of the response to browsing. In spite of
its age a i years, its height was only 61 c
Fic. 12. Anelm that had been ied é o grow only 32cm. in 17 years. The
close branching and abundance of dead twigs indicate that it is well along in the in-
verted cone or second stage.
that an axis dies after certain amount of pruning, precludes the
possibility of the formation of the spheroidal or of the erect conical
form. ;
The elms are not in general able to reach the independent stage
unaided. This is doubtless because they have the habits of growth
of the haw but not the haw’s vitality and ability to stand pruning.
Though the elms are not able to reach independence unaided it
must not be inferred that there are no elms which have obtained the
independent stage in the pastures. An elm reaching up through
460 Scotr: A Stupy, oF PASTURE TREES AND SHRUBBERY
a clump of wild roses, a haw or a spice bush or even from the center
of a group of less fortunate elms is a common sight.
In one plot of about 1.5 hectares sixty elms were counted, all of
which were unprotected. Of these 90 per cent., or fifty-four, were
less than 80 cm. in height and none were beyond the reach of the
cattle. The tallest was 1.5m. Inanother plot 50 m. square, I2I
elms were found, seventeen of which were either protected at the
Fic. 13. Anelmatast int liate bet th h in Fics. 11 and 12.
But since it was growing in the open, i.e., was unprotected by sedges, it was only 20
cm. in height though it was 15 years of age.
time or had been at a time which permitted them to escape from the
cattle. Of the 104 which were unprotected only two exceeded
I m. in height and none were beyond the reach of the cattle.
80 per cent. of all the elms in this plot were below 1 m. in height
and 23 per cent. enjoyed protection. In the same plot fifty oaks -
were found with only 56 per cent. below I m. and 12 per cent.
unprotected.
SUMMARY AND CONCLUSIONS
1. The haws, apples, oaks, and elms are able to withstand
constant browsing by cattle for a prolonged period of time.
2. In general other trees in pastures, as the ash, are killed with-
in a few years; or they are avoided by the cattle, as the hickories
and other nut-bearing trees.
Scott: A StupY OF PASTURE TREES AND SHRUBBERY 461
3. When the tree is able to withstand the cropping for a long
period it has good chances of reaching the natural size of the
species. This is attained in different ways by the different species:
(a) The oaks throw the stress of growth into a single series
of buds and so prolong the main axis at the expense of the lateral
branches that the tip of this axis may be lifted beyond the reach of
the cattle in comparatively few years.
(b) The haws and apples put out a large number of approx-
imately equal branches forming a close network about the center
of the tree, which is finally so far removed from the attacks of the
cattle that it may grow unmolested. This is a much slower method
than the one adopted by the oaks but seems to be somewhat surer
as judged by the number of successful individuals.
(c) Theelms have given up the attempt to reach independence
without the help of some other plant but because of their per-
sistence many of them are in time given the necessary protection.
4. While thorns are undoubtedly a factor in the escape of
some species from the cattle their importance may easily be over-
estimated; the apples are apparently quite as successful as are the
haws but are not provided with this armament.
* * *
Kerner has described similar observations on the oak and
beech but especially on the larch, as browsed by goats. Doubt-
less many others have observed related phenomena in pastural
regions.
New York City.
Dae
a
ih
nee
wie
Pat
oi
:
Notes on Rosaceae—X
P. A. RYDBERG
Ruspus Hysrips
Hybrids are rather common in the genus Rubus, especially
among the blackberries. Numerous hybrids have been admitted
in Europe. The only really good attempt made to segregate
hybrid forms in America, was that by Mr. Bicknell.* Mr. Bicknell
studied these forms in the field and in most cases found the
supposed parents growing in the immediate vicinity. I also have
studied blackberry hybrids two summers in the Adirondacks, the
Catskills, the mountains of southwestern Virginia, western North
Carolina, and eastern Tennessee, and around Ottawa, Canada.
It is a pity that it never occurred to Mr. Blanchard, who did so
much valuable work on the blackberries, to explain the numerous
forms as, partly at least, due to hybridity. It would not have
been very hard to determine whether or not many of his new species
probably had arisen through hybridizing. Mr. Bicknell, without
having seen many of Mr. Blanchard’s species in the field, reduced
nearly all of them into hybrids or into synonyms of older species.
I think that his interpretation was correct in the majority of
cases, but that he went too far in his reduction in some others. It
is risky to make pronunciations without seeing the species in the
field, and my course in the North American Flora was perhaps not
so wise. It may have been better to leave many of the supposed
species as such than to pronounce them as hybrids, without any
better proofs than the finding of intermediate and intergrading
characters. No definite conclusion can be had, however, with-
out experimental work in crossing. The only alternative course to
pursue would have been to admit all the proposed species as such
and to describe twice as many more, most of them without any
definite distinctive characters. In such a case, a key would have
been impossible to make.
Re
* Bull. Torrey Club 38: 103-133. I9II.
463
464 RYDBERG: NOTES ON ROSACEAE
The following ninety-three tentative hybrids have been re-
corded. Their essential characters have been pointed out in the
North American Flora, but in that work no specimens are cited,
except the types of new species and the first records of new hybrids.
It is, however, valuable to students of the brambles to know what
specimens the monographer regarded’as such hybrids. I have,
therefore, prepared the following list of specimens.
Rubus acaulis X stellatus.
ALASKA: 1885, Huff; Copper Center, Heidman 88.
Rubus acaulis X pubescens. Rubus propinquus Richardson is
regarded as a synonym of this.
MACKENZIE: Hay River, July 1, 1903, Preble & Cary 25;
wooded country, Richardson.
Rubus arcticus * pubescens. .
QuEBEC: Mt. Albert, Gaspé County, 1906, Fernald & Collins
6234; Table-top Mountain, Fernald & Collins 617.
Rubus arcticus X saxatilis (R. castoreus Frees). No locality
has been recorded on this side of the Atlantic, but it may be expect-
ed in northeastern Canada.
Rubus idaeus X spectabilis. This is unknown except as a
garden plant, and it is natural that it should be so, as Rubus idaeus
and R. spectabilis have no common territory where both grow wild.
Indigenous hybrids of R. spectabilis and species related to R. idaeus
have been collected in Alaska, viz., the two following:
Rubus spectabilis X viburnifolius..
ALASKA: Lake Clark, 1902, Gorman.
Rubus spectabilis X subarcticus.
ALASKA: Camp 53, 1899, Shrader.
Rubus occidentalis X rosaefolius. Known only as a garden
hybrid.
Rubus idaeus X ursinus. Only known asa spontaneous garden
hybrid, which is under cultivation as the Logan Berry.
Rubus idaeus X phoenicolasius. R. Paxii Focke. Only ite
as a garden hybrid.
* In answer to a letter to Dr. Fernald vtigmenting that the plant collected by him
and Mr. Collins was a hybrid, I received the following reply: “I have no doubt that
the plants you refer to are hybrids of Rubus deseo Seger pero: Such specimens
C ae ie ee oe * 1 + t Ge * wT q et erties v4 ahs) dor
wherever R. arcticus occurs.”
RYDBERG: NOTES ON ROSACEAE 465
Rubus occidentalis X strigosus (R. neglectus Peck; Melano-
batus neglectus and M. michiganus Greene). Thisisa rather com-
mon hybrid and it would be superfluous to cite specimens.
Sometimes it resembles more one parent, sometimes the other.
The original R. neglectus is most like R. occidentalis, while Melano-
batus michiganus approaches R. strigosus. |
Rubus occidentalis * procumbens. The very interesting plant,
which I referred to this hybrid, was collected by Commons nearly
forty years ago. Commons recognized it as a hybrid of R.
occidentalis but made no suggestion of the other parent. This is
evidently a member of the Eubatus section, apparently to be sought
among the dewberries, and most likely R. procumbens. Mr.
Commons remarks: “I tried to perpetuate this by sowing the
seeds at various times, but not one came up.”
Rubus leucodermis * melanolasius.
BriTIsH COLUMBIA: near international boundary between
Kettle and Columbia River, J. M. Macoun 64002; Chilliwack.
Valley, J. M. Macoun 34805; also 7035 and 69970 from British
Columbia.
Montana: Belton, 1903, Umbach 698; McDonald’s Lake,
1909, M. E. Jones.
OrEGON: Eastern Oregon, 1897, Cusick 1729.
WASHINGTON: 1897, Horner 163.
Rubus macropetalus X viburnifolius. No other specimen but
the original one, Mr. & Mrs. C. Van Brunt 121, from Glacier,
British Columbia, has been seen of this hybrid.
Rubus idaeus X strigosus. This is not uncommon in cultivation,
but no record of it in a wild state has been made.
Rubus argutus X cuneifolius. This was first recognized as a
hybrid by Mr. Bicknell. The same form had also been collected
by Britton & Hollick, who did not venture to name it. Their
specimens were determined as R. Andrewsianus by Mr. Blanchard,
but the dense grayish pubescence of the under leaves is not found
in that species. All the specimens seen are from the sandy coast
strip.
NEw JERSEY: Adanue Highlands, 1906, Hollick & Bride
Toms River, 1900, Bicknell go.
466 RYDBERG: NOTES ON ROSACEAE
Rubus cunetfolius * floridus. What I regard as this hybrid is
one much resembling the last hybrid but a weaker plant with more
curved prickles. Judging from the analogy, it is not improbable
that this plant is such a hybrid, as it differs from R. argutus X cunet-
folius in the same characters as does R. floridus from R. argutus,
and both R. floridus and R. cuneifolius are found in the district
where the supposed hybrid has been found. Mr. Ashe distributed
it mixed with R. cuneifolius.
VIRGINIA: Ocean View, Norfolk, 1898, Kearney 1212.
NortH CAROLINA: Chapel Hill, Ashe, mixed with R. cuneifolius.
Rubus cunetfolius X lucidus. Mr. Nash collected specimens
near Eustis, Florida, which resemble R. cuneifolius in many
respects, but the leaves are larger, more acute and more coarsely
serrate. He determined them doubtfully as R. cuneifolius. Simil-
ar specimens were also collected by Chapman, in Duval County,
Florida, together with R. cuneifolius. He also has collected R.
lucidus in the same county.
FLoripa: Duval County, Chapman 2832, mostly; Eustis,
1894, Nash 7o.
Rubus cuneifolius X trivialis. Somewhat similar specimens
were collected by Coker at Hartville, South Carolina, but in these
the plant is prostrate, the flowers mostly solitary, and in habit it
resembles more R. trivialis. These specimens I regard as R.
cuneifolius X trivialis.
Rubus canadensis X sativus. The specimens which I regard
as this hybrid were originally labeled R. canadensis, but Mr.
Blanchard has afterwards written on the sheets ‘A var. or form
of R. can. W. H. B.” It is evidently not pure R. canadensis for
the inflorescence is glandular and the leaflets of the new shoots are
broadly cordate and abruptly acuminate as in R. sativus.
VERMONT: Windham, June 6, 1903, Blanchard 265.
Rubus nigricans X sativus. 1 have no positive evidence that
R. frondisentis Blanchard is such a hybrid, but the plant combines
characters of the two species, and is found where the ranges of the
two species overlap. It is also a very local plant and has not been
collected except at the original station.
Rubus argutus X nigrobaccus. This hybrid was at recog-
RYDBERG: NOTES ON ROSACEAE 467
nized by Mr. Bicknell, but he used the name R. allegheniensis X
argutus, following Blanchard and Fernald in interpreting R.
allegheniensis. The plant is not uncommon and the following
specimens are referred here.
New York: Lynbrook, Long Island, 1901, Bicknell 36;
Elizabethtown, Essex County, 1900, Macy; Bronx Park, 1896,
Nash; Highland Falls, 1910, Mearns 5o.
MASSACHUSETTS: Quaise, Nantucket Island, 1910, Bicknell 33;
Shawaukemmo Spring, Nantucket Island, Bicknell 35; Canton,
near Boston, 1906, Blanchard 99, set 9.
VIRGINIA: Rosslyn, 1897, Kearney.
CONNECTICUT: Southington, 1904, Andrews 3.
PENNSYLVANIA: Tannersville, 1901, Britton.
NEw Jersey: Clifton, 1890, Nash goo; Paramus Swamp,
Bergen County, Nash 947; also Mackenzie 2086.
Rubus frondosus X nigrobaccus. This was also first recog-
nized as a hybrid by Bicknell, although he included in it specimens
which I do not think belong to it but rather to R. nigrobaccus X
recurvans. It was not at all unnatural that he should do so,
as he did not recognize R. recurvans as a distinct species.
MASSACHUSETTS: Shawaukemmo Spring, Nantucket Island,
1908, Bicknell 20.
NEw York: Jamaica, Long Island, 1901, Bicknell 25; Port
Washington, Bicknell 21, 23; Franklin Square, Bicknell 109.
PENNSYLVANIA: north of Philadelphia, 1906, Blanchard 100,
Set 6; Lancaster County, 1894, Ely.
NEW JERSEY: Succasunna, 1909, Mackenzie 4123.
Rubus canadensis X nigrobaccus. This hybrid was first recog-
nized by myself when I studied the blackberries in the Catskills,
1906, at Tannersville, where it grew together with R. nigrobaccus
and R. canadensis. I also saw it in the Adirondacks and in south-
western Virginia.
New York: Tannersville, 1906, Rydberg 7974; Little Moose
Lake, 1906, Rydberg 7820a; Stanford, 1909, Taylor 712; Tanners-
ville, 1892, Vail; North Harpersfield, Topping 132.
MASSACHUSETTS: Tyringham, Berkshire County, 1897, Vail.
VERMONT: Grafton, 1902, Blanchard 177; Simonsville, Blanchard
183; Rutland, 1902, Eggleston 2884.
468 RYDBERG: NOTES ON ROSACEAE
VirRGINIA: White Top Mountain, 1908, Rydberg 8104; be-
tween Marion and White Top Mountains, Rydberg 8138.
NortH CAroLina: Graham County, 1897, Biltmore Herbarium
1307¢.
New Brunswick: Grand Falls, Macoun 21181.
Rubus amicalis X nigrobaccus. Comparing the specimens re-
ferred here with R. canadensis X nigrobaccus, I came to the con-
clusion that this must be a hybrid of the species mentioned.
They were collected by Arthur at Isle au. Haut, Maine.
Rubus nigrobaccus X Randvi. This was first recognized by
myself while collecting in the Adirondacks. Both parents grew
in the vicinity. Subsequently I thought it was the same as R.
frondisentis Blanchard, but I have found it isnot. None except
the original specimens have been seen.
Rubus nigrobaccus X pergratus. The following specimens I
think belong to this hybrid. They were collected near the type
locality of R. amicalis X nigrobaccus. Rubus pergratus is also
common in the region.
MatneE: Isle au Haut, 1909, Arthur 55, a
Rubus heterophyllus * nigrobaccus. 1 have no evidence that
the specimens referred here belong to a hybrid except that the plant
combines the habit and glandular pubescence of R. nigrobaccus
with the leaf cut of R. heterophyllus. The original specimens are
the only ones seen. .
Rubus nigricans X nigrobaccus. This I collected at Kings-
mere Lake, Quebec, where both R. nigricans and R. nigrobaccus
were growing. I have no doubt but my specimens 'were of hybrid
origin. I cannot distinguish them from specimens of R. glandi-
caulis distributed by Mr. Blanchard. R. biformispinus Blan-
chard I think is also a hybrid of the same two species, though
approaching R. nigricans more than R. nigrobaccus. The following
specimens belong here:
QuEBEC: Kingsmere Lake, 1906, Rydberg 70940.
NEw Brunswick: Frederickstown, Blanchard 606.
MaInE: Yarmouth, 1909, Blanchard 507; Brownville, Blanchard
508; Bar Harbor, Blanchard 509; Calais, Blanchard 510; Isle au
Haut, 1909, Arthur 52; Kennebunk, 1905, gietigee 74. Sets 7.
2; Kennebunkport, 1905, Blanchard 79.
RYDBERG: NOTES ON ROSACEAE 469
Nova Scotia: Granville, 1909, Blanchard 718, 727; Yar-
mouth, Blanchard 728.
Rubus nigrobaccus X vermontanus. If R. glandicaulis is a
hybrid between R. nigrobaccus and R. nigricans, R. frondisentis
is very likely a hybrid between the former and R. vermontanus.
VERMONT: Townshead, 1903 and 1904, Blanchard 243, 244;
Huntington, 1903, Blanchard 257. :
Rubus hispidus X nigrobaccus. R. permixtus Blanchard stands
in the same relation to R. hispidus as R. glandicaulis does to
R. nigricans, I therefore do not hesitate in regarding it as a hybrid
between R. nigrobaccus and that species.
VERMONT: Westminster 1902 and 1903, Blanchard 225, 226,
227, 229; Putney, 1902, Blanchard 228; Wee tannnis: 1903, Eg-
gleston 3245.
New HaMPsHIRE: Walpole, 1903, Blanchard 233.
Nova Scotia: Macoun 81438, 81439.
Rubus argutus X frondosus. This was first recognized by Mr.
Bicknell.
NEw York: Rockville Center, Long Island, 1906, Bicknell 38;
between Yonkers and Mount Vernon, 1902, Burnham.
CONNECTICUT: Southington, 1905, Blanchard 9g, set I; 1904,
Andrews 27.
Rubus argutus X recurvans. This was included in R. argutus
< frondosus by Bicknell, who did not regard R. frondosus and
R. recurvans distinct.
MASSACHUSETTS: Quaise, Nantucket Island, 1909, Bicknell
34-
New York: Cedarhurst, Long Island, 1910, Bicknell 37.
Rubus argutus X Baileyanus. This was also segregated out
by Mr. Bicknell.
MASSACHUSETTS: below the “Cliff,” Nantucket Island, 1910,
Bicknell 4o.
New York: East Rockaway Bog, Long Island, 1908, Bicknell
41; also at Hawlett, Bicknell 42.
Rubus argutus X Enslenii. This was also recognized by Mr.
Bicknell, although he did not publish the same. His specimens
closely match the type of R. ostryifolius Rydb., which I now do
not hesitate in regarding as this hybrid.
470 RYDBERG: NOTES ON ROSACEAE
New York: Hempstead Plains, Long Island, 1910, Bicknell
65; Lynbrook, Bicknell 62, 64, 66.
New Jersey: Highlands, Britton; Ogdensburg, 1910, Macken-
zie 4625; South Amboy, 1905, Mackenzie 13706.
VircIniA: Norfolk, 1898, Kearney 1365.
Rubus argutus X carpinifolius. Both Engelmann and Bailey
recognized this as a hybrid, regarding one of the parents as R.
trivialis from which R. carpinifolius was not separated at that
time. Bailey thought that the other parent was R. argutus.
Engelmann gave it as R. villosus, under which name both R.
argutus and R. nigrobaccus were then known.
Missouri: St. Louis, 1863, Engelmann; Cahokia, 1863,
Engelmann.
Rubus floridus X lucidus. In the description of R. persistens
Rydb. two different plants were included. Unfortunately, it
happened that the specimen which was assigned as the type evi-
dently belongs to a hybrid of R. floridus and a species then un-
known. R. persistens, therefore, must pass into synonymy.
Rubus carpinifolius X floridus. Dr. Engelmann regarded the
original specimen of this hybrid as a variety of R. trivialis. It is
more likely that it is a hybrid of R. carpinifolius, which at his
time was not distinguished from R. trivialis, and R. floridus, which
is also found in Texas.
Texas: Houston, 1842, Lindheimer.
Rubus floridus X frondosus. Some specimens collected at
Bluff Lake, Missouri, by Eggert probably belong to a hybrid
between R. floridus and R.frondosus. The only reason for believ-
ing so is that they combine characters of both and were found in
a region common to both. ©
Missourt: Bluff Lake, Eggert.
TENNESSEE: Knoxville, 1896, Ruth.
Rubus Enslenti X frondosus. This was first recognized by
Mr. Bicknell and well described by him.
New York: Hempstead Plains, Long Island, 1906, Bicknell,
48, 49, 70.
MassacuusEtTts: Nantucket Island, 1906, Bicknell 47.
Rubus frondosus X procumbens. I have no definite evidence
RYDBERG: NOTES ON ROSACEAE 471
that R. Rossbergianus is a hybrid of R. frondosus and R. procum-
bens, but the habit is intermediate between the two; the leaves
resemble in form those of R. frondosus, but are harsher; and the
stem is light colored, tough and prickly asin R. procumbens. Both
are found in the region.
CONNECTICUT: Southington, 1905, Blanchard 89, set 1-3;
Graniteville, Blanchard 89, set 3; Canton, Blanchard 89, set 5;
Southington, 1904, Andrews 21, 22, 23, 31, 33, 35-
NEw Jersey: Tuckerton, Mackenzie.
Rubus frondosus X hispidus. Also one of Mr. Bicknell’s
hybrids, collected only at the original station and vicinity, Bick-
nell 43, O69.
Rubus canadensis X recurvans. The specimens from which
R. recurvans subrecurvans was described approach R. canadensis
L. in so many respects, such as the form of the leaves, the inflores-
cence, and the weakly developed armature of the stems, that I
have regarded it as a hybrid with that species.
MaIneE: Kennebunkport, 1905, Blanchard 71; York Beach,
1909, Blanchard 516.
Rubus Brainerdi X vermontanus. The specimens referred to
this hybrid were collected at the type station of R. Brainerdi
and together with it. R. vermontanus is common in the region.
VERMONT: Weybridge, Brainerd; also 1902, Eggleston 2804.
Rubus amicalis X pergratus. No specimens except the origi-
nal ones have been seen.
Rubus argutus X canadensis. This was first recorded by
myself from southwestern Virginia, but similar specimens have
also been collected in the north.
VIRGINIA: road between Marion and White Top Mountain,
1908, Rydberg 8099; Chattam Hill Road, 1892, Small.
Nortu CAROLINA: Biltmore, 1909, Crafton & Eggleston, 4416.
New Jersey: Greenwood Lake, 1907, Mackenzie 2675;
Ogdensburg, 1910, Mackenzie 4625.
Rubus canadensis X Randii. This was first recorded y
myself from the Adirondacks, where both parents grow
New York: Little Moose Lake, 1906, Rydberg 7818; 7820;
Twin Lakes, Rydberg 7842; East Lake, Rydberg 7861.
472 RYDBERG: NOTES ON ROSACEAE
Nova Scotia: Pictou, 1906, C. B. Robinson 528.
Rubus canadensis X pergratus. Also first recorded by myself
at Tannersville, New York, 7960, 7961.
Rubus Baileyanus X canadensis. The specimens originally
recorded as this hybrid were collected near an artificial lake or
reservoir above Sparta, New Jersey. R. canadensis was growing
in the neighborhood but R. Baileyanus was not collected there.
The habit of the plant and the poor fruiting indicate a hybrid of
R. canadensis and, probably, R. Baileyanus. ‘To this belong:
New Jersey: Morris Lake, Sparta, 1906, Rydberg 79092;
Montague Township, Sussex County, 1909, Nash; Midvale,
1908, Rydberg So16.
Nova Scotia: Macoun 81433, 81431.
Rubus elegantulus X nigrobaccus. Rubus flavinanus Blanchard
evidently represents this hybrid. The plant is decidedly inter-
mediate between the two. None except Blanchard’s specimens
have been seen.
Rubus Baileyanus X nigrobaccus. This was first recognized
as a hybrid by Mr. Bicknell, who, however, did not publish the
same. The following specimens belong here:
New York: Smithtown, Long Island, 1910, Bicknell 17, 24.
PENNSYLVANIA: Presque Isle, Erie, Otto Kuntze 2671.
New JrErsEy: Midvale, 1908, Rydberg 8017.
Rubus Baileyanus X procumbens. This has been known for
fifteen years or more and is even in cultivation, but it was first re-
garded as a variety of R. procumbens, and later as a distinct
species under the name R. roribaccus. Mr. Bicknell was the first
one to recognize it as a hybrid. Some of his specimens match
very closely the original ones distributed by Professor Bailey.
Massacuusetts: Nantucket Island, 1910, Bicknell 8.
New York: Cedarhurst, Long Island, 1910, Bicknell 6;
Aqueduct, 1905, Bicknell 7.
New JERsEY: Ledgewood, 1909, eshsaaiicd 4127; Succasunna,
Mackenzie 4120.
ViRGINIA: Glade Springs, 1908, Rydberg 8148; Fairfax, 1906,
Blanchard 101, set 3.
PENNSYLVANIA: Willow Grove Park, 1906, ‘Blanchard rol,
set 2; Lancaster, Blanchard Tor, set I.
RYDBERG: NOTES ON ROSACEAE 473
CONNECTICUT: Southington, 1906, Blanchard ror, set 51%.
Missour!: Little Blue Tank, 1897, Mackenzie 210.
Iowa: Papoose Creek, 1894, Mackenzie 681.
Rubus Baileyanus X frondosus. First segregated by Mr.
Bicknell.
MASSACHUSETTS: Quaise, Marcy’s Pond, below the “ Cliff,’’ and
Squam, Nantucket Island, 1910, Bicknell 85, 86, 88, 87, 84.
New Jersey: Budd’s Lake, 1905, Eggleston 0; Mt. Arlington,
1905, Mackenzie 1402; Stanhope, 2085; Berkeley Heights, 2047.
Rubus Baileyanus X hispidus. This was at first included in
R. hispidus X procumbens by Bicknell. He, however, determined
his Long Island material as R. Baileyanus X hispidus.
New York: Aqueduct, Long Island, 1905, Bicknell 73.
New Jersey: near Musconetcony River, 1906, Mackenzie
2107.
Massacuusetts: Reed Pond, Nantucket Island, 1910, Bick-
nell 75; Reading, 1897, Kingman.
Rubus Baileyanus X carpinifolius. Another plant collected
by Eggert at Bluff Lake, Missouri, in 1878 must be, on account
of the combination of characters, a hybrid between R. Baileyanus
and R. carpinifolius, which both grow in the region.
Rubus arenicola X hispidus. The original specimens are the
only ones seen of this hybrid. ,
Rubus nigrobaccus X plicatifolius. The specimen referred
to this hybrid (Blanchard 516, sheet 1), was labeled by Mr. Blan-
chard R. recurvans var. subrecurvans, which variety I regard as a
hybrid between R. recurvans and R. canadensis; but this number
evidently does not belong there. The glandular pubescence of the
inflorescence and the cut of the leaves in the specimens mentioned
indicating R. nigrobaccus blood. The pubescence and the plica-
tion of the leaves suggest R. plicatifolius. —
Rubus nigricans X plicatifolius. 1 | regard R. multiformis
Blanchard as a hybrid of R. nigricans and R. plicatifolius.
Nova Scotia: Yarmouth, 1909, Blanchard 724; -Annapolis
Blanchard 726; Kentville, Blanchard 725. . .
Matne: Bar Harbor, 1909, Blanchard 512; Kennebunkport,
1905, Blanchard 80. | ;
474 RYDBERG: NOTES ON ROSACEAE
QuEBEC: Macoun 67904.
Rubus canadensis X plicatifolius. The almost unarmed stem
of Rubus multiformis delicatior Blanchard indicates a hybrid of
R. canadensis. The plicate leaves suggest R. plicatifolius.
MAINE: Kennebunkport, 1905, Blanchard 81.
Nova Scotia: Halifax, Macoun 81426.
Rubus hispidus X plicatifolius. Rubus semierectus Blanchard
I take as a hybrid. The small flowers, bristly stem and broad
leaflets suggest R. hispidus, the pubescent strongly veined and
subplicate leaflets, R. plicatifolius.
MAINE: Kennebunk, 1905, Blanchard 67.
NEw York: vicinity of Tannersville, 1891, Vail.
Nova Scotia: Macoun 5792, 19076.
Rubus heterophyllus * recurvans. No other specimens than
those cited in the North American Flora, viz., Blanchard 256,
from Huntington Hills, Massachusetts, have been seen which
may be referred to this hybrid. !
Rubus argutus X flagellaris. This as well as all the other
hybrids of R. flagellaris were discovered by Mr. Bicknell. For
specimens and notes see his paper in the Bull. Torrey Club 38:
103-133, 1911. The other hybrids are Rubus flagellaris < frond-
osus, R. flagellaris X procumbens, R. Baileyanus X flagellaris, R.
flagellaris X hispidus and R. flagellaris * Enslenit.
Rubus nigrobaccus X procumbens. This was first recognized
by Mr. mo but under the name R. allegheniensis * procum-
bens.
New York: Merrick, Long Island, 1910, Bicknell 30; Hew-
lett, Bicknell 28; Lynbrook, Bicknell 26.
MASSACHUSETTS: Wauwinet, Nantucket diet 1910, Bick-
nell 27.
Rubus argutus X OE This was ee first recognized
as a hybrid by Bicknell. Rubus multispinus Blanchard matches
Bicknell’s specimens very closely.
New York: Rockville Center, Long Island, 1909, Bicknell;
Hewlett, Bicknell 68; Lawrence, Bicknell 67.
MassAcausetts: Nantucket Island, 1899, Bicknell 29; 1910,
Bicknell, 82, 83; Reheboth, 1906, Blanchard 96, set 2; jisians om
1906, Blanchard 96, set 3.
RYDBERG: NOTES ON ROSACEAE 475
RHODE ISLAND: Rocky Point, 1906, Blanchard 96, set I.
Rubus procumbens X recurvans. Many specimens from Con-
necticut resemble R. procumbens closely, but are more pubescent
and with the leaves of the floral branches more incised. These
I have regarded as hybrids with R. recurvans. They are:
CONNECTICUT: Southington, 1904, Andrews 29, 28, 41, OI,
37,34, 30; Plainville, 1903, Blanchard 222; Winsted Blanchard, 259.
MASSACHUSETTS: Southwick, 1903, Blanchard 253.
Rubus pergratus X procumbens. Rubus recurvicaulis Blanchard
seems to me a hybrid of R. pergratus and R. procumbens, com-
bining characters of both. I have seen no specimens except
those distributed by Mr. Blanchard.
Rubus canadensis X procumbens. Rubus recurvicaulis inar-
matus stands to R. canadensis as R. recurvicaulis does to R.
pergratus, and I believe also this to be a hybrid.
MAINE: Kennebunk, 1905, Blanchard 67, 69.
Nova ScotTta: North Sidney, 1907, C. B. Robinson 616.
Rubus hispidus X procumbens. This was first recognized by
Mr. Bicknell. The following specimens belong here:
New York: Cypress Hill, 1905, Bicknell 74.
VERMONT: Londonderry, 1903, Blanchard, 246; Newfane,
1902, Blanchard 186.
Nova Scotia: Boylston, 1890, Chas. Hamilton, 190985.
New Jersey: Mt. Arlington, 1905, Mackenzie 1404; also
1904, Mackenzie 903.
Rubus Baileyanus X Enslenii. Also one of Mr. Bicknell’s
hybrids.
MassacuusEtts: Nantucket Island, 1908, Bicknell 52, 54, 55.
ConnEcticuT: New Haven, 1895, Helen Ingersoll.
New York: Woodmere, 1904, Bicknell 56.
District oF CoLuMBIA: Riverdale, 1901, Steele.
VIRGINIA: Marion, N. L. & E. G. Britton & A. M. Vail.
Rubus Enslenii X procumbens,
Rubus Baileyanus X Enslenit,
Rubus Enslenii X flagellaris, and
Rubus Enslenii X hispidus. Of these four bybeids, I have
seen none but Mr. Bicknell’s specimens. See Bull. Torrey Club
_ 38: 130-131. I9II.
476 RypBEerG: NOTES ON ROSACEAE
Rubus argutus X nigricans. 1 cannot distinguish R. adscend-
ens Blanchard from Mr. Bicknell’s hybrid. The following speci-
mens belong here:
New York: Smithtown, Long Island, 1808, Bicknell 89.
NEw. JERSEY: Allamuchy Mountains, 1906, Mackenzie 2108,
2300; Succasunna, 1909, Mackenzie 4153.
- ConnectIcuT: Southington, 1906, Blanchard go; 1903, Blanch-
ard 262,224; Londonderry, Blanchard 246; sess 53,57, 50.
MASSACHUSETTS: Essex County, Oakes.
Rubus frondosus X nigricans. From this I cannot separate
Rubus abbrevians Blanchard. The following specimens belong
here:
MAssACHUSETTs: Long Pond and Trot’s Swamp, Nantucket
Island, 1910, Bicknell 31, 32.
CONNECTICUT: Southington, 1906, Blanchard 95.
VERMONT: Windham, 1904, Blanchard 57.
PENNSYLVANIA: Germantown, Meehan 8670.
Rubus nigricans X recurvans. R. semisetosus Blanchard I
regard as a hybrid between R. nigricans and R. recurvans. The
following specimens belong here:
CONNECTICUT: Southington, 1903, Blanchard 201, 202, 225;
Londonderry, Blanchard 247; Plainville, 1906, Blanchard g1, set I;
also 1903, 261, 203.
RHODE ISLAND: pe Point, 1906, Blaschard Ql, set 3.
Nova Scotia: Sable Island, Macoun aEtOSi 21183 (these
specimens are doubtful).
Rubus amicalis X nigricans. Rubus tardatus Blanchard is prob-
ably also a hybrid, the parents being R. amicalis and R. N1igricans.
Mr. Blanchard’s specimens are the only ones seen.
Rubus nigricans X pergratus. In my opinion R. peculiaris
Blanchard is also a hybrid, viz., of R. nigricans and R. pergratus.
Besides Mr. Blanchard’s specimens the following belong here:
QUEBEC: vicinity of weiss: Falls, 1905, Macoun 67904,
67905.
Rubus canadensis X nigricans. Rubus junceus Blanchard is
probably also a hybrid. Besides the specimens distributed by
Mr. Blanchard, the following are referred to it.
RYDBERG: NOTES ON ROSACEAE 477
Ontario: Cranberry Lake, Algonquin Park, 1900, Macoun
24690a.
MAINE: Sangerville, 1895, Fernald 244.
Rubus nigricans X Randii. Only the specimens of the
original collection have been recorded.
Rubus Baileyanus X nigricans. This was first recognized
by Mr. Bicknell. C. C. Curtis’s specimens have much narrower
leaflets and are doubtfully referred to it.
~New York: Hempstead Plains, 1910, Bicknell 15; between
Yonkers and Mount Vernon, 1903, C. C. Curtis; Van Courtlandt
Park, 1896, Bicknell.
Connecticut: Plainville, 1904, Andrews 26; Southington,
‘ 1903, Blanchard 237 (determined as R. semisetosus).
Rubus flagellaris X nigricans. This was collected by Mr.
Bicknell, who included it in R. nigricans. The following speci-
mens so named I refer to the hybrid instead.
Massacuusetts: Little Neck, Millbrook Swamp and Long
Pond, Nantucket Island, 1910, Bicknell X, XX, A
Rubus nigricans X procumbens. This was also recognized as
a hybrid although he did not describe the same. The following
specimens belong to it:
New York: East Hempstead, Long Island, I910, Bicknell 16;
Jamaica, 1906, Bicknell 18.
Connecticut: Plainville, 1904, Andrews 55; Southington,
Andrews 54, 57, 58, 59, 02: :
Vermont: Londonderry, 1903, Blanchard 245 (referred doubt-
fully to R. vermontanus).
Nova Scotia: Macoun 81425; Sable Island, Macoun 21193.
Rubus nigricans X vermontanus. Rubus vermontanus viridifolius
has glandular hispid inflorescence and I therefore believe it
contains some R. nigricans blood in it. I therefore regarded it
asa hybrid. Many specimens referred to R. setosus by Blanchard
belong here: :
VERMONT: Marlboro, 1902, Blanchard 191; Woodstock, 1910,
Britton; Stockbridge, 1897, Eggleston; West Windham, 1904,
Blanchard 59, set 1; Stratton, 59, Blanchard, set 2; Rutland, 1899,
Eggleston 1228.
478 RypBERG: NOTES ON ROSACEAE
New York: Tannersville, 1906, Rydberg 7054.
New HampsuireE: Alstead, 1903, Blanchard 236, 234, 235.
MaInE: Biddeford, 1905, Blanchard 86.
QuEBEc: East Templeton, 1903, Macoun 59963.
Rubus hispidus X nigricans. Mr. Bicknell found this very
variable, sometimes approaching one parent, sometimes the other.
My observations agree very well with his. Often the plant
resembles a robust R. hispidus, but with the leaflets of the new
shoots having a tendency often to be five instead of three. It
is then R. hispidus major Blanchard. Sometimes it resembles
more R. nigricans in habit. If the leaflets of the new shoots
happen to be only three, it is R. trifrons Blanchard. The following
represent R. hispidus major:
ConneEcTIcUT: Southington, 1904, Andrews 60; Botsford,
1897, Eames.
MAINE: Kennebunkport, 1905, Blanchard 84; Kennebunk,
Blanchard 85.
Nova Scotia: Cape Breton, 1901, Howe & Lang 748; Pictou,
1907, C. B. Robinson 574, 404.
MASSACHUSETTS: 1879, Perkins.
PENNSYLVANIA: Pocono Plateau, 1893, Britton.
New York: Tannersville, 1908, Rydberg 8037.
Specimens resembling R. trifrons are the following:
VERMONT: West Wadsboro, 1904, Blanchard 55; Stratton,
1902, Blanchard 198.
NEw York: Tannersville, 1906, Rydberg 7057, 7054 a
sheet of this number Mr. Blanchard has determined as R.
pidus, the other as R. setosus. Both were from the same ee
The following are intermediate or cannot be referred to either
form:
VERMONT: North Windham, 1903, Blanchard 251 (named
R. setosus); Stratton, 1902, Blanchard 199.
New JERSEY: Musconetcong River, Mackenzie 2108, 2300.
NEw York: Tannersville, 1906, Rydberg 7056.
CONNECTICUT: Southington, 1904, Andrews 60.
Rubus hispidus X vermontanus. In my mind there is very
little doubt but R. jacens Blanchard represents this hybrid. A
‘RYDBERG: NOTES ON ROSACEAE 479
few specimens referred to R. trifrons by Blanchard apparently
belong here also. The following specimens have been recorded:
VERMONT: Townsend, 1903, Blanchard 241; Stratton, 1904,
Blanchard 55, set 2; South Londonderry, 1903, Blahchard 250.
New HampsuireE: Alstead, 1903, Blanchard 232, 231.
Rubus cubitans Blanchard seems to me to be a mixture.
The flowering branches and the new shoots were not col-
lected at the same time. Do they belong to the same plant?
The floral branches seem to me as typical R. hispidus, though
with more flowers than usual. The leaflets of the new shoots
have the texture of those of R. hispidus but the form of those of
R. vermontanus. They may very well represent a hybrid.
Rubus argutus X hispidus. This was first recognized by Mr.
Bicknell, who, however, did not characterize the same. Only
Bicknell’s specimens are known.
Rubus canadensis X hispidus. The only specimens seen,
which may be referred to this hybrid, are the original ones cited
in the North American Flora.
Nova Scotia: Le Have River, Macoun 81436a.
New York BoraNicaL GARDEN
INDEX TO AMERICAN BOTANICAL LITERATURE
1912-1915
The aim of this Index is to include all current botanical literature written by
Americans, published in gee or based upon American material ; the word Amer-
ica being used in the broadest sen
Reviews, and papers that natin exclusively to forestry, agriculture, horticulture,
_ manufactured products of vegetable origin, or laboratory methods are not include
attempt is made to index the literature of bacteriology. An occasional exception is
ais in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not mentioned unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, ted furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
subscriber must ike all cards published dialed the term of-his subscription, Corr
spondence relating to the card issue should be addressed to the Treasurer of the Torrey
Botanical Club,
Anastasia, G.E. Araldica nicotinae. Boll. Tecnico Coltiv. Tabac.
13: irate. pl.1-82. Autg14._ [lIllust.
A ix to this number containing 60 plates additional, variously labeled
A-Z, ies easel S 1914
Appel, O. Disease resistance in plants. Science II. 41: 773-782. 28
My I915.
Appel, O. The relations between scientific botany and phytopathology.
Ann. Missouri Bot. Gard. 2: 275-285. 17 My 1915.
Atkinson, G. F. Phylogeny and relationships in the Ascomycetes.
Ann. Missouri Bot. Gard. 2: 315-376. f. 1-10. 17 My 1915.
Bailey, L. H. The Standard Cyclopedia of Horticulture. 3: 1201-
1760. pl. 40-60 + f. 1471-2047. New York. 12 My 1915.
Belling, J. Inheritance of pod pubescence and partial sterility in
Stizolobium crosses. Rep. Florida Agr. Exp. Sta. 1914: Ixxxi-cv.
f. 1-9. Ap 1915.
Benedict, R. C., and others. Where Ophioglossum grows (A multiple
report). Am. Fern Jour. 5: 42-50. My 1915.
Bessey, C. E. The phylogenetic taxonomy of flowering plants. Ann.
Missouri Bot. Gard. 2: 109-164. f. 72. 17 My 1915.
Blake, S.F. An Aéripiex new to North America. Rhodora 17: 83-86.
f.t, 2. AaB Ap 1915.
481
‘
482 INDEX TO AMERICAN BOTANICAL LITERATURE
Britton, N. L. The vegetation of Mona Island. Ann. Missouri Bot.
Gard. 2: 33-58. pl.1, 2. 17 My 1915.
Includes Pedilanthus latifolius Millsp. & Britton, Tabebuia lucida, Riccia Brittonit
Howe, and Riccia violacea Howe, spp. nov.
Cameron, F. K. Potash from kelp. U.S. Dept. Agr, Soils Rep. 100:
1-122. pl. 1-40 +f.1 + maps A—G, 1-61. 10 Ap 1915.
Includes chapters by Crandall, Rigg, and Frye here indexed separately.
Chrysler, M. A. The medullary rays of Cedrus. Bot. Gaz. 59: 387- |
396. f.1-7. 15 My tIois.
Collins, F. S. Some algae from the Chincha Islands. Rhodora 17:
89-96. My I915.
Condit, I. J. The Loquat. Calif. Agr. Exp. Sta. Bull. 250: 251-284.
f. I-11. Mr 1915.
Conzatti, C. The botanical garden of Oaxaca. Ann. Missouri Bot.
Gard. 2: 165-174. pl.3 +f.1,2. 17 My 1915.
Coulter, J. M. The origin of monocotyledony. II. Monocotyledony
in grasses. Ann. Missouri Bot. Gard. 2: 175-183. f. 1-9. 17 My
1915.
Crandall, W.C. The kelp beds from Lower California to Puget Sound.
U.S. Dept. Agr. Soils Rep. 100: 33-49. 10 Ap I915.
Czapek, F. Recent investigations on the protoplasm of plant cells and
its colloidal properties. Ann. Missouri Bot. Gard. 2: 241-152.
17 My 1915.
Davis, W. T. Union of an oak and a birch. Proc. Staten Island
Assoc. Arts and Sci. 5: 10. pl. 1 +f.37. 24 Ap 1915.
Dorsey, M. J. Pollen development in Vitis with special reference to
sterility. Minnesota Agr. Exp. Sta. Bull. 144: 1-60. pl. 1-4. S 1914.
Dorsey, M. J. Pollen sterility in grapes. Jour. Heredity 6: 243-249.
f. 1-6. 25 My 1915.
Edson, H. A. Seedling diseases of sugar beets and their relation to
root-rot and crown-rot. Jour. Agr. Research 4: 135-168. pl. 16-20.
15 My 1915.
Eriksson, J. The control of plant diseases in Sweden. Bull. Foreign
Agr. Intelligence 5: 187-192. Mr 1915.
Evans, A.W. The genus Plagiochasma and its North American species.
Bull. Torrey Club 42: 259-308. f. 1-8. 22 My 1015.
Includes Plagiochasma Landii sp. nov.
Fairchild, D. Green leaf in a cherry blossom. Jour. Heredity 6: 262,
263. f. 7. 25 My 1015.
INDEX TO AMERICAN BOTANICAL LITERATURE 483
Farwell, O. A. Notes on the Michigan species of Polygonatum. Bull.
Torrey Club 42: 247-258. pl. r2-18. 22 My 1915.
Includes Polygonatum ellipticum sp. nov. and several new varieties and combina-
tions,
Fernald, M. L. Altitudinal limits in Connecticut, a correction. Rho-
dora 17: 88. 28 Ap I19gI5.
Fernald, M. L. Botrychium angustisegmentum (Pease and Moore), n.
comb. B. lanceolatum var. angustisegmentum Pease and Moore. Rho-
dora 17: 87, 88. 28 Ap. 1915.
Fernald, M. L. Two variations of Silene antirrhina. Rhodora 17:
96,97. My 1915.
Frye, T. C. The kelp beds of southeast Alaska. U.S. Dept. Agr.
Soils Rep. 100: 60-104. 10 Ap 1915.
Gable, C. H. The wild tomato. Jour. Heredity 6: 242. frontispiece.
25 My 1915. [Illust.]
Gates, R.R. On the origin and behavior of Oenothera rubricalyx. Jour.
Genetics 4: 353-360. Ap 1915.
Giddings, N. J., & Berg, A. Apple rust or cedar rust in West Virginia,
W. Virginia Agr. Exp. Sta. Circ. 15: 1-16. Mrig15._ [Illust.]
Goodspeed, T. H. Quantitative studies of inheritance in Nicotiana
hybrids. II. Univ. Calif. Publ. Bot. 5: 223-231. 21 Ap 1915.
Harms, H. Uber einige von P. Preuss gesammelte Arten der Gattung
Inga Scop. Repert. Sp. Nov. 13: 419,420. 31 D 1915.
Hieronymus, G. Eine neue Selaginella (S. Herteri) aus Uruguay.
Repert. Sp. Nov. 13: 421, 422. 31 Drigrq.
Hill, A. W. The history and functions of botanic gardens. Ann.
Missouri Bot. Gard. 2: 185-240. pl. 6-12 + f.1. 17 My 1915.
Hoehne, F. C. Plantae novae brasilianae. I. Repert. Sp. Nov. 13:
423-438. 31 D 1914.
(Ex. comm. Linh. Telegr. Estrat. de Matto Grosso ao Amazonas. Annexo no.
5. Botanica Parte I. Dec. 1910, 71 pp., 63 tab.’’)
Hollick, A. Union of an oak and a beech. Proc. Staten Island Assoc.
Arts and Set. §: 11, 12. $1.1 -- 7. 2. 24 Ap 1615.
Hopkins, L. S. An interesting specimen of the fragrant shield fern.
Am. Fern Jour. 5: 41, 42. pl. 2. My 1915.
Horne, W. T. Oak fungus or Armillaria mellea in connection with
nursery stock. Monthly Bull. Calif. State Comm. Hort. 4: 179-184.
f- 31-33. Ap Igis.
Howe, I. A. New botanical finds for St. Johnsbury and vicinity.
Vermont Bot. and Bird Bull. 1: 33, 34. Ap 1915.
484 INDEX TO AMERICAN BOTANICAL LITERATURE
Hubbard, F. T. Mibora minima (L.) Desv. at Plymouth, Massachu-
setts. Rhodora 17: 97. My 1915.
Hubbard, F. T. A taxonomic study of Setaria italica and its immediate
allies. Am. Jour. Bot. 2: 169-198. 13 My 1915.
Jennings, O. E. Notes on the Pteridophytes of northwestern Ontario.
Am. Fern Jour. 5: 33-39. My 1915.
Kinman, C. F. Citrus fertilization experiments in Porto Rico. Porto
Rico Agr. Exp. Sta. Bull. 18: 5-33. pl. 1, 2. 14 My 1915.
Land, W. J. G. Microtechincal methods. Bot. Gaz. 59: 397-401.
15 My 1015.
Lipman, C. B., & Sharp, L. T. Effect of moisture content of a sandy
soil on its nitrogen fixing power. Bot. Gaz. 59: 402-406. 15 My
IQI5.
Loeb, J. The stimulation of growth. Science II. 41: 704-715. 14
My 1915.
Lorenz, A. Additions to the Vermont hepatic list for 1914. Vermont
Bot. and Bird Bull. 1: 28, 29. Ap 1915.
Lorenz, A. New Vermont Hepaticaein1913. Vermont Bot. and Bird
Bull. 1: 29, 30. Ap 1915.
Lutman, B. F. Plant quarantine laws. Vermont Bot. and Bird Bull.
I: 38,10... Ap: tors:
MacDougal, D. T. The experimental modification of germ-plasm.
Ann. Missouri Bot. Gard. 2: 253-274. f. 1-4. 17 My 1915.
Maxon, W. R. Note upon Polypodium subtile and a related species.
Am. Fern Jour. §: 50-52. My 1015.
Maxon, W. R. Report upon a collection of ferns from western South
America. es epee onpate meee 65: 1-12. 3 My I9Is.
Includes Polyp , C. incarum, Notholaena
Brackenridget, N. ‘meddisenits. and Dryopteris Rosei, spp. nov.
Meyer, R. Echinopsis cinnabarina Lab. Monats. Kakteenk. 25:
20-23. Mr 1915.
Meyer, R. LEchinopsis Meyeri Heese. Monats. Kakteenk. 24: 178,
179. 15 D 1914.
Meyer, R. Einiges iiber den Echinocactus Grusonii Hildm. Monats.
Kakteenk. 25: 15, 16. F 1915.
Miller, W. H. Tree fruits and flowers. Am. Forestry 21: 617-625.
My rog15._ [Illust.]
Newman, O.W. Russian thistle. Salsola kali L. var. tenuifolia G. W.
F. May. Monthly Bull. Calif. State Comm. Hort. 4: 193-196.
f. 34. Ap 1915.
INDEX TO AMERICAN BOTANICAL LITERATURE 485
Osterhout, W. J. V. Antagonism between acids and salts. Jour. Biol.
Chem. 19: 517-520. 1914.
Osterhout, W. J. V. The effect of acid on permeability. Jour. Biol.
Chem. 19: 493~501. f. 1-3%. 1914.
Osterhout, W. J. V. The effect of alkali on permeability. Jour. Biol.
Chem. 19: 335-343. f. I-4. 1914.
Oswald, L. W., & Boss. A, Minnesota weeds—II. Minnesota Agr.
Exp. Sta. Bull. 139: 1-47. f. 1-25. My 1914.
Pammel, L.H. Introduced plants of the Clear Creek Canon, Colorado.
Proc. Iowa Acad. Sci. 21: 119-121. pl. 15, 16. 1914.
Perry, F.E. The inheritance of size in tomatoes. Ohio Nat. 15: 473-
495. pl. 22-24 + f. 1, 2.. Ap 1915.
Petry, L. C. Branching in the Ophioglossaceae. Bot. Gaz. 59: 345-
365. pl. 20, 2 +f.2-6. 15 My 1915.
Pierce, R. G. The present status of the chestnut bark disease in
Massachusetts. Arborea 1: 8,9. My 10914.
Pool, V. W., & McKay, M. B. Phoma betae on the leaves of the sugar
beet. Jour. Agr. Research 4: 169-177. pl. 27. 15 My I915.
[Popenoe, P.] Development of a cherry. Jour. Heredity 6: 278, 279.
f. 8, 9. 25 My 1915.
Redles, G. Some experiences in fern-hunting near Philadelphia. Am.
Fern Jour. 5: 52-54. My 1915.
Reed, G. B. Evidence for the general distribution of oxidases in
plants. Bot. Gaz. 59: 407-409. 15 My 1915.
Reed, H. S., & Grissom, J. T. The development of alkalinity in
Glomerella cultures. Jour. Biol. Chem. 21: 159-163. My 1915.
Rigg, G. B. The kelp beds of Puget Sound. U.S. Dept. Agr. Soils
Rep. 100: 50-59. f. 2. 10 Ap 1915.
Rigg, G.B. The kelp beds of western Alaska. U.S. Dept. Agr. Soils
Rep. 100: 105-122. 10 Ap 1915. 7
Schlechter, R. Asclepiadaceae novae bolivienses Herzogianae. Repert.
Sp. Nov. 13: 438-443. 31 D 1914.
Includes eleven new species and one new genus (Corollonema).
Seely, H. M. Why does the gilliflower apple fail to protect its seed?
Vermont Bot. and Bird Club Bull. 1: 13-15. Ap 1915.
Setchell,W.A. The law of temperature connected with the distribution
“* of the marine algae. Ann. Missouri Bot. Gard. 2: 287-305. 17
My I915.
486 INDEX TO AMERICAN BOTANICAL LITERATURE
Shapovalov,"M. Effect of temperature on germination and growth of
the common potato-scab organism. Jour. Agr. Research 4: 129-
133. pl. 15 + f.1. 15 My 1915.
Shive, J. W. A three-salt nutrient solution for plants. Am. Jour. Bot
2: 157-160. 13 My 1915.
Skottsberg, C. Notes on Pacific coast algae. I. Pylaiella postelsiae
n. sp., a new type in the genus Pylaiella. Univ. Calif. Publ. Bot. 6:
153-164. pl. 17-19. 7 My 1915.
Smith, E. F. A conspectus of bacterial diseases of plants. Ann.
Missouri Bot. Gard. 2: 377-401. 17 My 1915.
Spaulding, P. Forest fungiof Bethel. Vermont Bot. and Bird Bull. 1:
24,95. Ap iors.
Spoehr, H. A. Variations in respiratory activity in relation to sunlight
Bot. Gaz. 59: 366-386. f. 1-10. 15 My 1915.
Stewart, F. C., & Sirrine, FA. The spindling-sprout disease of pota-
toes. N. Y. Agr. Exp. Sta. Bull. 399: 133-143. pl. 1-3. Mr 1915.
St. John, H. Elymus arenarius and its American representatives.
Rhodora 17: 98-103. My 1915.
Includes Elymus strigatus sp. nov.
St. John, H. Rumex persicarioides and its allies in North America.
Rhodora 17: 73-83. pl. 113. 28 Ap 1915.
Stone, G. E. The pin oak (Quercus palustris Muench.). Arborea 1:
28, 29. Au 1914. [Illust.]
Stuart, W. Potato breeding and selection. U.S. Dept. Agr. Bull. 195:
I-35. pl. 1-16. 20 My 1915.
Studhalter, R. A., & Heald, F. D. The persistence of viable pycno-
spores of the chestnut blight fungus on normal bark below lesions.
Am. Jour. Bot. 2: 162-168. 13 My 1915.
Thompson, W. P. Preliminary note on the morphology of Gnetum.
Am. Jour. Bot. 2: 161. 13 My 1015.
Townsend, C.O. Field studies of the crown-gall of sugar beets. U.S.
Dept. Agr. Bull. 203: 1-8. pl. z,2+f.1. 30 Ap 1915.
Urban, I. Sertum antillanum. I. Repert. Sp. Nov. 13: 444-459.
31 D 1914.
Includes new species in Piper (1), Phoradendrum (1), Coccoloba (4), Portulaca
(1), Magnolia (1), Phyllanthus (7), Croton (6), and Rhamnidium (x). :
Vaupel, F. Peruanische Kakteen. Monats. Kakteenk. 24: 154-157.
15 N 1914; 24: 161-177. 15 D 1914. [Illust.]
BuLL. TORREY CLUB VOLUME 42, PLATE 26
STOUT: HIBISCUS OCULIROSEUS
BULL. TORREY CLUB VOLUME 42, PLATE 27
STOUT: HIBISCUS OCULIROSEUS
Vol. 42 No. 9
BULLETIN
OF THE
TORREY BOTANICAL CLUB
SEPTEMBER, 1915
Studies of West Indian plants—VII
ee Sameer Lorp een
30. Slee GENUS SCLERIA Brerc. IN CUBA
The last previous treatment of the Cuban species is that of
Mr. CB. Clarke in Urban, Symb. Ant. 2: 8-169. 1900.
A. Hypogynium present.
. Hypogynium not covered with a white rough crust.
a. Margin of the hypogynium neither ciliate nor fim-
briate [see S. cubensis].
* Roots fibrous; rootstocks none or very shor
Achene reticulated or irregularly ida
plants slender. i. S. sclacee.
Achene smooth; plants stout. 2. S. lacus
** Perennials, with rootstocks. [Rootstock in S. -
setuloso-ciliata short, or perhaps none e.]
h
- Achene /
Ligule sre its margin scarious or callous.
Climbing; branched; pilose. 3. es SCANS.
Erect; ne —— : 4. S. catalinae.
’ Ligule sho: t di
Culms sens ‘eaves relatively enneth
oe red-purple; achene purple to
gichia e. 5. S. melaleuca.
Panicle brown-green; achene white.
Achene depressed-globose. 6. S. plerota.
Achene rt ick
Leaves 2-4 mm. wide. 7. S. Wrightiana.
Leaves 8-15 mm. wide. 8. S. setuloso-ciliata.
Culms stout; leaves very rough. 9g. S. scindens.
Achene verrucose, reticulated or papillose. :
Achene papillose. 10. S. ciliata.
[The BULLETIN for August (42: 429-486. pl. 26, 27) was issued September 22, 191 5]
487
488 BRITTON: STUDIES OF WEST INDIAN PLANTS
Achene verrucose or reticulated.
Cc.
porting 3 deeply 3-lobed tubercles. 11. S. Curtissii.
Achene_ verrucose-reticulated; hypogy-
nium 3-lobe
Glabrous or penel 12. S. stereorrhiza.
Leaves and sate detat pilose-
pubes 13. S. pilosissima.
b. Margin of the Rupa ciliate, ciliolate or
fimbriate [nearly or quite eciliate in S. cubensis].
Margin of the hypogynium ciliate or ciliolate.
Achene 2 mm. long or less. 14. S. microcarpa.
Achene 2.5-3 mm. lon
: tame black, persistent; achene ellip-
15. S. mitis.
divin pale, deciduous; achene sub-
globose.
Leaves about 3 mm. wide. 16. S. phylloptera.
Leaves 8-16 mm. wide.
argin Pus the hypogynium ciliolate
or eci 17. S. cubensis.
Margin oy ‘ae hypogynium densely
long-ciliate. 18. S. Grisebachit.
Margin of the ce eae: 3-lobed, the lobes
laciniate or fimbria 19. S. porphyrorrhiza.
2. Hypogynium covered wa a white rough crust. 20. S. havanensis.
B. Hypogynium none, or obsolete.
Annuals with fibrous roots; inflorescence glomerate-
spicate.
Awl
ch s 21. S. distans.
Achene verrucose, reticulated or cancellate
Bracts strongly ciliate; achene cancellate. 22. S. pinetorum.
Bracts glabrous; achene verrucose or verrucose-
culated.
Inflorescence simply glomerate-spicate. 23. S. verticillata.
florescence branched. 24. S. Liebmanni.
Perennial by rootstocks.
Inflorescence glomerate-spicate; achene smooth. 25. S. hirtella.
Inflorescence not glomerate-spicate.
icle loose. 26. S. lithosperma.
oo subcapitate.
mm. ein with 2 pits on each side
of the 3-angled base. 27. S. gracilis.
A 4mm, acs without pits. 28. S. Baldwinii.
I. SCLERIA SETACEA Poir. in Lam. Encycl. 7: 4. 1806
Scleria hemitaphra Steud. Syn. Pl. Cyp. 169. 1855.
(Scleria Torreyana Walp. Ann. 3: 696. ees
¢
BRITTON: STUDIES OF WEsT INDIAN PLANTS 489
Scleria dictyocarpa Griseb. Cat. Pl. Cub. 259. 1866.
Scleria debilis C. Wright; Sauvalle, Anales Acad. Habana 8: 154.
1872.
S. reticularis pubescens Britton, Ann. Lyc. N. Y. 3: 232. 1885,
Savannas and along streams, Pinar del Rio:—southern United
States; Porto Rico; tropical continental America.
2. SCLERIA LACusTRIS C. Wright; Sauvalle, Anales Acad. Habana
S: 152. “1872
Lagoons near Pinar del Rio. Recorded by Clarke from French
~ Guiana and from Java.
3. SCLERIA SECANS (L.) Urban, Symb. Ant. 2: 169. 1900
Schoenus secans L. Syst. ed. 10, 865. 1759.
Scleria reflexca HBK. Nov. Gen. 1: 232. 1815.
Mountain woodlands, Oriente :—Haiti to Martinique and Trini-
dad; Jamaica; tropical continental America.
My examination of the type specimen of Schoenus secans L.,
in the herbarium of the British Museum of Natural History,
some years ago, showed that it was the same as Scleria reflexa
HBK., and not the same as Scleria Flagellum-nigrorum Berg.
4. Scleria catalinae sp. nov.
Perennial by thick rootstocks; culm stout, sharply 3-angled,
roughish on the angles, glabrous, about 1.3 m. high. Leaves
glabrous, slightly roughish-margined, 3-5 dm. long, I~-2.3 cm.
wide, 3-nerved, attenuate-acuminate, the ligule triangular-ovate
to triangular-lanceolate, acute or acuminate, many-striate, nar-
rowly callous-margined, 2 cm. long or less; panicle about 5 dm.
long, of few, distant, slender branches, the staminate and pistillate
spikelets intermixed; staminate spikelets oblong, 3-4 mm. long;
achene ovoid, purplish, smooth, shining, acute, 2 mm. long, the
style-base persistent; hypogynium obconic, 1 mm. long, glabrous,
its base rounded, its 3 rounded lobes appressed.
Santa Catalina, Pinar del Rio (Van Hermann 3272).
By its very large ligule related to S. arundinacea Kunth, but
the achene and inflorescence are quite different, resembling those
of S. cubensis Boeckl.
490 BRITTON: STUDIES OF WEST INDIAN PLANTS
5. SCLERIA MELALEUCA Schl. & Cham. Linnaea 6: 29. 1831
Savannas and banks, Camaguey; Havana; Pinar del Rio:—
Jamaica; Porto Rico; St. Vincent to Trinidad; continental tropical
America. The West Indian specimens here referred are not very
different from the following species.
6. SCLERIA PTEROTA Presl in Oken, Isis 21: 268. 1828
Scleria pratensis Nees in Mart. FI. Bras. 2!: 179. 1843.
Scleria Ottonis Boeckl. Linnaea 38: 490. 1874.
Woodlands and banks, Santa Clara; Havana; Pinar del Rio;
Isle of Pines:—Haiti to St. Thomas and Barbadoes; Jamaica;
continental tropical America.
7. SCLERIA WRIGHTIANA Boeckl. Flora 64: 79. 1881
Scleria elaita C. Wright; Sauvalle, Anales Acad. Habana @: 153.
1872. Not Thwaites.
Pine-lands and savannas, Pinar del Rio and Isle of Pines.
Endemic.
8. SCLERIA SETULOSO-CILIATA Boeckl. Flora 65: 30. 1882 |
Wet situations, Matanzas; Havana; Isle of Pines:—Guatemala.
9. SCLERIA SCINDENS Nees, Linnaea 9: 303. 1834
Mountain woodlands, Oriente:—Tortola; St. Kitts to St.
Vincent.
10. SCLERIA CILIATA Michx. Fl. Bor. Am. 2: 167. 1803
‘Scleria Elliottii Chapm. F1.S. U.S. 531. 1860.
_ Barrens and pinelands, Santa Clara; Matanzas; Pinar del Rio;
Isle of Pines:—southeastern United States; Santo Domingo.
II. ScLERIA CURTISSII Britton; Small, Fl. S. E. U. S. 200, 1328.
1903
? Sclerta pauciflora shouts Clarke in Urban, Symb. Ant. 2: 143.
1900.
- Savannas, Pinar del Rio and Isle of Pines:—Florida. Referred
by Wright, by Clarke and formerly by me to S. pauciflora Muhl.
BRITTON: STUDIES OF WEsT INDIAN PLANTS 491
12. SCLERIA STEREORRHIZA C. Wright; Clarke in Urban, Symb.
Ant. 2: 147. 1900
Scleria bracteata Griseb. Cat. Pl. Cub. 249. 1866. Not Cav.
Scleria bracteata angustata Griseb. loc. cit. 1866.
Scleria stereorrhiza major Clarke in Urban, loc. cit. 1900.
Along streams and on hillsides, Santa Clara; Havana; Pinar
del Rio. Endemic.
13. Scleria pilosissima sp. nov.
Perennial by short, branched rootstocks; culms rather slender,
but stiff, sharply 3-angled, loosely pilose, simple, 3-4 dm. high.
Leaves roughish-margined, 1-2 dm. long, 2-4 mm. wide, acu-
minate, stiff, densely pilose, the midvein prominent beneath;
clusters of spikelets 1 or 2, pilose, 2-4 cm. long, the staminate and
pistillate spikelets intermixed; scales lanceolate, acute, pubescent;
achene subglobose, 2—-2.5 mm. in diameter, transversely verrucose,
pubescent, obtuse, abruptly apiculate; hypogynium with
rounded, appressed lobes.
Sunny hillsides among rocks, between Punta Gorda and Wood-
fred, Oriente, Cuba, 400-500 m. alt. (Shafer 3090).
Related to S. stereorrhiza C. Wright.
14. SCLERIA MICROCARPA Nees, Linnaea 9: 302. 1834
Scleria foliosa C. Wright; Sauvalle, Anales Acad. Habana 8: 154.
1872. Not A. Rich.
Scleria microcarpa foliosa Clarke in Urban, Symb. Ant. 2: 149.
1900.
River banks, Pinar del Rio and Isle of Pines:—Porto Rico;
Guadaloupe; Jamaica; Trinidad; continental tropical America.
15. SCLERIA MITIS Berg. Vet. Akad. Handl. Stockh. 26: 145.
1865
Scleria Eggersiana Boeckl. Cyp. Nov. 2: 41. 1890.
Pinal de Sta. Ana collected by Eggers, according to Clarke:—
Porto Rico; Guadeloupe; Martinique; St. Vincent; Trinidad:
continental tropical America. —
16. SCLERIA PHYLLOPTERA C. Wright; Griseb. Cat. Pl. Cub. 248.
1866
Scleria microcarpa angustifolia Boeckl. Flora 64: 79. 1881.
492 BRITTON: STUDIES OF WEST INDIAN PLANTS
Moist savannas and borders of lagoons, Santa Clara; Pinar del
Rio. Endemic.
17. SCLERIA CUBENSIS Boeckl. Cyp. Nov. 2: 42. 1890
Scleria microcarpa subeciliata Clarke in Urban, Symb. Ant. 2:
149. 1900
Banks and woodlands, tains of Oriente and Santa Clara:—
Hispaniola; Porto Rico; Jamaica.
18. SCLERIA GRISEBACHII Clarke in Urban, Symb. Ant. 2: 150.
1900
- Bogs, Santa Clara; Havana:—Hispaniola; Antigua; Dominica;
Martinique; Jamaica.
19. SCLERIA PORPHYRORRHIZA C. Wright; Sauvalle, Anales Acad.
Habana 8: 155. 1872
Pinelands, Pinar del Rio. Recorded by Clarke from Brazil.
20. Scleria havanensis sp. nov.
Rootstocks stout, horizontal, 3-4 cm. long, about 6 mm. thick.
Culms many, slender, trigonous, nearly smooth, about 5 dm.
long; leaves nearly smooth, narrow, I-1.5 mm. wide; inflorescence
a single small cluster of intermixed staminate and pistillate spike-
lets; achene white, ridged-reticulated, globose-oblong, rounded,
2 mm. long; hypogynium low, obscurely 3-lobed, covered with a
rough whitish crust.
Coca Hill, Campo Florida, province of Havana (Brother Leon
4731).
21. SCLERIA DISTANS Poir. in Lam. Encycl. 7: 4. 1806
Scleria tenella Griseb. Cat. Pl. Cub. 249, in part. 1866. Not
Kunth.
? S. hirtella pauciliata Britton, Ann. N. Y. Acad. Sci. 3: 236. 1885.
Cuba, collected by Wright:—Hispaniola; Porto Rico; St.
Thomas; Guadeloupe.
22. Scleria pinetorum sp. nov.
Annual, with em finely fibrous roots; culms slender, erect,
2-4 dm. high. es grass-like, pubescent with spreading hairs,
I-2 mm. wide; eieeare glomerate-spicate, a cm. long;
BRITTON: STUDIES OF WEsT INDIAN PLANTS 493
bracts linear-lanceolate, ciliate, 6 mm. long or less; achene
globular, white, deeply reticulated, about 1.5 mm. in diameter,
with a row of pits at its base, the short base 3-angled.
Pine-lands and savannas, Pinar del Rio. Type collected be-
tween Candelaria and Artemisa (Wilson 1724). |
Previously referred by me (Ann. N. Y. Acad. Sci. 3: 236) to
S. interrupta A. Rich., of Guiana, which now seems to me to be
distinct.
23. SCLERIA VERTICILLATA Muhl.; Willd. Sp. Pl. 4: 317. 1805
Pinelands, Pinar de! Rio; Isle of Pines:—eastern United States;
New Providence, Bahamas.
24. SCLERIA LIEBMANNI Steud. Syn. Pl. Cyp. 179. 1855
Scleria tenella Griseb. Cat. Pl. Cub. 249, in part. 1866. Not
Kunth.
Sclerta luzulaeformis C. Wright; Sauvalle, Anales Acad. Habana
8: 156. 1872.
Savannas near San Juan de Buena Vista, jurisdiction of
Bayamo, according to Sauvalle (Wright 3418, in part); Mexico
to Brazil.
25. SCLERIA HIRTELLA Sw. Prodr. 19. 1788
Scleria nutans Kunth, Enum. 2: 352. 1837.
Moist grounds, Oriente; Santa Clara; Pinar del Rio; Isle of
Pines,—southern United States; Haiti; Porto Rico; Jamaica;
Trinidad; continental tropical America; tropical Africa.
26. SCLERIA LITHOSPERMA (L.) Sw. Prodr. 18. 1788
Scirpus lithospermus L. Sp. Pl. 51. 1753-
Scleria filiformis Sw. Prodr. 19. 1788.
S. lithosperma filiformis Britton, Ann. N. Y. Acad. Sci. 3: 231.
188
Woodlands and thickets, all provinces:—Florida; Bahamas;
West Indies; tropical continental America; Old World tropics.
27. SCLERIA GRACILIs Ell. Bot. S. C. & Ga. 2: 571. 1824
Moist savannas, Pinar del Rio:—southeastern United States.
494 BRITTON: STUDIES OF WEST INDIAN PLANTS
28. SCLERIA BALDWINII (Torr.) Steud. Syn. Pl. Cyp. 175. 1855
Hypoporum Baldwini Torr. Ann. Lyc. N. Y. 3: 382. 1836.
Border of lagoon, Laguna Santa Maria, Pinar del Rio:—south-
eastern United States.
FURTHER STUDIES OF BADIERA DC.
Since the publication of my former paper on this genus (Bull.
Torrey Club 37: 360-363. 1910), additional specimens which
have come to hand afford the following notes and descriptions.
1. BADIERA PENAEA (L.) DC.
Professor Urban has informed me that my opinion that this
is identical with Badiera domingensis (Jacq.) DC. is correct,
although he has distributed Santo Domingo specimens under the
name B. domingensis.
1a. Badiera portoricensis sp. nov.
A tree up to 6 m. high, the trunk sometimes 1.5 m. in diameter,
the slender branches ascending, the young twigs appressed-
pubescent. Leaves subcoriaceous, obovate, 3 cm. long or less,
7-15 mm. wide, bright green and shining above, paler and rather
dull green beneath, both surfaces with short, scattered hairs when
young, becoming glabrate above, the midvein slightly impressed
above, rather prominent beneath, the lateral veins few, distant,
obscure, the apex rounded or obtuse, the base narrowed or cuneate,
the pubescent petioles 1-2.5 mm. long; inflorescence few—several-
flowered, 4-6 mm. long; bracts ovate or ovate-lanceolate, acute,
pubescent, 0.5 mm. long; pedicels appressed-pubescent, 1.5-2-5
mm. long; sepals broadly ovate, obtuse, ciliolate, the larger about
I mm. long; petals white; keel appressed-pubescent, 2 mm. long,
narrowed below; fruit about 6 mm. wide, about 5 mm. high,
finely pubescent, narrowly margined, notched at the apex, sub-
truncate at the base, its stipe about 0.6 mm. long.
Rocky slopes and hillsides, western Porto Rico, especially
on serpentine, from sea-level to 700 m. altitude. Type collected
at Guanajibo, near Mayaguez (Britton, Cowell & Brown 4349).
Recorded by Bello from my type locality as B. domingensis DC.
[Polygala domingensis Jacq.; B. Penaea (L.) DC.] of Hispaniola,
to which species it is nearest related, but which differs in pilose-
pubescent twigs, rather densely pubescent leaves, and much more
: pubescent, shorter-pedicelled flowers.
BRITTON: STUDIES OF WEsT INDIAN PLANTS 495
2. BADIERA VIRGATA Britton
The specimen referred to by me under B. oblongata Britton
(Britton & Wilson 6066) from Santa Clara, Cuba, is now, in the
light of additional material, put with B. virgata, as a broad-leaved
race.
4. BADIERA DIVERSIFOLIA (L.) DC.
This, the only Jamaican species of the genus, forms a tree
up to 6 m. high, as at St. Ann’s Bay (Britton 2408).
5. BADIERA CUBENSIS Britton
In the original description of this species (Bull. Torrey Club
37: 362), I included too many citations of specimens. C.
Wright’s Cuban r973 consists, apparently, of three species; it is
to the specimen with abruptly acuminate leaves cuneate-narrowed
at the base, 4-6 cm. long, that the name should be restricted. As
shown by specimens from Bahia Honda, Pinar del Rio (P. Wilson
9429) the leaves vary to obovate. The fruit is glabrous when
mature, its slender stipe 2 mm. long. .
5a. BADIERA FUERTESII Urban, Symb. Ant. 7: 244. I912
A tree of the mountains of Santo Domingo, with glabrous
leaves, strongly reticulate-veined on the upper surface.
6. BADIERA MONTANA Britton
Described from fruiting specimens collected in the Trinidad
Mountains, Santa Clara, Cuba. Flowering specimens with ap-
parently the same foliage and pubescence come from limestone
hills in the vicinity of Sumidero, Pinar del Rio (Shafer 13819), but
flowers from the Trinidad Mountain tree are needed to make the
identity wholly satisfactory.
8. Badiera propinqua sp. nov.
Twigs slender, densely appressed-pubescent.. Leaves ovate to
ovate-elliptic, 2-4.5 cm. long, 8-25 mm. wide, bluntly acute or
obtuse and emarginate at the apex, narrowed or obtuse at the
base, sparingly short-pubescent on both sides or becoming glab-
rous, the midvein impressed above, prominent beneath, the slender
lateral veins 6 to 8 on each side of the midvein, the appressed-
pubescent petioles 1.5-3 mm. long; inflorescence few-flowered,
496 BRITTON: STUDIES OF WEsT INDIAN PLANTS
4-5 mm. long; bracts ovate, densely pubescent, scarcely 0.5 mm.
long; pedicels slender, appressed-pubescent, 1.5-2.5 mm. long;
sepals suborbicular, rounded, pubescent, the larger about 1.3 mm.
long; petals white; keel pubescent on the sides, about 2.5 mm.
long; fruit (from Wright 1913, in part) 10 mm. wide, 7 mm. long,
deeply notched, scarcely margined, glabrous, decurrent on its
stipe, which is about 2 mm. long.
Cuba. Type collected between Los Palacios and San Juan
de Zayas, Pinar del Rio (Shafer 11818). The description is
drawn to include the part of Wright 1913, and Wright 3496, pre-
viously referred by me to B. cubensis. The part of Wright 115,
included by me in the original description of B. cubensis, is perhaps
referable here also.
9. Badiera punctata sp. nov.
A shrub about 1 m. high, with slender, ascending, or nearly
erect branches, the twigs densely appressed-pubescent with short
hairs. Leaves ovate to ovate-elliptic, firm in texture, 1.5-2.5 cm.
long, 12 mm. wide or less, obtuse or emarginate at the apex, obtuse
at the base, the midvein impressed above, prominent beneath,
the lateral veins 3 or 4 on each side, the upper surface strongly
punctate, shining and glabrous or nearly so, the under surface
dull, the petioles about 1.5 mm. long; inflorescence few-flowered ;
bracts scarcely 0.5 mm. long; fruiting pedicels slender, nearly
glabrous, 2—2.5 mm. long; sepals ovate, obtuse, ciliate, about I
mm. long; fruit (not quite mature) ‘‘green and red,’’ 7 mm. wide,
about as long as wide, shallowly notched at the apex, obliquely
subtruncate at the base, narrowly margined, slightly pubescent,
its stipe 1-1.5 mm. long.
Near astream, Arroyo del Medio above the falls, Oriente, Cuba
(Shafer 3644). The leaves droop at night, a phenomenon also
observed in B. virgata.
10. Badiera heterophylla sp. nov.
An upright shrub 6 dm. high, the twigs short-pubescent with
somewhat spreading hairs. Leaves of two quite different forms, .
(1) narrowly ovate, 3-4 cm. long, ang mm. wide, bluntly acute
at the apex, (2) elliptic, 2-2.5 cm. long, 12-14 mm. wide, rounded
or obtuse at the apex; both forms uaa at the apex, nar-
rowed or obtuse at the base, sparingly short-pubescent and dull
on both sides, the midvein impressed above, prominent beneath,
the slender lateral veins only 2 or 3 on each side of the midvein,
BRITTON: STUDIES OF WEsT INDIAN PLANTS 497
the appressed-pubescent petioles about 2 mm. long; inflorescence
about 5 mm. long; bracts about 0.5 mm. long; pedicels pubescent,
2 mm. long or less; sepals rounded, pubescent, the larger about
1 mm. long; “flowers yellow”’; petals slightly pubescent in the
bud; fruit 7-8 mm. wide, about 5 mm. high, margined, rather
deeply notched, sparingly pubescent when young, glabrous when
mature, slightly decurrent on its stipe which is about 1.5 mm. long.
Deciduous woods, Sierra Nipe, near Woodfred, Oriente, at
450-550 m. altitude (Shafer 3070).
40. THE GENUS TRICERA Sw. IN CUBA
I. Styles coherent, at least at the base
ae oblong to ‘sihaaeslminiiictileds 2-3 cm. wide; styles
coherent below 1. T. Muelleriana.
Leaves broadly ovate to ovate-lanceolate, 5 cm. wide or
less; en coherent pearly throughout. 2. T. marginalis.
2. Styles separa d distinct.
* Leaves 4-10 cm. long.
Venation of me leaves manifest.
Leaves broadly ovate, subcordat 3. T. cubana.
Leaves pompeees to elliptic or cesusie narrowed
at the bas
pe soon 4. T. acuminata.
Capsule ovoid, a - 5. 2. brevipes.
Paws of the leaves Ie or indistinct.
tioles very stout, 10 mm. long or less; leaves very
pes 6. T. crassifolia.
Petioles slender, 2-6 mm. long; leaves firm, but not
very thick.
Twigs yellow. 7. T. flaviramea.
os ae
b io 8. T. retusa.
Capsule lobes linear-spatulate. 9. T. gonoclada.
** Leaves I-3.5 cm. long.
Leaves ee to obovate or elliptic, not more
hi wice as long as wide
Inflorescence and parse petit 3 10. T. rotundifolia,
Inflorescence and ca) . T. Shaferi.
Leaves oblong to ite eesti 2-5 times as long
wide.
Leaves oblong, 2-3 times as long as wide.
Leaves densely imbricated, less than I cm. long,
the in es very short.
Leaves not densely imbricated, 1.5-3.5 cm. long.
Staminate flowers sessile or very nearly so. 13. T. glomerata.
Staminate flowers manifestly pedicelled.
Leaves 2-3.5 cm. long, the lateral venation
obsolete. 14. T. bahamensis.
2. T. vaccinoides.
im
498 BRITTON: STUDIES OF WEST INDIAN PLANTS
Leaves I-1.5 cm. long, the lateral vena-
tion wholly obscure. 15. T. microphylla.
Leaves linear-oblanceolate or linear-oblong, 4-5
times as long as wide
Capsule parte 16. T. Leoni.
astern pubesc
' Leaves cea revolute-margined, abruptly
ucronate. 17. T. revoluta.
Leaves scarcely revolute-margined, acute or
obtuse. 18. T. foliosa.
1. Tricera Muelleriana (Urban)
Buxus Muelleriana Urban, Symb. Ant. 5: 400. 1908.
TYPE LOCALITY: Cuba.
DisTRIBUTION: Known only from the type specimens (C.
Wright 1920).
Referred by Grisebach to Tricera laevigata Sw., of Jamaica,
and by Mueller to Buxus subcolumnaris Muell. Arg. = Tricera
subcolumnaris (Muell. Arg.) Britton, of Martinique.
2. Tricera marginalis sp. nov.
A glabrous shrub about 3 m. high. Leaves broadly ovate to
ovate-lanceolate, subcoriaceous, 4-8 cm. long, 2-5 cm. wide,
reticulate-veined on both sides, with 2 distance “yeins 1.5-3 mm
rom the margins, the midvein impressed above, prominent be-
neath, the apex acute, the base obtuse or subtruncate, the rather
slender petioles 1 cm. long or less; cymes several-flowered, longer
than the petioles, glabrous; bracts ovate-lanceolate, acute, 1.5
mm. long; pedicels of the staminate flowers about 4 mm. long,
the white calyx about 3 mm. long, its segments oblong, obtuse;
stamens about as long as the calyx, the filaments about twice as
long as the anthers; ovary ovoid; united styles somewhat longer
cen the ovary; stigmas recurved, about one-third as long as the
styles.
Alluvial valley of Rio Yamaniguey, Oriente (Shafer 4226).
3. TRICERA CUBANA A. Rich. in Sagra, Hist. Cub. 11: 217. pl. 71.
1845
Buxus cubana Baill. Mon. Bux. 71. 1859.
TYPE LOCALITY: Monte Libau, Oriente.
-ILLustraTIon: A. Rich. loc. cit.
DisTRIBUTION: Known only from the type locality.
BRITTON: STUDIES OF WEsT INDIAN PLANTS 499
4. Tricera acuminata Griseb. Nachr. Gesell. Wiss. Goetting. 1865:
162. 1865
Buxus acuminata Muell. Arg. in DC. Prodr. 16: 15. 1869.
TYPE LOCALITY: Cuba.
DISTRIBUTION: Known only from the type specimens (C.
Wright rozro, in part).
5. Tricera brevipes (Muell. Arg.)
Buxus citrifolia brevipes Muell. Arg. in DC. Prodr. 16'!: 15. 1869.
Buxus brevipes Urban, Symb. Ant. 5: 400. 1908.
TYPE LOCALITY: Cuba (C. Wright 1919, in part).
DISTRIBUTION: the label with Wright rorg in the herbarium of
the New York Botanical Garden gives the locality as San Diego
de Tapis; the following specimens appear to agree with it: upper
valley of the Rio Navas, Oriente (Shafer 4402); Rio San Miguel
below Mal Paso, Pinar del Rio (P. Wilson 9309).
Referred by Grisebach to T. fasciculata Griseb., of Jamaica.
I am not confident that the species is distinct from 7. acuminata.
6. Tricera (?) crassifolia sp. nov.
A glabrous shrub about 3 m. high, the twigs stout. Leaves
elliptic-obovate, 6-10 cm. long, 2.5-4 cm. wide, very thick, obtuse
' or emarginate and mucronulate at the apex, narrowed at the base,
the midvein impressed above, prominent beneath, the ee
venation faint and obscure, the stout petioles about 1 cm. lon
‘ young inflorescence axillary, about as long as the petioles, both
staminate and pistillate one apparently sessile, the staminate
about 2 mm.
Between Camp Toa and Camp La Barga, 400-450 m. alt.,
mountains of northern Oriente (Shafer 4163).
7. Tricera flaviramea sp. nov.
A glabrous shrub, 1 m. high, the young twigs and leaves yellow-
ish green, the branches slender, somewhat ridged. Leaves elliptic
or elliptic-obovate, coriaceous, revolute-margined, 3-5 cm. long,
I~2 cm. wide, rounded and minutely apiculate or emarginate at
the apex, narrowed at the base, the midvein impressed above,
rather prominent beneath, the lateral venation obsolete, or faintly
discernible on the upper surface, the petioles 1.5-3 mm. long;
inflorescence glabrous, about 1 cm. long; bracts ovate, acute, 1.5
500 BRITTON: STUDIES OF WEsT INDIAN PLANTS
mm. long; young staminate flowers about 2 mm. long, on pedicels
about as long; capsule ovoid, obtuse, glabrous, 7-8 mm. long,
5-6 mm. thick, the persistent styles somewhat recurved, linear,
5 mm. long, separated from each other about 1 mm. at the base.
Palm-barren, Santa Clara City, Santa Clara (Britton & Cowell
13324, type; Britton & Wilson 6211).
8. TRICERA RETUSA Griseb. Nachr. Ges. Wiss. Goetting. 1865:
163. 1865
Buxus retusa Muell. Arg. in DC. Prodr. 16': 17. 1869.
TYPE LOCALITY: Monte Verde, Oriente.
DISTRIBUTION: Mountains of northern Oriente.
g. TRICERA GONOCLADA C. Wright; Griseb. Cat. Pl. Cub. 282.
1866
Buxus gonoclada Muell. Arg. in DC. Prodr. 16': 16. 1869.
TyPE LOCALITY: Savanna near Guamacaro, western Cuba.
DIsTRIBUTION: Known only from the type locality.
10. Tricera rotundifolia sp. nov.
A glabrous shrub, 6 dm. high, much branched, the internodes
mostly shorter than the leaves. Leaves suborbicular to obovate,
coriaceous, 2 cm. long or less, rounded and mucronulate or
slightly emarginate at the apex, narrowed at the base, the margins
thickened and somewhat revolute, the midvein rather prominent
beneath, the lateral venation obsolete, the petioles 1.5-2 mm.
long; inflorescence glabrous, few-flowered, in the uppermost axils;
pedicels of the staminate flowers about 3 mm. long; bracts ovate,
acute, I mm. long; staminate flowers 1.5 mm. long, their segments
oblong-lanceolate id, about 6 mm. long, glabrous,
the persistent styles contiguous, about 2 mm. long.
Between Camp La Barga and Camp San Benito, about 1,000
m. alt., northern Oriente (Shafer 4114).
11. Tricera Shaferi sp. nov.
A much-branched shrub up to 3 m. high, the twigs slender,
glabrous, t ter than the leaves. Leavesobovate to
elliptic, coriaceous, £.5-3.5 cm. long, rounded or retuse at the
apex and minutely mucronulate, narrowed or obtuse at the base,
the midvein impressed above, prominent beneath, the lateral
venation wholly aoe: the upper surface shining, dark green,
BRITTON: STUDIES OF WEsT INDIAN PLANTS 501
the under surface dull and pale, the petioles 1-3 mm. long; inflores-
cence axillary, puberulent, short-stalked, much shorter than the
leaves; bracts ovate, acute, about 0.5 mm. long; pedicels of the
staminate flowers rather stout, 2-3 mm. long; staminate flowers
about 1.5 mm. long; capsule subglobose, densely puberulent,
about 4 mm. in diameter, the separated persistent styles about
2 mm. long.
Wet mountains of northern Oriente. Type collected between
Camp Toa and Camp La Barga, 400-450 m. alt. (Shafer 4167).
12. Tricera vaccinioides sp. nov.
A low, glabrous, much-branched shrub, 3-6 dm. high, the
branches stout, the twigs erect-ascending, densely leafy, the inter-
nodes 3-6 mm. long. Leaves oblong to oblong-obovate, 5-8 mm.
long, about twice as long as wide, coriaceous, faintly 3-veined, the
secondary venation almost wholly obscure, the apex acute an
mucronate, the base narrowed, the stout petioles scarcely I mm
long; inflorescence terminal, glabrous, few-several-flowered ; pedi-
cels of the staminate flowers rather stout, rigid, about 2 mm. long;
bracts lanceolate; staminate flowers about 1 mm. long; capsule
globose-ovoid, glabrous, dull, about 6mm. long, about three times
as long as the persistent styles.
Stony pass, between Camp La Barga and Camp San Benito,
mountains of northern Oriente, Cuba, about 1000 m. alt. (Shafer
4122).
13. TRICERA GLOMERATA Griseb. Mem. Am. Acad. II. 8: 157.
1860
Buxus glomerata Muell. Arg. in DC. Prodr. 16': 17. 1869.
TYPE LOCALITY: Cuba.
DISTRIBUTION: Dry rocky hillsides, Oriente; Santa Clara.
This is a shrub up to 3 m. high, abundant near the mouth of
Cienfuegos Bay; I am indebted to Professor Fernald for a com-
parison of a specimen collected there, at Castillo de Jagua (Britton,
Wilson & Earle 4597), with the specimen of C. Wright in the
Gray Herbarium; he pronounced it “‘a perfect match.”
14. TRICERA BAHAMENSIS (Baker) Britton, Bull. N. Y. Bot.
Gard. 4: 139. 1906
Buxus bahamensis Baker in Hook. Ic. Pl. pl. 1806. 1889.
Type LocaLity: Long Cay, Bahamas.
502 BRITTON: STUDIES OF WEsT INDIAN PLANTS
DISTRIBUTION: CayoRomano, Camaguey :— Bahamas; Jamaica.
Closely related to Tricera Vahlii (Baill.) Britton [Buxus Vahlu
Baill.] of Porto Rico and St. Croix.
15. TRICERA MICROPHYLLA Griseb. Nachr. Ges. Wiss. Goetting.
1865: 163. 1865
Buxus Wrightit Muell. Arg. in DC. Prodr. 611: 17. 1869.
TYPE LOCALITY: Western Cuba.
DISTRIBUTION: Known only from the type specimens collected
by Wright.
16. Tricera Leoni sp. nov.
A shrub, or small tree, up to 4 m. high, glabrous throughout,
the slender twigs densely leafy. Leaves linear to linear-oblong,
coriaceous, shining, 2 cm. long or less, 2-4 mm. wide, spinulose-
acute at the apex, narrowed at the base, the midvein faintly dis-
tinguishable above, rather prominent beneath, the lateral vena-
tion wholly obscure, the petioles 1-1.5 mm. long; inflorescence
terminal, short-stalked; bracts ovate, acute, about I mm. long;
pedicels of the staminate flowers 1.5-2 mm. long; staminate
perianth-segments oblong-lanceolate, acute, 2-2.5 mm. long, ©
filaments a little longer than the anthers; capsule globose-oblong;
glabrous, 3 mm. long, the separated, persistent styles 2 mm. long.
Hill north of Monte Catalina, northeast of San Diego de los
Banos, Pinar del Rio (Brothers Leon & Charles 4874, type);
barren specimens from the southern slope of Monte Cajalbana,
Pinar del Rio (Brothers Leon & Charles 4943), with narrower
leaves than the type, are provisionally referred to the same species.
17. Tricera revoluta sp. nov.
A much-branched shrub, 3-10 dm. high, the twigs densely
leafy, the internodes only 1-2 mm. long. Leaves linear-oblanceo-
late, coriaceous, 12-20 mm. long, 3-5 mm. wide, strongly revolute-
margined, sharply cuspidate-mucronate at the apex, attenuate
at the base, the midvein elevated beneath, the venation otherwise
wholly obscure, the petioles about 2 mm. long; inflorescence few-
flowered in the upper axils, pubescent; staminate flowers nearly
2 mm. long, and nearly as long as their stout pedicels; capsule
subglobose, 5-6 mm. long, . esingaeste shorter than the persistent,
separated styles.
Dry, rocky situations, serpentine hills near mouth of Rio
Yamaniguey, Oriente (Shafer 4247, type); between Camp La
BRITTON: STUDIES OF WEsT INDIAN PLANTS 503
Barga and Camp San Benito, about 1,000 m. alt., Oriente (Shafer
4472).
18. Tricera foliosa sp. nov.
A low, much-branched shrub, the slender twigs densely leafy.
Leaves linear-oblanceolate, subcoriaceous, glabrous, 10-15 mm.
long, 2 mm. wide or less, obtuse and often minutely mucronulate
at the apex, long-attenuate to the base, the midvein impressed
above, slightly elevated and channeled beneath, the lateral vena-
tion wholly obscure, the margins slightly or scarcely revolute, the
petioles 1-2 mm. long; inflorescence axillary, shorter than the
leaves; staminate flowers white, about 1.5 mm. long, as long or
longer than their pubescent petioles; bracts lanceolate, about I
mm. long; capsule subglobose, densely and finely pubescent,
about 3 mm. in diameter, the separated persistent styles about
2 mm. long.
Among rocks near water, Camp La Gloria, south of Sierra
Moa, Oriente (Shafer 8256, type); on rocks, vicinity of Camp San
Benito, Oriente (Shafer 4073).
41. THE GENUS PLUMIERA L. IN CUBA
Leaves narrowly linear, 1.7 mm. wide or less. 1. P. filifolia.
Leaves oblong to elliptic-oblanceolate or obovate, 1-8 cm. wide.
Leaves densely lanate or pubescent beneath.
Corolla-lobes owly obovate 2. P. sericifolia.
Corolla-lobes meee elliptic-obova 3. P. lanata
Leaves glabrous, or sparingly pubescent Besekth:
Petioles short, I cm. long or
Leaves Resale or 9 ORDA the lateral venation
obscure. 4. P. clusioides.
Leaves narrowly oblong, the lateral venation promi- ’
nent. 5. P. nipensis.
Petioles slender, mostly 2—4 cm. lon
abrous beneath, their margins not involute.
Corolla-lobes 2.5-3.5 cm. long, I-I.5 cm. wide;
venation widely spreading. 6. P. emarginata.
mipemcating I.5-2.5 cm g.
on of the ‘are regen tae not prominent
ou is corolla-lobes obova
Venation of the leaves widely diab promi-
nent eeu: corolla-lobes oblong-oblance
1
. P. obtusa.
~s
te. 8. P. venosa.
Leaves pubescent on the veins beneath, their margins
involute. 9. P. trinitensis.
504 BRITTON: STUDIES OF WEST INDIAN PLANTS
1. PLUMIERA FILIFOLIA Griseb. Mem. Am. Acad. II. 8: 519.
1862
River cliffs, foothills of the Sierra Maestra, Nouvelle Sophie
and Ensenada de Mora, Oriente. Endemic.
2. PLUMIERA SERICIFOLIA C. Wright; Griseb. Cat. Pl. Cub. 171.
1866
P. emarginata sericifolia Maza, Anales Soc. Esp. Hist. Nat. 23:
273. 1894.
Coastal thickets, Oriente; Matanzas; Havana; Pinar del Rio:
—Inagua.
Perhaps, as suggested by Professor Gomez de la Maza, a
pubescent race of P. emarginata Griseb., which has a similar dis-
tribution in western Cuba, although I have not seen the two
growing in close proximity.
3. Plumiera lanata sp. nov.
A shrub or.small tree up to 3.3 m. high. Leaves broadly
elliptic to elliptic-obovate, firm in texture, 10-18 cm. long, 5-9
cm. wide, rounded, emarginate or subtruncate at the apex,
narrowed or subtruncate at the base, dark green and glabrous
above, densely white-lanate beneath, the midvein channeled
above, prominent beneath, the lateral veins widely spreading,
_averaging 6 or 7 mm. apart, united close to the margin, the stout,
somewhat pubescent petioles 1-2.5 cm. long; peduncle 3-11 cm.
long, glabrous; cymes many-flowered, dense; pedicels 10 mm.
long or less, glabrous or pubescent; calyx subtruncate, 2-3 m
long; corolla-tube about 1.5 cm. long; 6étollaclobes onde
elliptic-obovate, white, rounded, 1.5-2.5 cm. long, 1.2-2 cm. wide;
follicles 8-10 cm. long, about 1 cm. thick, short-pointed.
Coastal rocks, vicinity of Baracoa, Oriente (Underwood &
Earle 1442).
4. PLUMIERA CLUsIOIDEs Griseb. Cat. Pl. Cub. 171. 1866
? P. obtusa parvifolia Griseb. Mem. Am. Acad. II. 8: 519. 1862.
? P. clusioides parvifolia Maza, Anales Soc. Esp. Hist. Nat. 23:
273. 1894.
Rocky hillsides and palm-barrens, Oriente; Camaguey; Santa
Clara, Havana. Endemic.
Fruiting specimens collected by Dr. Shafer in the gorge of the
BRITTON: STUDIES OF WEST INDIAN PLANTS 505
Rio Yamuri, Oriente (7800), tentatively referred to this species,
have spatulate leaves 12 cm. long.
5. Plumiera nipensis sp. nov.
A slender tree, up to about 6 m. in height. Leaves narrowly »
oblong, 9-12 cm. long, I.5-2.3 cm. wide, obtuse at the apex,
narrowed at the base, glabrous on both sides, shining above, dull
beneath, the midvein channeled above, prominent beneath, the
lateral venation widely spreading, impressed, but distinct, above,
the rather slender petioles I cm. long or less; peduncle stout, 6 cm.
long; follicles 13-15 cm. long, about 2 cm. thick, acutish; seeds
10 mm. long, 6-7 mm. wide, the wing about 15 mm. long, 8-9
mm. wide.
Rocky thicket, between Piedra Gorda and Woodfred, Sierra
Nipe, Oriente, at 500 m. alt. (Shafer 3089). Perhaps the same as
P. obtusa laevis Griseb. (Mem. Am. Acad. IT. 8: 519) from Monte
Verde, Oriente.
6. PLUMIERA EMARGINATA Griseb. Cat. Pl. Cub. 171. 1866
Rocky cliffs and hillsides, Matanzas; Pinar del Rio; Isle of
Pines. Endemic.
7. PLUMIERA OBTUSA L. Sp. Pl. 210. 1753
Coastal hills, keys and thickets, Oriente; Camaguey; Santa
Clara:—Bahamas; Hispaniola; Mona. Attributed to Jamaica, but
some specimens which have been so referred represent a distinct
species.”
* Plumiera confusa sp. n
A tree, up to 6 m. high little agers sggleths throughout. Leaves narrowly
elliptic or pee et shining, 1-3 dm. long, wide or le se obtuse, emarginate or
cute at the apex, narrowed at the base, ‘he midvein channeled above ret
beneath, the lateral —- widely ae the stout ‘ps les 4 :
peduncles stout, 7-I - long; cymes lar. y-flowered; pedic 2 co stout,
ckened upward, 9-12 elie tee vith aie, abo mm. long;
white with a pale yellow eye, its tube 12-15 long, its lobes broadly oblong,
rounded, 2.5—3 cm. long, about I cm. wide; follicles 812 cm. long, 10-12 mm. thick.
ky situations, southern side of Jamaica. Type collected on Great Goat
Island ee 9309);
pecies more Tepe es, PF. ener erate than P. obtusa. I amunable to deter-
mine, iu th time, whether P. obtus amaica or
not. Other ee referable to P. pees are: Healthshire Hills near Salt Island
r)
8
(Brition 3061); base of Healthshire Hills near Salt Ponds (Britton 3036); Great Goat
me ( arris 10168; Britton 1874).
e Plumiera of Pedro Bluff, a locality about thirty miles west of Great Goat
ae but still within the xerophytic region, may be P. obtusa, but the specimen
.
506 BRITTON: STUDIES OF WEST INDIAN PLANTS
8. Plumiera venosa sp. nov.
A tree up to 6 m. high, glabrous throughout. Leaves oblong-
oblanceolate to narrowly obovate, 7-15 cm. long, 3 cm. wide or
less, obtuse or emarginate at the apex, narrowed or cuneate at
the base, the midvein impressed above, prominent beneath, the
lateral veins numerous, widely spreading, united near the margin,
2-3 mm. apart, prominent beneath; petioles slender, 2—3 cm. long;
cymes several—many-flowered; peduncles 6-10 cm. long; pedicels
8-15 mm. long, upwardly somewhat thickened; calyx about 3
mm. long, its teeth very broad and short; corolla-tube about I. i
m. long; corolla-lobes narrowly oblong-oblanceolate, 2—-2.7 c
ie follicles 10-15 cm. long, about 12 mm. thick; seed bbe
I cm. long, its wing about 1.5 cm. long.
Coastal cliffs and hillsides near the coast, southern Oriente.
‘Type collected in the basin of Rio Guama, Sevilla Estate near
Santiago (Taylor 93).
Shoots from cut stumps bear nearly linear leaves up to 2 dm.
long, 10-15 mm. wide, as seen by me in a coastal thicket near the
mouth of Santiago Harbor (Britton & Cowell 12632).
9g. Plumiera trinitensis sp. nov.
A tree 5 m. high. Leaves oblong to oblong-oblanceolate, II
cm. long or less, 1.5-3.5 cm. wide, obtuse or emarginate at the
apex, gradually narrowed hi the base, glabrous above, pubescent
on the veins beneath, the midvein broad, channeled above,
prominent beneath, the lateral veins rather widely spreading,
2-4 mm. apart, united near the revolute margins; petioles 1.5-2.5
cm. long; follicle slightly curved, pointed, 18 cm. long, about
1.5 cm. thic
Dry hillside, La Vigia, Trinidad, Santa Clara (Britton & Wilson
5514). Similar to P. jamaicensis Britton, which is a glabrous
species up to 10 m. high, with broader leaves.
(Harris 9996) is not complete. In 1895, E. Campbell collected a Plumiera at Con-
stant Spring per which may be P. obtusa.
Gr P. confusa on Great Goat Island there is a Plumiera which has
leaves over 3 Keen long, narrowly oblong to linear-oblong, some of them not more
3 cm. wide, and all long-acuminate at the apex (Britton 1870). It was not in
flower at the time of my visit, March 4, 1908, but I obtained its fruit, which is much
like that of P. confusa, and its leaf-venation resembles that of P. confusa also; it
may be that P. confusa is sufficiently heterophyllous to have these narrow, elongated,
long-pointed leaves.
BRITTON: STUDIES OF WEST INDIAN PLANTS 507
THE GENUS GUETTARDA L. IN CUBA
A. Leaves minute, nearly sessile, 6 mm. long or less. 1. G. Echinodendron,
B. Leaves larger, 1 scm . long or more.
a, Leaves spinulose-tipped.
Leaves ovate to elliptic, strongly reticulate-veined
beneath
neath, 2. G. Coxiana,
Leaves oblong, scarcely Se beneath. 3. G. cueroensis.
bv. I spinulose-tipp
* Leaves small, be at cm. ie or less; flowers axillary,
mostly so
Leaves ener ivcadiis beneath. 4. G. rigida.
Leaves glabrous or somewhat pubescent beneath.
Flowers sessile or nearly so; leaves ovate or
blong.
Venation inconspicuous on upper leaf-
surfaces.
Corolla ahd 2 cm. long; calyx
spathac 5. G. camague yensis.
Corolla 1-1.5 cm. long.
yx spathaceous; corolla-tube
little longer than the li mb. 6. G. clarensis.
ee truncate or oblique; corolla-
3-4 timesaslongasthelimb. 7. G. undulata,
Venation patie on upper leaf-surfaces. 8. G. xanthocarpa.
Peduncles nearly as long as the fruits; leaves
ona 9. G. sciaphila,
* * Leaves larger —15 cm. lon
T Inflorescence ict or ETE or flowers
solitary at the ends of twigs, or some flowers
also peduncled in the 2a r axils.
ee capitate-clustered.
: Leaves densely oe or puberulent
pie ath.
Leaves densely puberulent beneath. 10. G. densiflora.
Leaves villous beneath, at least on the
veins. :
Fruit 2-3 cm. in diameter. 11. G. macrocarpa.
Fruit 6-15 mm, in
=
°
.—_i
3 ye
5
c
n
a
re
one i ecw larenCe
| «il
o
=
|
a
i
J
r " 19 21 23 25
Fic, 2. Regression 0 of barren wd Ween came on eee of pistillate, and of
number of pist
A. proboscideum.
represent the deviation of the number of staminate flowers per
inflorescence from the probable number on the assumption that the
proportion of the two types, s and #, is unaffected by the total
number produced per inflorescence. If the correlation between
HARRIS: DISTRIBUTION AND CORRELATION OF SEXES 671
t, = s + p, or total flowers per inflorescence, and z be positive it
indicates that as the total number of flowers per inflorescence
becomes larger the staminate flowers are relatively more numerous.
Using the formula suggested to me* by Professor Karl Pearson
I find:
For A. vulgare:} ;
Tim = + .964 + .002,
fiz = — .230 + .026.
For A. proboscideum:
Yim = + .931 + .002,
Tig = — .234 + .O12.
In both cases the correlation between total flowers and number
of staminate flowers is high. This is as one would expect it to be.
The interesting feature of the result is that in both species there
is a significant negative correlation between the total number of
flowers and the deviation of the number of staminate flowers from
their probable value. In short, the male flowers while absolutely
more abundant in the larger inflorescences are there relatively
distinctly less abundant. Or in other words the larger inflores-
cences tend to produce a larger proportion of pistillate flowers.
Consider these relationships on the basis of the individual
collections.
TABLE V, in which the constants for the individual collections
of A. proboscideum are given, shows that with a single exception,
in which WN is very small, the correlations for total flowers and
staminate flowers are very high.
The critical values derived from these are given in the sixth
column. Of these 10 coefficients, 7 are negative in sign, and 5
may be considered unquestionably significant in comparison with
their probable errors. Neither of the 3 positive values may be
looked upon as certainly trustworthy. Thus while the constants
for the individual sub-series are very irregular in magnitude they
fully substantiate the conclusions to be drawn from the constant
* Harris, J. Arthur. The Correlation between a Variable and the Deviation
of a Theiskident Variable from its Probable Value. Biometrika 6: 438-443. 1909.
+ Calculations from ungrouped data.
672 HARRIS: DISTRIBUTION AND CORRELATION OF SEXES
for the whole material. The average (weighted) value is
72 = — .223. This value is not significantly different from the
Tig = — .234 obtained from the combined materials.
Ill. SumMMARY AND CONCLUSIONS
As a suggestion for more adequate investigation of variation
and correlation in the specialized inflorescence of the aroids this
note presents the results of a statistical analysis of two rather large
series of data collected for the genus Arisarum by Cannarella.
The following points may be emphasized:
In both A. vulgare and A. proboscideum, the coefficient of
variation for number of pistillate flowers is far higher than that
for number of staminate flowers. The interpretation of this
result presents considerable difficulty.
In A. vulgare the inflorescence not only produces a larger
number of both staminate and pistillate flowers than does that of
A. proboscideum, but shows a far higher variability, both absolute
and relative.
Apparently the collections of A. proboscideum came from a
wider range of habitats than those of A. vulgare. If this be true,
the variability which they show is in part due to heterogeneity
arising through environmental differentiation. This probability
renders more significant the observed lower variability in A. pro-
boscideum. Any subsequent investigation should if possible take
into account this factor in drawing comparisons between the
variability of these or any other forms. :
The inflorescence of A. vulgare is not only more variable, but
shows a higher degree of correlation of its parts than does that of
A. proboscideum. In both cases the correlation between number
of staminate and pistillate flowers is rather small. The relation-—
ship between the numbers of the two types of flowers may be fairly
well described by a straight line. The fit of the straight line
equations is not altogether satisfactory, but it seems idle to carry
the statistical analysis farther on the present data. The effort
would much better be expended in securing more adequate bio-
logical material.
The correlation between the total number of flowers produced
by the inflorescence and the number of staminate flowers is neces-
HARRIS: DISTRIBUTION AND CORRELATION OF SEXES 673
sarily positive and high. The correlation between the total
number of flowers and the deviation of the staminate flowers from
their probable value, on the assumption that the character (sex)
of the flowers is independent of the size of the inflorescence, is
significantly negative. This shows that the larger inflorescences
have relatively more pistillate flowers.
CoLp SPRING HARBOR,
EW YORK
Dictyophlois reticulata gen. et sp. nov.
AucG. F, FOERSTE
(WITH PLATE 33)
The name Stigmaria ficoides Brongniart at present does service
in designating the so-called roots of both of the genera of stems
known as Lepidodendron and Sigillaria. Not only have these
roots been found in direct connection with the Lepidodendron
and Sigillaria stems but it has been found impossible so far to
discover any apecific or generic differences in the roots so con-
nected.
The genus Lepidodendron makes its appearance in the Lower
Devonian and dies out in the Permian. Recently Dr. David White
has called my attention to the fact that in Mississippian or Sub-
carboniferous specimens of Stigmaria ficoides the characteristic
round scars, indicating the points of attachment of the so-called
rootlets, are distinctly smaller than in Pennsylvanian or Coal
Measure representatives passing under the same name.
Students of the microscopic structure of Stigmaria and of its
rootlet-like appendages, regard the latter as homologous with the
monarch roots of certain living plants, so that the main axes of
the so-called roots known as Stigmaria should be known as rhizo-
phores rather than roots. There is a possibility that these rhizo-
phores represent underground stems, but this still is a debatable
question.
In the vicinity of eee two miles east of Stephensport,
Breckenridge County, western Kentucky, the rhizophores of both
Stigmaria ficoides and of an undescribed but related genus occur
in one of the sandstone layers belonging to the lower part of the
Chester division of Sebcarhopierees rocks.
emble those of Stigmaria ficoides
in possessing small ring-like areas of attachment of the monarch
roots, the latter being arranged in diagonally intersecting rows.
In several of these ring-like areas a minute central point could be
recognized, but the smaller ring, which in Stigmaria intervenes
675
676 FOERSTE: DICTYOPHLOIS RETICULATA SP. NOV.
between this central point and the larger ring, could not be iden-
tified with certainty in any case.
These undescribed rhizophores differ from those of Stigmaria
in having the areas between the ring-like scars of attachment of
the monarch roots marked by irregular ridge-like elevations and
depressions presenting the appearance of an irregular network.
In the immediate vicinity of the ring-like scars the ridge-like
elevations of this network tend to be arranged in radiating series.
The network evidently is a characteristic of the outer cortex, hence
the name Dictyophlois reticulata, here proposed for the rhizophores
in question. No explanation based on evidence can be assigned
at present for the reticulate appearance of the surface of this outer
cortex. But, if these rhizophores are homologous with subterranean
stems rather than with roots, the reticulate structure may be con-
nected with the former presence of more or less appressed scales,
of which, however, there is no positive evidence in Dictyophlois; in
Stigmaria the former presence of appressed scales seems impos-
sible since the outer cortex of the rhizophores in the latter genus
is smooth,
The specimens of Dictyophlois here described and figured were
discovered about ten years ago and were retained in the hope that
further evidence regarding their structure might be obtained.
Since this delay has not been attended by any additional informa-
tion the accompanying illustration is published in the hopes that
it may arouse interest in other paleobotanists. If Dictyophlois
is indeed an underground scale-bearing stem then these scales
should be found sooner or later still attached to the rhizophores.
Dictyophlois reticulata gen. et sp. nov.
Original diameter of rhizophores varying from 10 cm. to 15
cm. The figured specimen (type) is a flattened fragment 10 cm.
in length and 13 cm. in width, exposing a part of one side of a
rhizophore, the original diameter of which is unknown. Round
areas of attachment of the so-called rootlets usually 3.5-4 mm.
in width, but not infrequently attaining a length of 5 mm.;
with a small central pit about 0.75 mm. in diameter. These :
areas are arranged in diagonally intersecting series, five areas
occupying a length of 60-80 mm. measured along the diagonal
rows. For adistance of 3-4 mm. from the margins of these round
areas of attachment the raised ridges forming the intermediate
FOERSTE: DICTYOPHLOIS RETICULATA SP. NOV. 677
network tend to be arranged in a more or less radiating manner.
In the more distant parts of the intermediate area these ridges
form a more irregularly reticulated network. Measured along
the crests of the Rie Ses the meshes of the network frequently
attain a length of 3 mm. and a width of 2 mm.; the included
space frequently attains a depth of I-1.5 mm., narrowing toward
the bottom, so as to form an irregular funnel- like pit.
& Type specimen in the possession of the writer.
KENTUCKY: Sample, Breckenridge County.
Dayton, OHIO
Explanation of plate 33
DICTYOPHLOIS RETICULATA Foerste
Type specimen, collected at Sample, Breck idge County, Kentucky. Natural
size.
a 2 ; : i
oats » Be:
: ar %
"| 4 7 eal
we iB i
cis ; : /
= i
ais ™ ae K,
; %
mq at * 2 a!
INDEX TO AMERICAN BOTANICAL LITERATURE
1915
The aim of this Index is to include all current botanical literature written by
Americans, eerie in veneers or based upon American material ; the word Amer-
ica being used in the broadest sen
a i and papers that ene a to forestry, agriculture, horticulture,
manufactured products of Ap die origin, or laboratory methods are not included, an
attem mpt is made to index the literature of bacteriology. An occasional exception is
n favor of some paper appearing in an American periodical which is devoted
hcl to botany. Reprints are not mentioned unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciate
This ae ge is reprinted pti: on cards, and farnished { in this form to subscribers
at the e of one cent for each card. Selections of cards are not permitted ; each
ena must take all aan published during the term of his subscription, Corre-
spondence relating to the card issue should be addressed to the Treasurer of the Torrey
Botanical Club.
Aase, H. C. Vascular anatomy of the megasporophylls of conifers.
Bot. Gaz. 60: 277-313. f. I-196. 15 O 1915.
Andrews, A. L. Odontoschisma Macounii in Iceland. Bryologist 18:
ST, $2... 21-5. 1915.
Andrews, A. L. Calypogeia arguta from The Faroe Islands. Bryologist
18: 94,95. 200 1915.
Andrews, F. M. Die Wirkung der Zentrifugalkraft auf Pflanzen.
Jahrb. Wiss. Bot. 56: 221-253. pl. r-+f. 1, 2. 1915.
Atwell, C. B. Abnormality in Lilium philadelphicum. Torreya 15:
231, 232. 7. 3.4. IOEm
Babcock, D. C. Mushrooms—edible and poisonous. Ohio Agr. Sta.
Circ. 153: 89-92. f. 1-3. 1 My 1915.
Babcock, E. B. Walnut mutant investigations. Proc. Nat. Acad. Sci.
I? 535-537. O 1915.
Bailey, F. D. Powdery scab of potatoes in Oregon. Science II. 42:
424, 425. 245 1915.
e, A. L. The index of foliar transpiring power as an indicator
of permanent wilting in plants. Bot. Gaz. 60: 314-319. f. I.
15 O 1915.
Balfour, F. R. S. The trees and shrubs of the Pacific coast. Jour.
Roy. Hort. Soc. 41: 21-27. f. 13-24. Au 1915.
Bartholomew, E. T. A pathological and physiological study of the
black heart of potato tubers. Centralb. Bakt. Zweite Abt. 43:
609-639. pl. 1-3. 4 Je 1915.
680 INDEX TO AMERICAN BOTANICAL LITERATURE
Berry, E. W. Pleistocene plants from Indian Head, Maryland. Tor-
reya 15: 205-208. f. 7. 28S 1915.
Blake, S.F. Anew form of Polygala polygama. Rhodora 17: 201, 202.
O 1915.
Polygala polygama Walt. forma obovata, n. forma.
Blake, S. F. Two new Zexmenias. Jour. Bot. 53: 306, 307. O 1915.
Bovie, W. T. A tropism chamber. Bot, Gaz. 60: 320-322. f.I, 2.
15 O 1915.
Brinkman, A. H. Rambles in the Canadian Rockies.—II. Banff.
Bryologist 18: 49-51. 21S 1915.
Britton, N. L. Studies of West Indian plants—VII. Bull. Torrey
Club 42: 487-517. 13 N I9I5.
Includes description of 36 new species in Scleria (4), Badiera (4), Tricera (9),
Plumiera (4), Guettarda (7), Coccolobis (1), Portulaca (1), Chamaecrista (1), Rham-
nidium (3), Nashia (1), and Exostema (1).
Buller,A.H.R. Die Erzeugung und Befreiung der Sporen bei Coprinus
sterquilinus. Jahrb. Wiss. Bot. 56: 299-329. pl. 2,3 +f.1, 2. 1915.
The Thelephoraceae of North America—IV. Ann.
Missouri Bot. Gard. 2: 627-658. pl. 2z. S 1915.
Carpenter, C. W. Some potato tuber-rots caused by species of Fusar-
ium. Jour. Agr. Research §: 183-210. pl. a, b, r4-19. 1 N 1915.
Chivers, A.H. A monograph of the genera Chaetomium and Ascotricha.
Mem. Torrey Club 14: 155-240. pl. 16-17. 10 Je 1915.
Clinton, G. P. Report of the botanist for 1914. Chlorosis of plants
with special reference to calico of tobacco. Ann. Rep. Connecticut
Agr. Exp. Sta. 1914: 357-424. pl. 25-32. Au 1915.
Cockerell, T. D. A. Some plants from the vicinity of the Arapahoe
glacier. Torreya 15: 202-205. 28S 1915.
Cockerell, T. D. A. Specific and varietal characters in annual sun-
flowers. Am. Nat. 49: 609-622. f. 1, 2. O 1915.
Davis, B.M. Additional evicenee of mutation in Oenothera. Am. Nat-
49: 702-706. N 1015.
Dodge, B. O. The effect of the host on the morphology of certain
species of Gymnosporangium. Bull. Torrey Club 42: 519-542.
pl. 28, 29. 13 N 1915.
Duggar, B. M. Rhizoctonia crocorum (Pers.) DC. and R. Solani Kiihn
(Corticium vagum B. & C.) with notes on other species. Ann.
Missouri Bot. Gard. 2: 403-458. f. 1-9. S 1915.
Dunham, E.M. Dicranwm fuluum with double setae. Bryologist 18:
95. 200 10915.
Durfee, T. Lichens of the Mount Monadnock Region, N. H.—No. 6.
Bryologist 18: 51. 21 S 1915.
INDEX TO AMERICAN BOTANICAL LITERATURE 681
Edgerton, C. W. Effect of temperature on Glomerella. Phytopathol-
ogy 5: 247-259. f. 1-4. O 1915.
Elmer, A. D. E. Notes and descriptions of Zingiberaceae. Leaflets
Philip. Bot. 8: 2885-2919. 25 Au 1915.
Twenty-two new species are described.
Evans, A. W. Notes on North American Hepaticae.—VI. Bryologist
18: 81-91. pi. 1+f. I-g. 200 1915.
Includes Jamesoniella heterostipa and Cololejeunea tuberculata, spp. nov.
Fernald, M. L., & Wiegand, K. M. The genus Euphrasia in North
America. Rhodora 17: 181-201. O 1915.
Includes Euphrasia disjuncta, and E. hudsoniana, spp. nov.
Fletcher, W.F. The native persimmon. U.S. Dept. Agr. Farm. Bull.
O88: 1-28. J. 1-17... 12.0 10s
Francis, M.S. Double seeding petunias. Jour. Heredity 6: 456-461,
j. 6-8. 27 S 1615.
Frost, H. B. The inheritance of doutienee’t in Matthiola and Petunia.
—I. The hypotheses. Am. Nat. 49: 623-636. f. 1. O 1915
Gates, F.C. Relation of sunshine to the habitat of Rottboellia sealiali
(Poaceae). Torreya 15: 209. 285 1915.
Gleason, H. A. Botanical sketches from the Asiatic tropics. III.—Java.
Torreya 15: 187-202. f. 18-22. 28 S 1915.
Gloyer, W. O. Ascochyta clematidina, the cause of stem-rot and leaf-
spot of Clematis. N.Y. Agr. Exp. Sta. Tech. Bull. 44: 3-14. Pl.
I-5. Au 1915.
Greenman, J. M. Monograph of the North and Central American
species of the genus Senecio—Part II. Ann. Missouri Bot. Gard. 2:
573-626. pl. 17-20. S 1915.
Grout, A. J. Some common mosses. Nature-study Rev. 11: 204-208.
Ap 1915;—II. The hair caps. Nature-study Rev. If: 229-232,
f. 1-4. My 1915;—III. Three hypnums. Nature-study Rev.
Ir: 336-340. O 1915. [Illust.]
Harper, R.M. Vegetation types. Ann. Rep. Florida Geol. Survey 7:
135-188. f. 54-70. I915. :
Harris, J. A. An extension to 5.99° of tables to determine the osmotic
pressure of expressed vegetable saps from the depression of the
freezing point. Am. Jour. Bot. 2: 418, 419. O 1915.
Harris, J. A. The value of inter-annual correlations. Am. Nat. 49:
707-712. N 1915.
Hasse, H. E. Additions to the lichen flora of southern California—
No. 11. Bryologist 18: 92-94. 20 O 1915.
Hawkins, L. A. The utilization of certain pentoses and compounds of
682 INDEX TO AMERICAN BOTANICAL LITERATURE
pentoses by Glomerella cingulata. Am. Jour. Bot. 2: 375-388.
O 1915.
Hedgcock, G. G. Parasitism of Comandra umbellata. Jour. Agr.
Research §:133-135. 18 O 1915. :
Hedrick, U. P., & Anthony, R. D. Inheritance of certain characters of
grapes. N. Y. Agr. Exp. Sta. Tech. Bull. 45: 3-19. Au 1915.
Henry, J.K. A hybrid rose. Ottawa Nat. 29: 78. O 1915.
Rosa gymnocarpa Nutt. X R. nutkana Presi.
Hibbard, R. P. The question of the toxicity of distilled water. Am.
Jour. Bot. 2: 389-401. O 1915.
Howe, M. A. Report on a visit to Porto Rico for collecting marine
algae. Jour. N. Y. Bot. Gard. 16: 219-225. 2N I915.
Howe, M.A. The Twentieth Anniversary Celebration of the New York
Botanical Garden. Jour. N. Y. Bot. Gard. 16: 203-219. pl. 163.
2N 1915.
Howe, R. H. Jr. The genus Cetraria as represented in the United
States and Canada. Torreya 15: 213-230. f. r-ro. 4N 1915.
Howe, R.H. Jr. The Usneas of the world, 1752-1914, withcitations, type
localities, original descriptions, and keys. Part I1—South America.
Bryologist 18: 38-43. 15 Je 1915; 52-63.f. I. 215 1915.
Jones, D. F. Illustration of inbreeding. Jour. Heredity 6: 477-479.
dy FO; Tdi ae oe S08 6,
Keitt, G. W. Simple technique for isolating single-spore strains of
certain types of fungi. Phytopathology 5: 266-269. f. zr. O 1915.
Kraus, E. J. Germinating pollen. Science II. 42: 610, 611. 29 O
Lewis, C. L. Plant breeding problems. Jour. Heredity 6: 468-470.
IQI5.
Lewis, I. M. The trees of Texas. Univ. Texas. Bull. 22: 1-160.
f. 1-48. Ap 1915.
An illustrated manual of the native and introduced trees of the state.
Loeb, J. Rules and mechanism of inhibition and correlation in the
regeneration of Bryophyllum calycinum. Bot. Gaz. 60: 249-276.
f. Yogr. 15°O 1915:
Ludwig, C. A. Notes on some North American rusts with caeoma-like
sori. Phytopathology 5: 273-281.- O 1915.
Includes Caeoma dubium n. sp.
Lynch, J. J. Sambucus pubens var. xanthocarpba. Am. Mid. Nat. 4:
77-181, $i. 77,12. S 1935.
Mason, S.C. Dates of Egypt and the Sudan. U.S. Dept. Agr. Bull.
271: 1-40. pl. 1-16 +f. 1-9. 28S 1915.
INDEX TO AMERICAN BOTANICAL LITERATURE 683
Mattoon, W. R. The southern cypress. U.S. Dept. Agr. Bull. 272:
1-74. pl. r-12 +f. 1-7: 27 S 1915.
Meinecke, E.P. Spore measurements. Science II. 42: 430,431. 24S
1915.
Melchers, L. E. A new alfalfa leaf-spot in America. Science II. 42:
536, 537. 150 1915.
Merrill, E. D.- A contribution to the bibliography of the botany of
Borneo. Sarawak Mus. Jour. 2: 99-136. S I915.
Merrill, E. D. On the application of the generic name WNauclea of
Linnaeus. Jour. Washington Acad. Sci. §: 530-542. 19S 1915.
Merrill, M.C. Electrolytic determination of exosmosis from the roots
of plants subjected to the action of various agents. Ann. Missouri
Bot. Gard. 2: 507-572. f. 1-18. S 1915.
Merrill, M. C. Some relations of plants to distilled water and certain
dilute toxic solutions. Ann. Missouri Bot. Gard. 2: 459-506.
pl. 13-16 +f. 1-4. S 1915.
This paper is also published in Am. Jour. Phar, 87: 549-606. D ro1I5s.
Meyer, R. Die verschiedenen Formen des Echinocactus setispinus
Eng. Monats. Kakteenk. 25: 37-39. My 1915.
Mottier, D. M. Beobachtungen iiber einige Farnprothallien mit Bezug
auf eingebettete Antheridien und Apogamie. Jahrb. Wiss. Bot. 56:
65-83. f. I-3. I915.
Newcombe, F.C. Das Verhalten der Windepflanzen in der Dunkelheit.
Jahrb. Wiss. Bot. 56: 511-528. 1915.
O’Gara, P. J. A bacterial wasi of western wheat- “grass. Science
Il. 42: 616, 617. 29 0 19
First account of the occurrence i a new type of bacterial disease in America.
Osmun, A. V. Report of the botanist. Ann. Rep. Massachusetts
Agr. Exp. Sta. 27: 55a-59a. Ja 1915.
Osmun, A. V., & Anderson, P. J. Ring-spot of cauliflower. Phy-
topathology 5: 260-265. f. 1-4. O I915.
Oskamp, J. Soil temperatures as influenced by cultural methods.
Jour. Agr. Research 5: 173-177. f. I-4. 25 O 1915
Peirce, G. Ein multipler Klinostat. Jahrb. Wiss. Bot. 56: 330-336.
pee PE ee
Porsild, M. P. Naturfredning i danske Grénland. Meddelelser om
Grénland 51: 253-263. f. I, 2. 1915. -
Porsild, M. P. On the genus Antennaria in Greenland. Meddelelser
om Grénland 51: 267-281. f. I-7. 1915.
Includes A. glabrata and A. intermedia, spp. nov.
Rolfs, F. M.
ginata, 316, 317; Mariae, 316, 3
eeuctaes ae. 383 pendula, 316, oe
neice aa 316, 317; psittacorum,
316; pulverulenta, 326, 327;
¥7,4255 eg 316,
16, 317; revoluta,
minea, 316, , 327; stricta, 3
318, 325; orcteg ee 318; rilicns.,
316, a villosa, 316, 317; Wagneriana,
316,
gone Soine new species and varieties of,
Blepharineuron tricholepis, 638
Boehm
ria, cylindrica, 188
pera g A
Borrera, 579
B kia, 117-119
Botrydium, 194
gpoweies eee 638, 640; pros-
sivechythoion oe 189
Brasenia Schreberi, 1
Britton, E. G. ope of Bermuda, i :
West Indian mosses—II. osses
so —— West Indies and we
Islands,
BRITTON, 'E. G., & sprees, A. A new
American fossil m
Keren: No Le St a of West Indian
s—VI, 365; —VII, 487
PF breviaristatus, 638; Flodmanii,
INDEX 687
638; lanatipes, 639; hordeaceus, 642;
polyanthus; 632, 638; Porteri, 632,
633, 638, 639; Pumpellianus, 633, 638
639; secundus, 642; Richardsonii, 632,
3, 638
Bryum bimum, 189; blandum, 104; capil-
74; concavum, 402; cp 6; 74;
decursivum, 5; dichotomu 43 -
de ens, ike Pee Riis 6; ripense, 6;
cur
a
Bucegia, 265
Sarrieita: 112; leptalea, 112; microglossa,
II
Buxus acuminata, 499; bahamensis,
501; brevipes, 499; citrifolia brevipes,
oe
clada, 500; Mueller na, 498; retusa,
500; subcolumnaris, pee Wri ehtii, 502
eth as blanda, 636; canadensis,
185, 186, 192
ha, 636 ;
lecta, 636; purpurascens, 634, 640; ‘Scrib.
neri, 636; Suksdorfii, 641
Calla palustris, 204
Callicostella Beccariana, 574; papillata,
74; Belangeriana, 7
pees Afzelii, 1; Breutelii, 35
Dozyanum, 572; hexagonum, 3; Hoo
eri, 3; : lonchophy iim, 3 clacetei.
572; Richardi, 1,
Campanula a aaa 185; rapuncu-
Campylopus bermudianus, 71, 72; in-
Pp us, 393
34 ‘
Capse Bursa-pastoris, 661; Heegeri,
661, 662
Cardiobatus, 1
Carex abdita, 621; acuta, 408, 412, 413,
mutata Ee. cri nita, 188;
198, 199; Goodenovii, 417; marie. Coe
616; grisea, 617; grisea globosa, STs
aminea 603; _
pa naa Bi ye rian tene
Haydeni 406,° 407, 410; segs waar
408; Le avenworthii, 618; laev
19; Muhlenbergii, 618; onusta, 618
pauciflora, 04; reula, ;
Pseudo-Cyperus, 593; trata, 204;
rugos a, 621; scoparia, 192; Shel-
donii, 618; stellulata, 192; stipata, ra
I ve ee, Bie 02) 004, 005, (G51:
minea
a, 606;
stricta, 171, 184-192, 405-417; aioe
angustata, 406, : 19; ricta
urtissima, 406, 415; stricta decora
410; stri oryi, tricta
Haydenii, 410; stricta xerocarpa, 407,
umbellata, 621; vitpintana, 408; virg rs
ex, Notes on, — VIII, 405; — IX, 603
arex straminea Willd. and some of its
nearest allies,
03
Carex stricta Lam. and its allies, 405
Ca 3
rpinus caroliniana, I
ore cordiformis, 193 _
1 A 515
astalia odorata, 171, 180, “181
ica be
ephalanthus icra te 187, 3
zia connivens, 189; flui snag 204
Cephalozi
Gcusonbee this m de bi abi 179, 180
ee
86
ochloa
glauca, 642
Chactomitim papillifolium, 574
amaicensis,
nan aaa 519-530; Sis: 188
Chamaedaphne, 209, 212; calyculata,
201-207
ee 123; anglica, 123, 124;
mes
Chats a ép.. 179, 195, 1
hiloscyphu
55
sosplenium americanum, 188
a
choriaceae, 549
Cichorium Intybus,
Cicuta bulb: Spier vit; mae maculata, 186
Ci
nilaaes. 1893 latifolia, 640
183
aru
Cladium mariscoides,
688 INDEX
Claopodium hawaiense, 576; priono- | Cypripedium acaule, 210
. phyllum,
Claytonia megarrhiza, Dactylis nie gs cane
ane alnifolia, 187, jet hyalina, 261 | Dalibarda, 117; repens
Climacium americanum, 189 | Danthonia californica, pide 636; inter-
Clintonia borealis, 210 media, 631, 40
Coccolobis armata, 365, 366; benitensis, Datura Stramonium, 334; Tatula, 334
366, 370; brevipes, ers 371; calobotrys, | Davallia
367; colomensis, 365, 369; Per ecaden verte, 199, 201
365, 367; costata, 366, 360; C ii, | Delphinium, 62
65, 368; cubensis, 3 ‘ divers Denearie di iphy ylla,
— eo ; flavescens, 367; ri- | De eschampsia par 631; atropurpurea,
i
dan + 306) fie 631; caespitosa, 631, 636; curtifolia,
aula, 65 368; signet cor- 631, 633; elongata, 641; pungens, 642
ngs
5; paucifior
368; pallida, 366, 3715 screens 365, grees flaccidus, pes ‘hematin: 579;
367; cox, 366, 370; reflexa, 3 Seky nn gee
etusa, 365, 368; nage Fon 370; Purana angustiflorum, 93; Blumei.
‘cue longifolia a, 370; .rumicifolia, ; grossialare, 94
514; Shaferi, 366, 360; tenuifoia 371; Dicranum aucklandicum, 94; —
pea ph 3 eer oodfredensis,| 95; flagellare, 189; pumilum gg
vs Weightil, 4 ae m, 9 ium, 190, 2
Piette sa: in Cuba, ihe genus, 365 Dictyophlois reticulata, 675—
Com oo 123;%
Cansecticnt: The vegetation of, —IV.
Plan
Conocephalum
panini “angustfolium, 248; bifolia
7 “s
Dictyophlois reticulata, gen. et sp. nov.,
75
n Didymodon saree 95; orientalis,
chem in lowlands, 169
oss 652
Dioscorea conehyulacta, 592; grandi-
canaden: 356; biflora, 248, 249, 254; folia,
pan tee en 248, oe ae utata, Dipterceteinon, 585, 590; capitatus, 589};
ae 256; hirta, multiflora, 248, pauciflorus, 588, 589
6; par arvitlora, tr ape onsen 248-257 oe O23
vated code x e, 37 Di stichlis stricta, 637
Convolvulus americanus, 38; arvensis, | Distichophyllum microcarpum, 107; Os-
38; repens, 38; sepium, 37 e ai, 74
Coptis trifolia, 188, 210 Drxon, H.N. New and rare Australasian
Cordiobatus, 12! mosses, mostly from Mitten’s her-
Cornicularia, 579 barium, 93
Cornus Amomum, 187, 188 Donce, B. QO. The effect of the host on
Corsinia lamellosa, 278 the morphology of certain species of
Coumarouna, 62 Gymno: osporangi
m,
Cratae, sae = 2I, 24, 230; mollis,.521; lg eria Session 568; umbellata, 568
pun 2I
Coil ere as
Cuba, The genus U-paishs in, 365; T 202
int
ra ae ‘202, 204; rotundifolia,
» 204
= s Guettar . . in, 507; The ee Dulichium arundinaceum, 171, 183, 204
Plumiera L. in, 503; The genus tinge et atpaearinem: r2t
Berg. ry spre The genus Tricera S
497 Eatonia intermedia, 636; pennsylvanica,
Cucurbitaceae, 349 636, 637
( bata, i Echium vulgare, 4
Cuscuta Gronovii, 38 pero tecrye det cece: 575; Moritz-
se re ls sonii, 77 $75
Cyan ‘w Hawaiian, 77 Effect. of the host on the morphology of
¢ Lorain 7% ~~ 74 certain species of. Gymnosporangium,
Cylactis, 120, ; lyncimontana, 120,) The, 519
125; ebfce ete eS Eleocharis acicularis, 191, 194; palustris,
Cymbalaria Cymbalaria, 3 204; Robbinsii, 179; tenuis, 185;
Cyperu: appt deci 3873 eee thermalis, 642
fates: 387; strigosus, 186, 191, oh Elodea canadensis, 179
INDEX
Elodium paludosum, 189
Elymus wit da us, 641; canadensis, 639;
condens. _ 638; glaucus, 632, 637,
641; in us, 640; simplex, 632, 637;
sca ritieiton 638
Bndephy llum poiee sag Abe
rips ae Pa 58, 59,
Endophy llum, A new Neca American,
55;
5
Epilobium homey 186; sp., 204
dense,
Erigeron annuus, 560; can 570;
shitedetniiede: 569; ssaleheltie 569;
pusillus, 570; ramosus, pons
Yrioeauled septangulare, I9I
Eriocoma cuspidata, 641
— si um sp., 204; virginicum, 186,
éythehen Pickeringii, 30; spicata, 30
his rsa agia: americanum, 193
II2
Eubaeria
Eucladium, 71; bichrpareniair 73
Eupatorium aromaticum, 557; hyssopi-
maculatui, 556; perfoli-
86, - pubescens, 557
purpureum 556; rotundi-
folium, $57 ve ena um, 557
Eupolypodium,
Euthamia stated te +P 564; minor, 564;
nar 564
vans, A.W. The genus Plagiochasma
and its North American species, 2
Evernia, 579
pele stenophyllum, 517
Fabronia andina,
ene sranitttedia” ae
; mea occidentalis, 593
cba ike 13 ie A otes on Michigan
Liliaceae, 351: Notes on ip Michigan
a4 of Polygonatum, 2
Ferns and flowering plants of Abacus.
The. —XI V, 275 V, 331; —— V1,
549
Festuca — oe 633, 639; ceeehsy-
phylla, 640; pestris, 638, 640
confinis, 639; scr ie 634; elatior,
637, 641; rhe ay 637, 639; ingrata,
ae 633, 6393 pres 640; Sa umien:
; octoflora montana, 633,
en 639, 640; Thurber, 632, 633, 638,
poet 190; elegans,
Kegeli-
3; ri, as Ete 5723
anus, 2; minutulus, 73; Mitteni, 5723
palm atus, at tesitvliak | 3; Zippelianus,
572
Floribundaria floribunda, 574
ForerstE, A. F. Dictyophlois reticulata
gen. et sp. nov., 675
Fossil moss, A new American, 9
Fraxinu: eri , 20s -arpomiala, Is
689
reat a 30; nigra, 188; pennsylvanica,
Fone, F.D., & ArtHur, J.C. Anew
rth American reread um, 55
unaria, 71; calvescens, 401; flavicans,
| hygrometrica, 14 401; macrospora,
401; suber
Further th ao Badiera DC., 494
Galeopsis ce 42; Tetrahit, 42
Galium Claytoni, 189
mg fee baccata, 204, 210, dumosa
oviana, 204
nog "30
Gentiana crinita, 192
Genus Guettarda L. in
59
Genus Plumiera in Cuba, The, 503
Genus Teloschistes in North America,
he, 579
Genus Tricera Sw. in eee The, 497
Georgia pellucida, 190, 2
Gerardia decemloba, re paupercula,
338; maritima, 338; pedicularia, 337;
Legsbiteabars 337; Skinneriana, 339
um rivale, 186
Glaus iarieiealk 28; maritima obtusi-
folia, 28
pes eda oar 42
eee ra cages fag 185; nervata, 192;
feiyphncaentan Piitieleiene: 9
Goch cifolia, 38 :
Ba
347; palustre,
tinctorium, 346;
florum, 345
Gian fluminensis, fs limbata, 222,
223; linearis, ; marginella, 221;
quaerenda, 0
ge inc nghei 635; paerienn 631, 636;
Shearii, 636; Wolfii, 631, 6
Gratiola aurea, I9I, 33
GR IGGS RF, Some new species and
trifidum, 346; tri.
Re 5
ta, ~ saxatilis, 400
Grimmia micr
GROSSENBACHER 1. G. edullary spots
their cause, 227
Guettarda ambigua, 514; bracteata, Ria:
brevinodis, 508, 513; icola, 508,
513; calyptrata, 508, 512; camagueyen-
sis, 507, 509; ¢ 8, 507, 510;
ina, 507, 508; Combsii, 508, 514;
crassipes, 507, 511; cu i 507,
509; a, 507, 5 ino-
ferruginea, 507, 511; havanensis, 514;
ocarpa, 509; inaequipes, 508, 513;
690 INDEX
lanuginosa, ee 511; Lindeniana, $08, | | | Hordeum boreale, 636; jubatum, 631,
513; rele Sit: oc 4 636, 638; nodosum
508, 512; pone and atl He S533 sere a Shebe> coerulea, 343; longifolia, 344
5133 retusa, 508, dg rigida, 507, | Howe, R.H., Jr. The genus Teloschistes
509; rugosa, 514; scabra, 508, 514; in North America, 579
sciaphila, 507, 5143 undulata, 507, 500; | | Hypericum mutilum, 191; virginianum,
Valenzuelana, 508, 512; xanthocarpa ue ites 20
507, 510; zygophlebia, 50 = ise shane nn 651
Guettarda L. in Cuba, The genus, 507 eas omum "Breutelii, I, 3; cubense,
Gymnoconia, 55; interstitials, oe 59, 60 | 3 as m
aem
fe) a
80; refracta, 85 pega, 86: eon commutata, 572; uliginosa, 4
Warcewiczii, 80, 8 |/Hypnum Antillarum, 2; Breutelii, 2;
Gymnosporangium Blasdaleanum, 520 neckeraides, 1053 avadcian, 107;
521; biseptatum, 519-542; biseptatum | Schreberi
foliicolum, 520; clavip 51 2 at pothierts idee
1 a, 549
529; Ellisii, 523, 528, 530; | Hyp opterygium SAN 574
; 25;
macropus, 519, 521; nidus-avis, 519 , Idaeobatus, 126, 5
gare Gear The effect of the host | [ex vertic aha se , 188, 205
n the morphology of certain species | Ilysanthes dubia, <% gratioloides, 336
wri 519 si eseepae oe a, 185, 1
Gymnostomum Breutelii, 3; senocarpum, Index to American Piet literature,
49, red 161, 241; 300, 423; 481, 543,
ynoxys, 380 595, 643,
Gyroweisia, 72; Barbula, 73; boliviana, reaping nails, 568
394 | Ipom a purpurea, 37
Iris venir. ree 186
SaaS rp ataatace tit 204 puting: » 725 oe iy foes
Hag ci
‘ans, 75; Somer 7i,
H . Notes on Baeria and} Iva paspiaion 5543 oeital 554; xanthi-
Tesiieahe Itr folia, 554
Se virginian:
Hance, R. T. Pollen gin rate and | Jacandra Cowellii, 392
ieenere in Zebrina pendula, with | Juncus acuminatus, 185; canadensis, 185,
special reference to the chromosomes, effusus, 185, 192
63 ungermannia australis, 278; limbata,
Haplocladium microphyllum, 8, 74 278, 281
agian Jamesoni, 401; seriolum,
Kalmia angustifolia, 204, 210; latifolia,
Hawnas oa as Pte the patito and 7
the staminate | Koeleria gracilis, 638, 640; flexuosa, 43;
ont gold haven | in the inflores- mutica, 44 Torreyi, 44; peaticilats.
cence of aroids Arisarum vulgare and 44; virginiana, 44
Arisarum proboscideum, 663 igia virginica, 549
Labiatae, 41
angusta barqueta, | Lacinaria scariosa, 558
3245 a Foe zi _braslenss, 330; Lactu canadensis, 551; hirsuta, $53;
51
30; 330; pul
lenta, 326; hae + pee stricta, 325 Lamiu Oa dete
Hesperopeuce Mertensi
ana, Lappula Lappul
Hibiscus Moscheutos, 431-4493 oculi- | Larix lari ; i Lyallii, 314, 17;
roseus, 429-449 occidentalis, 14, 24
Hieracium canadensis, 551; aurantiacum, | Lasthenia, 111, 116; conjugens, 114, 1T5;
552; Gronovii, rs mMarianum, 552, glaberrima, 116; mi I16
553; scabrum, 552, 591; venosum, | Lasthenia, Notes on Baeria ae rt
552, 553 Ledum groenlandicum, 204
HOLtick, a? att bg E.G. A new) Leiberg, Mosses of the Philippine and
Ameri ossil mo: Hawaiian seausge collected by the late
Saieioes ik um, 95 John B.,
Ho maliodendron microdendron, 574 Lemna, 182
INDEX
Lentibulariaceae, 3
Leo nhs cops er ee A 550; Tarax-
» 550
Pecan Cardiaca, 43
he tana ane 675
Leptamnium virginianiuin
Teco camera i ie, as pu usillum, 570
Leptobry yriforme, 401; Wilsoni, 401
Leptodon me gracile, ae
Leskea gracillim
Leucobryum, ns davis ucum, 72, 189, a
javense Hii Polakowskyi, 2; sanctum
572; sericeum, 572
eucslema Riedlei, 2
Libocedrus, 520
Lichen, 579; chrysophthalmus, 579, 581;
flavicans, 580
Liliaceae, Notes on Michigan, 351
Lilium canadense, 352, 253; Mic
ganense, 353; michiganense sinbelli-
f » 353; michiganense uniflorum,
353; peramoenum, 354
Lilium, New species of, 352
imonium carolini > 2
ai tenuifolia, 3
Lin ach ag ae " Gilbsieth: 3343
n 335; vulgaris, 335; vulgaris
ceieeti. 4, we
Linnaea americana, 34
ae. communis,
45; membranace ie. 453 rubellus, 453
sessliflorus, 46; uniflorus, 45; virgini-
S, 45
poems asters ere 34
Lyonia ligustrina, 187, 1
Lysimachia N eg? 27; quadrifolia,
27; terrestris, 27, 185, 186, I9I, 204;
vulgaris, 27
MACKENzIE, K. K. Notes on Carex—
VIII, 405; —IX, cae
Macromitrium abbreviatum, 100; erosu-
lum, 97-101; ep ances 100, I01I;
ligulifolium, x m, 99; ortho-
stichum, 573; papillifolium, 100; per- | Munro
papi iei,
losum,
es-
um, 574;
canadense, 188, 210, 354,
356; canadense interius, 354, 355
691
Sea O ty moat 276
Man
cea a. fae agg 273
Ma mea ium vulgare, a
Ww. R “The orth American
peas of Siieeane 79; Poly-
emp marginellum and its immediate
allies, 219
Mayepea cuben
Mecosorus aE OTE 22
Medullary spots a their cause, 227
Melampyrum line
Melanobatus, rog as bernardinus, 129;
ne oe 120; — — 122;
122; niger 0
eo bulbose a, 636; Patol ‘se ans
spectab am nk subulata, 637, 6
Melissa ere
entha cai sone 46; 473
pape: 46; piperita, 46; rotundifolia,
46; spicata, 46
‘ensanthen: 34; trifoliata, 34, 204
Mertensia, 62
Micrampelis lobata, 349
rob:
Mic
Mielichhoferia inl Seg 401
Mikani ndens, 557
utivum,
» 344
Mittenothamnium dimin 7
oe 189; punctatum,
nium cinclidioides,
Monopterys, hoe 625; alge oey 623,
6 24-626; Uau 25
Monopteryx henaa On the ha
sia relationships of the genus, 623
Mosses from the west coast of South
(ex erica, 393
Nseacs. mostly from oe s herbarium,
tralasian, 93
a, 72
Philippine and Hawaiian
Ssrwes collected by the late John B.
g, 571
ie "West Indian, II. Mosses me
Danish West Indies and Virgi
lands, I
Muhlenbergia aristata, ne comata, 631,
636; ae 634, 640; filiformis, 636;
gracl odin: en
baci oar; racemosa, 631, 636;
Bir hardsonis, 636, 638; simplex, 637;
uarrosa, 636, 638; subalpina, 633,
yi 639, 640; Thurberi, 641; Wolfii,
631, 637; Wrightii, 636
a squatrosa, 641
Myosotis — 40; palustris, 40;
scorpioides
Myrica pagar 205, 210
Nabalus serpentarius, 553; trifoliolatus,
553
692
Najas flexilis, 179, 1
gies eee fee fhe and pveabten pong
aaa —XV, 331; XIV, 5
Be? capiias S, 517
ha disticha, 7; jamaicense, 7
s crinita, 5743 t cenicha: 574
Ss
Nemopanthus mucronata, 205
Nepeta Cataria, 42
New merican fossil eee A,9Q
Ne are Australasian ——
ines soo acc s moar , 93
ew Hawaiian Cyanea, A, 77
New North pret hee ‘Endophyllum, A,
nee species of Lilium,
ew species of Uredineae—-IX, 585
New varieties of Allium canadense, 351
Nicandra a hnnaaai 331
CHOLS, G. e vegetation of Con-
necticut —1V. Plant societies in low-
lands,
North haces species of Psilogramme,
The, 79
Notes on Baeria and ar iagiey III
Notes on Carex—VIII, 405; —IX, 603
. vanes on Re ister 35I
on Rosaceae—IX, 117; —X, 4
Note = shines “Michigan species of valde
gona
Notas 6 on nT chanel AK. The identity
Tri es pyxidiferum, 651
Nyssa, 193; sylvatica, 193
Conon ae albidum, 2, 572
Oenothera alta, 442; biennis, 443; Cock-
Oueae sna tse
Onosmodiu anum
bers ane relationship of
ryx ce,
mee ae
of
the sexes (staminate ane pistillate
flowers) in the inflorescence of the
aroid and
120; deliciosus, 119;
mexi $, I19; rubicundus, 119;
trilobus, 120
Origin of dwarf ones as shown in a sport |
i i Th
eae e, 429
Orobanchace
Orobanche aloes 342
INDEX
Orontium aquaticum, 18
Orthomnium stolonaceum 574
a hess ns tone 640; exigua, 639,
641; antha, 637, ae
Osmnunda cinnamome 188; regalis,
I
OsrENL, . H. Ruppia anomala s
ig eat saci type of the Pota-
getonaceae, 65
Cntige ah iiea. T4, 25
Otionia, 261; italica, 278; rupestris, 278
Pallavicinia sage 189
Panax trifolium, 193
Panicu laria peanx 630, 635; borealis,
635; elata, 635, 637, 641; Holmii, 637;
nervata, 631-6 pauciflora, 630-641
Panicum agrostoides, 191; barbipul-
vinatum, 641, ; capillare, 642;
pacificum, 636; thermale, 642
Paramena, 122, 126; Menziesii, 126
Parmelia, 579; villosa, 582
Pasture trees and shubbery, A study of,
Pedicularis canadensis, 340
Pelekium velatum, 575
Pellia epiphylla, 189
Peltandra rhea 183
Petunia axillaris, 334
ae arundinacea 63
Phascum Floe num, 5; sessile,
Philonotis catiole 104; pasar: urcu-
Bt ss se Pine ana, 189; fragilis,
arpa, 6; tenella,
Phiedat er 631, 636; cata 631,
Phragmites Phragmites, 635
Phrygilanthus Sonorae, 587
Phyllothallae, 579
Physalis naire aek Hi ee Liters 331
Physalodes ysalodes, 3
Physcia, ane
Physcomitrium Rose
00
o- he fit ntact token on
Mo
the R
qa Forests of ou
Su sara ‘ead Montane Zones, II;
—V, Grasslands hes “ne Subalpine and
ontane Zones, 629
Picea albertiana, 14, 22, 24; canadensis,
22, > has - Seinggeiaia 13-25; P ana,
13-2 ariana, 202, 203; rubra, 2
Pinus aipiecwie rat aristata, 13-193
edulis, 13; 7 ticola,
389
he characters and rela-
INDEX 693
Plagiochasma, 2593 abyssinicum, 262 +
pai ia, 259, 2 278; algericum, 262, |
287; tinea: 250-305; a |
latum, 262, 263, 270-276; asm
4 ilie a: Vebl
262; brasiliense, 262; rocarpum, |
260; Colsmannianu 260; cordat
2590; 305; crenulatum, 260, ag
05; cuneatum, 247; dschallanum, 287; |
elongatum, 262, 277, 85
9, 305; eximium, 27 305;
fissisquamum, 261; intermedium, 60,
262, 260, 278, 280 maicen
86, 2 1,28
nepalense, 261, 287; naratoninis 260;
pedicellatum, 260; ede ere 259,
TOS mh 36s:
2 ru 2 te)
Schimperi, 262; subplanum, 287; tenue,
261, tes 288; validum, 260, 287;
Wrightii, 260, 277, 294-30
Plagiochasma nd its North American
s, The genus, 259
We isiucals — 10; Scudderi, 9, 10
, aes
Plantago arista 343; decipiens, 3435
halophila, 343; end 343; major,
Plumiera tices nts sie 504; confusa,
wh emarginata, , 505; emarginata
cifolia, 504; alifous. 503, 504;
a onic vg 506; lanata, 503, 504;
» 503, 505; obtusa, 503, 505;
obtusa yer 505; obtusa tooo
505; sericifolia, 503, 504; trinite:
osa
3, 506; venosa, 503
Plumiera L. i ube; The genus, 503
Pneumaria sepstrere!
Poa alpina, 631, 636; annua, 642; aperta,
633, 639; Buckiyes i 632, 638;
callichroa, 631; compressa, 640;
fusa, 633, 634, 638; crocata, 632-640;
ot ke _ 634, pe epilis, 633, 634,
Fendleriana, 639, 640; in-
poi as ogres pees gg "638, 639;
leptoco ghee — 636; longiligula, 632
39; longi nculata, 633, 639; longi-
pila, 6s6r, terete 632, 63 8; macroclada,
639; nervosa, 631, 636; aioasenaie
dentali i:
6:
pudica, 631; en ane 636; rupicola,
ae Epebgepnen es 633," 639; Sue aan
Het triflora, 636; Vaseyana, 631
ks 03
HP sheep polycarpum, 402
Pogonia ophioglossoides, 185, 204
Pohlia, 6; elongata, 103; rg es 102;
nova ~seel ndiae, 102; nutans, 102;
i
| 74
, Pollen development and degeneration in
ebrina pen abate prvi special reference
to the chro mes, vn
olgeeaaiaer aa americanum, 240;
biflorum, 247-257; biflorum ovatum,
352, 355; biflorum Segara, 262.
254; boreale, 249-253 trale,
252,. °253; canalicatum, 249-2553
canaliculatum am num, 252, 256;
caniculatum ene Coe :
comm t
futatum, 249, 251, 256; cuneatum,
Aq. 2 I, 253; ellipticum, 252,
255; giganteum, 249, 251, 256; hirtum,
oct, 282: ifolium, 249, 251, 256;
miultiflorum, 249, 251, eeasld, parviflorum,
251; pubescens, 249, 25 0-253; pubes-
-cens cuneatum, 252, 253; virginicum,
249, 251, 254
Po phates Notes on the Michigan
of, 247
oly foaieis arifolium, 188
Polypodium ebeninum, 221, 224; flumi-
nense, 223; gramineum, 219, 220, 205;
Hessii, 221-224; leptopodon, 220, 222;
limbatum, 220-224; yer ioe 219-
225; nigrolimbatum, 2
Po weap seg liane” pes! its im-
iate allie
Po apogee monepeliensi, 642
Polytrichum co ne, 189, 207
ene niga cordata, 182, 183
Popul minata, 25; ‘angustifolia, 2I-
253 Saeed 21-2 ; Besseyana, I4,
24; Fremo 13; hastata, 14, 24
panera 630; sablitellind. ¥78, T7992
nei 181; pois phyllus, 178,
oa es 179; natans, 179; pecti-
natus, "179; pasting 178; Richardsonii,
2
Proserpinaca otha 183, 1
Prunella pennsylvani a ere » 42
Prunus, 230; avium, ai domestica, 231;
mel at
Psedera quinquefolia, 188
| Pse
ocrossidium, 396; Pac mia 20.7%
chilense, 396; excava
Pseudoleskeopsis decurvata, ae
694
Pseudotsuga mucronata, 13-24
Psilocarya nitens, 387; portoricensis, 387
Psilogramme conges esta, 79, 81; chiapensis,
sa, 85;
glaberrima, 83; ;
Dasmatoder, 80, 84; ‘hi
6; Watecewiczii, 70, 80, 81; villosula,
Palgrammne, The North dsaceeas species
papi: 585; physodes, 587; Purshii,
Psychrobatia, 123, 124
Pterobryum angustifolium, 6
Pterocarpus, 625, 626
on, ae:
Pterogoneurum Roseae
Siepecnioiiia: dstichonhtoides, 106
hata MS Sesh s
Ptilomeris,
Puccinelli iroides, 637
Pucci holy 590; Andropogonis,
338; ingle 587; Claytoniata,
591; scham mpaine, sue Dondiae, 585,
592; ieee 521, 538; Fraseri, 585,
591; Hieracium, 585, 591; Lojkojana,
590; malvacearum, 60; nodosa, 588,
5890; Feneacenas: 585, 589; lee.
585, 59
Pucciniosira, 5
Purdi
velutin
Pycnan the std incanum, 44
Pyrus americana, ak Been ve nmearg 187,
205
eget rae 93; Gambellii, 13; macro-
5; palustris, 193; undulata,
Racelopus pilifer, 574
Ranunculus aquatilis, 180; septentrio-
nalis, 186
pest 271-274; oMyperge sagen 288,
293; maderensis, 278, 2
ie ty an 72: peshbesress ae tomen-
osum,
ee tae Pact 516; ellipticum,
6; Shafi
Rincei Hess Li
192
esac da mancisn iti, 188; viscosum,
aie sini
Ribes, 228
Ricciella fluitans, 189
Ricciocarpus natans,
Rock, J. F. A new acai Cyanea, 77
|
iaea Bicone, 380; Shaferi, 389;
INDEX
oo Botryapites, 520, 522, 528, 536;
Ellisii, 536; transfo mera 525, 520, 536
Rolland gle pi
aceae, Not oe 117; —X, 463
po carolina, eis
Rubiaceae, 3
ubacer, a Pea tomentosus, I19;
velutinus, 119
| Rubus aboriginum 156; abundus,
mck oi : 133; acaulis, 124; ne
X stellatus, 464; acaulis X pubescens,
464; a i I I 142
aegopodioides, 125; allegheniensis, 149-
I51; allegheniensis X argutus, 467; al-
nifolius, 144; alpinu 8, 142, 143; ama-
bilis, 153; americanus, 120, 125, 149}
amicalis, 153; amicalis XX nigricans,
476; amicalis XX nigrobaccus, 468;
amicalis x per us, : i
us X pu
ms, 464; arcticus XX saxatilis, 464;
arenicola X hispidus, 473; arenicola,
55; argut xX Bail us, 469; ar-
gutus X ensis, 471; argutus XX
earpinifolius, 470; x cunei-
folius, 465; argutus Enslenii, 469;
t flagellaris, 474; argut x
hispidus, 479; argutus X nigricans,
= ; epee nigrobaccus, 466;
x id 4743 ss gsi
473; Bailey x A773
Pano x asdtated pe Ba iley-
mbens, 472; bernardinus,
120; betulifolius, £SEs Boyn ton mi, 1553
er
» 1373 a an
ngeo cu
os
; X frondosus, 470;
Enslenii x aolies: 475; Enslenii X
INDEX
procumbens, 475; eriocarpus, 28;
fagifolius, 121, 130; 1433 ferruginens,
136; spate Seth I sti Lapa X nigri-
cans, 4773 , 472; floribundus,
142; flori ie spe floridus, 151;
florulentus, 14As 145; Fockeanus, 1243
466
Hie oa sonny apn frondosus x h
u 47, I51
glabratus, eae: caaticastae ea glauci-
folius, 129; glauc us, 127, 128; leah
bachii, 144; Groutianus, 158; Gun
anus, 124; guyanensis, oe TA3; Heller
146; hesperus, 125, 129; heter piyii,
157; rerhaiede gh nigrobaccus ~
hetero hyllus 4743
pidus X n
plicaifotius,
ntan 478; h 8). 15450-3551
huimistratu, 154, 158; pel 121-135;
oenicolasius, 464; idaeus pec-
tabilis, 464; idaeus xX strigosus, 465;
X ursinus, 464; invisus, 155;
Linkianus, 1473 lucidus, 152, 159, ee
6, 147; macropetalus x viburnifolis,
‘is 5; melanolasius, melan
35
grobaccus, 4! cans pli-
catifolius, 473; nigricans X procum-
ns, 477; nigricans X Randii, 477;
nigricans recurvans, 476; nigrican
X sativus, 466 ni ns X vermon
tanus, 477; nigrobaccus, 147-156;
nigrobaccus x pergratus, 468; nigro-
baccus X Randii, 468; oeabeen ect x
esha es 473; nigro! Sm ver
us, 469; nivalis, 792." 190:
dace IIo difl
122; occidentalis, 125-130; occidentalis
xX procum 465; occidentalis xX
rosaefolius, 4643 occid seals X strigo-
sus, hg oratus, 117, 119; oligo-
oli
spermus, 154; ostryifolius, 469; Pal-
695
meri, 143; pacificus, Sap yoo. tee,
118; parviflorus, 118, Paxii, 464;
ete rpc 476; :
ramoenus, 134, 135; Dp
vcralemie 59,470; piadelphicus 52;
philyr oe llus, #395 colasius,
27; illatus, ree 141;
olicatifoline, 148-157; ae eet 138,
Pringlei, 122, 128;
oc
75; recurvicaulis, 475; recurvicaulis
inarmatus, pubescens, I 4
pumilis 6; Randii, 150, 151, 1543
recurvans, 152, 467; Roezli, 119; Ross-
bergianus, 471; rosaefolius, 122, 126;
rhodophyllus, 155; rubrisetus, 160;
sativus, 1473 saxatilis, 121, 2S;
sapidus, 139-144; s e 8
spectabilis,
viburnifolius, aba: atus, I2I
124; strig
35; subun
trichomallus, ete: ar us guatemalen-
sis, 120; trifrons, 478, 4793 trivialis, I od
s,
143
a. australis, 278; eeeeatk. 278;
ntermedia, 301; italica, 278; lichen-
277; limbata, 278; mexicana,
278; rupestris, 278
Ruppia anomala, 659; maritima, 659-
662; maritima pe tots 660
Ruppia anomala sp. nov., an abberant
type of the Poi tceae, 659
RypBErG, P. A. Noteson Rosaceae—IX,
pis theedraphical
montane zones.
of the a as Nous montane zones, 629
Boone ra alba, 201, 204; borinquen-
suas. ‘aati 387; glomerata, 192
Sabbatia campanulata, 30-32; gracilis,
30, ep serene
bina 37 scopeiorums 13,
20, sh ples ss
Sagittaria latifolia, we
Salix, 230; discolor, 187; Lyallii, 24;
sac rege eae aa re 24;
7; sessilifolia, 2.
696
Ss biflora, 253; commutata, 254,
Salvin officinalis,
Sambucu 5 cadaienala, 188, 347
dus, 27
INDEX
aa 675
isyrinchium arenicola,
a rad nga ~ 8395 oe
640; 639;
ri i. 634, 640
Sium cicutantoliasti, 183, I
SLosson, M. Notes on Trichom manes—lI.
The va mage 4 of Trichomanes pyxi-
diferu OSI
Ram ora peas canadensis
ovalis, 355, 3563 ice temede trifolia,
356; trifolia, 204
Solanacea I
Solanum Dulcamara, 333; nigrum, 331,
m
nigrum Dillenii, 333; pere-
m, 332
Solidago, 39; aestivalis, 561, 562; altis-
s.
ima, 563; aspera, 560; asperula, 560;
480; distans, 188, 492; dion: 40%; bicolor, 559; caesia, 558; canadensis,
elata, 490; Elliottii, 490; foliosa, 491;| 563; Elliottii, 562; juncea, 562, 563;
filifo 493; gracil 488, 493;| neglecta, 186, 562; nemoralis, 563;
Grisebachii, 488, 492; havanensis, 488, cana 5595 patula, ok gages hi
2 emitaphbra, 488; hirtella, 488, seve 560, 561
4 493; _interrupta, Pee lacustris, reve, 560; ee ras. ee Sat:
487, 489; Liebmanni , 493; litho- neues
sperma, 488, 493; sane 493; | Some new bas and varieties of Bihai,
melaleca, ik 490; microcarpa, 488, 315
491; microcarpa foliosa, 491; mitis, mchus arvensis, 550; asper, 5513
8, 4 = abel 493; Otton mis, 490;| oleraceus, 550
pauciflora effusa, 490; phylloptera, Sparganium, 630; americanum andro-
488, 491; pilosissima, 488, ; pine-) -cladum, 183; eurycarpum, 183; mini-
torum, 488, 492; porphyrorrhiza, 48 mum, 179
492; pratensis, 490; pterota, 487, 490; | Spartina gracilis, 637, 642
reflexa, 489; retic laris pubescens, 489; | Specularia perfoliata, 349
— » 489; scindens, 487, 490; pape 72, 180, 196-2 Si eveniiae
setacea, 44. 488; setulonarstiate, nah 2; mbifolium, 72; macrophyllum
, 488, stereorrhiz
mal or, 491; tenella, 492, 193; acoiwem
na, sh ohne a 488, 493; Wrighti-
ae sya Raa a3 na ai 92:
trinitense, 72
Spiraea latifolia, 187; tomentosa, 187
487, 4 ale . .
Siete Berg. fn Cuba, The genus, 487 bes —— fies cegeeenap he
Spirodela,
COTT, Buch uA eer of pasture trees Spo ee
and shrabbery, : "Cas, 6a — — an
Sere h levies lla, - 41; chryptandrus, 638; confusa,
phularia lepore la 335; occidentalis, I; cryptandrus, 638, 641
ne 334 acs hyssopifolia 43
Scutellaria galericulata, 42, 186; lateri- Stenochlamys, 315, 316
oe Stenolobium stans, 372
Sedg esis a a 278 Sterophyllum lsucseteeus, Yate
Selaginella apus, Stigmaria ficoides, 675
Selmorition, 121 gets comata, 638; Elmeri, sie Letter-
Sematophyllum, Rat admistum, 7; ad- , ; minor, 632-639; Nelsonii,
natum, het hyalinum, 575; serici- 633, 638, 639; Porteri, aes Richard-
folium 7 Pies ee Me Scribneri, 639;
7 o, + O32; Vaseyi, 639, 640;
pranindotss. AE 564; linifolius, omy
oe cleistogama, 341; subu
ai
Sherardia gut ensis, 347
Bienes angulatus, 349
Sidalcea, 57; candida, 57, 61;.malvaeflora,
59; neo-mexicana, 57, 59
Tw
a PG Pos pein! 638
Stout, A. B. The origin of dwarf plan
ies ny a sport of Hibiscus ecules
ros
Birehiontie Gin. 06
ares y of pasture trees and shrubbery,
451
INDEX
Studies of West Indian plants —VI, 365;
—Vil
Symphoricarpos Symphoricarpos, 349;
vulgaris, 34
te ec aspe errimum
Syrrhopodon, 72; albovaginatus, 572;
flavescens, 3; floridanus, 73; Muelleri,
572; subtubulosus, 572
Tabebuia acrophylla, 378; Dhetonentghy f
cee’ 378; cola,
‘han
bibracteolata,
uchii,
Ww
~
uw
379%
378
seh
377;
76; Cur-
» 3753 ensis, 378; er nen-
sis, 375; heterophylla 378 jamaicensis,
376; lepidota, 377; 74;
Sauvallei, 377;
Nsis, 373
Tabebuia Gotses: Additional species of,
b osoneaiiaag gr ath, 410
Tarax hee eae 550; erythro-
, 550
Kner alare, 575; instratum, 575;
, 2, 7; polystictum, 107
eee brevifolia, T4, 24
Tecoma acrophylla, 378; actinophyllum,
79;
377; bahamensis, . rii, by
Brittonii, 379; ittonii decussata,
379; Buchii, 377; bibracteolata, pay
€p
378; myrtifolia, 378; platyantha, bik
revoluta, 378; sanguinea, 377; sta
Teloschistes Sol tly seamen ay 580, 581;
= ‘
chryso almus flavicans, 580; flavi-
cans, 579, 580; villosus peas
Lgppeincwgen in rth America, The
genu
S, 579
Teucrium littorale, 41; occidentalis, 41
Thalesia uniflora, 342
oe polygam 85h eo mes: 92).
Tham arbusculans, 105, baculi-
crum, "en: 4; australe, ree spoclvea ra
lon
Thuidiam, pa delicatulum, 190; minutu-.
pl » 575
um, 74; plumulosum
Thuja plicata, 14, 23
Thymus yllum, 44
Tornabenia, 5
Tortula, 72; agraria, 1, 5, 74; confusa,
398; Kunzeana, 398; limensis, 398;
697
pate geste 71, 73; Minuscula, 399;
Pgtt en” 73
8, 550
re -
Tricera acuminata, 497, 499; brevipes,
497, meyer 497, 499; cubana
mensis,
micsheee ts Pee a vi
folios:
502; rotun
tee Ti, 497, 500; subcolumnaris, 498;
—— ste 501; ; 50
Tricera Sw a, The genus, 497
Trichachne sa accharatum, ve 639
Trichocolea tomentella,
Trichomanes, 651, neu leas. 651;
hymeno or ake 658; lepto-
mie m, : branaceum, 651;
pyxidiferum, 652
"658
bileectnena Notes on,—I. The iden-
y of Trichomanes pyxidi iferum, 651
ae hae hamatum, 575; papillatum,
¥ seeapeb pene dichotomum, 41
fg Paalay num,
um bermudan 9%,
nil, 065 orientale, 572
733
348
setum, 635; majus, 631, 633, 636, 639;
rear , 632, 641; subspicatum, 631,
633,
Trlonegistla lancifolia, 575
Tsuga canadensis, 190, 193, 214; hetero-
phylla, 14, 23; Mertensiana, 14
Typha latifolia, 182
Undescribed West Indian species, 387, 514
us americana, 14, 25, 188
Unifolium ee 355-358; tee
oe e, 356; bifolium kam
5: bi folium pained se
bitiom ovale, 356; bifolium trifolium
Uaiotiam — —- on, 354
redineae, New es of —IX, 585
Uredo tesbena. 303
Uromyces abbreviatus, ay 587; orna-
tipes, 586; Psoraleae, 5
no ia toniana, vg
Usnea,
Uericularta Seogemonse — cleistogama,
341; cornuta, 204; scapa, 340;
intermedia, 179, 340; gases a, 340;
’ subulata, 340; vulgaris americana, 340
Vaccinium, 247; corymbosum, 187, 188,
205, 210; macrocarpon, 204, 207; Oxy-
698
Vagnera Sythe a, 357; stellata ee
357; trifolia, 358; trifolia bifolia, 358
trifolia unifolia, 35
a
fi
of Con ecticut, The,—IV.
. Plant societies in enclave, 169
Veratrum viride, 189
Blattaria, 334; phlomoides,
Verbascum
334; Thapsus, 334
gee hastata, 41, 186, 192; urticifolia,
Vertethaens:
biisecat rae noveboraceni 192
Vero arve ie OS ager to
peregrina, ens Se vitifo lia, 337; T
m, 3373 Torraefectil S20; Ciainice,
Vesicularia ee 575; filicuspis,
5753 yeriana, 5
Viburnum dentatum, ans 188, 347, 348;
ssinoides, 205; Lentago, 188; molle,
ihe venosum, 347, 34
Vinca minor, 34
INDEX
Viola cucullata, 189; pallens, 189
Webera rupestris, 574
Weisia Breutelii, 3; Renee BSS longiseta,
- Pabstiana, 3; serioc: ; viridula,
West Indian mosses— Mosses of the
Soares West Indies ‘and Virgin Is-
lands,
= Taian plants, Studies of, —365;
~ uaa species, Vadeusibed, 384,
Watt Mosses from the west
coast pr south America, 393; Mosses of
the Philipp waiian Islands
collected i an ‘ray Pes B. mais g 571
oodwardia virginica, 186, 202,
Xanthium echinatum, 556
Zebrina pendula, 63
Zonotrichia, 196
Zygodon pa 400
VOLUME 42, PLATE 30
BULL. TorREY CLUB
TRICHOMANES PYXIDIFERUM L.
BULL, TORREY CLUB VOLUME 42, PLATE 31
TRICHOMANES HYMENOPHYLLO:DES VAN DEN BoscH
BuLL. TORREY CLUB VOLUME 42, PLATE 32
RUPPIA ANOMALA OSTENFELD .
BuLL. TorREY CLUB : VOLUME 42, PLATE 33
DICTYOPHLOIS RETICULATA FOERSTE