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i

THE MARINE ALGAE OF THE PACIFIC COAST
OF NORTH AMERICA

PART III

MELANOPHYCEAE

BY

WILLIAM ALBERT SETCHELL

AND
NATHANIEL LYON GARDNER

University of California Publications in Botant
Vol. 8, Part III, pp. 383-898, plates 34-107

UNIVERSITY OF CALIFORNIA PRESS

BERKELEY, CALIFORNIA

1925

UNIVEESITY or CAUFOKNIA PUBLICATIONS

Note. — The University of California Publications are offered in exdiaiig'e for the publl-
cationa of learned societies and institutions, universities and libraries. Complete liafts of
all the publications of the University will be sent upon request. For sample copies, lists
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Publications of the University of California Press may also be obtained from THE
CAMBRIDGE UNIVERSITY PRESS, FETTER LANE, LONDON, B.C. 4, ENGLAND, to
which orders originating In Great Britain and Ireland should be sent.

BOTANY.— W. A. Setdiell and B, M. Holman, Editors. Price per volume, $S.50 (voL 5,
$6.00) ; beginning with volume 6, $5.00. Volumes I, II, HI, IV, V, Vi; VH, and IS
completed. Volumes Vm, X, XI, xn, and XIH in progress.

Cited aa Univ. Calif. PubL Bot.

VoL 6. 1912-1922.

I, by William Albert SetcheU. Pp. 1-86. December,

1. Studies in Nicotiana.

1912 .

2. Quantitative Studies of Inheritance in JV^tcof tana Hybrids. I, by Thomas

Harper Ooodspeed. Pp. 87-168, plates 1-28. December, 1912 ... .......

3. Quantitative Studies of Inheritance in Nicotiana 'Hybrids. U, by Thomas

Harper Goodspeed. Pp. 169-188, plates 29-34. January, 1913 .._.„..^ ._„_

4. On the Partial Sterility of Nicotiana Hybrids njade with N^ sylvestris as a

Parent, by Thomas Harper Gk>odspeed. Pp. 189-198. March, 1913 ...

5. Notes on the Germination of Tobacco Seed. 1, by Thomas Harper Good-

speed. Pp. 199-222. May, 1913 .... ... „... _..

6. Quantitative Studies of Inheritance In Nieotiana Sy'btii.n. Ill, by Thomas

Harper Goodspeed. Pp. 223-231. April, 1915 ...... „.....,...................™. ....

7. Notes on the Germination of Tobacco Seed. II, b7 Thomas Harper Good-

speed. Pp. 233-248. June, 1915 . .. ...... ..._._._„^ — ..„...._......_. ,

8. Parthenogenesis, Parthenocarpy and Phenospermy in Z^tcof tana, by Thomas

Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 ..._ _.„.... _. — .

9. On the Partial Sterility of Nicotiana Eyirids made with jV. sylvestris as a

Parent, n, by T. H. Goodspeed and A. H. Ayres. Pp. 273-29^ plate 36.

10. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent, m, An Account of the Mode of Floral Abscission In the F^ Species
Hybrids, by T. H. Goodspeed and J. N. Kend^. Pp. 293-299. November,

11. The Nature of the Fj Species Hybrids between Nicotiana sylvestris and

Varieties of Nicotiana Tabacum, with Special Reference to the Conception
of Reaction System Contrasts in Heredity, by T. H. Gh>od^eed and B. E.
Clausen. Pp. 301-346, plates 37-48. January, 1917 ^.. _

12. Abscission of Flowers and Fruits In the Solanaceae, with Special Reference

to Nicotiana, by John N. HendaU. Pp. 347-428, 10 text figures, plates 49-

13. Controlled Pollination in Nicotiana, by Thomas Harper Good^>eed and Plrle

Davidson. Pp. 429-434. August, 1918 ._.. -.. — ,..-...- . — — ™

14. An Apparatus for Flower Measurement, by T. H. Goodspeed and R. E.

Clausen- Pp. 435-437, plate 54, 1 figure in text. September, 1918 .„.

15. Note on the Effects of Illuminating Gas and Its Constituents In Causing

Abscission of Flowers in Nicotiana and Citriis, by T. H. Goodspeed, J. M.
McGee and R. W. Hodgson. Pp. 439-450. December, 1918 .. : .

16. Notes on the Germination of Tobacco Seed. HI, Note on the Relation of

Light and Darkness to Germination, by T. Harper Goodspeed. Pp. 451-

17. Inheritance in <^icottano Tabacum. I, A Report on the Results of Crossing

Certain Varieties, by William Albert Setchell, Thomas jHarper Goodspeed,
and Roy Elwood Clausen. Pp. 457-682, 2 figures in text, plates 56-86.

91.26
LOO
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University of California Publications in
t >. ^

BOTANY

\OLUME VIII
Part III

1925

WILLIAM ALBERT SETCHELL
R. M. HOLMAN

EDITORS

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY, CALIFORNIA

THE MARINE ALGAE OF THE PACIFIC
COAST OF NORTH AMERICA

PART III

MELANOPHYCEAE

BY

WILLIAM ALBERT SETCHELL
AND

NATHANIEL LYON GARDNER

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY, CALIFORNIA

University of California Publications in Botany

Vol. 8, Part III, pp. 383-898, plates 34-107

Issued June 26, 1925

The University of California Press
Berkeley, California

The Cambridge University Press
London, England

COxX TENTS

PAGE

Subclass 3. Melanophyceae 387

Series 1. Phaeosporeae 389

Order 1. Sphacelariales 390

Family 1. Sphacelariaceae 391

1. Sphacelaria 392

2. Chaetopteris 397

Order 2. Ectocarpales 398

Family 2. Ectocarpaceae 400

3. Pylaiella 401

4. Ectocarpus 407

5. Streblonema 440

Family 3. Myrionemataceae 453

6. Myrionema 454

7. Compsonema 473

8. Hecatonema 488

9. Microspongium 492

Family 4. Ralfsiaceae 493

10. Ralfsia 494

11. Haptcrophycus 499

12. Lithoderma 500

Family 5. Elachisteaceae 502

13. Elaehistea 503

14. Gonodia 505

Family 6. Leathesiaceae 507

15. Petrospongium 508

16. Leathesia 510

Family 7. Punctariaceae 513

17. Punctaria 515

Family 8. Asperococoaceae ' 522

18. Halorhipis 523

19. Soranthcra 525

20. Myelophycus 527

Family 9. Striariaceae 528

21. Striaria 529

22. Stictyosiphon 530

Family 10. Scytosiphonaceae 530

23. Scytosiphon 531

24. Ilea 535

25. Endarachne 538

26. Colpomenia 539

27. Hydroclathrus 542

Family 11. ^Egiraceae 543

28. ^gira 544

29. Meneghiniella 548

Familj' 12. Heteroehordariaceae 549

30. Heterochordaria 550

[383]

384

University of California Publications in Botany [Vol. 8

PAGE

Family 13. Chnoosporaceae 552

31. Chnoospora 552

Order 3. Desmarestiales 554

Family 14. Myriogloiaceae 555

32. Myriogloia 556

Family 15. Desmarestiaceae 558

33. Desmarestia 559

Order 4. Chordariales 570

Family 16. Chordariaceae 571

34. Chordaria 571

35. AnaHpus 575

36. Gobia 576

Family 17. Coilodesmaceae 577

37. Coilodesme 578

38. Phaeostrophion 585

Order 5. Dictyosiphonales 586

Family 18. Dictyosiphonaceae 587

39. Dictyosiphon 587

Order 6. Laminariales 590

Family 19. Chordaceae 591

40. Chorda 592

Family 20. Laminariaceae 593

Tribe 1. Laminarieae 594

41. Laminaria 594

Tribe 2. Cymathaereae 606

42. Pleurophycus 606

43. Cymathaere 608

Tribe 3. Agareae 609

44. Costaria 609

45. Thalassiophyllum 613

46. Agarum 614

Tribe 4. Hedophylleae 616

47. Hedophyllum 617

48. Arthrothamnus 619

Family 21. Lessoniaceae 621

Tribe 1. Lessonieae 621

49. Dictyoneurum 621

50. Nereocystis 623

51. Postelsia 624

Tribe 2. Macrocysteae 626

52. Macrocystis 626

53. Pelagophycus 629

Tribe 3. Lessoniopseae 631

54. Lessoniopsis 631

Family 22. Alariaceae 633

Tribe 1. Alarieae 633

55. Pterygophora 634

56. Alaria 635

Tribe 2. Ecklonieae 645

57. Eisenia 646

Tribe 3. Egregieae 647

58. Egregia 647

1925] Setchell-Gardncr: Melanophijceae 385

PAGE

Series 2. Aplanosporeae 649

Order 7. Dictyotales 650

Family 23. Dictyotaceae 650

50. Diftyota 651

GO. Xc'urocarpus 655

61. Taenia 656

62. Chlanidophora 658

63. Zonaria 659

64. Padina 661

Series 3. Cyclosporeae 662

Orders. Fucalcs 663

Family 24. Fucaceae ; 663

65. Fiicus 664

66. Pelvetia 700

67. Pelvetiopsis 702

68. Hesperophycus 703

Family 25. Sargassaceae 704

69. Blossevillea 705

70. Cystophyllum 706

71. Halidrys 707

72. Cystoseira 708

73. Sargassum...... 711

The third part of the Marine Algae of the Pacific Coast of North
America, comprising an account of the Melanophyceae or Brown
Algae, succeeding the accounts of the Myxophyceae and Chloro-
phyceae, is presented without introduction or explanation, pending
the publication of the final part, which is in advanced preparation.
It is intended to issue with the completed volumes a statement as to
the principles followed, methods used, territory covered, sources of
information, material, etc., as well as to make acknowledgment to the
various authorities, students and collaborators who have been of the
greatest assistance in carrying out the long and laborious task.

W. A. Setchell and N. L. Gardner.

to

CM

THE MARINE ALGAE OF THE PACIFIC COAST
OF NORTH AMERICA

PART III

BY

LIBRA pv

WILLIAM ALBERT SETCHELL \;ni,

AND

NATHANIEL LYON GARDNER

rjr

Subclass 3. MELANOPHYCEAE stiz.

.Multieelhilar thallophytes containing the green pigment, chloro-
phyll (including xanthophyll), and the brown pigment, phycophaein;
thallus varying from simple or slightly branched filaments of a single
row of cells and simple membranes varying from a single layer to
several layers of similar cells, to solid plant bodies of varying form,
e.g., membranous, erustaceous, globular, hemispherical, filamentous,
etc., and with differentiated internal and external tissues; cells with
distinct cell Avails, cytoplasm, at least the outer cells containing definite
chromatophores, carrying the pigments, and typically uninucleate,
although at times becoming multinucleate; assimilation products
various carbohydrates and oils, but never true starch ; internal cells
in more complex forms elongated and forming definite mechanical
conducting and storage systems as distinct from the outer tissues of
absorbing and assimilating cells ; multiplication by splitting and by
gemmae of more or less distinctive form ; reproduction by both motile
and non-motile cells, motile cells with one or two lateral, or at least
subapical, cilia; non-sexual reproduction by 2-ciliated zoospores or by
aplanospores (Dictyotales) : sexual reproduction by the fusion of
two equal, or of two unequal, motile gametes, or by the fusion of a
smaller motile 2-ciliated sperm and a larger non-motile egg (Lamina-
riales and Fucales) or by the fusion of a smaller 1-ciliated sperm and
a larger non-motile egg (Dictyotales) ; an antithetic alternation of
generations present in many, if not all, species; gametophytes usually
unisexual, similar to, up to extremely different from, the sporophytes.
Mclanophyceae Stizenberger, Dr. Ludwig Rabenhorst's Algen
22 Sachsens, 1860, p. 36; Rabenhorst, Flora Eur. Alg., vol. 3, 1868, p.
CO 393. Fucoideae Agardh, 8yn. Alg. Scand., 1817, p. IX. Melano-
CM spermeas Harvey, in Mackay, Flora Hibern., part III, 1836, p. 157;

388 University of California Puhlications in Botany [Vol. 8

Gen. So. African Plants, 1838, p. 393. Phaeophyceae Kjellman, in
Engler and Prantl, Die natiirl. Pflanzenfam., Teil 1, Abt. 2 (Lief. 60),
1891, p. 176.

The Melanophyceae, or Brown Algae, are, with very few exceptions,
marine and are, properly, all multicellular. AVe feel convinced that
the termination -phyceae should be retained for the subclasses of the
class Alg'ae, or Phycophyta. Since Stizenberger's Melanophyceae,
1860, applies to the same group of plants as does Kjellman's Phaeo-
phyceae, 1891, and since Stizenberger 's term seems equally applicable,
we have chosen it in preference to Kjellman's, even though Phaeo-
phyceae is at present in general use. The Melanophyceae are much
more complex in both structure and reproductive processes than are
the Chlorophyceae or the Myxophyceae. In size, they range from
minute disks to plants of considerable size, reaching among the kelps
to a length of possibly 100 meters or longer. It has seemed the more
natural to restrict the Melanophyceae to the multicellular forms than
to include the unicellular forms with chlorophyll and a brown pigment.
The Bacillariales, or Diatoms, and the Peridiniales, as well as certain
groups of the Flagellates, even if they may be regarded as belonging
to the vegetable kingdom, seem to have no characters that would
associate them with the Melanophyceae except the possession of a
similar brown pigment.

The Melanophyceae are, with few exceptions, viz., Phaeosaccion
Farlow and Omphalophyllum Rosenvinge, provided with specialized
zoosporangia or with gametangia or with both. In one order, viz.,
Dictyotales, the non-sexual spores are always motionless.

There is an antithetic alternation of generations in most of the
groups of the Melanophyceae, the Fucales being the only order in
which it is certainly absent, where the sporophyte generation of this
order is considered as represented by the products of the division of
the oogonial cell. In the Sphacelariales and the Ectocarpales it is not
so clearly proved as yet, but Kylin (1918) has demonstrated reduction
divisions of the primary nucleus of the zoosporangium in certain
species. The two generations in both of these orders are alike in size.
We find species, however, in which the gametophyte is microscopic, in
striking contrast to the large, often even gigantic, sporophj^te. The
same relation between gametophyte and sporophyte is suspected, but
not yet observed, in the Dcsmarestiales, Chordariales, and Dictyosi-
phonales. In the Cutleriales the two generations may be similar or
dissimilar, while in the Dictyotales they are similar in size and
vegetative structure.

19:25J iSetchcII-Ganhur: Melanophyceae 389

The classification of llic Melanophyceae is in a very unsatisfactory
condition, that of Kjrllnian (1891) and that of Oltmanns (1904)
seeming very iinpt rfcd, especially since the later studies of Sauvageau,
Kylin, and others have changed our views in so many important
respects. The more recent classifications of Kylin (1917), of Oltmanns
(1922) and of William Randolph Taylor (1922) have assisted greatly
toward a more natural arrangement. We have therefore adopted many
of the views of Kylin (1917) and of Oltmanns (1922) and have incor-
porated some additional ideas of our own. Oltmanns has divided the
suhclass into seven orders. In this, we are inclined to follow him, but
think it better to regard some of the other divisions also as of equal
rank with these. According to our scheme, the Melanophyceae of the
Pacific Coast of North America are arranged in the following manner :

Key to the Series

1. Reproduction sexual only 3. Cyclosporeae (p. 662)

1. Reproduction both sexual and non-sexual 2

2. Aplanosporcs present 2. Aplanosporeae (p. 649)

2. Aplanosporcs absent 1. Phaeosporeae (p. 889)

Series 1. PHAEOSPOREAE thuret

Thallus composed of simple or branching filaments either of a
single row of cells (monosiphonous) or solid with diverse tissues or of
simple or solid membranes, or cushion form, branched or unbranched,
sometimes hollow, variously aggregated; non-sexual reproduction
(where known) by zoospores with 2 lateral cilia produced in unilo-
cular zoosporangia ; aplanosporcs never present ; gametangia pluri-
locular, or in one order (Laminariales) unilocular, male gametes
motile, biciliated, female gametes motile and biciliated or non-motile ;
gametophyte and sporophyte similar or more or less dissimilar.

Phaeosporeae Thuret, Rech. sur les zoosp. I, 1850, p. 233 (p. 24,
repr.) ; Kjellman, in Engler and Prantl, Natiirl. Pflanzenfam., Teil 1,
Abt. 2 (Lief. 60), 1891, p. 180 (in part) ; Oltmanns, Morph. und Biol.
der Algen, vol. 1, 1904. p. 348. Phaeozoosporeae De-Toni, Syst.
Uebers. Fucoid., 1891, p. 175. Phaeo^porales Oltmanns, loc. cit., p. 348.

The Phaeosporeae of Thuret have generally been regarded as con-
stituting an order, but they seem to us rather to represent a series
under the subclass Melanophyceae composed of the orders Sphace-
lariales, Ectocarpales, Cutleriales, Dictyosiphonales, Desmarestiales,
and possibly also the Laminariales. The general characteristic is the
occurrence of the typical brown zoospores of the Melanophyceae. These
are, as yet, unknown in some genera and species of the series, while

390 University of Calif ornia PuMications in Botany [Vol. 8

in others they occur but seldom. What our increasing knowledge may
teach us concerning these seemingly exceptional forms must be left for
future consideration. The typical members of the Phaeosporeae possess
unilocular zoosporangia and plurilocular gametangia and have the
gametophyte and sporophyte of very closely similar size and struc-
ture. Some scattering genera and species are only known to produce
the unilocular zoosporangia, while others produce only the plurilocular
gametangia, but, considering seemingly close affinities in these puzzling
cases, it seems undesirable for the joresent, at least, to refer either
elsewhere than to the Phaeosporeae. They may be found to repre-
sent either imperfect, or perhaps even dimorphic forms, when their
development shall have been carefully studied. The Laminariales,
which we still include under the Phaeosporeae, were long known only
with unilocular sporangia producing biciliated zoospores, but have
recently been found also to possess a microscopic gametophyte with
decidedly modified (unilocular) gametangia producing either non-
motile egg cells (or oospheres) or biciliated sperms. The Phaeosporeae
may be distinguished from the Acinetosporeae by the lack of aplano-
spores, and from the Fucales by the possession of either known zoo-
sporangia or plurilocular gametangia or both.

Key to the Orders

1. Gametangia present on macroscopic plants (zoosporangia known or unknown). ... 2
1. Only zoosporangia present on macroscopic plants (gametangia known, or

suspected of being, on microscopic plants) 3

2. Terminal cell present and conspicuous 1. Sphaeelariales (p. 390)

2. Terminal cell, if present, at least not conspicuous. .2. Ectocarpales (p. 398)

3. Growth of zoosporangial frond, strictly terminal 5. Dictyosiphonales (p. 586)

3. Growth of zoosporangial frond not strictly terminal 4

4. Growth of zoosporangial frond trichothalHc 3. Desmarestiales (p. 554)

4. Growth of zoosporangial frond not trichothalHc 5

5. Growth apical, from subapical cells 4. Chordariales (p. 570)

5. Growth never apical, either intermediate or near the base .. 6. Laminariales (p 590 )

Order 1. SPHACELARIALES oltmanns

Filamentous, generally tufted and branched Phaeosporeae, rarely
monosiphonous throughout, usually monosiphonous near or at the tips,
becoming polysiphonous below through longitudinal walls, and in some
cases corticated slightly or considerably by descending filaments,
attached by horizontal rhizoids often cohering to form a disk, or, in
some cases, arising early and forming a more or less complex hori-
zontal thallus giving rise to the erect filaments ; growth of each erect

1925] Setchell-Gardncr: Melanophyceae 391

filament from a more or less conspicuous apical cell ; older cell walls
becoming dark colored when treated with eau de javellf ; chromato-
phores numerous in each cell, usually disk-shaped; zoosporangia uni-
locular, large; gametangia plurilocular, similar or with some distinc-
tion as 1() size and miiiilicr of loeuli (mega- and meio-gametangia) ;
zoosporangia and gametangia borne on separate (macroscopic) plants
differing little, if at all, in size or complexity and formed from the
transformed apical portions of usually short branchlets; propagula
present and characteristic in certain species of Sphacelaria, arising
from the transformation of lateral branchlets, producing a short
stalked body with longer or shorter radiate terminal branches, the
whole separating from the branch bearing it and developing into a
new plant.

Spliacelariales Oltmanns, Morph. und Biol, der Algen, vol. 2, 1922,
p. 83. Sphacelariaceae Reinke, Ber. d. deut. bot. Gesell.. vol. 8, 1890,
p. 203.

The ISphacelariales form a natural group, coordinate but distinct
from the Ectocarpales in that all the species included take their growth
from a conspicuous and very characteristic apical cell. The species
are, also, more or less polysiphonous through longitudinal division
walls, producing fronds unlike those of any of the species we have
included under Ectocarpales. The cell wall is 'said to differ in composi-
tion, or constitution, from that of any of the species of Ectocarpales, in
that, when older, it darkens when treated with eau de javelle, turning
black in mature cells. The members of the Sphacelariales resemble
those of the Ectocarpales in having unilocular zoosporangia and pluri-
locular gametangia on macroscopic plants closely resembling one
another in size and structure. In some species the gametangia are
differentiated into two types, megagametangia or oogonia with few and
large loeuli, and antheridia with numerous and small loeuli. Oltmanns
(loc. cit., p. 85) divides the order into three families; Sphacelariaceae,
Cladostephaceae, and Stypocaulonaceae, of which only the first, so far
as we know, is represented on the west coast of North America.

FAMILY 1. SPHACELARIACEAE REINKE, emend, oltmanns

Thallus of erect filaments from a discoid or thalloid base, mono-
siphonous at the tip or throughout, polysiphonous and. at tinu's, cor-
ticated below ; apical cell short, but conspicuous, not giving rise
directly to the initials of the branches, but these arising from the
apical portion of a cell already segmented, and usually situated at

392 University of California Publications in Botany [Vol. 8

some distance below the apical cell; branches variously arranged,
alternate and distichous or polystichous, or, at times, opposite and
distichous, occasionally distinguished into two sets, long (indefinite)
axes and short (definite) axes, but the latter never in regular whorls ;
otherwise as in the order ; propagula formed on certain species of
this family.

Spliacelariaceae Reinke, Ber. d. deut. bot. Gesell., vol. 8, 1890,
p. 203 (in part) ; Oltmanns, Morph. und Biol, der Algen, vol. 2, 1922,
p. 85.

On the west coast of North America are to be found few of the
Sphacelariales and none is frequent. The specimens which have been
found belong to the Sphacelariaceae. The Cladostephus verticillatus
(Liglitfoot) Ag., referred to the coasts of the Russian American
possessions on the North Pacific Ocean (Postels and Ruprecht, 1840,
p. 21) and apparently not in typical form, has not been seen since, so
that it does not seem best to continue to include it as belonging to our
flora.

Key to the Genera of the Sphacelakiaceae

1. Zoosporangia and gametangia borne on the regular branches from the axis

1. Sphacelaria fp. 392)

1. Zoosporangia and gametangia borne on special branches arising from the

corticating filaments 2. Chaetopteris (p. 397)

1. Sphacelaria Lyngb.

Fronds forming relatively small, usually profusely branched, more
or less spherical or irregular tufts attached to the surface of rocks or
to other algae by small disk-like or thalloid holdfasts, or penetrating
into the substance of other plant bodies; the main axes and the
branches terminated by an apical growing cell which divides cross-
wise, the resultant segments sooner or later dividing lengthwise and
more or less obliquely, building up a polysiphonous frond, the sur-
face layer of which consists of transverse bands of rectangular cells
obscured below in corticating species; reproduction by unilocular
zoosporangia and by pluriloeular gametangia, and vegetative, by the
formation and separation of propagula, or bulblet-like portions of the
frond, produced on different individuals from the sporangia ; zoo-
sporangia and gametangia arising from the regular branches from the
axes of the plant, not from special branches produced from corticating
filaments.

Lyngbye, Hydrophyt. Dan., 1819, p. 103.

1925] 8 etchell -Gardner : MeJanophyceae 393

The genus Sphacelaria is composed of species attached in various
"ways to the substratum or host, but which lacks any pronounced dimor-
phism such as is to be found in some genera of the Sphacelariaceae,
The erect filaments are always more or less branched and with one
exception show primary vertical partitions of the cells over the greater
extent of their axes and in certain species also show secondary trans-
verse partitions. Hairs may be present or absent and simple or
geminate. Rhizoids may, or may not, be present and may form corti-
cations. Certain species produce propagula of distinctive form while
others may lack them. The sporangia and gametangia are borne on
the axes, never on the rhizoids as in Chaetoptcris. The gametangia
are plurilocular and all alike or differentiated into oogonia and
antheridia.

Our species are few and not at all abundant or well known. Care-
ful search for additional material is much to be desired. We have
relied particularly upon Sauvageau's different papers collected under
the general title of Remarques sur les Sphacelariacees (1900-1914) for
guidance in the determination of our west coast forms. We have
arranged the material accessible to us under six species, awaiting
further opportunities for study and verification.

Key to the Species

1. Erect plants showing both primary transverse and longitudinal and regular

secondary transverse partitions 1. S. racemosa (p. 393)

1. Erect plants showing, regularly, only primary transverse and longitudinal

partitions 2

2. Propagula stout with short or no projecting rays 3

2. Propagula slender with elongated rays 4

3. Rays of propagula of 2-3 short cells, attenuated outwards.. 2. S. brevicomis (p. 394)

3. Rays of propagula usually of a single cell, broad 3. S. califomica (p. 395)

4. Rays of propagula simple 5

4. Rays of propagula bifurcate 6. S. didichotoma (p. 397)

5. Rays of projiagula strictly two 5. S. furcigera (p. 396 )

5. Rays of propagula in twos and threes in same individual... 4. S. subfusca p. 395)

1. Sphacelaria racemosa Grev.

Erect filaments forming tufts 1-7 cm. high, dark brown, irregularly
branched above, arising from a very small, prostrate thallus ; joints
showing several longitudinal and more or less regular secondary
transverse partitions 55-70;tt diam. below ; propagula unknown ; zoo-
sporangia 40-50ja long by about -iOfi diam. ("60-70/^ x 52-60ju"),
borne on short (1 or 2 celled) pedicels irregularly racemosely arranged
on small lateral branchlets; gametangia (not seen in our specimens)
ovate-cylindrical or cylindrical racemosely arranged.

394 University of California Puhlications in Botany [Vol. 8

Growing on rocks, shells and, at times, on other algae, in the lower
littoral or upper sublittoral belts. Alaska (Port Clarence, Kjellman ;
Prince William Sound, Yakutat Bay and Wrangell, Saunders) to west
coast of Whidbey Island, Washington, Gardner.

Greville, Scot. Crypt. Flora, vol. 2, 1824, pi. 96. Sphacelaria race-
mosa var. arctica Saunders, Alg. Harriman Exp., 1901, p. 419 ; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 239. Sphacelaria arctica
Kjellman, Beringh. Algfl., 1889, p. 51.

AVe have only the specimens from Whidbey Island to guide us as
to the occurrence of this species on our coast and these are scanty
with very few zoosporangia and covered by epiphytes of the Dermo-
carpa-type. The irregular (never pinnate) branching, the regular
divisions of the outer cells of the joints by a secondary transverse
partition as well as the irregularly racemose arrangement of the
sporangia indicate the position of the Whidbey Island specimen under
the species and not under the var. arctica (Kjellm.) Reinke. It may
be that the Alaska specimens belong to the var. arctica. The few
zoosporangia found in our specimens are small (as indicated in our
description) but are probably not well developed. The racemose
arrangement in our specimens is not so regular or luxuriant as that
shown in Greville 's figures (Joe. cit.).

2. Sphacelaria brevicornis S. and G. (Orothog. mut.)

Erect filaments 1-1.5 cm. high, sparselj" and strictly branched,
attached by small, penetrating filaments ; joints showing only primary
longitudinal partitions, 35/* diam. below, 22-30,/x near the tip ; hairs
arising near the tips composed of 6-8 cells ; zoosporangia and game-
tangia unknown; propagula stout, obovate, tricornute, about 120/i
long and about TOju, wide (below the rays), composed of a few large
cells and on 2-3-celled pedicels, rays ("horns") short, of 2-3 cells,
blunt, attenuate outwards.

Growing on Sargassum polyacanthum f. americanum. La Paz,
Lower California.

Sphacelaria hrevicorms Setchell and Gardner, Mar. Alg. Gulf of
Calif., 1924, p. 725, pi. 19, figs. 59, 60.

;S'. hrevicornis is very close to S. cornuta Sauv. of New Caledonia,
but the propagula are provided with shorter horns from a more
robust basal portion.

1925] SetchcU-Gardner : Melanophyceae 395

3. Sphacelaria californica Sauy.

Plate 37, figs. 23-27

Erect filaments forming light olive brown tufts, up to 3 em. or
more in height, naked for about 1 cm. below, attached by a small disk ;
joints with primary vertical and occasional, but irregularly placed,
secondary transverse partitions, up to 120/x diam. ; hairs and rhizoids
absent ; propagula broad, 140-160ja long, 80-120ja broad, without pro-
jecting rays or a single large cell projecting slightly at each of the
upper 3 corners; zoosporangia short, ellipsoidal to obovate, 75-150|a
long, 50-7 Oju. wide ; gametangia unknown.

Growing on rocks, in pools, in the middle littoral belt. Southern
California (San Pedro to San Diego).

Sauvageau, Remarques sur les Sphace., 1901, p. 92 (with ?).
Sphacelaria plumula var. californica Sauvageau, Joe. cit., p. 91, fig. 21 ;
Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1694,
1910. S. trihuloides Saunders. Phyc. Mem., 1898, p. 158, pi. 26 (not of
Meneghini).

Sphacelaria californica was suspected to be a distinct species by
Sauvageau w^ho described it {loc. cit.) under S. pluvmda var. cali-
fornica. He published, however, the binomial, Sphacelaria californica?
(loc. cit.), and it seems proper to attribute this combination to him.
The species, while close to S. plumiUa Zan., differs in having the lower
portion of the axis (for a centimeter or more) free from branches, in
having a basal disk and a straighter axis. Sauvageau 's plants were
more slender than those of S. plumula, but ours equal them in measure-
ment (up to 120j(x) ; many of our plants, however, have the smaller
measurements of those of Sauvageau. Secondary transverse partitions
occur in the joints of this species but they are occasional and irregu-
larly placed. The type locality is San Diego, California.

4. Sphacelaria subfusca S. and G.

Plate 37, fig. 28

Erect filaments forming brown tufts about 0.5 mm. high, or slightly
higher, alternately but irregularly branching, penetrating slightly
among the superficial cells of the host by short thick rhizoids, not
forming a distinct disk; joints with primary vertical partitions only,
24-40/x high, 40-50ja wide ; hairs uniformly present and abundant,
single, seriate, opposite or secund, up to 8 cells in length, about 15fi
diam. ; propagula slender, with 2 or 3 slender rays, pedicel 8-12 cells,

396 University of California Publications in Botany [Voh. 8

with 3-4 of the upper cells showing longitudinal septa, 24/x diam.,
slightly attenuate toward the point of attachment, rays 4-9 celled,
tapering slightly, if at all, towards the tip ; zoosporangia and game-
tangia unknown.

Growing on various algae. Alaska (Sitka) to southern California
(Redondo).

Setchell and Gardner, Phyc. Cont. VII, 1924, p. 1.

Sphacelaria fusca Collins, Holden and Setchell, Phyc. Bor.-Anier.
(Exsicc), no. 1693, 1910 (not of Agardh). Sphacelaria cirrhosa
Saunders, Alg. Harriman Exp., 1901, p. 419 (?).

Sphacelaria sudfusca might ordinarily be referred to S. fusca
(Huds.) Ag., to which, undoubtedly, it is closely related. It differs
from S. fusca in its basal portion, in its much lower stature, smaller
diameter, and in certain details of structure of the propagula. In the
last, the stalk tapers little and the rays are barely perceptibly atten-
uated toward their apices. In addition, the occurrence of frequent
bifurcate, along with the trifurcate, propagula, so characteristic of
8. fusca, leads us to consider our plant as a distinct species. It is
intermediate between S. fusca (Huds.) Ag. and 8. furcigera Kuetz.
While we have not seen Saunders' 8. cirrhosa from Annette Island,
his description of its habit corresponds so well with that of a specimen
from Sitka, that we refer it here provisionally,

5. Sphacelaria furcigera Kuetz.

Plate 37, fig. 29

Erect filaments a few millimeters (up to 5 mm. or perhaps more)
high, forming small tufts, arising from short, closely intertwined
creeping filaments often also penetrating the host, sparsely branched,
with lateral and subterminal hairs, the latter, at times, pushing aside
the terminal cell and taking its position; joints with 1-3 primary
vertical partitions, 16-30ju, (in ours, but up to 45/*, fide Sauvageau)
in diameter and up to 50/* or more high; hairs abundant, few-celled
(4-6), cells 10-14/* wide, up to 50/* long; propagula slender, bifurcate
with an umbo between the rays, stalk with 6-9 joints, the upper with
2-3 vertical partitions, slightly attenuated below, about 20/* diam. ; rays
with 5-8 cells, not at all attenuated toward the apices ; zoosporangia
spherical (fide Sauvageau), 50-70/* diam., short (1 cell) pedicellate;
gametangia (fide Sauvageau) of 2 sorts, antheridia, cylindrical, 45-65/*
long, 24-28/* wide, with loculi about 3/* diam. and oogonia more
irregular, 30-45/* (up to 60/*) long, 28-40/* broad, with large loculi.

1925] Setchell-Gardncr : MeJanopJtyceae 397

Epiphytic on other algae. Gulf of California.

Kuetzing, Tah. Phyc, vol. 5, 1855, p. 27, pi. 90, fig. 11; Reinke,
Beitr. zur vergl. Anat. u. Morph. der Sphac, 1891, p. 14, pi. 4, figs.
5-13 ; Sauvageau, Rem. sur les Sphace., 1901, p. 156 ; Setchell and
Gardner, Mar. Alg. Gulf of Calif., 1924, p. 724, pi. 19, fig. 58.

Kuetzing founded this species on a plant from the Island of Karak,
in the Persian Gulf. Reinke (loc. cit.) and Sauvageau {loc. cit.) have
interpreted it, the former confining it to the Indian and Pacific oceans,
the latter extending it to the Atlantic Ocean and adjacent European
seas. According to Sauvageau 's interpretation our scanty material
may very well be included under the form cycle of S. furcigera, but we
doubt its conforming exactly to the type of Kuetzing. Our specimens
show scanty propagula and seemingly young oogonia.

6. Sphacelaria didichotoma Saunders

Erect filaments up to 4 mm. high, forming dense tufts, with prob-
ably a penetrating base, irregularly branched ; joints with as many
as 3 vertical partitions, 25-35/a diam., and slightly longer than broad ;
hairs not known ; propagula slender, bifurcate, with each ray again
furcate, stalk slightly attenuated at the base, 200-300ja long, main
branches 100-200/x long ; sporangia and gametangia unknown.

On other algae, including Melobesieae. Central and southern
California.

Saunders, Phyc. Mem., 1898, p. 158, pi. 27. Sphacelaria variabilis
Sauvageau, Rem. sur les Sphace., 1901, pp. 160-162, fig. 37 ?.

The species noted above is known to us chiefly from descriptions
and figures. A specimen from Clifton, California, collected by Carl
Epling, seems referable here. It is very uncertain (cf. Sauvageau,
Joe. cit., p. 160) as to whether S. didichotoma is identical with S.
variabilis Sauv., or with 8. divaricata Mont., or is distinct from both.

2. Chaetopteris Kuetz.

Frond arising from a solid parenchymatous base, cylindrical,
filiform, solid, parenchymatous throughout, irregularly divided into
longer and shorter branches, which in turn are pinnately branched ;
reproduction by unilocular zoosporangia and plurilocular gametangia
borne on special branchlets arising from the corticating pseudo-
parenchyma ; perennial.

Kuetzing, Phyc. Gen. 1843, p. 293.

398 University of California Publications in Botany [Vol. s ,

Chaetopteris plumosa (Lyngb.) Kuetz.

Frond terete, 7-11 cm. high, the lower portion stipe-like and
nndivided, arborescent above, with the principal branches alternate
or opposite ; the stipe and principal branches nude ; the primary
branches closely beset with pinnate branches ; pinnae distichous, strict,
unbranched, 5-8 mm. long ; branches arising from the medulla ; color
of main branches dark, of the pinnae, olive brown, turning green on
drying ; unilocular sporangia spherical, plurilocular gametangia cylin-
drical, both borne on short lateral pedicels.

Growing in the sublittoral belt. St. Lawrence Island and Pt.
Clarence, Alaska.

Kuetzing, Phyc. Gen., 1843, p. 293 ; Tab. Phyc, vol. 6, 1856, pi. 6,
fig. I ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 239 ; Kjellman,
Om Beringh. Algfl., 1889, p. 51. Sphacelaria plumosa Lyngbye,
Hyclrophyt. Dan., 1819, p. 103, pi. 30c.

The occurrence of this species on our coast is known to us only
from the reference of Kjellman.

Order 2. ECTOCARPALES s. and g.

Confervoid (monosiphonous) or solid Phaeosporeae of varying
dimensions, habit and complexity, typically possessing unilocular zoo-
sporangia and plurilocular gametangia and with no portion of the cell
membranes turning black with eau de javelle; growth in length strictly
subapical, often ' ' trichothallic, ' ' or more or less intermediate between
typical forms of either ; both unilocular zoosporangia and plurilocular
gametangia in the same individual in some species, but, most commonly,
borne on different individuals, thus pointing toward an alternation of
generations, reduction division taking place in the primary nucleus of
the unilocular zoosporangium ; gametangia all similar or slightly differ-
ing in number and size of loculi ; gametophyte and sporophyte prac-
tically indistinguishable as to size and complexity.

Ectocarpales Setchell and Gardner, Phyc. Cont. VI, May 16, 1922,
p. 403 (lim. mut.) ; Oltmanns, Morph. und Biol, der Algen, vol. 2,
1922, p. 2 (lim. mut.). E ctocarpaceae Oltmanns, Morph. und Biol, der
Algen, vol. 1, 1904, p. 350 fin part). Ectocarpineae Tajdor, Recent
Studies of Phaeophyceae, etc., Dec, 1922, p. 436 (in part).

It seems more consistent with the present usage in other subclasses
of the thallophytes to consider the extended Ectocarpaceae of Oltmanns
as an Order rather than as a Family. Oltmanns in the last edition

1925] SetcJieU-Oardner : Melanophyceae 399

(1922) of his "Morphologie unci Biologie der Algen" (p. 2 ct seq.) ,
has come to the same conclusion. Our present conception of the
Ectocarpales differs from that of Oltmanns in limiting it to families
in "which plurilocular gametangia are present on gametophytes slightly,
if at all different from the sporophyte, or at least definitely macro-
scopic. While the families under Ectocarpales, as limited by us, pre-
sent a very considerable variety in the details of structural differences,
they have in common the possession of plurilocular gametangia giving
rise to nearly similar gametes (isogamy) and an identity in size and
structure of both gametophyte and sporophyte. The Ectocarpales are
closely related to the Cutleriales, but in the latter there is distinct
oogamy and in some species considerable differences between the two
generations. From the Sphacelariales, the Ectocarpales differ in the
absence of the conspicuous apical cell of the first mentioned and the
very different cell wall. The cell wall, or the older layers of it, changes
to black in the Sphacelariales when treated with eau de javelle, but
does not show this reaction in any of the Ectocarpales. The macro-
scopic plants of the members of the Dictyosiphonales, Desmarestiales
and Laminariales are known only with unilocular zoosporangia. The
first and last of these orders are known to have greatly reduced game-
tophytes. From analogy the Desmarestiales may be suspected of being
similar to the other two orders in this respect.

Key to the Families

1. Fronds prostrate, discoid, crustaceous, or pulvinate, never hollow nor carnose,

seldom gelatinous 2

1. Fronds erect, membranaceous or filamentous, when globular or prostrate, either

hollow or carnose 4

2. Fronds wholly discoid, composed of a basal disk of 1 or more layers of

cells giving rise to erect free filaments 3

2. Fronds crustaceous or pulvinate, consisting of a basal cushion of several
layers of cells with penetrating filaments below and erect, free fila-
ments above 5. Elachisteaceae (p. 502)

3. Fronds small, composed of a disk of 1 or 2 layers of cells all of which give rise

to free, erect filaments or reproductive cells

3. Myrionemataceae (p. 453)

3. Fronds larger (up to several cm. across), composed of a disk of more than 2

layers of cells which give rise to erect, free filaments in sori

4. Ralfsiaceae (p. 493)

4. Fronds globular or flattened, carnose 6. Leathesiaceae (p. 507)

4. Fronds elongated, erect when globular, not carnose 5

5. Fronds membranaceous or flattened, at times hollow 6

5. Fronds cylindrical, not membranaceous 7

6. Sori definite, entirely projecting 8. Asperococcaceae (p. 522)

6. Sori definite, superficial, at times partially projecting

7. Punctariaceae (p. 513)

6. Sori usually indefinite, superficial 10. Scytosiphonaceae (p. 530)

400 University of California Puhlications in Boiamy [Vol. 8

7. Fronds monosiphonous 2. Ectocarpaceae (p. 400)

7. Fronds more complex 8

8. Growth trichothallic 9

8. Growth subterminal, not distincth^ trichothallic 10

9. Sori distinct, more or less transversely arranged, superficial or projecting....

9. Striariaceae (p. 528)

9. Without distinct sori, sporangia and gametangia neither superficial nor project-
ing 11. .ffigiraceae fp. 543)

10. Fronds dichotomousty much branched 13. Chnoosporaceae (p. 552)

10. Fronds alternately branched on all sides

12. Heterochordariaceae (p. 549)

FAMILY 2. ECTOCAEPACEAE harvey

Thallns of monosiphonous filaments, occasionally partially poly-
siphonous, branched in various ways, arising from creeping, superficial
or at times penetrating filaments, occasionally arising from a small
superficial disk, usually entirely free from one another, occasionally
slightly intertwined or loosely united by a thin jelly ; growth proceed-
ing from a subapical cell, often situated at the base of a hair (tricho-
thallic) ; cells uninucleate with one or more parietal chromatophores
of fixed and definite form ; zoosporangia unilocular, terminal, either a
branchlet transformed wholly or in part, or intercalary ; gametangia
plurilocular, of various shapes, arising from a branchlet transformed
wholly or in part, or intercalary, in some cases differentiated as to
size and number of divisions (mega- and meio-gametangia) ; game-
tophyte and sporophyte identical in size and structure.

Ectocarpaceae Harvey, Ner. Bor.-Amer., part 1, 1852, p. 132 (in

part) ; Kjellman, in Engler and Prantl, Die natiirl. Pflanzenfam. Teil

1, Abt. 2 (Lief. 60), 1891, p. 182 (in part) ; Oltmanns, Morph. und

Biol, der Algen, vol. 2, 1922, p. 6 (in part). Ectocarpeae Agardh,

Syst. Alg., 1824, p. XXX (in part) ; Oltmanns, Morph. und Biol, der

Algen, vol. 1, 1904, p. 353 (in part). Kuetzing, Phyc. Gen., 1843,

p. 287 (in part) ; Oltmanns, Morph. und Biol, der Algen, vol. 1. 1904,

p. 353 (in part).

Key to the Genera

1. Zoosporangia and gametangia transformed branchlets or tips of branches or

branchlets, strictly terminal 2

1. Zoo.sporangia and gametangia catenate, intercalary 3. Pylaiella (p. 401)

2. Vegetative filaments superficial or penetrating the host merely by

rhizoids 4. Ectocarpus (p. 407)

2. Vegetative filaments deeply penetrating the host, projecting little, if
at all, beyond its surface 5. Streblonema (p. 440)

1^2o] Setchcll-Gardncr: MelanopJiyceae 401

1. Pylaiella Bory

Fronds monosiphonous, composed of a creeping, attaching portion
and an erect portion ; creeping filaments irregular, more or less pro-
f usety branched, not penetrating the uninjured host ; erect filaments
either unbranched or moderately to profusely branched, at times pili-
f erous ; chromatophores smooth, regular disks or irregular, more or
less united plates, or bands ; zoosporangia formed by direct transforma-
tion of assimilating cells of the erect filaments, mostly catenate, usually
intercalary, rarely terminal; gametangia similar to the zoosporangia
in origin and position, both zoospores and gametes escaping through
lateral pores.

Bory, in Diet, class., vol. 4, 1823, p. 393, vol. 13, 1828, p. 565
(original spelling Pilayella) ; Leman, "Pylaiella," in Levrault, Diet,
sci. nat., vol. 44, 1826, p. 127.

The genus Pylaiella was founded by Bory de Saint Vincent in 1823
and redescribed by him in 1828. In the first article, the description of
the seriate sporangia tends definitely to diagnose the genus, but in
the second article, the type species of the genus is designated as
Conferva littoralis L., and further indicated as being the Ectocarpus
littoralis 8 protensus Lyngbye (1819, pi. 42, C). In every way, there-
fore, there is little doubt as to the conception of Bory. Bory has the
spelling Pilayella which seems probably to be a purely orthographic
error. It seems best to us to keep the species of Pylaiella separate from
those of Ectocarpus since the seriate, and largely intercalary, zoo-
sporangia differ decidedly from the projecting and solitary zoo-
sporangia of Ectocarpus. The gametangia in Pylaiella are, in typical
form, also very distinct from those of Ectocarpus. We, at least, basing
our opinion on the development and dehiscence, are inclined to regard
the gametangia in Pylaiella as seriate, each separate gametangium
opening laterally. In Ectocarpus, the gametangia are single, since all
the loculi open inwardly and the gametes are discharged through the
apex, even dislocating the terminal hair in forms of Ectocarpus sili-
culosus which, otherwise, has gametangia at times resembling, super-
ficially, the seriate gametangia of Pylaiella. In attempting to arrange
the numerous forms of Pylaiella, as found on the Pacific Coast of
North America, we have come to the conclusion that at least four
species are represented.

Three subgenera are described under Pylaiella, viz., Eicpylaiella
Born., Bachelotia Born., and Panthocarpus Scottsb. The first and last

402 University of California PiMications in Botamy [Vol. s

are represented by species occurring within our territory, but Bache-
lotia does not seem to have been detected here as yet. The subgenus
EiipylaieUa, whose species are distinguished by their comparatively
more frequent opposite or alternate branching, is founded naturally on
Pylaiella littoralis, some forms of which are very tall, but also includes
low forms, such as P. curta Foslie and P. nnilateralis S. and G. Bache-
lotia and Panthocarpus, whose erect filaments are either unbranched
or sparsely branched, include only forms not over three centimeters in
height. In Eupylaiella, and probably also in Bachelotia (known cer-
tainly only with zoosporangia) , the gametangia develop simultaneously
throughout the series which is, therefore, limited and definite. In
Panthocarpus, the gametangia develop successively and from above
downward. Further and more careful study of gametangial develop-
ment in a large series of forms and species is necessary to fully sustain
these statements.

Key to the Species

1. Branching frequent, opposite, alternate or secund — (Eupylaiella) 2

1. Branching none or sparse and opposite — (Panthocarpus) 3

2. Upper branches opposite or alternate 1. P. littoralis (p. 402)

2. Upper branches regularly secund 2. P. unilateralis (p. 404)

3. Plants up to 15 mm. high, simple or with opposite ramuli above

3. P. Gardneri (p. 405)

3. Plants not over 1 mm. high, simple 4. P. tenella (p. 406)

1. Pylaiella littoralis (L.) Kjellman
Plate 37, fig. 32

Erect assimilatory filaments very profusely branched, either densely
caespitose or funiculate, 2 cm. to 6 dm. hig'h, creeping filaments richly
branched, somewhat irregular in diameter ; color varying from light
yellowish brown to dark brown; primary filaments 20-60jti diam., cells
1-6 times as long as the diameter, ramuli more slender, opposite or
alternate, strict, patent or even at times recurved, short and subulate
or long and gradually attenuated, occasionally pilif erous ; zoosporangia
in each series mostly formed simultaneously, 5-25-catenate, usually
one in each joint, rarely 2-4 formed by longitudinal divisions of a
single cell, intercalary or terminal, varying in shape from oblate to
prolate spheroidal, 25-45/x diam. ; gametangia intercalary, in a con-
tinuous series up to ISOOju, long, 20-35ju, diam. ; zoosporangia and game-
tangia usually on separate plants, but occasionally on the same plant.

Growing on other algae, on rocks and on wood in the littoral belt.
Bering Sea to southern California (San Pedro).

1^25] Setchcl I -Gardner: Melanophyceae 403

Kjellman, Skand. Eet. och. Tilopt., 1872, p. 99, Alg. Arctic Sea,
1883, p. 281 ; KSetchell and Gardner, Alg. N.W. Amer., 1903, p. 235 ;
Saunders, Phyc. Mem., 1898, p. 156, Alg. Harriman Exp., 1901, p. 418.
Conferva Uttoralis Linnaeus, Spec, PI. (ed. 1), 1753, p. 1165 (in part) ;
Dillwyn, Brit. Conf., 1803, p. 70, pi. 31 (in part) ; C. A. Agardh, Syn.
Alg. Scand., 1817, p. 65 (in part), Spec. Alg., vol. 2, 1828, p. 40 (in
part). Ectocarpus Uttoralis 8 protensus Lyngbye, Hydrophyt. Dan.,
1819, p. 131, pi. 42, C.

We have quoted both Linnaeus and Kjellman as authorities for the
name of this species, with a full sense of the uncertainty existing con-
cerning the real sponsorship. The Conferva Uttoralis of Linnaeus, so
far as the original founding of the species is concerned, is most uncer-
tain since no definite type specimen is to be found. According to C. A.
Agardh (1828, pp. 40, 41), Linnaeus seems to have changed his
opinion as the years went on, at certain times having in mind appar-
ently one of the Ectocarpaceae, and at other times a member of the
Rhodomelaceae. In the first edition of the Species Plantarum (1753,
p. 1165), Linnaeus quotes the first edition of the Flora Suecica (1745,
p. 371) and a certain figure of Dillenius (1741, pi. 4, fig. 19). In the
Flora Suecica, however, he quotes a figure of Dillenius (1741, pi. 3,
fig. 13) to which he did not refer especially in 1753. In the second edi-
tion of the Species Plantarum (1763, p. 1635), Linnaeus again excludes
the Dillenius figure of plate 3, figure 13, but, later, in the second edi-
tion of the Flora Suecica (1755, p. 436), he again quotes only figure 13
of plate 3 repeating the diagnosis of the species published in 1753.
The Eetocarpaceous view of the Linnaean species seems to have been
general among the earlier phycologists, but, even thus, it remained a
mixture of species. Dillwyn (1803, pi. 31) shows under figure C a
plant seemingly with seriate gametangia, while figure D has what seems
to be a projecting solitary zoosporangium. The idea of C. A. Agardh
1817, p. 65) is also of a mixture, as was that of Lyngbye (1819,
pp. 130, 131, pi. 42) as well. By the time the first edition of Harvey's
Manual (1841) was issued, the' present idea of the species had begun to
assume form as to the seriate gametangia being characteristic of it.
While it is, therefore, fairly certain that Linnaeus may have included
the species now usually assigned under his specific name, but that he
had no distinctly concrete conception of the same, it seems best to fol-
low custom and not only ta adopt the specific name Uttoralis but also to
continue to cite Linnaeus as the author of it. We have quoted Kjell-
man as authority for the binomial, although, in 1828, Bory distinctly
cited the Conferva Uttoralis L. as the type of Pylaiella. He did not

404 University of California Publications in Botany [Vol.8

use the binomial itself and we follow custom in citing as authority the
first writer known to us to do so. .

Pylaiella littoralis, as usually recognized, is a very variable and
widespread species. In height, it varies from a few centimeters up to
six decimeters or more ; in color, from light yellowish brown to very
dark brown; in habit from a feathery frond of loosely entangled fila-
ments, through tufted masses to rope-like aggregations; in branching
from very abundant, opposite or alternate, to comparatively scanty
and chiefly alternate ; and in angle of branching, from narrowly acute
to broadly obtuse, erect, ascending or even recurved. There is con-
siderable variation in the diameters of the primary filaments and in
the dimensions and position of the zoosporangia and gametangia. It
is difficult, or even impossible at present, to determine how much of
this variation is due to age and environment and how much may be
of genetic significance. There are several varieties described with
more or less definiteness, and few to several forms under each. Earlier
(1903), we attempted to refer as many of the plants of the north-
western coast as possible to their proper forms. At present, it seems
to us undesirable, as well as very unsatisfactory, to attempt this. The
var. opposita Kjellm., loose in habit and with abundant, chiefly
opposite, branching, may be taken as including the normal, or also
even the taxonomic, type. The var. firma (Ag.) Kjellm. is also lax
in habit but with less abundant and more scattered branching. Var.
divaricata Kjellm. is more commonly funiculose or ropy and with the
branches irregularly placed, often incurved and geniculate. The
forms of var. varia (Kjellm.) Kuck. seem the most distinct in their
peculiarly aggregated gametangia and contorted funiculose habit.
Forms of all four, and perhaps even of other varieties, occur on our
coast and a study of the same plants over a series of weeks or months,
or in cultures, would undoubtedly assist much in elucidating the
significance of the great variation credited to this species.

2. Pylaiella unilateralis S. and G.

Erect filaments attached by creeping, irregular, more or less
branched filaments, tufted, slightly clavate, 4-7 mm. high, 19-22^
diam, at the bases, 28-32^ diam. at the apices; branching mostly
alternate and sparse below, secund above, and arising from cells
divided longitudinally 1-2 times; cells 2-2.5 times as long as the
diameter and cylindrical at the base, more or less quadrate and doli-
iform above; ehromatophores thin, irregular plates more or less con-

1^-5] Setchell-Gardner: Melanophyceae 405

nec'ted ; zoosporangia formed simultaneously, 7-12 catenate, mostly
terminal in the ramuli, spheroidal, 24-28/x diam. ; gametangia unknown.

Growing on rocks in shallow pools near high-tide limit. Sunset
Beach, near the mouth of Coos Bay, Oregon.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 386, pi. 42, figs. 6-8.

The secund branching together with the longitudinal divisions of
the cells from Avhich the secund branches arise, forming a polysiphon-
ous region, are the chief distinguishing characteristics of this species.
The branching below is sparse, alternate or very rarely opposite, and
the branches are relatively long and attenuated. These branches may
give rise to terminal zoosporangia, or to a few short alternate ramuli
which in turn produce the zoosporangia. Many of the main filaments
become slightly enlarged towards their outer ends, the cells becoming
nearly quadrate, their walls thickened and their contents very dense.
An average of about thirty-five, although frequently as many as sixty-
five, cells are thus transformed. A few cells of the ends of the fila-
ments are not transformed and soon disintegrate, thus producing
decidedly clavate filaments. The region of transformation seems to
be a new meristem of a peculiar nature. Many of the cells divide
once or, less frequently, twice, by longitudinal planes as a rule, but
occasionally the dividing planes are at right angles separating the
cell into four parts. The unique feature resulting from these divi-
sions is the lack of further growth of the cells. Usually one of the
cells resulting from longitudinal division gives rise to a branch,
shorter or longer, but similar to branches arising in the regular man-
ner. The branches are typically secund, but may rarely come from
the opposite side of the filament. Many of the main filaments become
much curved in this branching region.

3. Pylaiella Gardner! Collins

Erect assimilating filaments slightly attenuated when young,
unbranched or sparsely forked near the base giving rise above to
numerous, short, obtuse, mostly opposite, rigid, patent branches,
10-12 mm. (up to 20 mm.) high; cells at base of filaments 12-15/x
diam., twice as long as broad, increasing in diameter slightly upwards,
moderately constricted at the joints; creeping filaments 6-10|a diam.,
cells 1-4 long; chromatophores thick, broken bands nearly filling
the cell ; zoosporangia unknown ; gametangia in terminal, more or less
clavate, series, either on the main filaments or on short lateral branches,
up to 24/i, diam.

406 University of California Puhlications in Botany ' [Vol. 8

Forming low patches on Postelsia pahnaeformis. Observed only
on the coast of central California, but it probably has a much wider
distribution.

Collins, in Collins, Holden and Setcheli, Phyc. Bor.-Amer.
(Exsicc), no. 1384. PylaieUa {Panthocarpus) Postehiae Skottsberg,
Notes on Pac. Coast Alg*. I, 1915, pp. 153-164, jjls. 17-19. Leptonema
fasciculatum Saunders, New and Little Known Brown Algae, Erythea,
vol. 7, 1899, p. 38, figs. 5-7 (not of Reinke).

Saunders (loc. cit.) reports the species as growing on the shells of
the mollusk, Mytilus, in the. vicinity of Postelsia. We have not seen
it growing elsewhere than on the above named host. The gametangia
are intercalary rather than terminal. There are always a few terminal,
assimilating cells which do not metamorphose into gametangia. The
creeping filaments do not seem to penetrate the uninjured host at
any point, but if the surface cells of the host become bruised or in
any way destroyed, the extending filaments follow the irregularities
of the surface, and at times even penetrate a short distance among
the cells beneath.

4. Pylaiella tenella S. and G.

Fronds widely diffused, attached by small, short, branched, con-
torted filaments, unbranched, 0,5-0.75 mm. high, long-attenuate
upwards, not piliferous; cells 7-lOja diam., 1-2.5 times as long as the
diameter ; chromatophores, when young, a parietal broken band nearly
filling the cell, later separating into several distinct pieces; zoo-
sporangia long-catenate, subterminal, numerous assimilating cells in
the series forming two or four zoosporangia by longitudinal divisions ;
gametangia subterminal.

Growing on Pleurophycus Gardneri Setcheli and Saunders, the
plants forming minute tufts, which later become confluent. Neali Bay,
near Cape Flattery, Washington.

Setcheli and Gardner, Phyc. Cont. V, 1922, p. 385, pi. 42, figs. 9-11.

This Pylaiella is the most diminutive of all the known species of
the genus, rarely attaining the length of one millimeter, its nearest
rival in this respect being P. nana of Kjellmaii, from tlie Norwegian
Polar Sea. It differs from that species in size, being only about one-
half as high, in not being branched and in having subterminal game-
tangia in a long series, instead of terminal branched ones as described
and figured by Kjellman for P. nana. The double and quadruple
zoosporangia observed in this species, formed by longitudinal and

1925] SetchcU-Gardner: Melanophyceae 407

cross-divisions of fructiferous cells have likewise been observed by
Borgesen (1920, p. 433) in P. fulvescens (Schousb.) Bornet, from the
Danish West Indies. We have also observed this condition highly
developed in forms of P. littoralis.

P. tenella evidently belongs to the Panthocarpus group of the
genus as established by Skottsberg (1915). The gametangia are
formed by the transformation of vegetative cells, usually beginning
relatively few cells back of the apices of the filaments, a considerable
number of cells transforming simultaneously. The transformation
continues in both directions until one-half lo three-fourths of the
filament is converted into gametangia. The gametangia vary greatly
in size, some producing as many as thirty-two loculi from a single
assimilating cell. They may be continuous or discontinuous, certain
assimilating cells in the series not being transformed. The cell walls
in the formation of loculi frequently are decidedly oblique. Zoo-
sporangia occupy very much the same position in the filaments as the
gametangia. They are subterminal, long-catenate, but seem to be
formed more nearly simultaneously than are the gametangia.

The plants were fairly abundant on the blade of the host-plant at
Neah Bay. They are, however, very inconsijicuous, forming small,
more or less continuous expansions, barely recognizable when the host-
plant is wet. We have not observed the species growing elsewhere,
although the host is abundant in the vicinity of the Strait of Juan
de Fuca.

4. Ectocarpus Lyngb.

Fronds monosiphonous, composed of a creeping attaching portion
and an erect portion; creeping filaments irregular, more or less well-
developed, and more or less richly branched, usually entirely or
largely external to the substratum ; erect filaments usually abundantly
branched, naked, or corticated by descending rhizoidal filaments,
piliferous, or terminating in an acute or rounded cell, growth apical,
subapical or trichothallic ; chromatophores discoid or band-shaped,
with or without pyrenoids ; zoosporangia on lateral branchlets, seldom
terminal, single, sessile or stalked ; gametangia terminal on lateral
branchlets, sessile or stalked, exceptionally seriate and seldom even
intercalary, in some species dimorphic or polymorphic (mega- and
meio- gametangia and antheridia).

Lyngbye, Hydrophyt. Dan., 1819, p. 130 (in part) ; Bory, in Diet,
class, d'hist. nat., vol. 6, 1824, p. 63. Colophermum Rafinesque, Precis
decouv. som., 1814, p. 49.

408 University of California Publications in Botany [Vol. 8

Lyng'bye established the genus Ecto-carpus on six species, the last
two mentioned of which are not even members of the Melanophyceae.
The other four are wholly, or in part, members of the genus as limited
by Bory, in which sense it has more recently been taken by many
writers. The only confusion has been with Pylaiella and there may be
a question as to the wisdom of segregating that. The first species listed
by Lyngbye is Ectocarpus littoralis but, as we have already indicated,
Lyngbye's conception of this species included a true Ectocarpus as
well as what we now separate as Pylaiella. littoralis. Otto Kuntze
(1891, p. 888 and 1898, p. 401) has attempted to revive the Rafines-
quian name Colophermum (1814) for Ectocarpus, but the latter name
has been included among the "nomina conservanda" by the Brussels
International Botanical Congress (cf. Briquet, 1912, p. 75).

We have already set forth the distinctive features of the genus
Pylaiella, separating it from the other genera of the family. Ecto-
carpus differs from Strehlonema in that the creeping basal filaments
either do not penetrate the host-plant at all, or do so only very slightly,
or, at least, the greater portion of the frond is free. It is frequently
a delicate matter to decide whether a certain species may better be
referred to Ectocarpus or to StreMonema, yet it seems best to keep
them separate.

In habit and stature, the species of Ectocarpus differ considerabh^,
some being very low and dwarfed, while others are tall; some are
tufted, some are feathery, while some have their filaments entangled
into ropy masses. The zoosporangia are either more rare or less fre-
quently found than the gametangia and are unknown in many species.
The gametangia differ much in the various species, in shape, size and
position. In some species they are dimorphic or even polymorphic
(Sauvageau, 1896, pp. 27 and 32, 1896a, 1896&, etc.). In some species
there are "megasporangia" (as Sauvageau calls them) Avith large
divisions (loculi), " meiosporangia " (Sauvageau) with medium sized
divisions (loculi), and antheridia with very small divisions (loculi).
It is supposed that the motile sperms are produced by the antheridia
and one or the other of the mega- or meio- gametangia produce the
egg cells (cf. Sauvageau, 1896c). Much more experience is needed
before we may be able to decide the question of the function of these
different gametangial structures. They are usually lateral and either
sessile or pedicellate. More rarely are they terminal and, at times,
even intercalary^ (e-g-, E. siliculosus f. varians Kuckuck, 1892, p. 256,
pi. 13).

1925] Setchell-Gardncr : MeJanophyceae 409

The species of Eciocarpus are numerous and often very puzzling.
The principal accounts, thus far published for our coast, are those
of Saunders (1898 and 1901) and that in our own report on the algae
of the northwestern coast (1903) as well as in our Phycological Con-
tributions VI (1922). We have subjected all of the material available
to us to careful scrutiny in the preparation of the following treatment
of our species.

Key to the Species

1. Chromatophores bftnd-shaped 2

I. Chromatophores discoid 20

2. Fronds forming hemispherical tufts, growth apical or subapical

21. E. chantransioides (p. 430)

2. Fronds not as above, growth chiefly intercalary 3

3. Gametangia on both creeping and erect filaments 4

3. Gametangia on erect filaments o'niy (except E. confervoides f. variabilis) 5

4. Erect filaments 8-10;u diam 8. E. Taoniae (p. 420)

4. Erect filaments 12-25^ diam E. confervoides f. pygmaeus (p. 415)

5. Fronds usually more than 1 cm. high 6

5. Fronds less than 1 cm. high 14

6. Fronds tightly twisted into branched, rope-like threads

4. E. tomentosus (p. 417)

6. Fronds more or less loosely tufted 7

7. Fronds over 5 cm. high 8

7. Fronds usually less than 5 cm. high 10

8. Terminal ramuli short and acute 3. E. acutus (p. 415)

8. Terminal ramuli long-attenuated 9

9. Gametangia not terminating in hairs 2. E. confervoides (p. 412)

9. Gametangia frequently terminating in hairs 13

10. Gametangia usually over lOO^i long E. confervoides f. parvus (p. 413)

10. Gametangia usually less than lOOyu long 11

II. Fronds never corticated 12

11. Fronds corticated 5. E. corticulatus (p. 418)

12. Gametangia usually 50-70m long 6. E. fructuosus (p. 419)

12. Gametangia usually 75-90ai long E. confervoides f. variabilis (p. 414)

13. Fronds loose, up to 20 cm. high 1. E. siliculosus (p. 410)

13. Fronds tufted, up to 6 cm. high E. siliculosus f. subulatus (p. 411)

14. Fronds penetrating the host more or less 15

14. Fronds not penetrating the host 18

15. Fronds more or less profusely branched 16

15. Fronds mostly simple 17

16. Gametangia long, attenuated 12. E. Mesogloiae (p. 423)

16. Gametangia short, blunt 13. E. commensalis (p. 424)

17. Fronds over 1 mm. high 14. E. eramosus (p. 424)

17. Fronds le.ss than 1 mm. high 17a

17a. Erect filaments terminating in a hair 15. E. luteolus (p. 425)

17a. Erect filaments not terminating in a hair 33. E. gonodioides (p. 439)

18. Fronds attached by creeping filaments and by rhizoidal filaments

from the lower cells 11. E. flagelliferus (p. 422)

18. Fronds attached by creeping filaments only 19

19. Fronds forming a velvety stratum 9. E. terminalis (p. 421)

410 University of Calif arnia Piiblications in Botany [Vol. 8

19. Fronds caespitose 10. E. simulans (p. 422)

20. Fronds 1 cm. or more high 21

20. Fronds less than 1 cm. high 26

21. Fronds twisted into dense rope-hke threads 20. E. breviarticulatus fp. 429)

21. Fronds more or less loose and caespitose 22

22. Ramuli mostly alternate 23

22. Ramuli mostly secund 25

23. Main filaments 25-40^ diam 24

23. Main filaments 40-60^ diam 19. E. oviger (p. 428)

24. Gametangia broadly fusiform 17. E. Parksii (p. 427)

24. Gametangia ellipsoidal, obtuse 18. E. Mitchellae (p. 428)

24. Gametangia broadty ovoid, mucronate 7. E. mucronatus fp. 419)

25. Gametangia 60-lOOyu long 16. E. granulosus (p. 426)

25. Gametangia 40-60m long 22. E. granulosoides (p. 431)

26. Fronds penetrating the host 27

26. Fronds not penetrating the host 30

27. Cells of the main filaments less than 30^1 diam 28

27. Cells of the main filaments more than 30^ diam 29

28. Gametangia fusiform, scattered on the upper parts of the fronds

30. E. socialis (p. 437)

28. Gametangia ovoid to ellipsoid, mostly near the base of the fronds

31. E. flocculiformis (p. 438)

29. Gametangia cylindrical, many opposite E. cylindricus f. codiophilus (p. 433)

29. Gametangia globular, mostly alternate 32. E. globifer (p. 438)

29. Gametangia fusiform, mostly alternate 34. E. Bryantii (p. 440)

30. Fronds less than 1 mm. high 23. E. affinis (p. 432)

30. Fronds more than 1 mm. high 31

31. Gametangia chiefly cylindrical 32

31. Gametangia not cylindrical 33

32. Fronds 1-2 mm. high 24. E. cylindricus (p. 432)

32. Fronds 7-10 mm. high E. cylindricus f. acmaeophilus (p. 433)

33. Fronds in dense hemispherical tufts 34

33. Fronds more or less caespitose 35

34. Fronds much branched, 2-4 mm. high 25. E. hemisphericus (p. 434)

34. Fronds mostly simple, 1-2 mm. high.E. hemisphericus f. minor (p. 434)

35. Cells over 20^ diam 26. E. Saundersii (p. 434)

35. Cells less than 20m diam 36

36. Gametangia long-acuminate, up to 400/xlong.27. E. acumdnatus (p. 435)

36. Gametangia not long-acuminate, up to 250/x long 37

37. Fronds 9-12/x diam 28. E. ellipticus (p. 436)

37. Fronds 14-15^ diam 29. E. chitonicola (p. 436)

1. Ectocarpus siliculosus (Dillw.) Lyngb.

Frondfs tufted, feathery, loose, more or less entangled below,
yellowish or brownish, pseudodichotomously branched below, alter-
nately secund above ; branches erect or, at times, somewhat recurved,
never patent, not fascicled ; sporangia ovoid to ellipsoidal, 30-65^
(usually 50ju,) long, 20-27/x broad, sessile and erect or diverging on
a 1-3-celled pedicel ; gametangia usually long subulate-conical, seldom

1925] Setchell-Gardner: MeJanophyceae 411

ovoid or curved, often ending in a longer or shorter hair, 500-600|U,
long- and 12-25/,i broad, seldom sessile, usually short pedicelled.

Growing in quiet pools attached to other algae. San Francisco
Bay, California.

Lyngbye, Hydrophyt. Dan., 181!), p. 131 (in part, as to pi. 43,
fig. C); Kjellman, Handb., I, 1890, p. 78; Kuckuck, Beitr. Kennt.
Ectocarp. Art., 1891, p. 65; Collins, Holden and Setchell, Phyc. Bor.-
Amer. (Exsicc), no. 2294. Conferva silicidosa Dillwyn, Brit. Conf.,
1809, p. 69, pi. E (excl. synonym).

We have followed Kjellman and Kuckuck in keeping Ectocarpus
siliculosus distinct from E. confervoides and in laying emphasis upon
the presence of elongated slender gametangia, some of which are
piliferous, as its chief characteristic. We know full well from our
own experience that it is difficult to draw a satisfactory line of
demarcation between the two species, but feel that the best we can
do is to keep tlieni distinct and draw the line somewhat arbitrarily.
The existence of such forms as Ectocarpus confervoides f. parvus
(Saunders) S. and G. indicates that there are perplexing forms whose
relationships and origins are perhaps impossible to formulate, but
otherwise, so far as the plants of our coast thus far discovered are
concerned, there seems little doubt that they conform to the original
idea of E. siliculosus.

The plants distributed in the Phycotheca Boreali-Americana
(Joe. cit.) are the only characteristic plants of the species known on
our coast. These were taken from a quiet pool in a salt marsh on
Bay Farm Island, Alameda, California.

Ectocarpus siliculosus f. subulatus (Kuetz.) S. and G.

Fronds 5-25 cm. high, light yellow, fleecy, much branched, not
constricted at the joints; branches long, attenuated above, many end-
ing in a long hair ; cells of the main filaments 30-36ja broad, 1-1.25
times as long as broad ; zoosporangia unknown ; gametangia elongated
subulate-ovoid, some stouter, some more slender, 200-600/^ long, 12-48/x
broad, the upper (and as a rule more slender) usually terminating
in a hair, on a 2-10 (or 12) celled pedicel.

In brackish pools, on sticks or grasses. Central California (San
Francisco Bay).

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 416. Ectocarpus
confervoides f. suhulatm Collins, Holden and Setchell, Phyc. Bor.-
Amer. (Exsicc), no. 1231. Ectocarpus subulatus Kuetzing, Actien,

412 University of California Puhlications in Botany ['Voi.. 8

1836, Phyc. Gen., 1843, p. 287. Sp. Alg., 1849, p. 454, Tab. Phyc,
vol. 5, 1855, p. 19, pi. 61, fig. II. Ectocarpus confervoides (3 subidatiis
Hauck, Meeresalg., 1884, p. 331 (excl. synonymy). Ectocarpus
amphihius Harvey, Phyc. Brit., vol. 2, 1848, pi. 183, Ner. Bor.-Amer.,
part 3, 1858, p. 125.

Our plant seems to agree very well with the figures and descrip-
tion of Ectocarpus ampkibius Harvey and also, although not so per-
fectly, with E. sutidatus Kuetz. They are both brackish water forms,
as are our plants. Hauck refers to his var. subulahis also Ectocarpus
drapar^naldiaeformis Kuetz. and E. macroceras Kuetz. Judging from
Kuetzing's illustrations, these two species may be forms of E. sili-
cuJosus but not to be included under f. subtdatus.

2. Ectocarpus confervoides (Roth) Le Jolis

Fronds gregarious, forming expansions or tufts, 0.5-10 cm. high,
light yellowish to dark brown, sparingly to frequently branched,
branching alternate or secund, never opposite, more or less cor-
ticated; branches long or short, gradually attenuated, sometimes end-
ing in a hair ; prostrate filaments irregular, branched, closely attached
to the substratum, but never penetrating it ; chromatophores band-
shaped, branched, often provided with pyrenoids; zoosporangia pre-
sent in some forms ; gametangia short-obovoid or longer or shorter
fusiform, suddenly or somewhat gradually attenuated toward the
blunt apex, never ending in a hair, 30-250/x (commonly 60-150ju,)
long and 12-35ju, thick, solitary or seriate, terminal or intercalary.

Growing on rocks, wood, and larger Melanophyceae, widely dis-
tributed in its various forms. Alaska (Yakutat Bay) to Washington
(Whidbey Island).

Le Jolis, Liste alg. mar. Cherb., 1863, p. 75 (in part) ; Kjellman,
Handb., I, 1890, p. 77 ; Kuckuck, Beitr. Kennt. Ect. Arten, 1891, p. 69 ;
Saunders, Alg. Harriman Exp. 1901, p. 418 ; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 237. Ceramium confervoides Roth, Cat.
Bot., I, .1797, p. 151, pi. 8, fig. 3. Ectocarpus silicidosus Lyngbye,
Hydrophyt. Dan., 1819, p. 131, pi. 43B.

Kjellman was the first to segregate Ectocarpus siliculosus from
E. confervoides and limit it in the sense in which it is understood
by Kuckuck and most writers since that time. Even in this narrow
sense, Ectocarpus confervoides includes some eight or ten reasonably
distinct forms, some of which may ultimately be found to deserve
specific rank. It is to be distinguished from Ectocarpus silicidosus

1925] 8 etchcU -Gardner: MeJanophyceae 413

by never having the gametangium prolonged into a hair, from E.
acutus by having blunt or piliferous branches, and from all the species
of Ectocarpus having band-shaped chromatophores by the shape and
dimensions of its gametangia.

Key to the Forms

1. Gametangia seldom over 100^ long 2

1. Gametangia usually over lOOyu long, even np to 400yu long f. parvus (p. 413)

2. Fronds frequently branched above, branches long f. typicus (p. 414)

2. Fronds sparsely branched above, branches short 3

3. Gametangia frequent above, seldom terminal f. variabilis (p. 414)

3. Gametangia occasional above, frequently terminal f. pygmaeus (p. 415)

Ectocarpus confervoides f. parvus (Saunders) S. and G.

Fronds gregarious, forming tufted or extended masses, 1-2 cm.
high, more or less alternately branched ; erect filaments 21-30/x broad,
slightly constricted at the joints, much attenuated at the tips or end-
ing in hairs ; cells 1-3 times as long as broad below, shorter above ;
chromatophores irregularly band-shaped ; zoosporangia ovoid or ellips-
oid, 35-55/x long, 20-27ja broad, usually sessile, sometimes on the same
filament with the gametangia; gametangia narowly lanceolate-conical,
gradually long-attenuate above, 120-400/i, long, 20-27fi broad, borne
on a longer or shorter pedicel, occasionally seriate and intercalary
(as in Pylaiella) .

On sand-covered rocks (type!), logs, or on the larger Melano-
phyceae. Central California (near San Francisco) and southern
California (San Pedro and the type locality, San Diego).

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 414. Ectocarpus
silicuJosus parvus Saunders, Phyc. Mem. 1898, p. 153, pi. 22, figs. 1-9.

The f. parvus, as defined above, presents certain resemblances to
both Ectocarpus silicidasus and E. confervoides. Its slender elongated
gametangia resemble those of E. siliculosus and much might be said
in favor of the opinion of Saunders in making it a " variety ' ' of that
species. On the other hand, very few, or none, of the typical game-
tangia end in a hair and we have therefore preferred rather to place
it with E. confervoides. There are often found in this form, as well
as in forms of E. siliculosus, very curious elongated intercalary game-
tangia which seem to be seriate, in that they open laterally in sections,
as do those of Pylaiella. Such gametangia are very puzzling, but
may possibly be regarded as indicating hybridization between Pylai-
ella and Ectocarpus or a mutation of some sort. Speculation, how-
ever, can scarcely explain such anomalies. The majority of the game-

414 University of California Publications in Botany [Vol. 8

tangia are typically of Ectocarpus and while approaching in shape
those of E. silicidosus, yet they are less slender and less elongated than
is typical for that species and are not ordinarily prolonged into
terminal hairs.

So far as we may judge from the scanty specimen in our copy,
no. 358 of Tilden's American Algae (from Pacific Grove) belongs
rather under f. parvus than f. variabilis, although the determination
as f . variahilis is attributed to Saunders,

Ectocarpus confervoides f. typicus Kuckuck (Orthog. mut.)

Fronds 1-10 cm. (ours 3-10 cm.) high, feathery, profusely
branched ; primary filament 25-32/1, diam., not constricted at the parti-
tions ; zoosporangia unknown ; gametangia abundant, alternate or
secund on the branches, sessile or upon a one to several celled pedicel,
40-80/x (up to 200jtt) long, 20-3 5ja diam.

On wood and rocks. Alaska (Juneau) to California (San
Francisco).

Kuckuck. Beitr. Kennt. Ect. Arten, 1891, p. 69, fig. 3; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 237 (excl. spec'n).

It may be doubtful whether the f. typicus of Ectocarpus confer-
voides is, as yet at least, to be included in our account. We have
referred here three collections, one of which (no. 5197, W. A. Setchell
and A. A. Lawson, Juneau, Alaska) was listed by us previously (1903,
p. 237) under E. siliculosus f. typicus. It is a poor and unsatisfactory
specimen, but seems to belong here rather than where we referred it.
The other specimens are from Oregon and California and are small,
but have the more abundant branching of this form.

Ectocarpus confervoides f. variabilis Saunders

Plate 46, fig. 7

Fronds forming more or less extended layers, 0.2-2 cm. high,
sparingly branched, branches alternate, distant, gradually attenuated
above ; main filaments up to 30/x diam., not constricted at the parti-
tions ; chromatophores large, band-shaped, few in each cell ; zoo-
sporangia ovoid, 60jU, long, 40ju, broad, single on 1-3-celled pedicels,
often on the same plant with the gametangia; gametangia abundant,
cylindrico-conical, fusiform to broadly ovoid, 75-225/x long, 16-35/a
broad, sessile or on 1 -few-celled pedicels, lateral or occasionally ter-
minal on a long pedicel from the basal filament, rarely seriate, terminal
or intercalary, on the long erect filaments, and up to 300/x. long.

^■•-•5] SetcheU-Gardncr : MeJanophyceae 415

On larger Melanophyceae {Dcsniarestia and the Laminariaceae).
Puget Sound to central California (Carmel).

Saunders, Pliye. Mem., 1898, p. 155, pi. 28 ; Sotchell and Gardner,
Alg. N.W. Amer., 1903, p. 238 (at least in part). Ectocarpiis con-
fervoidcs f. nanm Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsiec). no. 1737 (not of Kuckuck).

Saunders founded his f. variabilk on a plant common along the
coast from Puget Sound to central California. He named it, how-
ever, for the variable structures which he considered to be zoospor-
angia, but which are, most probably, undeveloped (parasitized or
abortive ?) gametangia such as Sauvageau (1896a, pp. 41-45) has
shown to exist in Ectocarpiis confervoides and other species of Europe.
The form, however, seems otherwise distinct and is fairly readily
recognized by the characters given above and ijidicated in the key to
the forms of E. confervoides.

Ectocarpus confervoides f. pygmaeus (Aresch.) Kjellm.

Fronds forming a more or- less extended layer, 1-12 mm. (mostly
1-3 mm.) high, unbranched, or slightly so, above, branches only
slightly tapering at the apices; erect filaments 12-25/* diam., not
constricted at the joints, cells 2-3 times as long as the diameter below,
shorter above; chromatophores few, irregularly band-shaped; zoo-
sporangia uncertain; gametangia terminal or lateral on the erect
filaments, sessile or on short pedicels and often abundant on the creep-
ing filaments, broadly conical or fusiform, abruptly narrowed at the
apex, 60-lOOjit long, 20-30/x broad.

On various larger Melanophyeeae, Alaska (Shumagin Islands)
to central California (Pacific Grove).

Kjellman, Handb., I, 1890, p. 77; Saunders, Phyc. Mem., 1898,
p. 154. pi. IT), figs. 5-9 ; Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc). no. 525. Ectocarpiis terminalis Collins, Holden and
Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1387. Ectocarpus confer-
voides variabilis Tilden, Amer. Alg. (Exsicc), no. 358. Ectocarpus
pygmaeus Areschoug, in Kjellman, Skand. Ectocarp., 1872, p. 85.

3. Ectocarpus acutus S. and G.

Fronds 5-9 cm. high, dark brown when dried, olive green when
living, feathery ; erect filaments more or less entangled and fasciculate
below, free above, profusely branched, main filaments densely cor-
ticated ; branches mostly alternate, strict ; ultimate ramuli often

416 University of California Publications in Botany [Vol. 8

secund, acute-subulate; cells slightly doliiform, 40-60;a diam., 0.4-2
times as long as broad (usually shorter than the breadth) ; chromato-
phores thick, irregularly branched bands, few in a cell, containing
several pyrenoids ; zoosporangia unknown ; gametangia numerous and
regularly placed, cylindrico-conical, 100-1 50/a (up to 230;U.) long,
20-35/x broad.

Growing, for the most part at least, on larger Melanophyceae.
Puget Sound to central California (Carmel).

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 404, pi. 48, figs.
36-39 and pi. 49, figs. 40, 41. Ectocarpus penicillatus Saunders, Phyc.
Mem., 1898, p. 155, pi. 21, figs. 3, 4 (not of Kjellman). Ectocarpus
confervoides f. acuminatus Collins, in Setchell and Gardner, Alg.
N.W. Amer., 1903, p. 237, Mar. Alg. Vancouver Is., 1913, p. 106;
Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), nos. 1033,
1127.

The type of this species is no. 2886 of Gardner, collected at Carmel,
California, in May, 1915, and growing on Desmarestia herbacea. It
seems to be the same as the plant figured (and described) by Saunders
(loc. cit.) under Ectocarpus penicillatus. It is undoubtedly the same
plant as that described by F. S. Collins under the name E. confer-
voides f. acuminatus which was founded on no. 235 of Gardner (Herb.
Univ. Calif., no. 99022) collected at Whidbey Island, Washington,
growing on a broad form of Desmarestia. Its affinities are closely with
Ectocarpus confervoides f. typicus, but it ha-s shorter cells, sharper
ramuli, decidedly constricted filaments, with the gametangia more
inclined toward cylindrical than is general in that form. The chro-
matophores are more distinctly band-shaped and more separated from
one another than in E. confervoides.

A few small but profusely fruiting specimens of plants seemingty
of this species have been found growing on Myriogloia Andersonii at
Neah Bay, Washington (Gardner, no. 3817). These were attached
to the host b}' means of penetrating rhizoidal filaments, particularly
by the descending corticating filaments. The specimens differ from
the typical form in being smaller, in having shorter and blunter game-
tangia and in having on separate individuals seriate 700sporangia( ?)
some of which are divided longitudinally into four loculi. Further
study of more material will be necessary to determine the status of
this form. For the present we are placing it with E. acutus. The
zoosporangia are very numerous, and they resemble in form those
figured by Sauvageau (1896&, p. 33) for Ectocarpus virescens. They,

1925] Setchell-Gardner : Melaiiophyceae 417

however, have a glistening appearance, as though they were abortive
or in a pathological condition, while those represented by Sauvageau
are producing zoospores. This condition in which we find ours is
quite common among our Pacific Coast species of Ectocarpus. We
have not yet seen a single specimen in which the so-called seriate
zoosporangia show any indication of producing zoospores.

4. Ectocarpus tomentosus (Huds.) Lyngb.

Fronds tufted or caespitose, consisting of masses of erect, simple
or branched, rope-like strands, yellowish brown to dark brown ; pros-
trate filaments short, irregularly branched ; erect filaments twisted
together into rope-like masses, 1 mm. to 20 cm. high ; main branches
long, divergent ; ultimate ramuli profuse, alternate, irregularly spaced,
never opposite, mostly short and patent, subpiliferous, often hooked
at the tips; cells of the main erect filaments 8-12^ broad, 1.5-2.5 times
as long as broad, especially below, mostlj^ quadrate above ; ehromato-
phores irregularly band-shaped, few in each cell ; zoosporangia ovoid,
terminal or lateral, 28-36,/x long, 20-26|U, broad ; gametangia lateral,
patent, often secund, straight or recurved, sessile or on short pedicels,
25-75ju, long, 10-15/a broad.

Growing on members of the Fucaceae. Extending from Alaska
(Harvester and Kadiak Islands) to southern California (Laguna).

Lyngbye, Hydrophyt. Dan., 1819, p. 132, pi. 44A 1-3; Greville,
Crypt. Flor., vol. 6, 1828, pi. 316; Saunders, Phyc. Mem., 1898, p. 155,
pi. 24, figs. 1, 2, Alg. Harriman Exp., 1901, p. 417; Setchell and
Gardner, Alg. N.W. Amer., 1903, p. 238. Ectocarpus tame^itosoides
Guernsey, Notes on Mar. Alg., 1912, p. 198. Conferva fomentosa
Hudson, Fl. Angl., 1762, p. 480 (in part). (Ed. 2, 1778, p. 594.)

Our idea of Ectocarpus tomentosus dates back definitely only to
Dillwj^n. His interpretation difiPers somewhat from that of botanists
earlier than his time, but is probably as reliable an interpretation of
the species of Hudson as may be possible. Much hinges on the ideas
as to the identity of the plant of Dillenius (1741, p. 19, pi. 3, fig. 19)
figured and described under the name of Conferva marine, tomentosa,
minus tenera et ferruginea which is quoted by Hudson among the
synonyms of his species. The figure of Dillenius is not reassuring
and Dawson Turner (Trans. Linn. Soc, vol. 7, 1804, p. 105), as
quoted by Dillwyn, states that the specimen in the Dillenian Her-
barium "is only a bad specimen of C. littoraU^." Dillwyn. however.

418 Vniversity of California Publications in Botany [Vol. 8

is convinced from the description of Dillenius, as well as by "the
original drawing in Sir Joseph Bank's Library" that he is correct
in his interpretation.

Ectocarpus tomentosus varies much in height and in the matter
of the greater or less branching of the rope-like masses into which the
erect filaments and their branches are intertwined. Our Pacific Coast
plants are all shorter than the average and more simple. We have
found, in certain specimens, long seriate intercalary gametangia of the
type of those of Pylaiella near the tips of some of the branches. The
occurrence of such gametangia in several species of true Ectocarpus
is worthy of note as well as extremely puzzling. They were par-
ticularly noticed also in specimens of E. confervoides f . variabilis.

The name Conferva tomentosa first appears in 1762 in the earliest
edition of Hudson's Flora Anglica (p. 480). Hudson quotes two
numbers of Dillenius's Historia Muscorum (1741), viz., no. 12 and
no. 13 (loc. cit., p. 19 and pi. 3, figs. 12 and 13). In the later editions
of his work (1778, etc.) Hudson quotes only the second (no. 13) of
these Dillenian descriptions as truly C. tomentosa, referring the first
(no. 12) to his Conferva alhida, now usually recognized to be Clado-
phora alhida Kuetzing. The actual founding of Conferva tomentosa
in the sense of Dillwyn, therefore, rests on the synonymy quoted in
the edition of 1778 which is, so far as we are aware, the only Hudsonian
reference thus far quoted for the species.

5. Ectocarpus corticulatus Saunders

Fronds 0.2-3 cm. high, tufted or feathery, from a small compact
mass of creeping filaments ; main filaments 90-120/x broad, usually
densely corticated, irregularly and frequently branched ; cells of main
filament 65-90/* broad, usually shorter than long, doliif orm ; cells of
branches and ramuli seldom as long as broad, terminal cells tapering,
blunt ; chromatophores band-shaped, few in each cell, often with
pyrenoids ; sporangia unknown ; gametangia narrowly to broadlj^
ovoid, 30-40/x long, 12-18/* broad, secund on the branches and ramuli,
or even on the corticating filaments, short-stalked or sessile.

On larger Melanophyceae and on Zostera. Alaska (Popof Island)
to southern California (San Pedro).

Saunders, Phyc. Mem., 1898, p. 152, pi. 20. Ectocarpus confer-
voides corticulatus Saunders, Alg. Harriman Exp., 1901, p. 418 ;
Setchell and Gardner, Alg. N.W. Amer., 1903, p. 238; Collins, Mar.

1925] Setchell-Gardner: Melanophyceae 419

Alg. Vancouver Is., 1913, p. 106, not Ectocarpus granu-latus f. coi^ti-
culatus Collins, in Collins, Holden and Setchell, Phyc. Bor.-Amer,
(Exsiec), no. 1590.

There are two species of Ectocarpus of fair height with broader
but short cells in the filaments occurring on our coast. One of these
is E. acutus S. and G. and the other E. corticulatus Saunders. The
differences between these two usually strongly corticated species are
to be found in the differing terminal cells of the branches and ramuli
and in the shapes and dimensions of the gametangia. After carefully
considering the different points of view as represented in the syn-
onymy, we have felt compelled to recognize the autonomy of this species
as well as its close relationship with Ectocarpus acutus.

6. Ectocarpus fructuosus S. and G.

Fronds tufted, profusely and alternately branched, up to 2.5 cm.
high, attached by a mass of relatively short, creeping filaments ; erect
filaments several times forked near the base, producing long branches
beset throughout with numerous, short, blunt ramuli of 2-3 orders;
cells of main erect filaments cylindrical to slightly doliiform, and
in part constricted at the cross-walls, 18-25/x diam. below, 2-4 times
as long, slightly wider and shorter above ; zoosporangia unknown ;
gametangia numerous, broadly conical, sessile or on 1-3-celled pedicels,
50-70(a long, 25-35ju, broad at the base.

Growing on the pneumatocyst of Nereocystis Liietkeana. Moss
Beach, San Mateo County, California,

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 410, pi. 45, figs. 1-4.

Ectocarpus fructiiosus comes within the E. confervoides group,
but seems amply distinct from any known forms to constitute a species.
This species is based upon the character of the branches, viz., numerous
long branches, producing throughout a great abundance of short,
rather blunt ramuli and upon these very numerous, rather short and
blunt, predominatingly sessile, gametangia. It seems to be a rare
species.

7. Ectocarpus mucronatus Saunders

Fronds loosely intertwined, 1-6 cm. high, light olive green, attached
by a few colorless creeping filaments; erect filaments profusely
branched in the upper parts, piliferous; primary branches sub-
dichotomous ; ramuli alternate, mostly short and pointed ; cells of

420 University of California Publications in Botany [Vol. 8

main filaments cylindrical, 30-40/x diam., 1-3 times as long as the
diameter below, 0.3-0.6 as long above; chromatoptiores nnmerous,
small, discoid ; zoosporangia unknown ; gametangia numerous, broadly
ovoid, short-acuminate, sessile, 50-100/x (up to 155/^) long, 20-30/x
broad.

Growing on other algae and on Zostera. Vancouver Island (Bay-
ard's Point) to southern California (San Pedro).

Saunders, Phyc. Mem., 1898, p. 152, pi. 19. Ectocarpus granulosus
Tilden, Amer. Alg. (Exsicc), no. 3596 (not 359ai).

Ectocarpios mucronaius seems to be a species of frequent occur-
rence in southern California. AVe have referred a very similar plant
from the Puget Sound region (no. 359& of Tilden) here also. It is to
be distinguished, from other larger feathery species of this genus
particularly by its broadly ovoid, abruptly and shortly mucronate
gametangia.

8. Ectocarpus Taeniae S. and G.

Plate 51, fig. 15

Fronds diffuse, 0.5-1.5 mm. high, attached by profusely branched,
superficial, creeping filaments ; erect filaments simple, narrowed slightly
at the base, gradually attenuated above, piliferous; cells of the creep-
ing filaments cylindrical, 4-6/1, diam., those of the erect filaments 8-lOju,
diam., quadrate below, 4-6 times as long as the diameter at the apices ;
chromatophores short, relatively thick, irregular bands; zoosporangia
unknown ; gametangia narrowly ellipsoidal, at times slightly curved,
mostly sessile on the creeping and on the erect filaments, 20-28/a
(up to 40ft) long, 15-20jU, broad.

Growing on Taonia Lennehackerae. San Pedro, California.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 413, pi. 46, fig. 15.

This species forms rather diffuse and even layers, mostly along the
margins of Taonia, over whose surface the creeping filaments spread
and give rise to gametangia or to erect filaments not over 1.5 mm.
high. In many ways it seems like a miniature Ectocarpus Padinae
(Buffh.) Sauvageau (1896c-1897a., p. 30 et seq.). Our species, how-
ever, is only about one-half as tall, the cells of the erect filaments about
one-half as broad, and the gametangia less than half as large as those of
E. Padinae. AVe have found gametangia only of the " antheridium "
type. Ectocarpus Padinae has its creeping filaments entirely endo-
phytic while E. Taoniae has them entirely epiphytic.

1925 J SetchcU-Gardner: Melanophyceae 421

9. Ectocarpus terminalis Kiietz.

Fronds 0.75-2 (or 4) mm. high, mostly forming* a continnons
velvety layer, brownish in color ; creeping filaments irregular,
branched, anastomosing and forming, at times, a partially pseudo-
parenchymatons layer; erect filaments simple or sparingly branched,
more or less attenuated above; cells of creeping filaments 8-24fi long,
10-18ft broad ; cells of erect filaments cylindrical, 8-12/a broad, up to
six times as long ; chromatophores short, band-shaped, few in each
cell ; zoosporangia terminal, ellipsoidal, 26-52ju, long, 20-30j«, broad ;
gametangia ovoid or ovoid-oblong, often curved, terminal, or lateral
and sessile or short-stalked, 48-120ju, long, 16-32/x broad.

On larger Melanophyceae. Alaska (Unalaska) to southern Cali-
fornia (Laguna).

Kuetzing, Phyc. Germ., 1845, p. 236, Tab. Phyc, vol. 5, 1855, pi.
74, fig. Ill; Kjellman, Skand. Ectocarp., 1872, p. 54, pi. 2, figs, la, lb;
Setchell and Gardner, Alg. N.W. Amer., 1903, p. 237 ; Collins, Holden
and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1034 (in part) (not
no. 1387).

We have accepted Kjellman 's interpretation of Kuetzing 's Ecto-
carpiis termmalis and refer here, although not without some hesita-
tion, no. 1034 of the Phycotheca Boreali-Americana found growing
on stipes of Alaria fistidosa in Unalaska Bay. There is intermingled
in our specimens, however, a plant which seems to be a form of
Ectocarpus confervoides. We have also referred here a plant found
growing on Fucus at Fort Ross, California.

The creeping filaments in this species anastomose more or less and
in some parts of the specimens form almost a parenchymatous basal
layer, strongly resembling that of the species of the Myrionemataceae.
Nearly every cell of the prostrate filaments gives off an erect filament,
a condition characteristic among the Myrionemataceae, but not gen-
eral among the Ectocarpaceae. The erect filaments are slender, with
rather long cylindrical cells, and bear terminal zoosporangia and either
terminal or lateral, sessile or short-stalked, gametangia. The chroma-
tophores are clearly band-shaped though short. No, 1387 of the
Phj'cotheca Boreali-Americana seems to be entirely made up of a
form of Ectocarpus confervoides.

422 University of California Pudlications in Botany [Vol.8

10. Ectocarpus simulans S. and G.

Fronds 1-2 mm, high, tufted ; prostrate filaments irregular, tor-
tuous, distinct ; erect filaments simple, tapering slightly upward, not
pilif erous ; cells cylindrical, not constricted at the dissepiments, ll-lSfi
diam. in the central part of the filament, 1-2.5 times as long as the
diameter ; chromatophores band-shaped, nearly covering the cell, few ;
zoosporangia unknown; gametangia lateral or occasionally terminal,
sessile, narrowly ellipsoidal, blunt, mostly slightly curved, 55-65/^,
long, 15-20ja wide.

Growing on Chaetomorpha aerea. Cypress Point, Monterey
County, California.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 412, pi. 45, figs.
9-11.

The present species resembles, in general, certain forms of Ecto-
carpus pusillios Griffiths (cf. Sauvageau, 1895), but the erect filaments
are much more slender, usually never branched, and with much
smaller gametangia. It forms tufts on Chaetomorpha aerea, whose
filaments are encircled by the prostrate filaments of the Ectocarpus
which form small cushions in their growth over one another. The
gametangia are rather blunt and often more or less curved.

11. Ectocarpus flagelliferus S. and G.

Fronds densely tufted, 3-5 mm. high, attached by a few creeping
filaments and by small, colorless, descending rhizoidal filaments from
the lower cells; erect filaments simple below, sparingly and alternately
branched above, tapering gradually upward and abruptly at the base,
pilif erous when young; cells of the main filaments slightly doliiform,
25-35//, diam., 0.25-1.5 times as long as the diameter ; chromatophores
small, irregular bands or plates, without pyrenoids; zoosporangia
unknown ; gametangia very variable in shape and size, fusiform to
cylindrico-conical, mostly lateral on long pedicels, occasionallj" ter-
minal on short, erect filaments, or long seriate-intercalary on the main
filaments, the lateral 125-200/x long, 28-40ja broad, the terminal and
intercalary up to 1.5 mm. long.

Growing on eel grass. Sitka, Alaska.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 408, pi. 47, figs.
25-27.

1^-5] SetcheU-Gardner: Melanophyceae 423

EctocarpuH flageUiferus, while resembling superficially some of the
shortest plants of Ectocarpiis confervaides f. variabilis, presents sev-
eral peculiarities which are characteristic. The plants of this species
are attached, so far as the adult specimens we have for study are
concerned, by rhizoidal filaments which originate from several of the
lower cells of the erect filaments and form complex basal masses which
almost completely obscure the creeping filaments. Neither the
rhizoidal filaments nor the creeping filaments, however, penetrate the
host. The chromatophores are small and irregular in outline, seeming
to be short bands rather than regular disks. They are generally so
closely placed in the younger cells as to seem almost like a continuous
band, but are separate in the older cells, with slender processes almost
connecting them to one another. The cells are short and the erect
filaments and their branches extend out into long hairs like M'hip-
lashes. The lateral gametangia are variable in shape, arranged much
as in Ectocarpics confervoides f. variabilis, Init are of different dimen-
sions. Besides the characteristic Ectacarpiis-tyiie of gametangia
which are lateral, there occur terminal seriate gametangia of the
PylaieUa-type W'hich reach an extreme length of 1.5 mm.

12. Ectocarpus Mesogloiae S. and G.

Fronds minute, 0.75-1.5 mm. high, attached by a mass of densely
intertwined, branched, rhizoidal filaments penetrating among the cells
of the host ; erect filaments sparingly and alternately branched ; main
filaments and ramiili tapering gradually upward, very acute, not
piliferous; cells cylindrical, slightly constricted at the dissepiments,
15-18/A diara. at the base, 1-2 times as long, 4— 6/x at the apices of the
filaments ; chromatophores thin, irregular bands, nearly covering the
cell ; zoosporangia" unknown ; gametangia narrowly cylindrico-conical,
120-1 GOjU, (up to 210ju,) long, 18-22/x broad, on short pedicels, rarely
sessile.

Growing on MyriogJoia Andersonii (Farlow) Kuck. Carrael Bay,
Monterey County, California.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 411, pi. 45, figs. 5, 6.

The general characters of this diminutive species ally it with the
E. confervaides group. We have deemed it best to consider it a dis-
tinct species on account of the small dimensions of all of its parts and
its penetrating habits. It has not been seen on any other host except
the one mentioned above.

424 University of California Publications in Botany [Vol. 8

13. Ectocarpus commensalis S. and G.

Fronds 200-400/* hig-h, densely fasciculate, attached by inter-
twined, slightly branched rhizoids penetrating among the utricles and
even well among the medullary filaments of the host ; erect filaments
alternately branched near the base, simple above, slightly attenuated
upward, not piliferous; cells of the erect filaments cylindrical, not
constricted, 12-18/* diameter at the base of the filaments, 1.5-2.5 times
as long as the diameter ; chromatophores band-shaped, few in each
cell ; zoosporangia unknown ; gametangia blunt-fusiform to cylindrico-
conical, short-pedicellate, mostly lateral near the base of the filaments,
rarely terminal, GO-lOO/i long, 15-20/i wide.

Growing on Codiiom Setchellii Gardner, Carmel Bay, and on
C odium fragile (Suring.) Hariot, Pacific Grove, Monterey County,
California.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 407, pi. 48, figs.
32-35.

This minute species of Ectocarpus is one of .several occurring on
species of Codium, both on this coast and on that of southwestern
Europe. The colorless rhizoidal filaments descend into the substance
of the host even to the central (or basal) medullary tissues. The
smaller diameter of the erect filaments and the more narrowly conical
or fusiform gametangia readily distinguish this species from the
others.

14. Ectocarpus eramosus S. and G.

Fronds diffused or in diminutive tufts, 1-3 mm. high, attached by
irregularly branched rhizoidal filaments penetrating deeply among the
utricles of the host ; erect filaments eramose or rarely producing short,
divergent branches usually terminated in gametangia, slightly atten-
uated toward the base and the apex, not piliferous; cells of mature
filaments 28-40/i diam., 0.5-1.25 times as long as the diameter, cylin-
drical, very slightly constricted ; chromatophores relatively thick,
closely crowded, irregular bands without pyrenoids; zoosporangia
unknown ; gametangia very variable in shape and size, lateral and
alternate along the entire filament or occasionally terminal and seriate
on the main filaments, mostly on few-celled pedicels, lateral forms
cylindrical to cylindrico-conical, 150-230/i long, 26-36/t diam., ter-
minal, seriate forms, 700-900/i long, 15-20/* broad.

1925] Setchell-Gardncr: Melmiophyceae 425

Growing on Codium fragile in the lower littoral belt. Near the
entrance to Tomales Bay, Marin County, California.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 407, pi. 47, figs.
18-23.

This very small species has the general appearance of a form of
Ectocarpus confervoides, but the cells are short and the filaments very
slightly constricted at the partitions. It seems, in these respects,
nearer to E. acutus and E. corticidatus, possibly bearing something
of the same relation to these species that the dwarf forms of E. con-
fervoides do to the typical form. Ectocarpiis eramosus, however, is
not readily to be referred as a dwarf form of either E. acutus or E.
corticidatus and is consequently to be kept separate, at least for the
present.

15. Ectocarpus luteolus Sauv.

Prostrate filaments moniliform, irregidarly branched, covering, or
even occupying, the injured cells on the surface of the host, even
occasionally penetrating slightly inward, not anastomosing, but form-
ing an irregular and confused layer of almost parenchymatous aspect
emitting tortuous rhizoidal filaments from its lower cells; erect fila-
ments very short, 100-300/x high, terminating in delicate hairs (or, at
times, in seriate gametangia), simple or with a few branches from
near the base ; cells of the erect filaments short, swollen, and nearly
globular below, cjdindrical and longer above, about 8/t broad, 1.5 times
as long as broad ; chromatophores few in each cell, band-shaped ; zoo-
sporangia terminal on short basal branchlets, oblong-ellipsoidal,
26-30/A long, 16-18/A broad ; gametangia narrowly cylindrical, con-
sisting of single or double rows of loculi, 30-80/x (in ours about 30-45|U,)
long, 7-13/A (in ours 11-13/a) broad.

Forming small expansions on the surface of the lower portion of
Pelvetiopsis limitata f. typica Gard. Central California (San
Francisco).

Sauvageau, Sur. quelq. alg., 1892, p. 79 (p. 42 in repr.), pi. II,
figs. 1-H9 ; Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc),
no. 1233. StrehJonema luteoliim De-Toni, Syll. Alg., vol. 3, 1895,
p. 575.

Ectocarpus luteolus Sauv. is one of the species which seems inter-
mediate between Ectocarpus and Strehlonema. It does penetrate the
host slightly and occasionally, but it seems to grow on the lower
abraded portion of the host, filling the hollows between and even the

426 Unwersity of California Publicaiions in Botany \yo-L. 8

cavities of the injured cells. Its prostrate filaments and the lower
portions of the erect filaments have moniliform, almost globular, cells
which, together with rhizoidal filaments issuing from the very basal
ones, while perfectly separate, form a thin layer or cushion, with
something of the appearance of that of Elachistea.

Our specimens seem to agree so closely with the figures and descrip-
tion of Sauvageau and to grow under such almost identical conditions,
that we must necessarily refer them to this species. Thus far it has
been detected only at Lands End, San Francisco, California, and
only on one occasion. Our specimens show zoosporangia (not hitherto
described) as well as gametangia.

16. Ectocarpus granulosus (Eng. Bot.) Ag,

Fronds tufted, rather coarse and rigid, 1-8 cm. (up to 16 cm.)
high, profusely branched ; primary branches opposite or whorled,
mostly corticated below; secondary branches opposite or at times
alternate, short, tapering upward, often recurved at the tip, ending
in hairs; ultimate ramuli secund, short, acute, ending in short hairs;
cells of the main filaments SO-lOO^u, diam., quadrate or shorter than
the diameter, constricted at the cross-walls; chromatophores small,
regular disks; zoosporangia "sessile, globose"; gametangia sessile,
broadly ovoid, asymmetrical, mostly secund on the terminal and
subterminal ramuli, 60-lOOfi long, 30-60/x broad.

Growing on various other algae and on stones in the lower littoral
belt. Washington (Puget Sound) to southern California (San Pedro).

Agardh, Sp. Alg., vol. 2, 1828, p. 45 ; Saunders, Phyc. Mem. 1898,
p. 156, pi. 24, figs. 3-5 ; Setchell and Gardner, Alg. N.W. Amer., 1903,
p. 238; Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc),
no. 1589. Ectocarpus gramiClosus f. corticidatus Collins, in Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1590. Conferva
granulosa English Botany, 1814, pi. 2351.

Ectocarpus granulosus was discovered by W. Borrer in the vicinity
of Brighthelmston (Brighton), East Sussex, England, and is now
reported from many coasts of Europe and both eastern and western
North America. There is considerable variation, especially as regards
dimensions both of the primary branches and of the gametangia. In
some of our specimens, the gametangia reach a length of 165/^, and
a breadth of 65;^, but the majority are 90-110/a long and 50-60;u, broad.
We have not seen zoosporangia in any of our specimens. The very

19-5] SetclicU-Gardner : MeJanophyceae 427

distinctly discoid chromatophores and the opposite arrangement of
most of the main branches clearly distinguish this species from any
other on onr coast.

17. Ectocarpus Parksii S. and G.
Plate 49, fig. 15

Fronds 5-7 cm. high, densely caespitose, flaccid ; main filaments of
erect fronds, 34— 40/^ diam. in the lower part, tapering very gradually
upward, profusely and alternateh' branched, very slightly corticated
at the base ; branches of succeeding orders reduced in diameter, long
and tapering very gradually upward, not terminating in hairs;
ultimate ramuli lO-lS/x diam., terminal cell 5-7/>i diam. ; cells cylin-
drical to very slightly doliiform, 40-70/a long in the main filaments,
quadrate to subquadrate above; chromatophores numerous, irregular
plates densely crowded together and more or less connected in the
ramuli, becoming more regularly disk-shaped and separate in the
lower parts of the main filaments ; gametangia relatively sparse, sessile,
straight or slightly curved upward, mostly on the subterminal and
terminal ramuli, secund, 4-10 in a group, occasionally solitary, blunt
fusiform to subconical, 35-45/x, long, 18-22/i, broad.

Growing on Gracilaria confervoides (L.) Grev. in a small cove,
San Francisco Bay, northeast of Tiburon, Marin County, California.

Setchell and Gardner, Phyc. Cont. VII, 1924, p. 1.

This species of Ectocarpus appears to be a very short lived spring
form. Within a month after it was first observed, apparently just
coming into fruit, it had largely disappeared. Its structure seems to
relate it to several of our west coast species. Its long, slender, grad-
ually tapering filaments, profusely branched, are similar to those of
E. siliculosus (Dillw.) Lyngb. Its sessile gametangia are similar in
form to those of E. Taoniae S. and G. The secund arrangement of the
gametangia and the disk-shaped chromatophores are characters belong-
ing to E. gramdosoides S. and G. The delicate fronds, with profuse,
alternate branching, the disk-shaped chromatophores, and the groups
of small, sessile, blunt, secund gametangia make a combination of
characters not found in any other species.

428 University of California Publications in Botany {'^o-l. 8

18. Ectocarpus Mitchellae Harv.

Fronds aggregated into dense feathery tnfts, 2-8 cm. high, decom-
poundly branched, attached by long creeping filaments; yellowish
green or olive green on drying; branching alternate, the branches
divaricate, attenuated, often ending in short hairs, ultimate ramuli
approximated ; cells in the main branches 25-40/a diam., 1-3 times as
long as the diameter below, shorter above ; chromatophores numerous,
small, discoid; zoosporangia imknown; gametangia ellipsoidal to nar-
rowly oblong, obtuse, 50-lOOja long, 18-35/a wide, sessile, mostly secund
on the upper side of the branches.

Growing on limpets and on Phyllospadix in the lower littoral belt.
Southern California.

Harvey, Ner. Bor.-Amer., part I, 1851, p. 142, pi. 12, G; Collins,
Notes on New England Algae V, 1891, p. 337 ; Saunders, Phyc. Mem.,
1898, p. 153, pi. 21, figs. 1, 2; Collins, Holden and Setchell, Phyc.
Bor.-Amer. (Exsicc), no. 671.

AVe have specimens from several collections from the coast of
southern California which agree so well with plants from the southern
New England coast referred to E. Mitchellae that we cannot satisfac-
torily separate them from it. We call attention to the views of Sau-
vageau (1896&, p. 39), Collins (1891, p. 337), Saunders (1898, p. 153)
and Borgesen (1914, pp. 159-162) as to the possible relation, or even
identity, of this species with E. indicus Sond., E. virescens Thuret,
and E. Duchassaingianus Grunow, but we have no additional facts to
present.

Our specimens show both mega- and meio-gametangia such as
Sauvageau (18966) describes and illustrates for E. virescens Thuret.

19. Ectocarpus oviger Harv.

Fronds somewhat densely matted together, 7-15 cm. (up to 22 cm.)
high, main filaments densely corticated, dark brown, decompositely
branched ; branches mainly alternate, in part secund, strict ; cells
cylindrical throughout the frond, not constricted, 60-70/* diam. in
the main filaments, 1.5-2.5 times as long as the diameter; chromato-
phores discoid, numerous ; zoosporangia uncertain ; gametangia
broadly ovoid to subspherical, asymmetrical, sessile or subsessile, alter-
nate .or in part secund, promiscuously scattered throughout the
branches, 35-45/* long, 30-35/* broad.

19-5] Setchell-Gardncr : Melanophyceae 429

Gro\Wng on Nereocystis Luetkeana. Puget Sound region to central
California.

Harvey, Coll. Alg. N.W. Amer., 1862, p. 167; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 238. Ectocarpus granulosus Tilden, Amer.
Alg. (Exsicc.), no. 359a (not no. 359&) ; Collins, Holden and Setchell,
Phyc. Bor.-Amer. (Exsicc.), no. 2242.

The Ectocarpus oviger Harvey has been a puzzle for phycologists
who, in general, have been inclined to refer it to E. granulosus, to
which latter species Harvey himself compared it. An examination of
the type specimen at Dublin and a careful study of Harvey 's descrip-
tion, lead us to refer here no. 359a, of Tilden 's American x\lgae and
no. 2242 of the Phycotheca Boreali-Americana and to keep E. oviger,
at least as thus interpreted, distinct from E. granulosus. Harvey
speaks of his plants as having the aspect of Pylaiella Uttoralis, and
the plants we refer to his species resemble certain varieties or forms
of that species more than they do any forms of E. granulosus. Harvey
also remarks that the "fruit" of his species is like that of E. granu-
losus, and it is true that, in our specimens, which show only game-
tangia, these resemble the gametangia of E. granulosus more than
those of any other species, but differ from them sufficiently in shape
and dimensions to prevent merging the one species into the other.
Our conclusion is that Ectocarpus oviger, while closely related to
E. granulosus, is sufficiently different from it in habit, in the length
of the main filaments, and in the shape and size of the gametangia, to
be retained as an independent species.

We have indicated that we are uncertain as to the zoosporangia in
this species, although it has been supposed that the ' ' fruit ' ' described
by Harvey is zoosporangial. The bodies seen on the portion of the
type specimen examined seemed to be empty zoosporangia. On the
other hand, the gametangia at maturity and about to discharge their
gametes show little or no trace of the internal partitions, so that
further experience with this species is very desirable.

20. Ectocarpus breviarticulatus J. Ag.

Fronds tufted, 2-4 cm. high, arising from creeping filaments ; erect
filaments about 27/x. broad (35-45ju,, fide Kuetzing and De-Toni), either
sparingly branched or more or less beset with short branchlets, both
branches and branchlets attenuated toward the apex and hooked,
intertwined into rope-like masses; cells usually 1.5-2 times as long as

430 University of California Piidlications in Botany [Vol. 8

broad ; chromatophores disk-shaped, several in each cell ; zoosporangia
unknown; gametangia from nearly spherical to broad ovoid, about
62fjL long, 57/A broad.

On Chnoospora pacifica. West coast of Mexico (St. Augustin).

J. G. Agardh, Nya Alger fran Mexico, 1847, p. 7, Sp. Alg., vol. 1,
1848, p. 16 ; Kuetzing, Spec. Alg., 1849, p. 453 ; Borgesen, Mar. Alg.
Dan. West Indies, pt. 2, 1914, p. 173, fig. 136. Ectocarpus hamatus
Crouan, in Maze et Schramm, Ess. Class. Alg. Guad. (ed. 2), 1870-77,
p. Ill (fide Borgesen) ; Vickers, Phyc. Barbad., pt. 2, 1908, pi. 29.

Ectocarpns hreviarticulatus J. Ag. is known to us as a member of
our flora only from the literature and particularly from Borgesen 's
{Joe. cit.) account of it. It seems closely related to E. oviger, from
which it differs chiefly in size and branching, and slightly in dimen-
sions of filaments and gametangia. Both Kuetzing and Borgesen
state that they have examined cotypes and may be, therefore, con-
sidered to have spoken with authority. We have been enabled to
study what certainly seems to be this species in material from
American Samoa (cf. Setchell, Veg. Tut. IsL, p. 171, fig. 37).

21. Ectocarpus chantransioides S. and G.

Fronds arising from contorted creeping filaments, forming dense
hemispherical cushions 4-8 mm. high; branching profuse, alternate
below, mostly secund above; main filaments and branches not atten-
uated; cells 8-10/x diam., 2-3 times as long as the diameter below^
quadrate above ; chromatophore band-shaped ; zoosporangia unknown ;
gametangia mostly sessile or on short pedicels, narrowly cylindrico-
conical, 80-IIOjM, long, 16-20/a broad at the base.

Growdng on boulders in the lower littoral belt. Three miles north-
west of Santa Monica, California.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 406, pi. 48, figs.
28-31.

The habit of Ectocarpus chantransioides distinguishes it at once
from all other species of the genus with the exception of E. hemi-
sphericus Saunders. The latter species is always found, so far as our
knowledge is concerned, epiphytic on Fucaceae, while the former
species is confined to rocks. Ectocarpios chantransioides has also more
slender filaments than has E. hemisphericus, not at all tapering, and
has distinct apical growth. The gametangia are differently shaped,
being longer and more slender. It is therefore very distinct even
from E. hemisphericus. It resembles the genus Choristocarpus of the

1925] Setchell-Gardner: Melanophyceae 431

family Choristocarpaceae, as diagnosed by Kjellman (1891, pp. 190,
191), in having apical groAvth. The groAvth in length of the erect
filaments in the Choristocarpaceae is by the division of the terminal
cell only, whereas in E. chant ransioides the meristem extends over
a number of cells at the outer or apical end of the filaments. These
cells, some 10 to 15 in number, are much richer in cell contents, the
terminal cell being the richest of all in the series. This is a very
unusual condition for an Ectocarpus. The nearly uniform diameter
throughout of the erect filaments and their method of branching
resemble to a remarkable degree those characters found in Chan-
transia. The gametangia are typically those of Ectocarpus and, not-
withstanding the other rather unusual characters, it seems to be most
closely related to that genus, but a very distinct species.

22. Ectocarpus granulosoides S. and G.

Fronds 2-3 cm. high, profusely branched ; main filaments sub-
dichotomously branched, all branches suddenly attenuated at the base,
corticated below, secondary filaments mostly alternate, in part secund,
strict, long-attenuate, acute, not piliferous ; ultimate ramuli mostly
secund, acute; cells of the main filaments 70-80jU, diam., 0.5-1 times
as long as the diameter, those of the secondary filaments 30-40ju, diam,,
0,3-1 times as long and those of the ultimate ramuli 14-20/x diam.,
0.3-1.5 times as long, all slightly constricted at the dissepiments ;
chromatophores numerous, discoid in the older cells, irregularly
angular in the ramuli ; cells of the corticating filaments 7-10;a diam.,
3-4 times as long as the diameter ; zoosporangia unknown ; gametangia
secund on the upper side of the ultimate and penultimate ramuli,
sessile, subfusiform, asymmetrical, 40-60/1, long, 12-20/x broad.

Growing on rocks (?). San Pedro, California.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 410, pi. 45, figs. 7, 8.

The very distinct species we have described under the name of
Ectocarpus granidosoides has the habit of a small E. grantdosus and
gametangia of the same general type as found in that species, but
considerably smaller. The branching, however, is never opposite,
and the acute ramuli gradually attenuated upward recall those of
E. acutus, from which our species is amply distinct in both chromato-
phore and characters of its gametangia. The branches, particularly
the larger, are very suddenly and considerably attenuated at the
base, giving the species a striking characteristic of its own, at least
within the group of species wdth discoid chromatophores.

432 University of California Pudlications in Botany [Vol.8

23. Ectocarpus affinis S. and G.

Plate 51, figs. 16, 17

Fronds diminutive, 500-700ja high, attached to the host by a few
branched, colorless, entwining, rhizoidal filaments; erect filaments
sparingly branched, gradually attenuated upward, pilif erous ; branches
alternate or opposite ; cells of the main filaments cylindrical, 24-30/^
diam. a.t the base, 9-11/t at the apex, varying in length throughout the
filament, 0.75-3 times as long as the diameter; chromatophores small,
thin, angular plates in young cells, nearly regular disks in older cells ;
zoosporangia unlvnown ; gametangia lateral, mostly sessile, occasionally
on 1-celled pedicels, solitary, secund, or in whorls from a single cell,
blunt-conical to spheroidal, 28-31;a long, 25-28/x broad, arranged
along the entire filament to the terminal hairs.

Growing on Callithamnion sp. Sitka, Alaska.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 405, pi. 46, figs.
16, 17.

Two collections from Sitka, Alaska, but both made on the same
day, show a small Ectocarpiis growing on a species of Callithamnion
which is so closely related to E. ovatus Kjellman (1877fl, p. 35) that
perhaps it may seem necessary, at some time, to refer it to that
species. The plants are, however, less stout than those of Kjellman and
with more nearly spherical gametangia. For these reasons and because
of its geographic remoteness and the somewhat different climatic con-
ditions, we feel that it is desirable to describe the Alaskan plant as a
distinct, but closely related, species.

24. Ectocarpus cylindricus Saunders

Fronds diffuse, 1-2 mm. high ; creeping filaments superficial ; erect
filaments very sparingly branched above, several arising from the
same creeping filament, tapering slightly at the base and apex; cells
of the creeping filaments 16-20|a diam. ; cells of the erect filaments
cylindrical, slightly constricted at the cross-walls, 2-3 times as long as
broad below and at the apex, 0.5-1 times in the middle of the filament ;
gametangia mostly on the middle and upper parts of the filaments,
usually alternate.

Growing on Egregia Menziesii and Cystoseira osmiondacea. Central
California (Pacific Grove).

1925] Setchell-Gardner: MelanopJiyceae 433

Saunders, Phyc. Mem., 189S, p. 150, pi. 16; Setchell and Oardner,
Phyc. Cont. VI, 1922, p. 415 (f. fypicHs).

The typical form, growing on the larger brown algae, differs some-
what from the variety found growing on C odium as is indicated below.

Ectocarpus cylindricus f. codiophilus S. and G.

Plate 51, fig. 14

Fronds densely caespitose, 3-5 mm. high ; rhizoidal filaments
densely intertwined, penetrating the host; erect filaments anastomos-
ing and forked at the surface of the host, unbranched above, pili-
ferous; gametangia, for the most part, near the base of the erect
fronds.

Growing on Codiiim fragile (Suring.) Ilariot and Codinni
SetchcUii Gardner. Carmel Bay, Monterey County, and La Jolla,
San Diego County, California.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 415, pi. 46, fig. 14
and pi. 49, figs. 42-45.

This form differs from the typical form in having penetrating
rhizoidal filaments and in having the gametangia in a dense zone at
or near the base of the erect filaments instead of being scattered.

Ectocarpus cylindricus f. acmaeophilus S. and G.

Fronds tufted, 7-10 mm. high ; creeping filaments superficial ;
erect filaments unbranched, nearly the same diameter throughout ;
zoosporangia unknown ; gametangia up to 270/x. long, mostly opposite
on the upper part of the erect filaments.

Growing on Acmaea sp. Carmel Bay, Monterey County, Cali-
fornia.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 415, pi. 49, fig. 46.

This form seems to be rare, at least but few specimens have been
detected up to the present time. It differs from the typical form in
being practically unbranched, except the fructiferous branches.
Where branches occur, they seem to arise a.s the result of an injury
to the main filament. Also it dift'ers in having the erect filaments
more nearly cylindrical throughout. And finally it differs in having
the gametangia more often opposite, and generally longer and
proportionally narrower.

434 University of California Publications in Botany [Vol. S

25. Ectocarpus hemisphericus Saunders

Fronds pulvinate, 2-4 mm. high, arising from a compact network
of creeping filaments rhizoidal in character, much attenuated and
crisped ; primary branches pseudodichotomous, divaricate, gTadually
attenuated upward; secondary branches numerous, short, clustered,
mostly long-piliferous; cells at the base of main filaments 22-26/x
diam., 1-3 times as long as the diameter, smaller above, slightly con-
stricted at the joints ; chromatophores numerous, small, discoid ; zoo-
sporangia cylindrical, 30-35/* long, 20-25/* diam., on 1-celled stalks, at
times on the same fronds with the gametangia (Saunders) ; game-
tangia fusiform to broadly ovoid, obtuse, 30-90/t long, 14-20/i diam.,
lateral on the upper parts of the fronds, short-pedicellate.

Growing mostly on Pelvetia fastigiata, but also on other species of
algae. California (San Diego, Laguna and San Pedro).

Saunders, Phyc. Mem., 1898, p. 151, pi. 17 ; Guernsey, Notes on
Mar. Alg., 1912, p. 198, fig. 108 ; Collins, Holden and Setchell, Phyc.
Bor.-Amer. (Exsicc), no. 528.

Ectocarpus hemisphericus f. minor Saunders

Very similar to the species, but forming less definite tufts and
with the erect filaments simple or but slightly branched, 1-2 mm. high,
16-21/x diam., and with gametangia and zoosporangia mostly sessile.

Growing on and near the receptacles of HesperopJiycus Har-
veyanus. San Pedro, California.

Saunders, Phyc. Mem., 1898, p. 151, pi. 18, figs. 1-3; Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), nos. 529 and 1794.

Ectocarpus hemisphericus Saunders, in typical form, can readily
be recognized by its habit and its host, but in some forms and par-
ticularly in f. minor is less distinct in habit, at least. Thus far, both
the species and the form seem confined to the coast of southern
California.

26, Ectocarpus Saundersii S. and G.

Fronds forming small tufts, 2-5 mm. high, attached by a mass of
branched filaments creeping on the surface of the host; branching
alternate ; main branches long and divergent, ultimate ramuli sparse,
short, acuminate; cells of the main filaments 25-40/i diam., quadrate
below, shorter above, slightly constricted at the dissepiments; chroma-

1925] SeicJieU-Gardner: Melanophyceae 435

tophores numerous, discoid ; zoosporangia on short stalks or inter-
calary, globose, about 30fi diam. ; gametangia cylindrical to ovoid,
obtuse or acuminate, lateral, borne on longer or shorter pedicels,
70-150;u, long, 25-50/A broad.

Growing on Fucus, in the middle of the littoral belt. Pacific
Grove, California.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 411. Ectocarpus
paradoxus var. pacificus Saunders, Phyc. Mem., 1898, p. 152, pi. 18,
figs. 4—7; Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc),
no. 530.

According to Saunders, his plant, the var. pacificus, differs from
the type of E. paradoxus Mont., in lacking any manifestation of
opposite branching as well as in having the gametangia longer and
more pointed. In these respects and also because it is a shorter plant,
it seems to us to be sufficiently distinct to be separated specifically
from the type, and to be different from any other species known to us.

27. Ectocarpus acuminatus Saunders

Fronds diminutive, attached by a network of delicate rhizoidal
filaments ; erect filaments uniform in size throughout, 1-2 mm. long ;
cells cylindrical, not constricted, 12-14/a diam., quadrate below, 2-5
times as long above ; chromatophores irregular or elliptical disks, more
abundant in the central part of the filament than elsewhere ; zoo-
sporangia unknown ; gametangia sessile on the creeping filaments or
on the base of the erect filaments or terminal on short filaments arising
directlj' from the creeping filaments, very long-acuminate, sometimes
tipped with a short hair, often more or less curved, 90-300/^ long,
20-30/t broad.

Growing within the conceptacles of Cystoseira osmundacea and
Halidrys dioica. Central California (Pacific Grove) to southern
California (San Pedro).

Saunders, Phyc. Mem., 1898, p. 149, pi. 14, figs. 1-5; Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), nos. 524 and 2142.

The shape of the gametangia would seem to ally this very incon-
spicuous species most closely with E. siliciUosus, but the chromato-
phores are described by Saunders as elliptical. We have only dried
material for examination and cannot, therefore, add anything to the
description by Saunders. It is a very curious and seemingly distinct
species. Its habitat, wathin the conceptacles of members of the
Fucaceae, seems to have reduced it almost to the state of parasitism,
at least one-half of the plant being unable to function in food making.

436 Universitii of California Puhlications in Botany [Vol. 8

28. Ectocarpns ellipticus Saunders

Fronds minute, about 1 mm. high, tufted, attached by numerous,
colorless, branched rhizoidal filaments ; erect filaments sparingly
dichotomous at the base, unbranched or with few unicellular branches
above, gradually attenuated upward, pilif erous ; cells 9-1 2/x diam.
and 1-2 times as long as the diameter at the base of the filaments, 3
times as long above ; ' * chromatophores thin branched bands, oval, ' '
numerous ; zoosporangia sessile, ovate or ellipsoidal, 30-40/a long,
12-18/A broad ; gametangia cylindrical to blunt-conical, 75-100;U, (up
to 180/x.) long, 18-25ju, diam., borne laterally at or near the base of the
erect filaments, or occasionally arising directly from the creeping
filaments, sessile or on short pedicels below ; loculi one to few seriate.

Growing in the conceptacles and cryptostomata of Fucus sp.
Pacific Grove, California.

Saunders, Phyc. Mem., 1898, p. 149, pi. 14, figs. 6-9; Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 527.

Ectocarpus ellipticus is a curiously localized species growing upon
the inner margins of the openings of the cryptostomata and con-
ceptacles of Fucus at Pacific Grove, California. Transverse sections
of the fronds of the Fucus show that this Ectocarpus simply forms a
ring of growth just within the outer opening of the cryptostomata and
conceptacles just outside the hair-forming tissues. The rhizoidal fila-
ments of the Ectocarpus penetrate into the apparently looser tissues of
the ostiole of the ca^aties it inliabits. The gametangia are all basal or
borne near the bases of the erect filaments. It seems to be a very
distinct species.

29. Ectocarpus chitonicola Saunders

Fronds small, 1-2 mm. high, tufted, creeping filaments numerous,
irregular, branched ; cells of the creeping filaments ll-15|ii diam., 1-2
times as long as the diameter; erect filaments mostly simple, 14/x diam.
at the base, somewhat narrower above, with cells 0.5-2 times as long
as the diameter below, 2-3 times as long above, not constricted at the
partitions ; chromatophores small oval disks, numerous ; zoosporangia
unknown ; gametangia subcylindrical to narrowly ovoid, obtuse at the
apices, 90-175/x (up to 250/^,) long, 25-35|U, diam., borne laterally on
the erect filaments, or occasionally on the creeping filaments, the
lower gametangia on pedicels, the upper sessile.

19^0] SetchcU-Gardncr: MelanopJnjreae 437

Growing- in minute tufts on the shells of Chitons and Limpets.
Pacific Grove and Carmel Bay, California.

" Ectocarpus vkiionkolus" Saunders, Phyc, Mem., 1898, p. 150,
pi. 15, figs. 1-4.

The illustrations of Saunders {Joe. cht.) show a diversity of forms
of the gametangia, ranging from long ones with many small loculi, to
short ones with few larg:e loculi. The species either possesses great
range of variation in this respect, or else there may possibly be two
species concerned. Another explanation of the phenomenon may be
the possibility that we have in this species the two forms of game-
tangia mentioned by Sauvageau (1896&, p. 17) found on Ectocarpus
viresccns. Saunders' figure 2 might well represent the "mega-
sporangia" and figures 3 and 4 the "meiosporangia" of Sauvageau.
The species needs further investigation to have the validity of these
points established.

In the specimens we have examined, both that of Saunders and the
single one of Gardner's collecting, the gametang^ia are long, and
large-celled. The species is to be distinguished from any other of our
loAV forms by its discoid chromatophores and its peculiar habitat.

30. Ectocarpus socialis S. and G.

Plate 51, tigs. 12, 13

Fronds eaespitose, 1-3 mm. high, attached by densely intertwined,
penetrating rhizoidal filaments; erect filaments several times forked
at the surface of the host, simple above or rarely alternately branched,
long-attenuate upwards, piliferous; cells cylindrical to slightly doli-
iform, not constricted, 22-28ju, diam., 1-2 times as long as the diameter
below, 4-6 times above ; chromatophores small, few in a cell, discoid ;
zoosporangia ellipsoidal, erect, lateral, alternate, mostly on 1-2-celled
pedicels, occasionally sessile near the base of the erect fronds, 60-95/x
long, 35-50/A broad ; gametangia fusiform, sparse on the upper parts of
the erect filaments; sessile or on 1-2-celled pedicels, alternate, 70-85ju.
long, 25-30/x broad.

Growing on CacUiim fragile (Suring.) Hariot, in company with
Ectocarpus glohifer Kuetz. and with various species of Myxophyceae
and Rhodophyceae. Redondo, California.

Setchell and Gardner, Phyc, Cont. VI, 1922, p. 412, pi. 46, figs.
12, 13.

Ectocarpus socialis seems to be a relative of the E. c ijlindricws
group. The chief distinctions between the two species lie in the shape,
size and position of the gametangia.

438 University of California Publications in Botany [Joi.. 8

31. Ectocarpus flocculiformis S. and G.

Fronds densely caespitose, 0.75-1.5 mm. high, attached by inter-
twined, penetrating, rhizoidal filaments; erect filaments more or less
forked at the base, simple above or the largest plants with few, short,
alternate, acute ramiili, tapering slightly at the base, long-attenuate
upwards, not piliferous; cells cylindrical to slight doliiform, 20-25/x
diam., 1-2.5 times as long ; chromatophores numerous, angular-discoid
in the younger cells, rounded in the older cells; zoosporangia ellip-
soidal, 56-66/A long, 35-40//, broad, on short pedicels near the base of
the erect filaments; gametangia ovoid to ellipsoidal, numerous, sessile
or short-pedicellate, lateral near the base of the erect filaments,
95-110/x (up to 150/x) long, 30-45ju, broad.

Growing on Codinni fragile, the rhizoidal filaments penetrating
deeply into the host, among and beyond the utricles. La JoUa,
California.

Setchell and Gardner, Phyc. Cont. VI, 1922, p. 409, pi. 47, fig. 24.

The nearest relative of E. floccidiformis apparently is E. cylin-
dricus f. codiophilus. One marked difference between it and any of
the forms of E. cylindricus is in the shape and size of the gametangia.
This difference, along with others of less importance perhaps, though
apparently constant, seems to be sufficient to warrant keeping it
separate.

32. Ectocarpus globifer Kuetz.

Fronds 2-5 mm. high, pulvinate, sparingly branched, attached
by a few branched, rhizoidal filaments penetrating among the cells
of the host ; branches opposite or alternate, divaricate, constricted at
the base, attenuated above, at times piliferous; cells cylindrical,
slightly constricted at the dissepiments, 45-50ya diam. in the main
filaments, 0.5-1.5 times as long as the diameter in the middle, up to
4 times the diameter above and below; chromatophores numerous,
small, irregular discoid ; zoosporangia lateral, spherical or subspher-
ical, mostly on 1-celled pedicels, 30-40/a diam. ; gametangia lateral,
broadly ovoid to spherical, mostly on 1-celled pedicels, 70-100/a long,
50-70/1 broad.

Growing on C odium fragile. Southern California (San Pedro and
La Jolla).

1925] SetchcU-Gordncr: Melanophyceae 4;}9

Kuetzing, Phyc. Gen., 1843, p. 289, Tab. Phyc, 1855, vol. 5, p. 16,
pi. 49, %. 2; Bornet, Note sur quelq. Ect., 1891, vol. 38, p. 358 (repr.
p. 6), figs. 6, 7; Collins, Ilolden and Setehell, Phyc. Bor.-Amer.
(Exsice.),no. 732.

This species has been distributed from southern California under
no. 732 of the Phycotheca Boreali-Americana. The specimens dis-
tributed were collected at La Jolla by Mrs. Snyder. Gardner has
collected it twice at Redondo. Both undoubted zoosporangia and
gametangia have been found. All of the specimens from our coast, so
far as is known, grow upon C odium fragile, into whose spongy sub-
stance they penetrate by long colorless rhizoidal filaments. The
specimens agree so well with the figure.^ and descriptions of Kuetzing
and Bornet that there seems to be little doubt that our plants are of
the same species as those from Europe.

33. Ectocarpus gonodioides S. and G.

Fronds minute, forming small tufts 500-550ju, high, attached by
long, more or less hyaline rhizoidal filaments penetrating the host ;
filaments sparsely branched at the surface of the host, tapering rather
abruptly at the base, long attenuated upward to a blunt apex, 18-24/a
diam. at the base, 10-14ja at the apex ; cells 1-2 times as long as broad ;
zoosporangia unknown ; gametangia narrowly fusiform on 1-2-celled
pedicels, near the base of the erect filaments, up to 125fi long, 20-28/a
diam. in the widest part.

Growing on Codmm cuneatuni. Smith Island, Gulf of California.

Setehell and Gardner, The Mar. Alg. Gulf of Calif., 1924, pp. 721,
722, pi. 17, fig. 44.

The small tufts which this species of Ectocarpus produces remind
one of the genus Gonodia {3Iyriactis) , but the penetrating part, which
extends relatively deep into the host, is composed of slender, almost
colorless, slightly branched, closely intertwined filaments, which, how-
ever, do not coalesce or form a false parenchyma as in the case of some
species of Gonodia. The plants, though sparse, are in excellent fruit-
ing condition. The chron)atophores are too much disorganized for
characterization.

440 University of California Puhlications in Botany [Vol. 8

34. Ectocarpus Bryantii S. and G.

Fronds intertwined, forming a more or less continuous stratum,
1-2.5 mm. high, attached by relatively short, penetrating, rhizoidal
filaments ; erect filaments forked more or less at the surface of the
host, with very few short ramuli above, nearly cylindrical, tapering
slightly above, uncorticated ; terminal cell blunt, 28-32/x diam., cells
1-2 times as long as broad ; chromatophores small disks ; zoosporangia
unknown ; gametangia narrowly to broadly fusiform, sessile or on
1-celled pedicels, 70-lOOjU, (up to 140ja) long, 25-35/x. broad, scattered
promiscuously along the whole length of the erect fronds.

Growing on Codium Brandegeei. La Paz, Lower California.

Setchell and Gardner, Mar. Alg. Gulf of Calif., 1924, p. 720, pi. 17,
fig. 45.

Ectocarpus Brya.ntii and E. gonodioides are evidently closely
related to each other and both have near affinities in the pusillus group
of Sauvageau (1895). They both differ from all of the forms pro-
posed in the method of branching and in having no hairs terminating
the erect filaments.

5. Streblonema Derb. and Sol.

Fronds composed of more or less branched, monosiphonous or in
part polysiphonous filaments, wholly or largely endophytic ; prostrate
primary filaments wholly endophytic, creeping among the cells of the
host, erect secondary filaments wholly or in large part endophytic,
simple or branched, hairs present or absent ; zoosporangia and g-ame-
tangia both present, terminal or lateral on the erect or on the prostrate
filaments, sessile or, more rarely, short-stalked.

Derbes and Solier, in Castagne, Supplem. Catal. Marseille, 1851,
p. 100.

The type species is >S^. sphaericum.

Key to the Species

1. Fronds causing noticeable distortions of the host 2

1. Fronds not causing noticeable distortions 3

2. Distortions in the form of pustules 13. S. scabiosum (p. 4.50)

2. Distortions in the form of extended rugose areas ...12. S. rugosum (p. 449)

3. Fronds producing noticeable patches or discolorations 4

3. Fronds inconspicuous 11

4. Patches large or extended 5

4. Patches small, usually orbicular (0.5 cm. or less diam.) 6

5. Patches orbicular 11. S. evagatum (p. 449)

1925] SetchcU-Gavdncr: MelanopJiyceae 441

5. Patches indefinite in outline and extent 6. S. penetrale (p. 446)

6. Patches definitely erumpent 7

6. Patches discolored, filaments only shghtly erumpent

7. S. transfixum (p. 446)

7. Gametangia pluriseriate 8. S. irregulare (p. 447)

7. Gametangia uniseriate 8

8. Parasitic on Zostera 4. S. vorax (p. 444)

8. Parasitic on other algae 9

9. Spots aecidioid 14. S. aecidioides f. pacificum (p. 450)

9. Spots not aecidioid 10

10. Gametangia cylindrical 15. S. myrionematoides (p. 452)

10. Gametangia narrowly elliptical 10. S. pacificum (p. 448)

11. Erect filaments not over 8m in diameter 12

11. Erect filaments over lOyu in diameter 15

12. Gametangia single 5. S. Porphyrae (p. 445)

12. Gametangia corymbose 13

13. Gametangia 25-40^ long 14

13. Gametangia over 60^ long 9. S. minutissimum (p. 447)

14. Erect filaments 4-5m in diameter 1. S. corymbiferum (p. 441)

14. Erect filaments 5-8yu in diameter 16. S. invesdens (p. 452)

15. Gametangia 50-100m long, 10-16m broad 2. S. anomalum (p 442)

15. Gametangia 90-130^ long, 28-36m broad 3. S. Johnstonae (p. 444)

1. Streblonema corymbiferum S. and G.

Plate 52, fig. 8

Fronds microscopic, composed of irregularly and alternately
branched filaments penetrating among the cells of the host ; fructi-
ferous ramuli decidedly aggregated, forming corymb-like clusters near
the surface of the host ; cells mostly cylindrical, in part irregular,
■ir-Ofx diam., 1.5-4 times as long; chromatophore band-shaped, not
covering the cell ; zoosporangia unknown ; gametangia cylindrical to
slightly fusiform, blunt, terminal on short pedicels, borne in clusters,
or sessile on filaments creeping near the surface of the host, 25-35/x
long, 4.5-5.5/i, broad; loculi uniseriate, dividing walls frequently
oblique.

Growing on Cwmagloia Andersonii (Farlow) S. and G., in com-
pany with Streblonema anomalum and *S^. Johnstonae. San Pedro,
California.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 391, pi. 43, fig. 8.

While studying material of Cwmagloia Andersonii (Farlow)
S. and G. collected by Mrs. H. D. Johnston at San Pedro, California,
in 1899 and deposited in the Herbarium of the University of Cali-
fornia under no. 94663, some specimens of Ectocarpaceae were inci-
dentally observed. When we came to study our material of this

442 University of California Puhlications in Botany [you 8

family, these specimens were given a careful scrutiny with the result
that three species new to science were detected on a single specimen
of the host, viz., Streblonema corymMferwrn, S. anomalum, and S.
Johnstonae, all of which have been published {lo<e. cit.).

The habit of each and the size of all the parts are so characteristic
that, notwithstanding their intimate association on the same host, they
can readily be distinguished. 8. corymhiferum is the most delicate
of the three. The very frequent grouping cf the gametangia into
corymb-like clusters is one of the most prominent distinguishing
characters of the species. Other species of this genus are as small or
even smaller. In size of parts, somewhat in habit and in habitat, it
resembles S. minuiissimnm Saunders found growing at Sitka, Alaska,
on " Liehmannia sp." (Saunders, 1901, p. 416.)

2. Streblonema anomalum S. and G.

Plate 52, figs. 1-3

Fronds microscopic, more or less confluent ; filaments of the
prostrate portion penetrating among the cells of the host, moderately
and irregularly branched, much contorted, up to 16/^ diam. ; erect
filaments unbranched, mostly cylindrical, blunt in part, and piliferous
in part, 200-250/x long, extending beyond the surface of the host ;
hair filaments sparse, arising on the creeping filaments ; cells of the
creeping filaments more or less divided 1-2 times lengthwise in the
oldest parts, producing a polysiphonous filament 18-24ja diam. ; other
cells somewhat irregular in form, 8-llfi diam., at times slightly con-
stricted at the cross-walls ; cells of the erect filaments cylindrical, 8-9/j.
diam., quadrate ; chromatophore a single thin parietal band, unbroken
in the erect filaments, more or less broken in the creeping filaments ;
zoosporangia ellipsoidal, ovoid or nearly spherical, 40-60/x long, 28-35ja
broad, sessile on the creeping filaments; gametangia cylindrico-conical,
mostly blunt, 50-100/a long, 10-16;u, broad, sessile or short-pedicellate
on the creeping filaments, single or secund.

Growing in Cumagloia Anclersonii (Farlow) S. and G. in com-
pany with Streblonema corymhiferum and ^S*. Johnstonae. San Pedro,
California.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 392, pi. 43, figs. 1-3.
Streblonema fasciculatum Saunders, Phyc. Mem., 1898, p. 148, pi. 13
(not of Thuret, in Le Jolis, Alg. mar. Cherb., no. 100, Liste, 1863,
p. 73).

1925] Setchell-Gardner: Melanophyceae 443

Examination of a small piece of authentic material of 8. fascicu-
latum Saunders collected at San Pedro, California, in August, 1896,
revealed the presence of specimens of the same three species mentioned
under S. corymhiferum, collected by Mrs. Johnston at the same place
and in the same month. S. anomalum is undoubtedly the one to which
Saunders refers as S. fasciculatum Thuret. It does not seem, how-
ever, to be Thuret 's species, if we have a correct conception of what
S. fasciculatum Thuret includes. His species was published in Le
Jolis, Algues marines de Cherbourg, no. 100, and in Liste, 1863, p. 73.
Thuret quotes as a synonym, ^S'. volubilis Pringsheim (Beitr. Morph.
Meeresalg., p. 13, pi. 3, fig. B, read in 1862). Pringsheim 's figure
of valiibilis shows the gametangia fasciculately branched, and he states
that this character is one of the chief characters of the genus Stre-
hlonema. This figure has been much quoted since. Pringsheim gives
no measurements of the parts, neither does Thuret.

Reinke (Algenfl., 1889, p. 41) recognizes StreMonema as a subgenus
of Ectocarpiis. He recognizes Pringsheim 's plant, quoting the fore-
going figure, but since voluhilis was already occupied by Crouan (1867,
p. 161) and fasciculatum was occupied by Harvey (Phyc. Brit., pi.
273), he renamed the plant, calling it Pringsheimii. Hauck (1884,
p. 323) seems to have been the first to give measurements of the
various parts of the plant. He lists it under Strehlonema, cites the
above literature of Thuret, and quotes S. voliobilis Pringsheim. Con-
sidering Pringsheim 's plant, recognized by Reinke and Hauck, as
being the same as Thuret 's, and taking Pringsheim 's figure and
Hauck 's measurements as being correct for Thuret 's S. fasciculatum,
then our plant is distinct and undescribed.

There is a little doubt in our minds at present whether the plant
we have described as Strehlonema anomalum is one or two species.
We do not find the filaments bearing the zoosporangia in the type
material examined to be like those figured by Saunders. His figures
show the main filaments as being like those of all the known Streh-
lonemas, monosiphonous, while those in both collections of material
which we have examined are uniformly polysiphonous, that is, the
cells of the main central parts of the thallus are divided once or twice
lengthwise. This never takes place in the plants which bear game-
tangia and which are intimately associated with them.

The measurements, method of branching of the two sets of plants,
and their chromatophore characters are practically the same. At
least two interpretations of this polysiphonous phenomenon appeal

444 University of California Publications in Batany [Vol.8

to us. They may represent a polymorphic state, in which a non-sexual
plant differs in form from a sexual plant of the same species, a unique
condition in the genus Strehlonema, or the polysiphonous condition
of the main filaments, found only in the non-sexual plants, may repre-
sent a character belonging to a wholly different genus, and hence it is
a new species of that genus. Until a more extensive study of fresh
material can be made, we feel that it is best to take the former view,
and place it in a new and polymorphic species of Strehlonema. We
have amended the family Ectocarpaceae to include species with this
polysiphonous condition.

3. Streblonema Johnstonae S. and G.

Plate 52, fig. -i

Fronds microscopic ; creeping filaments moderately branched,
branches alternate or opposite, at times slightly secund ; erect filaments
mostly simple, tapering slightly upward and at the base, extending
beyond the surface of the host, in part short-pilif erous ; cells of the
main creeping filaments more or less irregular in shape, chiefly doli-
iform, 12-lS/jL diam., 1.5-5 times as long; cells of erect filaments up to
24jti diam. in the widest part, nearly cylindrical, constricted at the
cross-walls; chromatophore a thin parietal band, nearly covering the
cell ; zoosporangia unknown ; gametangia cone-shaped to unsym-
metrically fusiform, 90-130ju, long, 28-36/a broad, mostly short-pedicel-
late on the creeping filaments extending to the surface of the host.

Growing in Cumagloia AndersonH (Farlow) S. and G. in company
with Strehlonema corymhiferum and S. anomahim. San Pedro,
California.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 394, pi. 43, fig. 4.

Of the three plants found ramifying among the filaments of the
above mentioned host, Strehlonema' Johnstonae is the most robust in
all of its parts. It can readily be distinguished from the other two
species with which it is associated by the large size of the gametangia.
No zoosporangia have been observed.

4. Streblonema vorax S. and G.

Fronds microscopic, prostrate filaments profusely branched, A'ery
tortuous, penetrating among the epidermal cells and spreading into
the interior of the host, decomposing the walls and filling the large

1^25] Sctchdl-Gardner: 3IeIanophyceae 445

parenchyma cells and devouring their contents; cells 6-10/i. long, ."i-S/x
diam. ; erect filaments 400-600ja long, 6.5-8|ii wide, attenuated ahove
into a hair, fasciculately branched at or near the surface of the host,
forming a compact mass of cells; zoosporangia broadly clavate,
GO-lOO/x long, 15-30ju, broad ; gametangia numerous, lateral on short
pedicels or sessile, narrowly cylindrical, blunt, 40-70//, (up to 100 fi)
long, 7-9ju, diam. ; loeuli mostly uniseriate.

Growing on the outer ends of the leaves of eel-grass, in the lower
littoral and upper sublittoral belts. Sitka, Alaska.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 389, pi. 44, figs. 1, 2.

Of all the species of Strehlanema thus far discovered on our coast,
S. vorax is the most destructive to the host. Although the penetrating
filaments have abundance of chromatophores, the plants seem to be in
a large degree parasitic. The cells of the host are closely compacted,
and have thick walls, yet they are devoured in large quantities. We
suggest the possibility of the secretion of enzymes with digestive
power which act upon the cell-wall and its protoplasm, after which
they are absorbed. This plant is found in company with several other
small Melanophyceae, Chlorophyceae and Rhodophyceae, none of
which penetrate the host.

5. Streblonema Porphyrae S. and G.

Fronds mostly endophytic ; prostrate filaments very tortuous, pene-
trating the cell-walls of the host between the cells, freely branched ;
erect filaments pushing slightly beyond the surface of the host, occa-
sionally branched ; hair filaments unknown ; cells of the creeping fila-
ments 3-4fi diam., irregular in shape ; zoosporangia unknown ; game-
tangia terminal in erect filaments, extending slightly beyond the sur-
face of the host, fusiform to irregular in shape, 25-35yu, long, 5-8/a
broad.

Growing in the parenchymatous base of PorpJiyra naiadum on
eel-grass. Pacific Grove, California.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 387, pi. 44, fig. 6.

Strel)Jonema Porphyrae is an exceedingly diminutive, though
apparently distinct, species, confined, so far as we know at present,
to the cushion-like bases of Porphyra naiadum. It ramifies deep into
the tissue, apparently not entering the cells.

446 University of California Publications in Botany [Vol. 8

6. Streblonema penetrale S. and G.

Fronds forming a continuous pulvinate stratum of indefinite shape
and size on the stipe of the host ; penetrating portion composed of
sparingly branched filaments extending relatively deep into the host
and mostly perpendicular to its surface; erect filaments fasciculately
branched at the surface of the host, 70-125/x long, tapering slightly
upwards, not piliferous; cells of the penetrating filaments cylindrical
to irregular, 6.5-8ju diam., 3-5 times as long; cells of the erect filaments
cylindrical to slightly doliif orm ; 6.5-8ju, diam., 1.5-2.5 times as long ;
chromatophores band-shaped ; zoosporangia unknown ; gametangia
cylindrical to blunt-fusiform, more or less irregular, 30-40;a long,
8-1 1/i broad ; loculi uniseriate.

Growing on the stipes of H esperophycus Harveyanus. Pacific
Grove, California.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 388, pi. 44, figs. 3, 4.

Streblonema penetrale, like S. myrionematoides below, is a difficult
species to classify. In its method of development and general struc-
ture it approximates S. myrionematoides closely. The penetrating
portion is much more highly developed than in that species, extending
into the host among the cells to a depth of four or five times as great
as the part which extends beyond the surface. The size of all the
parts is, in general, greater than in S. myrionematoides. No horizontal
filaments are present. The penetrating filaments are mostly per-
pendicular to the surface.

7. Streblonema transfixum S. and G.

Fronds forming patches 5-8 mm. diam. ; creeping filaments pene-
trating deeply among the cells of the host, distorted, irregularly
branched, 4—ojx diam., giving rise to scattered, short, erect, unbranched
filaments protruding from the surface and mostly terminated by game-
tangia; cells of the erect filaments cylindrical, 7-9/a diam., 0.75-1.5
times as long ; chromatophores band-shaped, nearly covering the cell
wall ; zoosporangia unknown ; gametangia cylindrico-conical, blunt,
40-60ju, long, 8-12/A broad ; loculi 1-2 seriate.

Growing on Desmarestia herbacea. San Pedro, California.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 391.

Streblonema transfixum forms definite, discolored areas on the
surface of the host, readily detectable. The species was described from

1925] Setchcll-Gurdner: Melanophijceae 4,4:1

dried material. The cells of the host were much collapsed and did
not straighten out when soaked and boiled, hence the details of the
species were not so thoroughly worked out as is desirable. The plants
penetrate among the cells of the host and seemingly pass all the way
through it. Just how much constitutes a single plant cannot at present
be ,made out.

8. Streblonema irregulare Saunders

Fronds consisting of irregularly branching prostrate filaments,
applied closely to the host plant, from wdiich arise numerous simple
or sparingly branched erect filaments 1-2 mm. high, 9-14/a diam., with
cells as long or twice as long as broad ; rhizoidal filaments arise from
the lower surface of the prostrate filaments and penetrate the host,
10-14/i, diam., longer or shorter than the diameter; zoosporangia
unknown ; gametangia narrowly ellipsoidal, terminal or lateral on
the erect filaments, 55-70ju long, 14-18/x diam.

Growing on the cysts of Nereocystis Luetkeana. Sitka, Ala.ska.

Saunders, Alg. Harriman Exp., 1901, p. 417, pi. 45, fig. 2 ; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 239.

We have not had an opportunity to examine the type material of
this species, nor have we seen any other material representing the
species since its discovery and first publication.

9. Streblonema minutissimum Saunders

Fronds composed of short, sparsely branched, creeping filaments,
l-2ju, diam., ramifying among the cortical filaments of the host, some-
times applied to them, and of erect filaments arising from the prostrate
filaments in the host, and intermingling with its peripheral filaments ;
erect filaments once or twice dichotomously branched, bearing above a
few short branches which become transformed into uniseriate game-
tangia, 20-30ja long, 3-5jx broad.

"Growing in the branches of Liehmannia.'" Sitka, Alaska.

Saunders, Alg. Harriman Exp., 1901, p. 416, pi. 45, fig. 3 ; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 239.

Nothing further has been learned of this species of Streblonema
nor of the host upon which it grew since its discovery at Sitka and
publication by Saunders (Joe. cit.). We have not seen any of the
specimens, but, judging from the figure and description, it seems
distinct from all other known forms.

448 TJyiiversitij of California Publications in Botany [Vol. 8

10. Streblonema pacificum Saunders (Orthog. mnt.)

Plate 41, fig. .53

Fronds occupying circular areas, 2-4 mm. diam. ; composed of a
network of profusely and irregularly branched horizontal creeping
filaments, penetrating among the paraphyses and sporangia of the
host, giving rise at right angles below to numerous acute branches, 3-4
cells long, and above to numerous fasciculately branched filaments
extending to the surface of the host and bearing the gametangia ; cells
of the horizontal filaments 3.5-4^ diam., somewhat irregular ; cells of
the rhizoidal filaments 1-2. S/a diam. ; cells of the erect filaments
variable in shape and size, 4-6/a diam. ; zoosporangia unknown ; game-
tangia numerous, terminating the erect filaments and projecting
beyond the surface of the host, blunt fusiform, 20-28fi long, 4-6/x
broad ; loculi uniseriate.

Growing on the sporophylls of Alaria. Alaska (Yakutat Bay) to
California (San Francisco Bay).

"Streblonema pacifica" Saunders, Alg. Harriman Exp., 1901, p.
417, pi. 45, figs, la, 15 ; Setchell and Gardner, Alg. N.W. Amer., 1903,
p. 239.

Superficially Streblonema pacificum Saunders appears like a small
Myrio'nema, the plants forming circular masses in the same manner
as that genus; The plants are exceedingly difficult to interpret in the
stage in which they begin to be visible to the unaided eye, that is
when the gametangia are forming. The sporangia of the host are at
this time elongating and more or less disturbing the arrangement of
the filaments. As far as can be ascertained from the study of mature
and nearly mature plants, the circular masses are occupied by a num-
ber of individuals instead of a single individual as in the case of a
Myrionema. This may not be the case, however. Against this con-
clusion the argument might be urged that the groups of plants are
too nearly uniform in size and too nearly circular in outline to be
composed of a necessarily variable number of plants intermingled,
yet this is the way they appear at maturity. More study of the
early stages in the life-history will have to be resorted to before the
question can be settled.

Saunders (loc. cit.) figures hair filaments but does not mention
them in his description. We have not been able to find any true hairs
in the material from California, nor in the portion of the type which
we examined. Neither have we been able to find anj^ anastomosing of

1925] SetcheJl-Gardncr: Melanoph\iveae 449

filaments such as he represents in figure Ih (Joe. cii.). These latter
conditions we may have overlooked in our study, as the plants are
exceedingly difficult to trace. The hair character is one that varies
considerably in any species.

11. Streblonema evagatum S. and G.

Fronds forming circular masses 1-2 cm. diam. ; creeping filaments
irregular, much branched, penetrating among the sporangia of the
host at their bases; erect filaments fasciculately branched, usually
within the host, tapering very slightly at the apex and at the base,
190-230/A high ; cells of the creeping portion 3.5-4ju, diam. ; cells of
the erect filaments 4/a diam. at the base, 2-3.5 times as long, 6.5-7. 5/i,
diam. in the widest part, 1-2 times as long, cylindrical, slightly con-
stricted at the cross-walls; chromatophores band-shaped, 1-2 in a
cell ; zoosporangia unknown ; gametangia numerous, cylindrical,
lateral, sessile or on short pedicels, 65-80/i, long, 5.5-6.5/x diam. ; loculi
uniseriate.

Growing' in the blades of Laminaria Anders&nii, upper sublittoral
belt. Cypress Point, Monterey County, California.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 390, pi. 42, figs. 1-5.

This species has a habit of growth very similar to that of Streh-
lonema pacificum, but the plants cover much more extensive areas
which are quite certainly not infested by a single plant as may be the
case in that species. Their presence seems to affect the host materially,
to a large extent destroying its sporangia and causing discoloration.
It is possibly partially parasitic.

12. Streblonema rugosum S. and G.

Plate 52, figs. 5-7

Fronds microscopic, producing decidedly rugose areas of consid-
erable extent on the surface of the host ; creeping portion penetrating
among the cells near the surface of the host, giving rise above to
fasciculately branched, erect, fructiferous filaments and hair fila-
ments ; cells of creeping and erect filaments very irregular in shape
and size ; chromatophores band-shaped ; zoosporangia unknown ; game-
tangia blunt, fusiform, terminal on most of the erect filaments, 18-24ju,
long, 5-6. 5/A broad ; loculi uniseriate, 4-6 in a series.

Growing on the blade of Alaria teniiifolm. Friday Harbor,
Washington.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 390, pi. 43, figs. 5-7.

450 University of California Publications in Botany [^""ol. 8

This species, like Strehlonema myrionematoides and S. scdbiosum,
has the larger part outside of the host. It penetrates to a considerable
depth among the cells of the uninjured host, later apparently mechanic-
ally causing the death of a few surface cells and an abnormal growth
of others, giving to the surface a rugose appearance. The plants
seem to associate in small groups of indefinite shape and size, often
confluent and although small, can readily be detected by the peculiar
appearance they give to the host.

13. Streblonema scabiosum S. and G.

Fronds forming circular to elliptical pustules of indefinite size
on the host ; penetrating portion profusely branched, at first pushing
between the cells causing their death, later taking possession of the
lumen of the cell, causing complete disintegration ; erect filaments
extending beyond the surface of the host, unbranched or more or less
f asciculately branched at the base, 50-80;U. high ; cells of the creeping
filaments verv^ irregular in shape and size ; cells of the erect filaments
cylindrical, 4.5-5.5ju, diam., 1-2.5 times as long ; chromatophores band-
shaped; zoosporangia unknown; g-ametangia cylindrical, seasile or
short-pedicellate, extending beyond the surface of the host, 40-60,u,
long, 4.5-6/A broad.

Growing on the stipe of Nereocystis Luetkeana. Cast ashore near
the "Cliff House," San Francisco, California.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 389, pi. 44, fig. 5.

Streblonema scabiosum forms marked scrofulous-like areas on the
lower part of the stipe of the host. It has a disastrous effect upon
the host, destroying its cells as far as it penetrates, and seems con-
stantly to spread by new infections around the margin of the affected
area. The cause of the death of the cells of the host has not been
investigated. In habit of growth and general form the species seems
related to aS'. myrionematoides and to S. penetrate.

14. Streblonema aecidioides f. pacificum S. and G.

Fronds microscopic, showing on the surface of the host as small
elevations, 75-150/x diam., the vegetative part forming a more or
less parenchymatous layer just beneath the surface laj^er of cells of
the host and giving rise below to a few rhizoidal filaments penetrating
deeper into the host ; all erect filaments fructiferous except a few
short hair filaments arising in the center of the frond; cells of the
hair filaments 4-5. 5/x diam., quadrate below, 5-8 times as long above.

19-5] SetchcU-Gardncr: MeJanopJnjceae 451

without a sheath ; zoosporangia ( ? ) narrowly clavate, sessile, 22-28|U.
long, 8-12ja broad at the apex ; gametangia numerous, closely crowded,
cylindrical, sessile on the horizontal layer, 45-55/1, long, 5-6.5/i, broad ;
loculi uniseriate.

Growing within the lamina of Hedopliylhim sessile (Aresch.)
Setchell, near the outer end. Neah Bay, Washington.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 395, pi. 44, figs. 8, 9.
Phycacelis fecunda Tilden, Amer. Alg. (Exsicc), no. 356 from
Victoria, B. C. (determined by De Alton Saunders).

Strehlonetna aecidiaides f. pmificuni seems very closely related to
the Ectocarpus aecklioides of Rosenvinge (1893, p. 894), found grow-
ing in Greenland on Laminarm longicruris and L. groenlandica. It
differs only in minor details as to the dimensions of the parts. Regard-
ing the zoosporangia we have to speak with uncertainty. As figured
by Rosenvinge, the Greenland plant has them well developed and
producing zoospores. They are in distinct "aecidia" and apparently
on distinct non-sexual plants. In our species they likewise appear to
be on non-sexual plants. In ours there is no indication of the produc-
tion of zoospores. The plants are possibly too young, or possibly they
bear abortive organs so commonly met with in various other genera
on our coast, the nature of which is still an open question. This
form seems to enter the host from the surface and after penetrating
to the second layer of cells spreads out horizontally between the sur-
face layer and the second layer. Later, filaments arise from the
under side of this layer and penetrate among the cells of the host,
apparently never entering them. Finally from the upper surface each
cell in the central region of the layer gives rise to a filament and the
mass acting together lifts up the surface layer of cells of the host,
forming a small blister which finally ruptures, as in the case of Rosen-
vinge's plants. The erect filaments are almost simultaneously trans-
formed into gametangia, except a few in the center which develop
into hairs.

Foslie (1894, p. 167, 23 in reprint) describes forms of the same
species found growing on Laminaria saccharincn at Kjelvik and at
Lyng0 near Troms0, Norway. These he listed under Ectocarpus
(Strehloncma) aecidiaides Rosenvinge. The measurements of the
parts of his forms average, in general, a little larger than those of
' Rosenvinge. De-Toni (1895, p. 577) cites these Arctic plants under
Strehlanema aecidioides (Rosenv.) Foslie. Foslie gives 80/i, as the
extreme length of the gametangia.

452 University of California Puhlicaiions in Botany [Vol.8

15. Streblonema mjrrionematoides S. and G.

Fronds microscopic ; prostrate portion very poorlj^ or scantily
developed, penetrating only sligHtly among the two or three outer
layers of cells of the host ; erect filaments more or less f asciculately
branched at the surface of the host, 65-80/a long, mostly fructiferous,
hairs few ; cells of the penetrating filaments 4— 5/a diam., irregular in
shape ; zoosporangia unknown ; gametangia numerous, cylindrical,
50-65ju, long, 4.5-6.5ju. broad ; loculi uniseriate.

Growing on the blade of Laminaria Andersonii Farlow. Moss
Beach, San Mateo County, California.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 387, pi. 44, fig. 7.

It is an extremely perplexing problem to decide upon the generic
position of forms such as the one described above. It has close affini-
ties with the three genera, Streblonema, Ectocarpiis and Myrionema.
The erect fronds exterior to the host and more or less branched, with
relatively scanty attaching portions, constitute the vegetative portion
of an Ectocarpns of microscopic size. The gametangia are strictly
those of a typical Myrionema, but there is lacking the disk-shaped basal
layer of filaments spreading out on the surface of the host, character-
istic of that genus. The extremely reduced character of the penetrat-
ing portion does not speak well for the genus Streblonema. On the
whole, we feel that with our present criteria for these genera it best
agrees with the characters of the genus Strehlonema, where we have
placed it. It penetrates the uninjured host, but only to a slight depth.
The plants, however, are usually so congested that their growth soon
crowds the surface layer of the host cells to such an extent that they
die and disintegrate, there being no evidence that the associate pene-
trates them and absorbs their material. The palisade-like stratum of
gametangia suggests very strongly the Myrionema^ character which is
the reason for the specific name.

16. Streblonema investiens (Collins) S. and G.

Fronds occupying indefinite areas on the host ; creeping filaments
irregularly branched, often curving outward and bearing on the out-
side short, simple, or sparsely branched filaments ; hairs sparse ; cells
of creeping filaments 5-8ja diam., 1-2 (up to 4) times as long, swollen
or cylindrical; cells of the ramuli G/x diam., 1-2 times as long; cells
of the hairs S^u. diam. ; chromatophores discoid, small, several in a

1925] Setchfll-Gardnrr: Melanophyceae 453

cell; zoosporangia ovoid, sessile or on 1-celled pedicels on both the
creeping filaments and the ramuli, 20/x long, 15/1, broad ; gametangia
cylindrical, 25-40/x long, 8-lOjU, broad ; loculi mostly uniseriate ; game-
tangia and zoosporangia growing on the same plant.

Growing in the fronds of Helmmthodadia calvadosii (Lamour.)
Setchell. San Pedro, California.

Setchell and Gardner, Phyc. Cont. V, 1922, p. 396. Strep^thalia
investieiis Collins, in Collins, Ilolden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. 738.

It seems that the chief technical distinction between the genera
Strepsithalia Saiivageau and Strehloiiema Derb. and Sol. is the secre-
tion by Strepsithalia of a rather copious gelatinous sheath investing
the entire plant, particularly the exposed portions, the ramuli. Since
we are not able to demonstrate the presence of such a sheath, even to
the slightest degree, we have thought it best to place Collins'
Strepsithalia investiens in the genus StreMonema.

FAMILY 3. MYEIONEMATACEAE foslif. (orthog. mut.)

Thallus two-fold, viz., (1) of a prostrate disk formed of radiating
filaments more or less closely united and one or two cells in thick-
ness, entirely superficial or penetrating the host by short rhizoidal
filaments, and (2) of erect, simple or branched, monosiphonous,
usually parallel filaments, either free or enclosed in a common jelly ;
growth peripheral in t'he basal portion and subapical or intercalary
in the erect filaments ; hairs present or absent ; erect filaments entirely
fertile, partially sterile or sterile and fertile intermixed ; cells uninu-
cleate, each with one or more parietal disk-shaped or band-shaped
chromatophores ; zoosporangia unilocular, terminal or lateral, sessile
or pedicelled, never intercalary; gametangia plurilocular, terminal or
lateral, with loculi uni- or pluriseriate ; gametophyte and sporophyte
identical in size and structure.

Myrionemaceae Foslie, Contr. Knowl. Mar. Algae Norway, I, 1890,
p. 91.

The members of the Myrionemataceae, as we understand them,
seem closely related to the Ectocarpaceae, on the one hand, and the
Elachisteaceae on the other. There seems to be little reason for plac-
ing the genera in Chordariaceae as Kjellman (1893, p. 225) has done,
nor even associating them closely with the Mesogloiaceae, as Oltmanns
(1922, p. 29) has done, since it seems to us that Strepsithalia, in
typical form, resembles StrehJonema in its basal portion, but agrees

454 Vniversiiy of Calif ornia Puhlications in Botany [Vol.8

better with the Elachistea and Myriactis-groi\i-)S, of the Elachisteaceae
in its erect filaments slightly contracted at the base. The various
Myrionemataceae are most closely allied to the species of Ralfsia,
particularly with those referred to Stragularia. The prostrate basal
portion of one or two layers of cells readily distinguishes the members
of the Myrionemataceae from those of all other families except those
of the simpler Ralfsiaceae.

IVEY TO THE Genera

1. Erect filaments free 2

1. Erect filaments enclosed in a jelly 9. Microspongium (p. 492)

2. Basal portion monostromatic 3

2. Basal portion distromatic 8. Hecatonema (p. 488)

3. Gametangia with locuU uniseriate 6. Myrionema (p. 454)

3. Gametangia with loculi pluriseriate 7. Compsonema (p. 473)

6. Myrionema Grev.

Fronds forming small cushions, circular to ellipsoidal, or even
quite irregular in outline, composed of a monostromatic basal stratum
and numerous pigment-bearing erect filaments with or without hairs
interspersed ; the erect filaments in part or wholty transformed into
gametangia, except at the margins ; basal stratum composed of closely
crowded filaments radiating from a common center, with dichotomous
branching by splitting of the terminal cells, rarely with a few, short,
subulate branches penetrating the host ; reproduction by unilocular
zoosporangia and by plurilocular gametangia mostly with uniseriate
loculi.

Greville, Scottish Crypt. Flora, vol. 5, 1827, pi. 300.

The type species of this genus is M. strangulans Grev. The type
locality is the shore of Appin, Scotland. It was first brought to
notice by Captain Carmichael, who found it growing on a small species
of Salenia (Enteromorpha) , which it usually completely encircles.
Greville says of his genus: "It evidently belongs to the Nostochinae
of Agardh. " Although it is undoubtedly known today to belong to
the Melanophyceae, there has been considerable diversity of opinion
concerning the family to which it properly has its closest affinities.
Harvey (1841, p. 124) placed it among the Chlorospermeae, under
the tribe Chaetophoroideae, probably on account of its "gelatinous"
nature as suggested by Greville. He states, however, that it is "a
genus of doubtful affinity," and "it rather belongs to the series
Melanospermeae. " J. G. Agardh (1848, p. 47) placed it in his order
Chordarieae under his tribe Mesogloiaceae. Kjellman (1890, p. 40)

192.')] Setchell-Gardner: Melanophyceae 455

emended the genus and placed it in the family Chordariaeeae, and in
1893 (p. 226) retained it in the same family. Foslie (1890. p. 91)
and Rosenvinge (1893, p. 901) and still later, Borgesen (1902, p. 419)
placed it in the "Myrionemaceae, " the first two designating the group
as a family, and the last as an order. De-Toni (1895, p. 178) retained
it in Chordariaeeae. Oltmanns (1904, p. 382) placed it in the group
Myrionemeae of the family Ectocarpaceae. In 1922 he arranged it
under the "Isogame Ectocarpales" in the family " Myrionemaceae, "
along with StrepsithaJia and Ralfsia.

After a careful study of a large assortment of material belonging
to the Ectocarpales, we have limited the genus to include forms
possessing the characters mentioned in the foregoing generic diagnosis
and placed the genus in the family Myrionemataceae, along with three
other very closely related genera; viz., Compsonema, Micraspongium,
and Hecatonema. We have, however, included under Myrionema the
species formerly assigned to Ascocyclus Magnus and Phycocelis
Stroemfelt.

In working over the members of the genus Myrionema found on
the Pacific Coast of North America, we have studied and separated
over twenty species or forms. This segregation must be considered
as provisional since no attempt at cultural criteria has been possible.
We have separated the species into two lines, the one in which only
zoosporangial forms are known, and the other in which only game-
tangial specimens are known, or those having gametangia and
"ascocysts, " the latter problematic structures. Possibly some of the
structures we have called "ascocysts" may be young zoosporangia,
but probably most or all of such structures, as we have seen them, are
hypertrophied gametangia. The possible genetic connection between
some of the one series of species and some of the other series cannot
be demonstrated except by cultures.

Key to the Species

1. With zoosporangia only 2

1. With gametangia only or with both gametangia and "aspocysts" 3

2. Plants forming definite circular thalH 21. M. strangulans (p. 471)

2. Plants forming indefinite rugose thalli 22. M. obscurum (p. 472)

3. Gametangia sessile 4

3. Gametangia pedicellate 9

4. All of the erect filaments transformed into gametangia 5

4. Part of the erect filaments transformed into gametangia

10. M. foecundum f. divergens fp. 463)

5. Plants with subulate rhizoids 7. M. foecundum f. subulatum (p. 462)

5. Plants without subulate rhizoids 6

6. Gametangia branched in part 8. M. foecundum f. ramulosum (p. 462)

6. Gametangia unbranched 7

456 University of California Fiiblications in Botany [Vol. 8

7. Gametangia up to 40/x long 6. M. foecundum f. simplicissimum (p. 461)

7. Gametangia over 40^ long 8

8. Gametangia up to 65/x long and 6m broad 1. M. primarium (p. 456)

8. Gametangia 80m long and 8m broad 9. M. foecundum f. majus (p. 463)

9. All of the erect filaments fructiferous 10

9. Part of the erect filaments fructiferous 12

10. Gametangia up to 130m long 12. M. balticum f. californicum (p. 465)

10. Gametangia up to 45m long 11

11, Pedicels 1-3-celled 3. M. corunnae f. uniforme (p. 458)

11, Pedicels up to 8-celled 11. M. minutissima (p. 464)

12. Sterile filaments longer than the gametangia 13

12. Sterile filaments not longer than the gametangia 20

13. Gametangia all terminal 14

13. Gametangia in part lateral on sterile filaments 19. M. setiferum (p. 470)

14. Sterile filaments partially setiferous 15

14. Sterile filaments not setiferous 16

15. Erect filaments unbranched 18. M. phyllophilum (p. 469)

15. Erect filaments branched 20. M. globosum f. alfine (p. 471)

16. Gametangia decidedly acuminate

2. M. primarium f. acuminatum (p. 457)

16. Gametangia cylindrical or nearly so 17

17. Cells of sterile filaments doliiform 17. M. attenuatum f. doliiforme (p. 468)

17. Cells of sterile filaments cylindrical 18

18. Creeping filaments with short subulate rhizoids ;

15. M. hecatonematoides (p. 467)

18. Creeping filaments without rhizoids 19

19. Gametangia up to 45m long 16. M. attenuatum (p. 468)

19. Gametangia up to 150m long 13. M. balticum f. pedicellatum (p. 466)

20. Erect filaments up to 65m high 5. M. corunnae f. sterile (p. 460)

20. Erect filaments over 65m high 21

21. Gametangia on 1-3-celled pedicels 14. M. compsonematoides (p. 467)

21. Gametangia on long pedicels, 7-10-celled.. .4. M. corunnae f. angulatum (p., 459)

1. Myrionema primarium S. and G.

Plate 55, fig. 12

Fronds forming circular cushions 0.5-1.25 mm. diam. ; prostrate
portion composed of regularly radiating, closely compacted filaments ;
erect filaments unbranched, densely crowded, arising from every cell
of the creeping filaments; true hair filaments numerous, scattered
promiscuously in the center of the frond ; cells of erect filaments 4-4.5/j.
diam., quadrate below, terminal cell, 2-3 times as long when young ;
cells of true hairs 4-5/i, diam., 4-6 times as long; zoosporangia
unknown ; gametangia sessile on the creeping filaments, arising by
transformation of all of each erect filament, except at the margin, very
compact, of nearly equal length in the center of the frond, shorter
toward the margin, cylindrical, 55-65/i, long, 5-6ju, broad, blunt or
tapering slightly at the apices, terminal cell acute-conical.

1925] Setchcll-Gurdncr: Melanophyceae 457

Growing on the outer end of the young blade of Costaria costaia
in the lower littoral and the upper sublittoral belts, Oregon (Coos
Bay) to central California.

Setehell and Gardner, Phyc. Cont. II, 1922, p. 334, pi. 34, fig. 12.

We consider Mijrionema priniaritiin to l)e a typical representative
of a group whose members are the most primitive of the genus when
considered from the standpoint of diiferentiation. The prostrate
basal layer is composed of long, regularly radiating filaments with
apical growth. Radial divisions of apical cells occur just often enough
to completely occupy all of the space between one another as the plant
continues to increase in diameter and in circumference. The branch-
ing is always dichotomous, and is accompanied by widening of the
apical cell, and the establishment of two growing regions on opposite
corners, which are subsequently separated from the remainder of the
cell by a wall, thus establishing two equal branches. Beginning in the
center, each cell successively toward the periphery gives rise by hori-
zontal divisions to erect filaments of nearly equal length, the only
exception or modification being that some cells give rise to long
filaments, the so-called true hairs, with the meristem at the base, and
the outer cells long and colorless. All other erect filaments are
transformed into gametangia. Only the cells of the basal filaments
remain sterile, or probably a few short erect filaments at the periphery
of the frond may never come to maturity. Thus starting with a single
cell the maximum of reproductive cells arises in this group. Starting
Avith such forms the course of evolution seems to have been in the
direction of the sterilization of tissue.

2. Myrionema primarium f, acuminatum S, and G,

Fronds microscopic, growing among other small algae, erect fila-
ments in part sterile, 80-100/x long, cells slightly doliif orm ; game-
tangia 4-5ju, diam., acuminate, in part short-pedicellate.

Growing on Macrocystis pyrifera. Carmel Bay, Monterey County,
California,

Setehell and Gardner, Phyc, Cont, II, 1922, p, 335, pi. 32, fig. 9.

There is but a slight difference between M. primarium and forma
acuminatum. It is worthy of note that many gametangia in the center
of the thallus are short-pedicellate, and that a few, about one in
twenty-five, of the erect filaments continue to grow, attaining a length
two to three times as long as the gametangia.

458 University of California Publications in Botany [Vol. 8

Myrionema corunnae Sauv.

Fronds forming- small, circular cushions, 2-3 mm. diam., basal
disk monostromatic, not attached by rhizoids from below ; erect fila-
ments longest in the middle of the thallus, gradually reduced in length
toward the margin ; hairs scattered, relatively short, 4— S.S^u. diam. at
the base, cells 25-40/a long at the apex, meristem at the base but all
cells elongating at maturity ; all erect filaments fructiferous ; game-
tangia on 2-4-cened pedicels at the center of the frond, becoming
sessile toward the margin, mostly simple but not infrequently bearing
3-5 branches, 25-120/x, long, 4.5-7jii diam. ; zoosporangia unknown.

Sauvageau, Sur quelq. Myrionem., 1897, p. 237 (repr. 1898, p. 77).

The type material of the species was found growing in minute,
closely associated cushions on the outer ends of Laminaria pallida
at La Corogne (Spain), in November, 1895. We have not yet dis-
covered in our region any specimen which we can refer to typical
31. corunnae as described and figured by Sauvageau. The material
collected by one of us (Gardner) in December, 1909, growing on
Nereocystis Luetkeana (Mert.) P. and R., was identified by F. S.
Collins as belonging to that species and distributed in Phycotheca
Boreali-Americana (Exsicc.) as no 1738. Careful study of this
material reveals a mixture of several genera belonging to Myrione-
mataceae and Ectocarpaceae, none of which, so far as the specimens
in our copies go, is M. corunnae. Experience in collecting and
critical study of the numerous forms found, growing upon the above
mentioned host, have shown the necessity of exercising extreme care
in selecting pure or uniform material. The presence of branched
gametangia in typical M. corunnae, as diagnosed by Sauvageau, is a
character not found in any of our Pacific coast forms.

The three following distinct forms have been segregated :

3. Myrionema corunnae f. uni forme

Fronds forming irregular cushions, 0.5-1.5 mm. diam. ; creeping
filaments forming a compact layer with peripheral growth ; hairs
absent; erect filaments unbranched, slightly attenuated at the apices,
80-100/x. (up to 120/a) high; cells of erect filaments 4.5-5. 5;li diam.,
1-1.5 times a.s long; "ascocysts" sparse, clavate; gametangia forming
a solid, compact, palisade stratum of approximately the same height

1925] SeicluU-Gardncv: Melanophyceae 459

in the center, g:radually diminishing toward the periphery, mostly
sessile, some on 1-3-celled pedicels, 80-100|U, high, 6-6.5ja diam. ; loculi
uniseriate.

Growing on the blades of Costaria costata, Lammaria Sinclairii
and Alaria marginata. Central California (Moss Beach, San Mateo
County, and Fort Point in San Francisco).

Setchell and Gardner, Phyc. Cont. II, 1922, p. 339, pi. 33, figs. 4, 5.

Forma uniforme ditfers from the species as described and figured
by Sauvageau (1898, pp. 77-82, repr., figs. 14 A-F) in having no hair
filaments, in having no branched gametangia, in having "ascocysts, "
and in having slightly different dimensions.

There are some differences in the size of the plants we have found
growing on different hosts in central California. The plants chosen
as the type of the form grew on the blades of Alaria marginata at
Fort Point, San Francisco. Very generally, this species of Alaria has
delicate grooves obliquely radiating from the midrib to the margin.
The presence of these grooves is often accentuated by the growth of this
minute Myrionema, usually so numerous as to be confluent, so that the
individuals cannot be detected except by microscopic examination.

4. Myrionema corunnae f. angulatum S. and G.

Fronds forming microscopic cushions, irregular in outline, among
other small algae on the host ; creeping filaments regularly radiating
when undisturbed ; erect filaments unbranched, densely crowded,
120-140/A long ; true hairs absent ; cells of creeping filaments 4-4.5/x
diam., mostly quadrate ; cells of erect filaments cylindrical, 4—4.5/*
diam., 1.5-3 times as long; zoosporangia and "ascocysts" unknown;
gametangia cylindrical, slightly attenuated at the apices, terminal on
6-10-celled pedicels, 35-45/a (up to 60/x) long, 4-6.5/x broad, many
partitions of the loculi oblique.

Growing on the cysts of Egregia Menziesii. Cast ashore near the
"Cliff House," San Francisco, California.

Setchell and Gardner, Phyc. Cont. II, 19-22, p. 340, pi. 33, figs. 6, 7.

This form is to be distinguished from the species as described by
Sauvageau (1898, pp. 77-82, repr.) by the absence of hairs, the
presence of sterile erect filaments, the gametangia mostly on longer
pedicels and by the very small size of the loculi, often produced by
perpendicular and slanting walls. In part, the gametangia seem pluri-
seriate, caused by the division of some of the original cells into small,
angular loculi.

460 University of California Puhlications in Botany [Vol. 8

5. Myrionema corunnae f. sterile S. and G.

Fronds forming circular cushions, 1.5-3 mm. diam. ; erect filaments
straight, unbranched, cylindrical, blunt, 55-65,;U. long; true hairs and
"ascocysts" absent; gametangia cylindrical, blunt, on 3-5-celled
pedicels, rarely sessile, 55-65/* long, 4.5-5. 5/x broad; gametangia and
sterile erect filaments diminishing in length very regularly and
gradually from the center to the circumference of the fronds.

Growing on the pneumatocysts of Nereocystis Lu-etkeana. Cast
ashore. Carmel Bay, Monterey County, California.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 340, pi. 33, figs.
13-15. Chilionema ocellatum Collins, Holden and Setchell, Phyc.
Bor.-Amer. (Exsicc), no. 1739.

This form is to be distinguished from the species as described by
Sauvageau (1898, pp. 77-82, repr.) by the presence of numerous
sterile erect filaments interspersed among the gametangia, about one-
fourth to one-fifth as many as the gametangia, by the absence of hairs,
and by the very gradual diminution in the length of the sterile fila-
ments and the gametangia from the center to the circumference of
the fronds. The sterile erect filaments are of the same length as the
gametangia.

Myrionema foecundum (Stroemf.) Foslie (Orthog. mut.)

Myrionema {Phycocelis) f&ecunda (Stroemf.) Foslie, New or
Crit. Norw. Algae, 1894, p. 17 (repr.). Phycocelis foecunda Stroem-
felt, in Notarisia, 1888, p. 383, pi. 3, fig. 5.

Foslie (1894, p. 15, repr.) states, "I have been unable to find any
real distinction between Myrionema, Phycocelis and Ascocycius,
and therefore I consider the two latter as subgenus of the former,
Ascocycius, however principally founded on the presence of para-
physes. " We likewise are unable to discern any fundamental dis-
tinctions among these three genera. Phycocelis was proposed by
Stroemf elt (1888, p. 383) with P. foecunda as the type. This species
has a unistratose base, and all of the erect filaments, except the hairs
and the marginal filaments, metamorphose into gametangia. These
are characters of typical Myrionema, as we interpret the genus.

The genus Ascocycius was founded by Magnus (1874, p. 73) based
upon J. G. Agardh's Myrionema orl)iculare, which, as figured by
Crouan (1867, pi. 25, gen. 163), has a unistratose base, plurilocular

1925] Setchell-Gardner : Melanophyceae 461

gametangia and hairs. Having found in addition to these certain
single celled sack-like hairs, he created the new genus based upon this
character. Two sizes of these are well illustrated by Kylin (1907,
p. 39, fig. 9), found growing on Zostera and Cladophora. These are
probably the structures which Foslie {loc. cii.) refers to as "para-
physes. " After having studied these structures as they occur in a
variety of species of different genera belonging to the Ectocarpales,
our experience leads us to conclude that they are not constant in any
species, and that they are probably caused by some pathological con-
dition, the nature of which we are not at present able to state. We are
taking the view, therefore, that neither of the two genera, Phycocelis
Stroemfelt and Asc&cyclus Magnus are valid genera, or even subgenera.
We have not seen either the type or any other authentic material
of the Phycocelis foecunda Stroemfelt, neither have we any authentic
measurements of the original material. De-Toni (1895, p. 582) gives
35/u, to 70/i as the height and Ijx to 12jLt as the width of the gametangia.
We are listing five forms under the species none of which, so far as we
are able to judge, corresponds to the typical material.

6. Myrionema foecundum f. simplicissimum S. and G.

Fronds forming circular cushions 0.5-1.5 mm. diam. ; prostrate
portion composed of regularly radiating, closely adhering filaments
with apical growtn and branching- ; erect filaments all fructiferous ;
true hairs unknown ; cells of creeping filaments 3-4. 5ju, diam., quadrate
toward the center, two times as long at the margin ; zoosporangia
unknown; "ascocysts" rare to numerous, clavate, 28-34/i. high; game-
tangia arising by transformation of all of each erect filament, except
near the margin of the frond, 30-40/x high, 7-9ju, broad.

Growing on the sporophylls of Nercocystis Luetkeama. Alaska
(Kadiak Island), to central California.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 336, pi. 33, figs.
9, 10. Myrionema strangulans Setchell and Gardner, Alg. N.W.
Amer., 1903, p. 249; Myrionema vulgare Collins, Ilolden and Setchell,
Phyc. Bor.-Amer. '^Exsicc), no. 924. Phycocelis reptans Tilden,^
Amer. Alg. (Exsicc), no. 357 (in part).

The Phycocelis reptans of Tilden, mentioned above, proves not to
be uniform, so far as the distribution in the copy at hand is concerned.
The material growing on the sporophyll of Ncreocystis seems clearl}'
to be 31. foecundum f. simplicissimum although young. The material

462 University of California Publications in Botany [Vol. 8

on the other part of the host is a mixture. In height of gametangia
forma simplicissimum approximates to forma seriata Reinke (1889,
p. 46), but has no true hairs so far as we have been able to observe
from the examination of many specimens from different localities. At
times the individuals are very numerous on the host, so much so as to
interfere with each others' growth. In one collection (Gardner,
no. 4651) they are completely confluent over several square inches,
and the outline of the individuals can be detected only by staining
and by microscopic examination.

7. Myrionema foecundum f. subulatum S. and G.

Basal filaments attached by numerous awl-shaped rhizoids, 2-4
cells long, penetrating between the cells of the hostj zoosporangia,
"ascocysts" and true hairs unknown.

Growing on the sporophylls of Nereocystis Luetkeana. Near the
"Cliff House, " San Francisco, California.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 337, pi. 33, figs. 1-3.

Forma suhulatum is very similar to forma simplicissimum. The
gametangia average slightly larger and are usually more blunt. There
are no so-called "ascocysts" present. The chief distinction is the
presence in this form of numerous awl-shaped rhizoids, found in no
other form of the species, so far as we have been able to ascertain.

8. Myrionema foecundum f. ramulosum S. and G.

Fronds forming small circular cushions, 1-3 mm. diam. ; erect
filaments slightly branched above, 55-65/a long ; cells of the erect fila-
ments 5.5-6.5fi diam., quadrate, terminal cell frequently 2-3 times
as long; zoosporangia, "ascocysts" and true hairs unknown; all of the
erect filaments transformed into gametangia.

Growing on the pneumatocysts of Nereocystis Lu-etkeam.a. Friday
Harbor, San Juan Island, Washington.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 337, pi. 33, fig. 8.

This form of M. foecimditm differs from the other forms of the
species in having secund ''proliferations" near the outer ends of
many of the gametangia.

1925] SetchfU-Gardnrr: Melnnophyceae 463

9. Myrionema foecundum f. majus S. and G.

Fronds forming circular cushions, 1-2 mm. diam. with convex
upper surface; prostrate portion composed of closely adhering, regu-
larly radiating filaments ; erect filaments densely crowded, unbranched ;
cells C3'lindrical, 5-7/^ diam. ; true hairs sparse, distributed promiscu-
ously over the frond, meristem at the base, surrounded by a sheath,
400-500/x long, cells 6/* diam., 5-8 times as long as the diameter at
the apices; "ascocysts" sessile on the prostrate filaments, cylindrical,
slightly larger at the distal end, 45-55/i, long, 8-12/i, diam., scattered
sparsely over the frond, occasionally in groups; zoosporangia un-
known ; gametangia sessile on the basal filaments, cylindrical, 60-80/a
long, 6-Six diam., arising by transformation of all of each erect
filament, cross-walls horizontal or oblique.

Growing on Laminaria ephemera Setchell. Port Renfrew, Van-
couver Island, Neah Bay, Puget Sound and central California (Carmel
Bay).

Setchell and Gardner, Phyc. Cont. II, 1922, p. 338, pi. 32, fig. 7.

This small epiphyte seems to be restricted in its habitat to the
above named host. The only known localities in which the host has
been observed are those mentioned above. The material from each
locality is richly infested by this epiphyte. The host, richly infested,
was distributed by Miss Josephine Tilden in her American Algae
(Exsicc), no. 609, under Renfrewia parvula Griggs.

It seems very closely related to Myrionema foecundum f. simpJi-
cissimum. The differences are in the larger measurements of all of
the parts, the presence of numerous, larger, so-called "ascocysts, "
occurring in groups, or scattered among the gametangia, in the
presence of well-developed hairs, and in the more convex upper
surface of the mass of erect filaments.

10. Myrionema foecundum f. divergens S. and G.

Plate 55, figs. 8, 9

Fronds forming circular cushions, 1.5-3.5 mm. diam. ; basal por-
tion composed of regularly radiating filaments closely compacted,
firmly adhering to each other and to the host ; sterile erect filaments
sparse, unbranched, blunt, scattered promiscuously among the game-
tangia, 300-400/A long; true hairs absent; "ascocysts" numerous,
scattered among and shorter than the gametangia, cylindrical to
clavate ; cells of the creeping filaments 5-6ju, diam., 1.5-2 times as

464 Universitij of California Puhlicaiions in Botany ['^''ol. 8

long"; cells of the erect filaments 8-9/t diam., 1.5-3 times as long,
cylindrical, not constricted; zoosporangia unknown; g:ametangia
variable in shape, cylindrical, clindrico-conical or fusiform, 50-60/a
long', 7.5-8. 5/A broad; loculi 1-2-seriate,

Growing on the sporophylls of Nereocystis Luetkeana. Pacific
Grove, California.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 338, pi. 34, figs. 8, 9.

This form of 31. foecundum manifests some slight differentiations
not present in any other form of the species. Scattered promiscuously
over the fronds are erect filaments two to three times as long as the
gametangia. The "ascocysts" are abundant. Some irregularities
are also shown in the creeping filaments. Typically the branching
seems to take place by the splitting of the apical cell, but it seems
that frequently one of the dichotomy fails to develop till later, pro-
ducing the appearance of lateral branching. There is also an occa-
sional oblique or longitudinal division of a cell of the creeping fila-
ments perpendicular to the host.

11. Myrionema minutissimum S. and G.

Plate 55, figs. 1-3

Fronds forming inconspicuous cushions, 0.75-1.5 mm. diam. ; pros-
trate portion composed of very closely compact regularly radiating
filaments; erect filaments densely crowded, cylindrical, all fructi-
ferous; true hairs unknown; cells of creeping filaments 4/x diam.,
quadrate in the center of the frond, slightly longer at the margin ; cells
of erect filaments the same in size as in the creeping filaments ; zoo-
sporangia and "ascocysts" unknown; gametangia on 4-8-celled
pedicels, cylindrical, densely crowded, 34-40/x long, 4-4.5;a broad.

Growing on the cysts of Egregia Menziesii. Cast ashore near the
"Cliff House," San Francisco, California.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 336, pi. 34, figs. 1-3.

Myrionema minutissimum may be distinguished by the absence of
sterile erect filaments and hairs, and by the small sized, very densely
crowded pedicellate gametangia. Thus far no "ascocysts" nor zoo-
sporangia have been observed. In its phylogenetic relationship it
would seem to stand equally close to M. corunnae and 31. foecundum,
differing from each, however, in details of measurements. The erect
filaments are of the same diameter at their bases as the length of the
cells in the creeping filaments. Since the lower part of each remains
sterile, it gives the mass of cells the appearance of being parenchy-
matous.

1925 1 Scfchcll-Gardncr: Melanoplujceae 465

Myrionema balticum (Reinke) Foslie

Myrionema (FJnjcoceJis) halticnm (Reinke) Foslie, New or Crit.
Nonv. Alg., 1894, p. 17. Ascoci/clus haltkus Reinke, Algenfl. westl.
Ostsee, 1889a, p. 45, Atlas, 1889, p. 19, pi. 16, figs. 1-4.

A discussion of the reasons for rejecting Ascocyclus may be found
under Myrionema foecundum. The chief character which Magnus
used in establishing his new genus AscocycJus, viz., the "ascocysts, "
does not appear in the diagnosis given by Reinke for A. balticus. It
seems to us clearly that Reinke 's plant is a Myrionema, and we are
grouping two distinct forms under the species which, to us, seem
amply different from the typical form, as described and figured by
Reinke.

12. Myrionema balticum f. californicum S. and G.

Fronds forming microscopic cushions, 0.5-0.75 mm. diam. ; pros-
trate portion composed of relatively straight, regularly radiating fila-
ments; erect filaments unbranched, attenuated slightly upwards and
at the base, blunt, 75-125/x long; true hairs 0.5-1 mm. long; cells
of prostrate filaments 6-7 fi diam., 1-1.5 times as long ; cells of erect
filaments 7.5-10/x diam., mostly quadrate; cells of true hairs 4-5. 5/x
diam,, 6-10 times as long above, 3-5 times as long at the base, quadrate
a few cells above the base ; chromatophores irregularly broken bands ;
zoosporangia unknown ; gametangia on 1-3-celled pedicels, either from
the creeping filaments, or lateral at the base of the erect filaments,
cylindrical, 90-130/i, long, 7-9/1, broad.

Growing on the outer ends of the leaves of Phyllospadix sp. Point
Carmel, Monterey County, California.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 341.

We have no authentic measurements of Ascoeyclus haJticus Reinke.
He does not mention the size of the gametangia either in the Atlas
(1889, pi. 16, figs. 1-4), where he first describes and figures it, or
in the Algenflora (1889a, p. 45) where he describes it again. Foslie
(1894, p. 17, repr.) made the combination Myrionema (PhyeoceJis)
haJticum, but he gave no measurements for any of its parts. De-Toni
(1895, p. 581) lists the plant as Phycocelis (?) baltica (Reinke) Foslie,
but gives no additional information concerning the size of the game-
tangia. Kylin (1907, p. 35) lists a plant under Myrionema 'balticum

466 University of California Publications in Botany [Vol. 8

(Rke.) Fosl. which he found growing on Delesseria sangtdnea at 15 m.
depth. He states that the assimilation filaments are 100-125,/i, long
and 4-6f(, broad, but gives no measurements for the gametangia.

The plant which we have diagnosed is doubtless very closely
related to Eeinke's Ascocyclus halticus. It may be a distinct species,
but we have listed it as a form of halticus until we can obtain more
knowledge of its variation, and more authentic information concern-
ing the type material of halticus. The particular distinguishing
character is the very frequent occurrence of lateral gametangia
arising very close to the base of the vegetative filaments.

13. Myrionema balticum f. pedicellatum S. and G.

Fronds forming microscopic cushions 150-400/i. (up to SOOju,) diam.,
very irregular in outline ; erect filaments short-pilif erous in part,
140-1 60jtt long; true hairs unknown; zoosporangia (?) narrowly
clavate, sessile on the creeping filaments, or terminal on the erect fila-
ments ; gametangia terminal on 10-15-celled pedicels, rarely sessile,
up to ISO/x long.

Growing on the outer ends of the leaves of Phyllospadix sp. Point
Carmel, Monterey County, California.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 341, pi. 33, figs.
11, 12.

This form differs from the species in the absence of hairs, in having
a few very short-piliferous, vegetative filaments, and in having the
gametangia on long pedicels.

In this form, as in many others, the correct interpretation of the
zoosporangia mentioned above in the diagnosis is an open question.
They are filled with dense cell contents and, although none has been
observed to produce zoospores, they appear as though they might
later. Whatever their nature may finally prove to be, they represent
another character in which this form differs from the species. Reinke
neither figures nor describes them in Ascocyclus halticus. They
appear on the same individuals with the gametangia or on different
individuals.

i*^-5J Setchell-Gardner : MeJanophyceae 467

14, Mjnrionema compsonematoides S. and G.
Plate 55, fig. 10

Fronds forming microscopic cushions irregular in outline, 200-400/a
diam., prostrate portion composed of distorted filaments; erect fila-
ments cylindrical, unbranched, 70-80ju, high ; hairs unknown ; cells of
creeping filaments considerably irregular in shape and size, 4-6jli diam. ;
cells of erect filaments cylindrical, 5-7jli diam., quadrate; "ascocysts"
sparse, terminal or at times lateral near the base of the erect filaments ;
gametangia fusiform to clavate, sessile or short-pedicellate, 50-65/x
long, 7-lOja broad ; loculi uniseriate below, biseriate above.

Growing on the blade of Laminaria complanata. Friday Harbor,
Washington.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 343, pi. 34, fig. 10.

Myrionema rompsonematoides seems to be a modification of M.
faecundum. There is a slight sterilization of tissue in that a few of
the erect filaments remain sterile and develop slightly beyond the
gametangia, and in that a part of the gametangia are pedicellate. It
seems to tend toward Compsonema in these respects, and in having
biseriate gametangia in part.

15. Myrionema hecatonematoides S. and G.

Plate 55, fig. 11

Fronds forming thin light brown cushions, more or less circular
in outline, up to 6 mm. diam. ; prostrate portion composed of distorted,
irregularly radiating filaments, mostly monostromatic, in part distro-
matic, producing a few penetrating peg-like rhizoids below ; erect
filaments unbranched, cylindrical, 240-260ju, long ; true hairs absent ;
cells of erect filaments 6.5-7.5/a diam., 2-3 times as long; zoosporangia
unknown ; gametangia sessile or on 1-2-celled pedicels on the creeping
filaments, cylindrical to fusiform, 45-55ju, long, 7-8. 5/a broad ; loculi
mostly uniseriate, biseriate in widest part of the gametangia.

Growing on thepneumatocysts of Nereocystis Liietkeana. Pacific
Grove, California.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 343, pi. 34, fig. 11.

As indicated by the partially distromatic base, the slight tendency
to biseriate loculi in the gametangia, and the sterilization of a part
of the erect filaments, this species of Myrionema seems to be differen-
tiating in the direction of the genus Hecatonema. We are retaining

468 University of California Puhlications in Botany [Vol. 8

it in the genus Myi'ionema. because of the preponderance in number
of uniseriate loculi and monostromatic filaments in the prostrate
portion.

16. Mjrrionema attenuatum S. and G.

Plate 55, figs. 6, 7

Fronds forming microscopic, more or less confluent strata on the
surface of the host ; prostrate portion composed of regularly radiating
filaments; erect filaments unbranched, attenuated gradually upwards
and at the base, 180-220/x long ; true hairs absent ; cells of creeping
filaments, 4.5-5. 5/a diam., nearly quadrate; cells of erect filaments 6-7 [x
diam., variable in length ; zoosporangia unknown ; gametangia sparse,
cylindrical to slightly fusiform, on shorter or longer pedicels, 30-45fi
long, 4.5-6|U, broad.

Growing on the stipes of Macrocystis pyrifera, Laminaria Farlowii,
and on the sterile base of Gigartina radula f. Carmel Bay, Monterey
County, California.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 34<1, pi. 34, figs. 6, 7.

It seems that sterilization has proceeded in this species of
Myrionema until about two-thirds of the erect filaments no longer
produce gametangia, and these filaments are several times longer than
the gametangia. The cells are slightly doliiform in some collections.

17. Myrionema attenuatum f. doliiforme S. and G. (Orthog. mut.)

Plate 55, figs. 4, 5

Fronds forming small, irregular tufts among other epiphytic algae ;
erect filaments up to 275/a long ; cells of erect filaments quadrate to
3 times as long as broad, decidedly doliiform ; gametangia occasion-
ally small and terminal on the long erect filaments, otherwise as the
species.

Growing on the stipe of Macrocystis pyrifera. Carmel Bay,
Monterey County, California.

Myrionema attenuatum f. doUiformum Setchell. and Gardner, Phyc.
Cont. II, 1922, p. 344, pi. 34, figs. 4, 5.

The chief distinction between this form and the species, 31.
attenuatum, is the decided doliiform character of the erect filaments.
Small terminal gametangia appear very frequently on the long, erect,
filaments.

19-i^] ISetchell-Gardncr: Melanophyccae 469

18. Myrionema phyllophilum S. and G.

Fronds forming cushions more or less circular in outline, 400-800/a
diam. ; creeping portion composed of straight filaments radiating from
the center, closely crowded ; erect filaments unbranched, cylindrical,
constricted at the base, llO-lSOfi long, attenuated upwards, piliferous
in part ; true hairs scattered promiscuously among the erect filaments
and gametangia; cells of creeping filaments cylindrical, 4-5|U diam.,
2-4 times as long; cells of erect filaments 8-9/i, diam., quadrate, to 2.5
times as long ; cells of the hairs quadrate at the base, 5-6fi diam., up
to 20 times as long in the upper part ; zoosporangia ( ?) broadly clavate,
sessile or terminal on longer or shorter pedicels, 50-70ju. long, 14-18;u,
broad at the outer end ; gametangia cylindrical, mostly on 1-2-celled
pedicels, on the creeping filaments, or more rarely lateral on the base
of the erect filaments, lilunt, 90-130/x long, 9-11/a broad ; loculi
uniseriate.

Growing on the outer ends of the blades of eel-grass, in the lower
littoral and upper sublittoral belts. Sitka, Alaska.

Setchell and Gardner, Phyc. Cont.- II, 1922, p. 344, pi. 32, figs. 1-5.

In this species about one-third of the erect filaments remain sterile,
and extend beyond the gametangia. The cells in the creeping filaments
are relatively long. The filaments are closely crowded, but distinct
and readily separable. All the erect filaments are abruptly narrowed
at the base. On account of the long creeping cells, the erect filaments
are very much less crowded than is usually the case in most Myrio-
nemas. The erect filaments do not arise successively toward the margin
of the nearly mature plants, cells here and there developing an erect
filament to the length of several cells before the intervening cells show
signs of farther growth. The gametangia are typical of the genus,
with uniseriate loculi and with mostly horizontal, cross cell walls. The
zoosporangia (?) are fairly abundant, and are either with the game-
tangia on the same individual or on separate individuals. We ques-
tion these structures as being functional since none of them has been
observed to produce zoospores. They have the appearance of being
abortive organs, either zoosporangia or gametangia, more likely the
latter, and are probably like the organs observed by Magnus which
induced him to establish the genus Ascocyclus.

470 University of California Publications in Botany [Vol. 8

«

19. Myrionema setiferum S. and G.

Fronds forming small, dense, irregular tufts or cushions, 0.5-0.75
mm. diam. ; prostrate portion composed of regularly radiating, closely
compacted filaments, 4r-5/x diam., 1.5-3 times as long toward the
margin ; true hairs unknown ; erect filaments numerous, unbranched,
gradually attenuated toward the apex, long-pilif erous, 700-800/x long ;
cells of erect filaments cylindrical, 4— 5/a diam., 2-3 times as long;
zoosporangia broadly clavate, 80-90/^ long, 18-24/x broad ; gametangia
cylindrical, 150-200ju, long, 7-8. 5/t broad, sessile, or on short pedicels
on the erect filaments or rarely on the creeping filaments ; loculi
uniseriate.

Growing on the outer ends of the leaves of eel-grass, in company
with other small algae, in the lower littoral belt. Sitka, Alaska.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 345, pi. 32, fig. 8.

Myrionema setiferum is undoubtedly very closely related to M.
phyllophilum on the one hand, and to 31. foecundi(.ni f. calif orniciim
on the other. From the former, it differs in having longer and nar-
rower erect filaments with longer cells and in having much longer
and narrower gametangia which are more frequently borne laterally
on the erect filaments, and often fasciculately branched from a short
pedicel. From the latter, it differs in having functional zoosporangia,
one of the few instances in which we have actually seen the undoubted
zoospores, in having longer and setiferous erect filaments, and longer
and slightly narrower gametangia.

Myrionema globosum (Reinke) Foslie

Myrionema {Phycocelis) glohosum Foslie, New or Crit. Norw.
Algae, 1894, p. 17. Ascocyclus glohosus Reinke, Algenfl. westl. Ostsee,
1889a, p. 46, Atlas, 1889, p. 20, pi. 17. Microspongium glohosum
Reinke, Die brauenen Alg., 1888, p. 20.

Typical M. glohosum, as we understand this plant, has not been
reported from our territorj^ but we have described a forma afflne
which seems very closely related to it.

1925] SetchcU-Gardncr: Melanopliyceae 471

20. Myrionema globosum f. aflfine S. and G.

Fronds microscopic 200-300/x diam., irregular in outline, inter-
mixed with other small algae; prostrate filaments much distorted,
freely branched; erect filaments fasciculately branched, 100-140/i,
long, tapering upwards, pilif erous in part ; true hairs 300-400/x, long ;
cells of prostrate filaments 4-6ja diam., variable in shape ; cells of
erect filaments cylindrical, 4-5ju, diam. ; zoosporangia unknown ; game-
tangia cylindrical, 35-45/x long, 4-4.5/* broad, sessile and lateral on
the erect filaments or rarely terminal ; loculi uniseriate.

Growing on the leaves of eel-grass. Sitka, Alaska, to central Cali-
fornia (Point Carmel, Monterey County).

Setchell and Gardner, Phyc. Cont. II, 1922, p. 342, pi. 33, fig. 16.

We have no measurements in detail for the parts of Myrionema
glohosum, this seemingly distinct species. Reinke (1889^, p. 46) states
that the plants are one-half to one millimeter in diameter. Our plants
seem much smaller, in general. It is difficult to make out the size on
account of their being so intimately intermixed with other small algae.
This is the case in both collections of plants which we have examined,
one from Sitka, Alaska, and the other from Point Carmel, California.

• 21. Myrionema strangulans Grev. {in scnsu strictiore)

Plate 35, fig. 12, and plate 40, fig. 51

Fronds forming very irregularly shaped cushions, 1-2 mm. diam.,
firmly adhering to the surface of the host, without rhizoids ; prostrate
portion composed of regularly radiating filaments with apical growth
and branching, producing dichotomous filaments, strictly monostro-
matic; cells of prostrate filaments 4-4.5/* diam., 1-2 times as long,
cylindrical or slightly doliiform ; hairs, 1-1.5 mm. long, cells of hairs
up to lOO/i long at the apices, 5/t diam., quadrate and with short sheath
at the base ; erect filaments clavate, longest in the center of the frond,
diminishing in length toward the periphery, 55-65/* (up to 110/t)
long, 4/x diam. at the bases, 8/t at the apices; basal cells cylindrical,
terminal cells subspherical ; chromatophores spherical, several in a
cell; unilocular zoosporangia numerous, interspersed among the erect
filaments, sessile or on 1-celled pedicels on the creeping filaments,
ellipsoidal to obovoid, 38-46/i (up to 60/*) long. 18-24/* broad ; game-
tangia unknown.

472 University of California Publications in Botany [Vol.8

Growing on the base of Ulva sp. in the lower littoral belt. Sitka,
Alaska, to central California (Carmel Bay).

Greville, Scot. Crypt. Flora, vol. 5, 1827, pi. 300; Harvey, Phyc.
Brit., vol. 3, 1851, pi. 280; Saunders, Alg. Harriman Exp., 1901,
p. 423, pi. 46, tig-s. 1, 2 ; Setchell and Gardner, Alg. N.W. Amer., 1903,
p. 249; Tilden, Amer. Alg. (Exsicc), no. 350. Not Collins, Holden
and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1795 (a mixture of
several Myrionemataceae).

Myrionema strangulans Greville has been variously interpreted
and the name has frequently been placed as a synonym under 31.
vulgare Thuret. In the attempt to determine the exact nature of
M. strangulans, specimens from Greville 's herbarium, from the western
isles of Scotland (but not from Appin, the technical type locality)
have been obtained (through the kindness of Professor Isaac Bayley
Balfour) and carefully studied. From these the description given
above has been drawn up. We find specimens on Ulva which agree
with these Greville specimens in every detail, except that the zoo-
sporangia in our plants average slightly larger.

22. ]V[3rrionema obscurum S. and G.

Fronds microscopic, distributed more or less irregularly on the
surface of the host ; creeping filaments much branched, very compact
and contorted, forming a pseudoparenchymatous layer ; erect filaments
sparse, unbrauched, 40-70jU. high ; true hairs unknown ; cells of creep-
ing filaments extremely irregular in shape and size, 4.5-10/x diam. ;
cells of erect filaments cylindrical, 6.5-7.5ja diam., quadrate or nearly
so ; zoosporangia very numerous, sessile on the creeping filaments,
ovoid to broadly clavate, 45-65|a long, 18-30^ broad ; gametangia
unknown.

Growing on the blade of Costaria eastata. Moss Beach, San Mateo
County, California.

Setchell and Gardner, Phyc. Cont. II, 1922, p. 346, pi. 32, fig. 6.

Myrionema ohscuruni seems decidedly distinct from all other
species of the genus. It seems to be a near relative of M. strangulans
Grev. The basal layer is very compact, and adheres very firmly to
the host, following closelj^ the irregularities on its surface. We are
unable to make out whether or not the epiphyte causes the death of the
cells of the host, which seems to be quite frequent in all of the plants
we have observed. The creeping filaments push among the dead cells
of the host at times. The zoosporangia are numerous and all sessile.

1925] Setclull-Gardncr: Melanophyccae 473

7. Compsonema Kiu-k.

Fronds forming' small cushions more or less circular in outline,
composed of a prostrate creeping, primary portion, gfiving rise to
numerous erect unbranched or branched filaments, with or without
hairs; prostrate portion monostromatic, composed of filaments quite
regularly radiating from a common center and usually with sub-
terminal branching, closely crowded ; reproduction by unilocular
zoosporangia and plurilocular gametangia with mostly pluriseriate
loculi.

Kuckuck, Beitrajje zur Kennt. Meeresalg., 1899, pp. 56-58 (90-92
inrepr.). pi. VT (12), figs. 6-9.

The type species of Compsonema is C. gracile and the type locality
is Rovigno, on the east coast of the Adriatic Sea, where it was found
growing on stones in water 1-2 meters deep.

The genus Compsonema was established by Kuckuck. We have
accepted the genus to include forms which, like Myrionema, start from
a single cell that, by divisions, soon originates a series of closely
crowded, radiating filaments, forming a monostromatic basal layer
more or less circular in outline ; and which, unlike Myrionema, pro-
duce on erect filaments numerous gametangia that develop pluriseriate
loculi. The genus is very closely related to the genus Myrionema on
one side and to Hecatonema on the other.

Key to the Species

1. Erect sterile filaments branched 2

1. Erect sterile filaments simple 5

2. Creeping filaments with few subulate rhizoids..7. C. fasciculatum (p. 478)

2. Creeping filaments without rhizoids 3

3. Sterile branches short, high up, secund 4

3. Sterile branches long, low down, not secund 13. C. intricatum (p. 482)

4. Erect filaments piliferous in part 17. C. ramulosum (p. 484)

4. Erect filaments not piliferous 12. C. sporangiiferum (p. 481)

5. Gametangia all terminal 6

5. Gametangia terminal and lateral or intercalary 14

6. Gametangia all sessile 7

6. Gametangia pedicellate or in part sessile ; 8

7. All erect filaments completely transformed into gametangia. .2. C. sessile (p. 474)

7. Gametangia intermixed with sterile filaments G. C. tenue (p. 478)

8. Prostrate filaments producing penetrating rhizoids 9

8. Prostrate filaments not producing rhizoids 10

9. Rhizoids abundant, straight, penetrating deeply into the host

11. C. streblonematoides (p. 481)

9. Rhizoids contorted, not penetrating deeply 5. C. pusillum (p. 477)

10. Gametangia very abundant, of uniform height ...4. C. fructuosum (p. 476)
10. Gametangia with pedicels very variable in height 11

474 University of California Puhlications in Bota-ny [Vol.8

11. Gametangia cylindrical or very slightly tapering 12

11. Gametangia decidedly fusiform 13

12. Gametangia mostly on 1-2-celled pedicels, 1. C. myrionematoides (p. 474)
12. Pedicels of gametangia very variable in height.. 3. C. nummuloides (p. 475)

13. Gametangia 24-30^ long, 6-8m broad 18. C. immixtum (p. 487)

13. Gametangia 45-65yu long, 15-18m broad 9. C. conifenun (p. 479)

13. Gametangia 50-75^ long, S-llju broad 8. C. dubium (p. 479)

13. Gametangia 60-130m long, 18-28/^ broad 10. C. serpens (p. 480)

14. Erect filaments in part piliferous 16. C. speciosum f. pilifenim (p. 485)

14. Erect filaments not pihferous 15

15. Terminal gametangia up to 400yu long 14. C. secundum (p. 483)

15. Terminal gametangia up to 110^ long 15. C. secundum f. terminale (p. 484)

1. Compsonema myrionematoides S. and G.

Fronds forming- small, circular to irregular cushions, 1-2.5 mm.
diam., prostrate portion strictly monostromatic, composed of closely
crowded, contorted, radiating filaments ; erect filaments unbranched,
cylindrical, llO-lSO/x. long ; true hairs absent ; cells of creeping fila-
ments cylindrical to irregular in form ; cells of erect filaments cylin-
drical, 7.5-8. 5/x diam., 1-2 times as long, terminal ofte^ several times
as long; chromatophore a slightly broken band in each cell; zoo-
sporangia ( ?) broadly clavate, sessile or on short pedicels on the
creeping filaments, 35-50ju, long, 20-28ja broad ; gametangia numerous,
cylindrical to fusiform, sessile or on short pedicels on the creeping
filaments, 50-7 O/x (up to 100/x) long, 9-12jU, broad.

Growing on the stipe of Nereocystis Luetkeana. Pacific Grove,
California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 361, pi. 36, fig. 1.

There is but little choice as to the generic position of this small
alga. It forms small brown tufts or cushions of loose filaments. Very
frequently the gametangia are wider in the middle than at the ends
and have decidedly biseriate loculi. Because of the more or less loose
tufted character of the erect filaments and the partially biseriate
gametangia we have placed it with the genus Compsonema.

2. Compsonema sessile S. and G.

Fronds forming inconspicuous cushions up to 1.5 mm. diam., cir-
cular to irregular in outline ; creeping filaments contorted, closely
crowded, adhering firmly to the host, without rhizoids; erect filaments
sparse, 20-24/a high; true hairs absent; cells of creeping filaments
irregular in shape and size, 9-12;U. diam. ; chromatophores band-shaped,
one in each cell ; zoosporangia unknown ; gametangia numerous, sessile

1925] Setchcll-Gardncr: Melanophyceae 475

on the creeping filaments, cone-shaped to fusiform, 16-22jli long, 11-14^
broad, many walls of the loculi oblique.

Growing on the lamina of Hedophyllum sessile near the outer end.
Neah Bay, Washington.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 358, pi. 39, fig. 6.

Compsonema sessile represents the extreme of the genus in the
direction of simplicity. There are no hair filaments, and it is doubtful
whether any of the erect filaments remain sterile. At times even the
cells" of the basal layer seem to become a part of the gametangia, at
least more than one cell in the basal layer may be at the base of a
single gametangium. There would seem to be onlj^ a single modifica-
tion of Myrionema of the simplest type to produce this species, viz.,
the change in the type of gametangium from uniseriate to pluri-
seriate. If we hold to the principal distinction between Compsonema
and Myrionema, it wall be necessary to ally this species with the
former. It was found growing in company with Strehlonenia aeci-
dioides f. padficum S. and G., which gave to the host a roughened
appearance, otherwise it probabh^ would not be noticeable without
microscopic examination.

3. Compsonema nummulcides S. and G.

Fronds forming thin circular cushions, 7-12 mm. diam. ; prostrate
portion composed of radiating, much crisped, closely crowded filaments
following closely the irregularities of the host ; erect filaments
unbranched, numerous, 300-400/x long, tapering slightly at both ends ;
true hairs absent ; cells of creeping filaments very irregular in shape
and size ; cells of erect filaments cylindrical, 6-7 fi diam. at the base,
3-8 times as long, 9-llju, diam. in the widest part, quadrate apical cell
6-7 /x diam., 3-8 times as long ; zoosporangia unJvnown ; gametangia
mostly sessile or on short pedicels, some terminal on the erect filaments,
70-90ju, (up to 140/x) long, 11-15/a broad; loculi mostly 2-seriate.

Growing on the pneumatocysts of Nereocystis Luetkeana. Moss
Beach, San Mateo County, California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 359, pi. 35, figs. 5, 6.

Compsonema nummidoides forms thin, light brown cushions on
the pneumatocysts of the host. The whole cushion is a single plant,
circular in outline and attaining a diameter of twelve or more milli-
meters, although the majority of the plants are much smaller. Like
a typical Myrionema the plant starts from a single cell and by divi-
sions forms a series of filaments radiating in all directions with apical

476 University of California Publications in Botany [Vol.8

gTOwtli and dichotomous branching, by the splitting' of the apical cell,
thus forming a continuous monostromatic layer of cells. Beginning
in the center, practically every cell in each radiating filament of the
basal lawyer gives rise successively to an erect filament, about two-thirds
of which are fructiferous, the others remaining sterile. A very large
majority of the gametangia, are sessile or on short pedicels, thus
occupying a zone near the creeping filaments, a myrionematoid char-
acter, but the remainder of the filaments continue to grow and attain
a length of SOO/x to 400/x. Some of these filaments are terminated by
relatively short gametangia. This is one of several species with this
general method of development which has been found growing only
on the above mentioned host, differing from each other and from the
type of the genus, C. gracile Kuckuck, in the size of the plant as a
whole, in details of dimensions of their parts, in the presence or absence
of zoosporangia and hairs, in the position of the gametangia, and in
the relative amount of sterile filaments. It seems to form a fairly
compact group but with overlappings, however, in th^ genera Stre-
hlonema, Myrionema, and Hecatonema.

4. Compsonema fructuosum S. and G.

Fronds forming circular or somewhat irregular cushions, 4-5 mm.
diam. ; prostrate portion composed of tortuous creeping filaments fol-
lowing closely the irregularities of the surface of the host ; erect fila-
ments unbranched, densely crowded, 190-230/x long, cells cylindrical,
8-9ju, diam., 2-4 times as long below, 1-2 times above ; chromatophores
band-shaped ; hairs absent ; zoosporangia sparse, broadly clavate,
terminal on short pedicels from the creeping filaments, 55-65/x (up to
100/x) long, 22-28/A broad; gametangia terminal on erect filaments,
cylindrical to slightly fusiform, blunt, 80-120/;. long, 12-16//, broad;
loculi 2-4-seriate.

Growing on the pneumatocysts of Nereocystis Luetkeana. Tomales
Bay, Marin County, California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 355, ph 36, fig. 2.

Compsonema fructuosum departs from the type of the genus, C.
gracile Kuckuck (1899, p. 90, pi. VI (12), figs. 6-9), in having a differ-
ent host, no hairs, fewer and shorter sterile filaments, more gametangia,
and in having differences in details of measurements. In the abun-
dance and position of the fruit and in the definite circular thallus
with compact monostromatic basal layer of filaments, it approaches

1925] Seirlu ll-G(irdner: MeJanophyceae -ill

very closely to a t^'pical Myrionema, but the multiseriate g-ametangia
represent a stage in development which we have not admitted into
that genus. The gametangia occupy a definite zone on the outer ends
of the erect filaments. Practically all of the erect filaments bear
gametangia.

At times the basal layer seems to be distromatic, or even poly-
stromatic, but careful investigation shows that this appearance is
occasioned by the overlapping creeping filaments which follow the
very irregular surface of the host, due to the death of its surface
layer of cells, probably before the epiphyte took possession. The
zoosporangia are very sparse, but specimens have been seen which show
undoubted zoospores.

5. Compsonema pusillum S. and G,

Plate 54, fig. 3

Fronds forming thin, light brown cushions, irregular in outline,
3-4 mm. diam. ; prostrate portion composed of very tortuous, branched
filaments following closely the irregularities of the host, producing a
few filaments from the under side penetrating among the cells of the
host ; erect filaments simple, cylindrical, 270— 300/x long ; cells of creep-
ing filaments very irregular in shape and size ; cells of erect filaments
cylindrical, 6-7 jx diam., 3-5 times as long; chromatophore a slightly
broken band ; true hairs absent ; zoosporangia spherical to ovoid, sessile
or on short pedicels on the creeping filaments, 18-25/x (up to 40/x)
long, 16-22jU, broad; gametangia cylindrico-conical to blunt fusiform,
sessile or on short pedicels on the creeping filaments, 40-60;a long,
10-12/i, broad ; loculi 1-2-seriate.

Growing on the pneumatocysts of Nereacystis Li(,etkeana. Carmel
Bay, Monterey County, California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 356, pi. 37, fig. 3.

Compsonema pusiUum seems to overlap each of several genera. Its
zoosporangia, gametangia, and penetrating filaments ally it with
Strehlonema. Sauvageau (1897) has figured a Myrionema with
rhizoids. The gametangia are very small and have about as many
uniseriate as biseriate loculi. It might be considered a very minute
creeping Ectocarpus. We have placed it with the genus Compsonema
on account of the scarcity of fruit in proportion to the sterile tissue,
and on account of the monostromatic base, believing that the penetrat-
ing filaments are more or less accidental on account of the injured
condition of the surface layer of cells of the host.

478 University of California. Publications in Botany [Vol. 8

6. Compsonema tenue S. and G.

Plate 54, fig. 6

Fronds microscopic, forming' a more or less continuous stratum on
the host among' other small algae; creeping filaments numerous,
branched, following closely the irregularities of the surface of the
host; erect filaments sparse, unbranched, tapering slightly at both
ends, 80-100/x high ; hair filaments sparse ; cells of creeping filaments
irregular in shape and size; cells of erect filaments cylindrical, not
constricted, 7-8/x diam., 1.5-2.5 times as long; cells of hair filaments
cylindrical, 5.5-6.5//, diam., quadrate or less below, 7-10 times as long
above; zoosporangia (?) clavate, 24-28jU, long, 8-10/x broad, sparse;
gametangia fusiform to irregularly conical, sessile on the creeping
filaments, numerous, 22-28jU, long, 7-llju, broad ; loculi mostly 2-seriate.

Growing on the fruiting fronds of Cystophyllum geminattini. Sitka,
Alaska.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 359, pi. 37, fig. 6.

Compsonema tenue, although scarcely typical of the genus, seems
to possess affinities as closely allied to Compsonema as to any other
genus, and hence we have placed it here for the present.

7. Compsonema fasciculatum S. and G.

Fronds forming microscopic tufts more or less confluent ; creeping
filaments tortuous, adhering to the host without rhizoids; erect fila-
ments branched near the base, 90-130/* long, tapering at the apex and
at the base, not piliferous, constricted at the cross-walls; true hairs
absent ; cells of erect filaments lO-14/i diam. in the widest part, 1-2.5
times as long ; zoosporangia unknown ; gametangia somewhat irregular
in form, conical to fusiform, 35-45/i long, 12-18/i broad, mostly lateral
on short pedicels.

Growing on Gigartina radula f . Pacific Grove, California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 360, pi. 38, figs.
7-9.

This species is allied here rather than with Ectocarpus because
of its extremely small size and its creeping fasciculate habit.

1925] SetchcU-Gardner: Melanophyceae 479

8. Compsonema dubium S. and G.

Fronds diminutive, forming small tufts among other microscopic
algae on the host ; prostrate portion composed of much contorted,
branched, creeping filaments; erect filaments unbranched, almost
cylindrical throughout, 275-350,;u, long, not piliferous; true hairs
absent ; cells of creeping filaments very irregular in form and size ;
cells of erect filaments cylindrical, l-^jx diam., slightly less at the base
and the apex of the filament, 2-3 times as long; zoosporangia (?)
ellipsoidal, terminal on short pedicels, 16-22ja long, 12-15fi broad ;
gametangia cylindrical to slightly fusiform, terminal on longer or
shorter pedicels from the creeping filaments, rarely terminal on the
long erect filaments, 50-75/a long, 8-llju, broad ; loculi mostly 2-seriate.

Growing on the sterile base of Gigartina ra-dula f . Cypress Point,
Monterey County, California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 365, pi. 38, fig. 6.

Compsonema duhium, like C. coniferum and C. serpens, has close
affinities with small members of the genus Ectocarpus. Its relatively
long and narrow gametangia distinguish it from the two species here
mentioned with which it is frequently associated.

9. Compsonema coniferum S. and G.

Fronds microscopic, more or less confluent, forming a velvety
stratum on the host among other minute algae ; prostrate portion pro-
fusely branched, much contorted ; erect filaments numerous, simple,
slightly attenuated at both ends, not piliferous, 150-200/x high ; true
hairs absent ; cells of creeping filaments irregular in shape and size,
6-7.5/1. diam.. 9-12/x, long; cells of erect filaments cylindrical to slightly
doliiform. 8-10/x, diam. in widest part, 1-1.5 times as long ; chromato-
phores band-shaped ; zoosporangia unknown ; gametangia sessile or on
short pedicels on the creeping filaments, narrowly conical, 45-65/i, long,
15-18/t broad.

Growing on the sterile base of Gigartina radida f. Cypress Point,
Monterey County, California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 365, pi. 38, fig. 3.

Compsonema coniferum seems to be a very close relative of C.
fasciculatum, found growing on the same host. The fronds are more
extensive, unbranched, and have larger gametangia. We consider this
species of Compsonema a very near approach to an Ectocarpus of

480 University .of California PuhJications in Botany [Vol. 8

extremely small size. We are inclined toward the genus Compsonema
as a more appropriate genus for this species than Ectocarpus on
account of its extensive creeping habit and prolific production from
the creeping filaments of nearly sessile gametangia and many sterile
erect filaments. It is to be found quite commonly associated on the
same host with several other species of Myrionemataceae.

10. Compsonema serpens S. and G.

Fronds forming a more or less continuous expanded stratum on
the surface of the host ; creeping portion composed of much contorted,
irregularly branched filaments; erect filaments unbranehed, or rarely
with a few subulate branches above, tapering slightly at the base,
gradually attenuated in the upper part, not pilif erous, 375-425/i, long ;
true hairs absent ; cells of creeping filaments cylindrical to irregular,
10-13ju, diam., 0.5-3 times as long; cells of erect filaments cylindrical,
not constricted at the dissepiments, 5.5-8.5/* diam. at the base, 10-17/x
diam. in the widest part, 1.25-2.5 times as long ; chromatophores band-
shaped, densely crowded in the young cells, more numerous and
separated in the old cells; zoosporangia unknoAvn; gametangia cylin-
drico-conical, terminal on shorter or longer pedicels from the creeping
filaments, rarely terminal on the long filaments, 60-130//, long, 18-28/x
broad.

Growing on the sterile base of Gigartina radula f ., in company with
several other small algae. Cypress Point, Monterey County, California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 363, pi. 39, fig. 7.

This diminutive member of the Melanophyceae is on the border
line between Compsonema. and Ectocarpus. Our comprehension of
these two genera, so far as the vegetative portion is concerned, is that
a typical Compsonema, starting from a single cell, develops an exten-
sive, creeping, attaching mass of filaments, radiating in all directions
from the origin, which later produces very numerous erect filaments,
some of which remain sterile and others become fructiferous ; and
that a typical Ectocarpus, starting likewise from a single cell, develops
a rather insignificant series of attaching filaments, more or less
rhizoidal in nature, and a relatively extensive system of erect filaments
finally producing the fruit. Starting with this conception concerning
the two genera, we are assuming that diflt'erentiation has proceeded
from Compsonema in the direction of the reduction of the creeping
portion, to a more extensive development of the erect portion, finally
resulting in an Ectocarpus.

1925] Setchcll-Gardner: Melanophyceae 481

The species under consideration has a relatively extensive system of
creeping filaments but very much less so than that of a typical Comp-
sonema. The creeping filaments do not form a solid disk, but those
from different plants are so intertwined that it is impossible to deter-
mine the limits of a distinct individual. In this character it resembles
an Ectocarpus. The gametangia are like those of a typical Ectocarpus,
but since they are mostly short-pedicellate and spring directly from
the creeping filaments, as is the case in a typical Compsonema, and
because of the relatively extensive system of creeping filaments, we
have placed it in the latter genus.

11. Compsonema streblonematoides S. and G.

Fronds forming thin, circular cushions, 3-5 mm, diam. ; prostrate
portion composed of filaments following closely the irregularities of
the host, giving rise below to numerous rhizoidal filaments penetrating
to a depth of 300-350jU, among the cells of the host, and above to sterile
•filaments and gametangia; erect filaments 140-160/x long; cells of
erect filaments cylindrical, 8.5-10/x diam., 1.5-3 times as long ; zoo-
sporangia unknown; gametangia mostly cylindrical, short-pedicellate,
55-70ju, long, ll-14ju, broad ; loculi mostly 2-seriate.

Growing on the pneumatocysts of NereocysUs Luetkeana. Mouth
of Tomales Bay, Marin County, California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 353, pi. 35, fig. 4.

Compsonema streblonematokles, like a Strehlonema, has a large
part of the vegetative portion of the frond within the host-plant, the
penetrating filaments extending into the host tM'o or three times as far
as the length of the external vegetative filaments. The whole frond,
however, is the result of the growth of a single cell and forms a com-
pact monostromatic basal disk. In the character of the gametangia it
is close to a Myrionema, but in a very large majority of cases the
loculi are biseriate.

12. Compsonema sporangiiferum S. and G.

Fronds forming circular cushions, 1-4 mm. in diameter ; prostrate
portion composed of compact, irregular, more or less contorted fila-
ments following the irregularities of the surface closely, monostro-
matic or in part distromatic ; erect filaments 300-700/x. long, un-
branched or bearing above a few short, mostly secund, fructiferous
ramuli ; cells of the erect filaments 6-8ja diam. at the base, and 2-3

482 University of California Publications in Botany [yoh. 8

times as long, 11-13/* diam. at the apex, and 1-1.5 times as long, many
cells in the upper part of the filaments dividing 1-2 times longitudin-
ally ; chromatophores band-shaped, few in a cell ; zoosporangia very
abundant, very variable in shape and size, narrowly ellipsoidal, clavate
to broadly ovoid, 40-130ju, long, 20-35/x wide, sessile or on short pedicels
from the prostrate filaments, or lateral or terminal on the erect fila-
ments ; gametangia rare, cylindrical, sessile or mostly short-pedicellate
on the creeping filaments, rarely terminal and tuberculate on the erect
filaments, 80-130ja long, 11-15/* broad.

Growing on the pneumatocyst of Nereo'cystis Luetkeana. Neah
Bay, Washington.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 357, pi. 36, figs. 3-8.

The unusual condition in Compsonema prevails in C. sporangi-
iferum. The gametangia seem to be of rare occurrence. Examination
of specimens of different sizes from different parts of the host revealed
the presence of a large number of zoosporangia in all stages of develop-
ment, many being empty. The gametangia are prevailingly near the
base, although occasionally a complex one appears terminal on an
erect filament. A very common character is the division of the cells
in the upper parts of the filaments into 2-4 parts by longitudinal walls,
producing a slight clavate condition in such filaments. We do not
know, at present, the morphological significance of this condition. The
zoosporangia and gametangia, so far as observed, develop on distinct
individuals.

13. Compsonema intricatum S. and G.

Fronds 1.5-2.5 mm. high, more or less confluent, forming a con-
tinuous velvety stratum of indefinite expansion; creeping filaments
numerous, profusely branched, crooked; erect filaments numerous,
unbranehed or with a few short branches near the base, straight, grad-
ually attenuated at the apices, pilif erous in part ; true hairs absent ;
cells of creeping filaments irregular in shape and size ; cells of erect
filaments cylindrical, 8-9/* diam., 1-2.5 times as long, slightly narrower
at the base of the filaments ; chromatophores band-shaped ; zoo-
sporangia ovoid to ellipsoidal, terminal on short filaments from the
creeping filaments or lateral and sessile or pedicellate on the erect
filaments near the base, 25-33/x long, 18-22/* broad ; gametangia ter-
minal on short erect filaments from the creeping filaments, lateral and
sessile or pedicellate near the base of the erect filaments or intercalary
in the main erect filaments, 80-120/* (up to 175/*) long, lO-14/i broad ;
terminal and intercalary ones up to 600./* long ; loculi mostly 2-seriate.

1^-5] Setclu'U-Gardncr: MeJanophyceae 483

Growing' on Fucus furcatus f. Carmel Bay, Monterey County,
California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 354, pi. 35, figs. 1-3.

Compsonema intricatum has diverged very far from a typical
Compsonema. The size of a single individual at maturity cannot be
ascertained, since the creeping filaments, and to some extent the erect
filaments, are thoroughly and inextricably intertwined, forming con-
tinuous velvety strata on the surface of the host for several inches in
expanse. The gametangia have, to a considerable degree, moved to
the lateral position, the majority, however, remain terminal on long
pedicels from the basal filaments. The lateral position is characteris-
tic of Ectocarpus. The intercalary gametangia are typical of the
genus Pylaiella. The creeping filaments are very numerous, although
they do not radiate regularly side by side, thus forming a regular
circular disk typical of the genus Myrionema. The zoosporangia ( ?)
are numerous and develop with the gametangia on the same individual.
It is exceedingly doubtful whether or not these structures are func-
tional. At the stage in which the material was found, they were prac-
ticallj^ all of the same size, and some of the accompanying gametangia
were empty. There is no indication of the formation of zoospores.
They are filled with what seem to be densely crowded, angular chro-
matophores. The apical end is slightly beaked, and the outer wall
dissolved, except a thin inner membrane. Their position on the same
plants with gametangia, the peculiar open beak, and the fact that they
all seem to be of the same age lead us to suspect that they are some
sort of abnormalities, or pathological conditions of the gametangia,
which will require much more investigation to interpret. We have
retained this in the genus Compsonema on account of the prevailing
simplicity of the erect filaments and the abundance of creeping fila-
ments, practically every cell of which gives rise to an erect filament.

14, Compsonema secundum S. and G,

Plate 54, figs. 1, 2

Fronds very small, usually more or less confluent ; prostrate por-
tion monostromatic, composed of much crisped, branched filaments;
cells of creeping filaments irregular in shape, 7-8ju, diam,, variable in
length ; erect filaments up to 1.25 mm. long, unbranched, or rarely with
a few short fructiferous ramuli; cells of erect filaments cylindrical,
5.5-6.5ju, diam. at the base, 4.5-6 times as long, 9,5-10,5/x, diam,, in

484 University of California Puhlications in Botany [Vol.8

widest part, terminal cells up to 9 times as long ; chromatophores band-
shaped ; zoosporangia broadly clavate, sessile or on short pedicels from
the creeping filaments, 60-90/x long, 22-28;a broad ; gametangia very
variable, rarely on short pedicels on the creeping filaments, at times
formed by transforming the upper portion of a long erect filament,
at other times by developing lateral secund projections from part or
all of the groups of loculi in the long, terminal gametangia, up to
400/x long, 11-15/x broad, blunt, clavate on the ends of the long
filaments.

Growing on the pneumatocyst of Nereocysiis Luetkeana. West
coast of Washington (Moclips) to central California (Carmel Bay).

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 361, pi. 37, figs. 1, 2.

Like Compsonema ramulosuni, C. secundum has gametangia occupy-
ing a variety of positions and assuming a diversity of forms. The
form of gametangium which marks the species is the long terminal
form which develops numerous, short, blunt-conical, seriate, secund,
sessile gametangia, thus forming a complex composed of these game-
tangia and the metamorphosed cells in the upper part of the filament.
The end of the filament becomes reflexed, at times almost scorpioid.
The specimens from Moclips, Washington, were taken in May and
the gametangia were practically all empty, ^%e other parts of the
plants manifested signs of old age conditions. They seem to belong
to this species, but the filaments have a considerably greater diameter.
Typical zoosporangia with well-formed zoospores have been observed
in this collection.

15. Compsonema secundum f. terminale S. and G.

Plate 54, figs. 4, 5

Fronds forming a continuous velvety stratum of indefinite expanse,
attached by very much distorted, branched, creeping filaments with
numerous, penetrating, subulate rhizoids from the under surface ;
erect filaments 700-800/x (up to 1 mm.) long, unbranched, or produc-
ing a few short secund fructiferous ramuli, straight, slightly tapering
toward the base, cylindrical above ; cells of the creeping and rhizoidal
portions irregular, variable in size ; cells of the erect filaments
■cylindrical, not constricted, 6-8/i. cliam. at the base of the filaments,
2-3.5 times as long, 9-12/x diam. above, 0.5-2 times as long-; chromato-
phores irregular bands, few in a cell; zoosporangia ( ?) broadly clavate,
sessile or on short pedicels from the creeping filaments, 50-60^ long,

l''--5] Setchell-Gardncr: Melanophj/ceae 485

18-22ja broad, or terminal and seriate with lateral seeund prolonga-
tions; gametangia sessile near the outer ends of the erect filaments,
or on short pedicels from the creeping filaments, 90-110/a long, 15-18ju,
broad, nearly cylindrical, or terminal on the erect filaments, clavate,
often with lateral, tiiberculate, seeund prolongations, variable in
size.

Growing on the pneumatocysts of Nereocystis Luetkeana. Central
California (San Francisco and Pacific Grove).

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 366, pi. 37, figs. 4, 5.

This form of Compsonema is to be distinguished by the great
predominance of relatively short terminal gametangia, and many
relatively large, lateral, seeund, sessile gametangia which are curved
upwards. There are a few sessile or short pedicellate examples at the
base and a few lateral forms on short ramuli. Specimens taken in
December had the larger number of gametangia empty.

16. Compsonema speciosum f. piliferum S. and G.

Fronds forming cushions, more or less circular in outline, 5-8 mm.
diam. ; creeping filaments very irregularly and profusely branched,
following closely the irregularities on the surface of the host, at times
short branches penetrating among the injured cells of the host ; erect
filaments numerous, unbranched or bearing short, seeund, fructiferous
branches or an occasional piliferous branch, 500-800ja long, tapering
at the base, piliferous above ; cells of creeping filaments very irregular
in shape and size; cells of erect filaments cylindrical, 6-7. S/a diam. at
the base, 9-12/* diam. in the widest part, 1-3 times as long; cells of
hairs 5-6.5/1 diam., up to 15 times as long; chromatophores band-
shaped, few in a cell ; zoosporangia obovoid, 55-65/(, long, 28-32/x broad,
rare ; gametangia sessile or on shorter or longer pedicels from the
creeping filaments or lateral, seeund, sessile or pedicellate on the erect
filaments or, more rarely, intercalary ; terminal gametangia 60-100/a
long, 14-18/t broad, seeund gametangia variable, much smaller.

Growing on the pneumatocysts of Nereocystis Luetkeana. Moclips,
Washington.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 356, pi. 38, figs. 1, 2.

Compsonema speciosum f. piliferum seems undoubtedly to be very
closely related to Compsonema speciosum (Borg.) S. and G. comb, nov.,
(Myrionema speciosum Borgesen, 1902, pp. 421-424, which is the
Hecatonema speciosum (Borgesen) Cotton, 1912, p. 121 and the

■iS6 Vniversiii) of California Puhlications in Botany [Vol.8

Hecatonema cliff usuni Kylin cf. Cotton, Joe. cit.), at least the two
sets of plants agree very closely in general habit and structure as
described by Borgesen. Judging alone from the description and
figures the chief differences are that our plant has short penetrating
rhizoids, many terminal and no lateral hairs, and larger gametangia.
"We are unable to say that the plant would produce rhizoids on an
uninjured host. The surface cells of the host are destroyed under the
epiphyte, and it is an open question whether the penetrating rhizoids
are or are not the cause of their death. Borgesen does not give the
dimensions of the whole plant, but he states that the plants form short
dense mats on the conceptacles of Himanthalia lorea, which would
indicate that they were quite small. Our plants form circular cushions
or mats 5-8 mm. in diameter. Typical zoosporangia were found at
the base of the erect filaments. The lateral secund gametangia, stand-
ing nearly perpendicular to the filaments, the cells of which become
the basal part of the gametangia, is a character not found in Comp-
sonema gracile Kuckuck.

17. Compsonema ramulosum S. and G.

Fronds forming circular cushions 5-7 mm. diam. ; prostrate por-
tion monostromatic, composed of more or less contorted and irregularly
branched radiating filaments; erect filaments piliferous in part,
unbranched or bearing above short, mostly fructiferous and secund
branches and occasional hairs, tapering slightly at the base ; cells of
creeping filaments 6-7/a diam., quadrate or slightly longer than the
diameter ; cells of erect filaments 6-7. 5/x diam. at the base, slightly
broader above, 1-3.5 times as long; cells of the branches and hairs
4^6,/^ diam. ; chromatophores band-shaped, more or less broken and
angular; zoosporangia (?) broadly clavate, terminal on longer or
shorter pedicels, lateral and secund or whorled near the ends of the
erect filaments; gametangia exceedingly variable in shape, size and
position, sessile or pedicellate on the creeping filaments or lateral on
the erect filaments or intercalary or terminal either on short lateral
ramuli or on the main filaments, clavate, cylindrical to cylindrico-
conical, up to 140/x long, 16ja broad, loculi mostly 2-seriate.

Growing on the pneumatocysts of Nereocystis Lioetkeana. Carmel
Bay, Monterey County, California.

Setchell and Gardner, Phyc. Cont. Ill, 1922, p. 362, pi. 39, figs. 1-5.

There exists in this species extreme variation in the form, size, and
position of the gametangia. Some are sessile on the creeping filaments,

19-5] SeichflJ-Gardncr: Melanophticeae 487

thoiigrh they are more frequently to be found on the pedicels up to
fifteen cells long;. At times sessile forms are to be found on the erect
filaments about t^vo-thirds of the way up to the apex, but more gen-
erally they are terminal on small lateral rarauli on the erect filaments
arising from the middle to near the apices. Rarely there are small
lateral forms on the ramuli. Finally, they may rarely lie terminal or
intercalary' and seriate on the erect filaments although the erect fila-
ments are usually piliferous. The terminal forms on pedicels, and
particularly those on the main erect filaments, are often quite blunt,
even clavate. Others on the prostrate filaments are narrow and
sharply attenuated, at times terminated by a sterile pointed filament,
in effect intercalary. Many of the gametangia, terminal on the ramuli,
are composed of a single series of loculi. Most of the others are two
or more seriate. The hair filaments do not seem to be abundant. They
arise laterally on the main filaments, usually above the center. The
meristem in these hairs is at the base which is surrounded by a sheath
similar to that described by Sauvageau (1898, p. 47) for Myrionema.
The ramuli are decidedly curved upwards. The species is similar to
Myrionema speciasum from the Faeroes, described by Borgesen (1902,
p. 421). Our plant is more slender in all of its parts, has much longer
gametangia, and possesses, in addition to the sessile, secund gametangia
figured by Borgesen, occasional intercalary ones and many terminal
ones on the numerous clustered ramuli. These ramuli are in turn
occasionally branched, usually arise in groups, sometimes whorled, two
or three arising from the same cell, in other instances they are secund.
The species often grows in association with other members of the
Myrionemataceae and the Ectocarpaceae. As a rule, the main erect
filaments do not enter into the formation of gametangia, but give rise
either to sessile gametangia or to the ramuli. No zoospores have been
observed in the so-called zoosporangia. We suspect that these may
represent pathological conditions.

18. Compsonema immixtiim S. and G.

Thallus inconspicuous, the basal filaments creeping among the
gametangia of the host ; erect filaments very numerous, all bearing
gametangia ; hairs and zoosporangia unknown ; gametangia narrowly
ellipsoidal, 24-30/a long.

Growing on Colpomenia sinuosa f. deformans. Isla Partida, Gulf
of California.

488 Vniversity of California PuMications in Botany [Vol. 8

Setchell and Gardner, Mar. Al^. Gulf, of Calif., 1924, p. 724, pi. 17,
fig. 49.

This organism is on the border between Myrionema and Comp-
sonema as Ave interpret these two genera (Setchell and Gardner, Phyc.
Cont. II-VI, 1922). Under the present condition of the host it is
impossible to learn much of the basal portion. It appears that the
basal layer was largely developed before the gametangia of the host
started to develop. Later the latter pushed out among the basal fila-
ments and leaving them behind developed along with the gametangia of
the epiphyte. The gametangia project beyond the general surface of
the host nearly their entire length. On account of the pluriseriate
nature of the gametangia, we have placed it in the genus Compsonema
rather than in Myrionema.

8. Hecatonema Sauv.

Fronds starting from a single cell, soon developing a series of
closely crowded filaments radiating in all directions, forming a com-
pact disk; the branching of radiating filaments often subterminal;
the cells of the radiating filaments wholly or in part dividing once hori-
zontally, thus creating a distromatic layer, the lower cells of which
may or may not produce short, penetrating, subulate rhizoids; the
upper layer of cells producing erect assimilating filaments, wholly
or in part producing gametangia, and hyaline hairs ; gametangia
pluriseriate.

Sauvageau, Sur quelq. Myrionem., 1897, p. 248 (p. 88, Repr.).

The type species of the genus is H. maculans, collected at Cher-
bourg, France, growing on Rhodymenia palmata Grev. The specific
name wa.s taken from Phycocelis maculans Collins (Bull. Tor. Bot.
Club., vol. 23, 1896, p. 459), the plants of which were distributed
in Collins, Holden and Setchell's Phycotheca Boreali-Americana
(Exsicc), no. 274.

Key to the Species

1. All of the erect filaments transformed into gametangia ...1. H. Lawsonii (p. 489)

1. Part of the erect filaments transformed into gametangia 2

2. Erect sterile filaments clavate 3. H. clavatum (p. 491)

2. Erect sterile filaments cylindrical 2. H. variabile (p. 490)

1925] Setchcll-Gardnrr: Melanophyceae 489

1. Hecatonema Lawsonii S. and G.

Plate 53, figs. 5-7

Fronds formings microscopic cushions, 250-500/i. diam., irregular
in outline ; prostrate portion composed of very much contorted, pro-
fusely branched filaments with subterminal forking; erect filaments
forming a compact, palisade-like stratum interspersed with numerous
hair filaments, 0.75-1.5 mm. long; cells of creeping filaments dividing
radially as well as tangentially, forming a pseudoparenchymatous
tissue in the center of the thallus, irregular in shape, 4-5/a diam. ; cells
of the hair filaments quadrate at the base and surrounded by a sheath,
up to 25 times as long as the diameter above ; zoosporangia unknown ;
gametangia cylindrical to slightly fusiform, 25-80/* long, 5-6.5/* broad ;
many loculi formed by longitudinal and by oblique walls.

Growing on the sporophylls of Nereocystis Luetkeana. IJyak Bay,
Alaska.

Setchell and Gardner, Phyc. Cont. IV, 1922, p. 379, pi. 40, figs. 5-7.
Myrioncmo vnJgare Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. 924 (not of Thuret). Myrionema strangidans Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 249 (not of Greville).

This plant superficially resembles \evy closely Myrionema foecun-
ditm f. simpJicissimuni S. and G. Microscopic examination, however,
reveals several important differences. The creeping filaments are
unique. The branches are very numerous, come off at wide angles and
seem, for the mast part at least, to be subterminal, or if the terminal
cell splits, one of the dichotomy very frequently fails to develop till
much later. Many of the cells in the center, and even toward the
periphery, divide radially and perpendicular to the surface of the
host, forming a pseudoparenchymatous layer. Thus the species,
strictly speaking, cannot be said to be distromatic, since the distro-
matic condition of a frond is really brought about by radial divisions
of the cells of the filaments, but parallel to the surface of the host.
The character of the gametangia and of the creeping filaments will
not permit of its being placed with Myrionema vnlgare Thuret as
further described and figured by Sauvageau (1897, p. 186, et seq.).
On account of the radial divisions of the creeping filaments and the
prevailing biseriate condition of most of the gametangia, we have
placed this species with the genus Hecatonema.

490 University of Calif omia Puhlications in Botany [Vol. 8

2. Hecatonema variabile S. and G.

Fronds forming circular cushions, 4-7 mm. diam. ; prostrate por-
tion composed of regularly radiating filaments with subterminal
branching, attached by numerous short rhizoids, 1-3 cells long ; erect
filaments not densely crowded, not arising successively from the center
toward the periphery, unbrauched, 400-500ju, long ; hairs unknown ;
cells of creeping filaments irregular in form, T-Oju, diam., 1-3 times as
long as at the margin ; cells of erect filaments 4.5-5jU, diam. at the base
and 2-5 times as long, 8-10/x diam. and 1-3 times as long toward the
distal end ; zoosporangia ( ? ) broadly clavate, sessile or on short
pedicels on the prostrate filaments, rarely lateral or terminal on the
erect filaments, 50-65jti long, 20-24jti broad ; gametangia cylindrical,
tapering slightly at the apices, sessile or on short pedicels on the
prostrate filaments or rarely terminal on the erect filaments, 70-120,ti
long, 9-12/x broad ; loculi 1-2-seriate.

Growing on the pneumatocysts of Nereocystis Luetheana. Carmel
Bay, Monterey County, California.

Setchell and Gardner, Phyc. Cont. IV, 1922, p. 377, pi. 41, figs.
1-12.

From the standpoint of the structure of the gametangia, the type
of this species of Hecatonema^ could scarcely be separated from a
Myrionema such as M. phyllophilum S. and G. They, in part, possess
uniseriate loculi, but have many perpendicular and slanting walls,
making a partial biseriate condition. The basal layer, particularly in
the center of the thallus, is distinctly distromatic, developing numerous
peg-like rhizoids, which serve to anchor it more firmly to the host, and,
for the most part at least, the branching of the filaments is subterminal,
whereas that of a true Myrionema' is terminal and brought about by
the splitting of the apical cell.

The size of the plants as a whole varies considerably. The material
of the type varies in this respect from 4 mm. to 7 mm. in diameter.
In some collections of specimens which we have included under the
species, the specimens are as small as 2 mm. in diameter, in others
they are up to 10 mm. in diameter. The length of the erect filaments
A^aries from 150/x to 500/^. Mature gametangia vary from 55jj. to
120/A long. The relative number of gametangia terminal on the
erect filaments, as well as their size and complexity, is exceedingly
variable in difi'erent collections. Two or three cells only may meta-
morphose in some filaments, in others eight or ten. Occasionally small

1^25] Setchdl-Gardner: Melanophyceae 491

lateral outgrowths from the metamorphosed cells may occur. The
number of erect filaments in proportion to the number of gametangia
is a character which seems to be very unstable. In some collections
they exceed the number of gametangia almost two to one, in others
they do not occur more than in proportion of one to five. They are
verj'^ irregular in origin along the radiating creeping filaments. Some
near the margin of the frond may be nearing maturity before others
near the center of the frond have started.

3. Hecatonema clavatum S. and G.

Plate 53, figs. 1-4

Fronds forming circular cushions 2-3 mm. diam., with relatively
wide margin free from erect filaments; prostrate portion attached
by numerous, short, subulate rhizoids, and composed of regularly
radiating, closely crowded filaments; erect filaments mostly clavate,
unbranched, 190-210/i, long; cells of creeping filaments 5-6ju, diam.,
1.5-2 times as long; cells of erect filaments cylindrical below and
5.5-6.5/A diam., 2-2.5 times as long, slightly constricted at the dissepi-
ments above and 8-11/i, diam., 1.5 times as long; zoosporangia (?)
sessile or subsessile on creeping filaments, clavate, 60-80ju, long, 10-14|U,
broad ; gametangia cylindrical to slightly fusiform, sessile or on short
pedicels on the creeping filaments, 55-85/a (up to 111|U,) long, 8-12/x
broad ; loculi mostly 2-seriate.

Growing on the pneumatocysts of Nereocystk Luetkeana. Mouth
of Tomales Bay, Marin County, California.

Setchell and Gardner, Phyc. Cont. IV, 1922, p. 378, pi. 40, figs. 1-4.

The structures designated zoosporangia (?) in the foregoing diag-
nosis of this species were found interspersed among the gametangia.
No indication of the formation of zoospores has been observed, which
leads to the suspicion that they may be abortive gametangia. The
terminal cell of the erect filaments is often longer than the cells
below it, and in these filaments in which the cells are divided length-
wise, producing the clavate condition, the terminal cell degenerates in
a similar manner as do the hairs in other species. The longitudinal
division of cells in the upper parts of the filaments is undoubtedly
connected with the formation of the short, lateral, often secund, game-
tangia characteristic of nearly related species in Hecatonema and
in Compsonema, for in a few instainces such gametangia were observed
in old specimens in which the gametes had, to a large degree, escaped
from the characteristic gametangia.

492 University of California Publications in Botany [Vol. 8

9. Microspongium Reinke

Fronds forming small, cushion-like, rounded, gelatinous masses of
various shapes and sizes, composed of a distromatic basal layer the
upper cells of which give rise to erect, compact, cylindrical filaments
branched or unbranched, among which are intermixed a few hyaline
hairs; reproduction by zoospores formed in single (or in seriate?)
zoosporangia and by gametes formed in plurilocular gametangia.

Reinke, Die braunen Algen der Kieler Bucht, 1888, pp. 16 and 20,
Atlas, 1889, p. 11, pis. 7, 8, Algenfl. westl. Ostsee, 1889a:, p. 46.
Hapalospongidion Saunders, New and little known brown Alg., 1899,
p. 37.

The type species is Microspongium gelaiinosum Reinke. In our
judgment there are no fundamental differences between Microspong-
ium Reinke (1888, p. 20) and Hapalospongidion Saunders (1899, p.
37), consequently we are adopting the earlier generic name. Both
genera form diminutive gelatinous cushions and have a distromatic
basal disk which gives rise to erect "hairs" and filaments bearing chro-
matophores. Unilocular zoosporangia and plurilocular gametangia are
present in both, although there are specific differences. In the first
place, the erect filaments in Hapalospongidion gelaiinosum are simple.
In the second place, the suspected zoosporangia are seriate and inter-
calary, probably always so since we doubt Saunder 's statement of their
being terminal (in complete filaments). In the third place, the game-
tangia are pluriseriate and always intercalary, although there may be,
at times, only one sterile, enlarged cell with dense contents at the
very apex of the gametangial filament. The arrangement and char-
acter of both zoosporangia (if normal?) and of the gametangia
resemble those of Pylaiella, but we find similar structures occasional
among the Myrionemataceae, but generally on plants showing the
terminal types of gametangia. It is for these reasons that it seems
best to merge Hapalospongidion in Microspongium.

Microspong'ium Saundersii S. and G.

Erect filaments 250-750/* long, embedded in a gelatinous matrix ;
cells ii;i the lower part of the filament doliiform, 8-10/* long, 4-5ft
broad, in the center of the filament cylindrical, 24-36/i long, 5-6/*
broad, in the distal end, 4-5/* long, 7-1 0/x broad ; hairs scattered, one
or more millimeters long ; chromatophores discoid ; zoosporangia (?)

1925] Setchcll-Gardner : Melanophyceae 493

terminal and seriate, ellipsoidal; gametangia terminal, with loculi
1-2-seriate, 10-14|U, broad.

Growing on rocks, forming small, irregular, gelatinous or sponge-
like cushions, separate at first, but later many coalescing into a common
mass. Pacific Grove, California.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 12. Hapalo-
spongidion gelatiTWsum Saunders, New and little known brown alg.,
1899, p. 37, pi. 1; Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. 534.

Microspongium Saundersii differs from M. gelatinosum Reinke
in having the erect filaments unbranched, the zoosporangia ( 1) seriate
instead of single, the gametangia with loculi 1-2-seriate and inter-
calarv instead of lateral and uniseriate, and in details of measurements.

FAMILY 4. EALFSIACEAE k.jellman

Thallns crustaceous, flat, discoid and rounded or crenate at the
margins, or deeply lobed and dissected, consisting of a flattened mono-
stromatic, discoid, basal layer with marginal growth, soon becoming
few to many layered by horizontal partitions, with hairs, single, or
in small groups, in depressions of the upper surface, with or without
rhizoids from the lower surface ; unilocular zoosporangia, either borne
at the base of more or less free multicellular paraphyses, collected into
more or less distinct patches or "sori" or terminal and forming naked
' ' sori ' ' ; gametangia uniseriate or pluriseriate, terminal or borne on
lateral branchlets near the tips of the more or less coalescent vertical
rows of the thallus.

Kjellman, Ilandbok Skand. Ilafsalg., I, 1890, p. 29 (lim. mut.) ;
Stragulariaceae Stroemfelt, Om Algenveg. vid Islands Kuster, 1886,
p. 49. Lithodermataceae Kjellman, loc. cit., 1890, p. 17.

The Ralfsiaceae, in the somewhat extended sense in which we con-
sider the application of the name, i.e., as including the Lithoderma-
taceae of Kjellman, is closely related to the Myrionemataceae (where
Reinke places Ralfsia and its segregates) and the Elachisteaceae, with-
out being conveniently placed under either. The frond seems clearly
a further development of that of the simpler Mtjrionemaspecies,
retaining the flattened circular habit as a fundamental type. The
coalescence of the vertical filaments, the greater or less separation of
fertile areas from sterile, the position of the sporangia and gametangia
as to whether terminal or lateral, find close resemblances among the
Myrionemataceae or may readily be considered as progressive develop-

494 University of California Puhlications in Botany [Vol. 8

ments from them. It does not seem desirable to separate the Litho-
derma series of genera and species from the Ralfsia series since both
may be considered as derived from the Myrionemataceae, the sporangia
being transformed vertical filaments in the Ralfsia series and only the
transformed terminal cells in the Lithoderma series, while the game-
tangia may be either terminal or on short lateral branches in the
Lithoderma series, but terminal in the Ralfsia series, so far as known.

Key to the Genera

1. Zoosporangia included, borne at the bases of the paraphyses 2

1. Zoosporangia superficial, terminal on the erect filaments. .12. Lithoderma (p. 500)

2. Thallus entire, or with rounded simple lobes 10. Ralfsia (p. 494)

2. Thallus deeply and narrowly lobed 11. Hapterophycus (p. 499)

10. Ralfsia Berk.

Thallus forming flat expansions of indefinite size and more or less
circular outline, at least at first, only slightly and broadly lobed at
the margin, never dissected into deep and narrow lobes ; either attached
firmly to the substratum or later separating largely from it, composed
at first of a single basal layer of few to many cells in thickness, but
later developing closely packed vertical filaments more or less tightly
cohering and covered by a common cuticula, those in fertile areas,
"sori, " later becoming free and rupturing the cuticula; hairs in
minute tufts, scattered, projecting beyond the upper surface of the
thallus, generally at least in funnel-shaped depressions; zoosporangia
lateral, at the bases of the free filaments, "paraphyses"; gametangia
terminal or subterminal, on the erect filaments or lateral from near
their tips.

Berkeley, in Engl. Bot., Suppl., vol. 3, 1843, pi. 2866 ; Stragularia
Stroemfelt, Om Algenveg. vid Islands Kuster, 1886, p. 49. Mesospora
Weber van Bosse, Not. sur quelq. gen. nouv., 1910, p. 27.

We have taken the genus Ralfsia in the original sense and as includ-
ing the genera Stragularia Stroemfelt and Mesospora: Weber van
Bosse, relying on the habit chiefly to characterize the genus and assign-
ing the variations in thickness, position of the gametangia, and aggre-
gation of the zoosporangia to distinguish the species, or at most, the
sections of the genus. We have been influenced to this point of view
by our experience with the variability in these respects among the
species of the Ectocarpaceae and the Myrionemataceae. The type of
the genus is RaUfsia fungiformis (Gunn.) S. and G. (Fueus fungi-
formis Gunn.).

19-5] SetcheU-Gardner : Melanophyceae 495

Most specimens collected show only the unilocular zoosporang:ia,

but although gametangia have been seen in the majority of the species,

we have not detected them, as yet, with certainty in any of our west

coast specimens.

Key to the Species

1. Thallus looseh' attached, free at the margin 5. R. fungiformis (p. 499)

1. Thallus firmly attached, not free at the margin 2

2. Thallus up to 2 mm. thick, 12 cm. diam 3

2. Thalkis less than 0.5 mm. thick, 5 cm. diam 4

3. Paraphyses up to 130yu long : 3. R. verrucosa (p. 497)

3. Paraphyses up to 360m long 4. R. hesperia (p. 498)

4. Zoosporangia 80-95^ long 2. R. californica (p. 497)

4. Zoosporangia 60-70^ long 1. R. clavata (p. 495)

1. Ralfsia clavata (Carm.) Crouan

Fronds flattened, crustaceous, circular in outline at first, later
becoming more or less lobed, firmly applied to the substratum by the
whole lower surface with occasional rhizoids, 2-20 mm. diam., about
150jU, thick ; zoosporangia narrowly ovoid to pyriform, distributed over
the whole upper surface of the frond except near the margin, 40-60//,
long, 13-18/A broad, borne at the bases of the paraphyses ; gametangia
short, terminal on the erect filaments, pluriseriate, paraphyses 80-lOOju,
long, 12-15/t broad, slightly clavate; hairs unknown.

Growing on rocks, wood, etc., in the upper sublittoral belt, and in
tide pools in the littoral belt. Unalaska Bay, Alaska.

Crouan, Alg. mar. du Finistere, Fasc. 1, 1853, no. 56 (excl.
specim.) ; Reinke, Atlas Meeresalg., 1889, pi. 6, figs. 14-20, Algen-
flora, 1889a, p. 48; Kuckuck, Bemerk. zur mar. Algenveg., 1894, p. 244,
fig. 14 ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 253, Myrio-
nema clavatum Harvey, Man. Brit. Alg., 1841, p. 124. Linkia clavata
Carmichael, MS., in W. J. Hooker, British Flora, vol. 2, 1833, p. 391.

The measurements given above for the zoosporangia are taken from
Kuckuck (lac. cit.) and from De-Toni (1895, p. 313). The material
from Unalaska cited above is scarcely mature but seems to come
within the range of measurements as given in the diagnosis above.
We are unable to harmonize the measurements given by Farlow (1881,
p. 88) for this species, with the material which we have examined from
the Atlantic coasts of America and Europe. His measurements for
the zoosporangia are 150-180/i, long, and 60-70/x broad. These measure-
ments are larger than those given for any other species except for
R. hesperia S. and G.

496 University of California Publications in Botany [^"ol. 8

Kuckuck (Bemerkimgen, 1894, p. 244 et seq.) figures and describes
''plurilocular sporangia" (gametangia) in R. clavata and in R.
Bornetii. These were found on the same individual and often in the
same sorus with zoosporangia. The material showing these organs
was collected in the winter. The study of the genus made from fresh
material and from herbarium material at our disposal, unfortunately
has been made on summer collections only. Study of the winter mate-
rial may reveal the presence of this mefhod of reproduction in the
species on our coast. Borgesen, however (1902, p. 482), reported hav-
ing discovered " plurilocular sporangia" on R. verrucosa collected in
May and July.

The Linkia clavata of Carmichael was proposed for specimens from
Appin on Loch Linnpe in the west of Scotland. Harvey adopted the
specific name of Carmichael, but removed the species to the genus
Myrionema, without, however, having seen a specimen. The Crouan
brothers referred the species to Ralfsia, quoting Harvey and thus
establishing the present binomial. The specimen referred by the
Crouans to their Ralfsia clavata seems to be R. verriwosa (cf. Reinke,
1889a, p. 48, as well as Crouan, Florule du Finistere, 1867, pi. 26,
fig. 168). Farlow (Mar. Alg. N.E., 1881, p. 88) used the correct
binomial and states that "American specimens resemble perfectly the
No. 56 of Crouan 's Algues marines du Finistere," but, as noted above,
he gives dimensions for the sporangia which are twice as long and
several times broader than those which we have assumed to belong to
the type of the species. No. 418 of the Phycotheca Boreali-Americana,
collected at Spectacle Island, Maine, seems perfectly in agreement
with European material from Reinbold and Kylin and the mature
sporangia in all average about 60ja high and 18ja in diameter. Our
Alaskan specimens agree with these in every detail. We have seen
no New England specimens with sporangia more nearly approximating
the dimensions given by Farlow than the European and Alaskan
specimens, although an occasional sporangium may be as long as 90/*,
but such is a very rare occurrence.

1925] Setchell-Gardner: Melanophyceae 497

2. Ralfsia californica S. and G.

Plate 36, fig. 22

Thallus flattened, crustaceous, circular to irregular in outline,
adhering firmly without rhizoids to the substratum by the whole lower
surface, 280-350/x thick, 2-4 cm. diam., dark brown in color ; basal
parenchymatous layer composed of 8-12 layers of cells in vertical rows;
zoosporang'ia numerous, clavate, borne laterally at the base of the
paraphyses, distributed uniformly over the central and larger part
of the frond, 80-95yii (up to 14(V) long, 16-22/x (up to 32/x) thick;
gametangia unknown; paraphyses cylindrical to slightly clavate,
180-220ju, long, 9-llja broad at the outer end ; hairs unknown.

Usually found growing on smooth, flattened pebbles and on rock
ledges in the upper sublittoral belt, and in tide pools in the littoral belt.
Central California.

Setchell and Gardner, Phyc. Cont., VTI, 1924, p. 2.

Ralfsia californica is very similar and is probably closely related to
R. clavata (Carm. ) Crouan and has passed under that name locally
for some time. Careful comparison, however, with material of that
species from the Atlantic coasts of America and Europe shows several
differences in detail, sufficient to entitle it to specific rank. The speci-
ments are, in general, much larger in diameter and are thicker. The
paraphyses are generally longer and narrower. The zoosporangia are
longer and relatively narrow. The cells of the basal portion are
slightly smaller,

3. Ralfsia verrucosa (Aresch.) J. Ag.

Frond coriaceous, adhering firmly throughout to the substratum,
juvenile state circular in outline, becoming more or less distorted,
especially on coming in contact with each other and with other crus-
taceous algae, 3-12 cm. diam., 1-2 mm. thick, with concentric zones
and radiating lines ; color olive brown ; erect vegetative filaments
curved towards the zone of active growth; paraphj'-ses up to 130/x
high, composed of 6-9 cells, the lower 2-3 times as long as broad ;
zoosporangia ovoid to pyriform, 65-80ja long, 15-25/^ broad ; game-
tangia short, terminal ; both reproductive organs on the same indi-
vidual, disposed in sori.

498 TJniversiiy of California PiiblicaUons in Botany [Vol. 8

Growing on rocks, shells, etc., in the middle and lower littoral
belts. Reported from various localities from the Bay of Unalaska,
Alaska, to Monterey Bay, California.

J. Agardh, Sp. Alg., vol. 1, 1848, p. 62 ; Reinke, Atlas Meeresalg.,
1889, pis. 5, 6, Algenflora, 1889a;, p. 48 ; Setchell and Gardner, Alg.
N.W. Amer., 1903, p. 253 ; Collins, Holden and Setchell, Phyc. Bor.-
Amer. (Exsicc), no. LV (?). Cruoria verrucosa Aresehoug, in
Linnaea, 1843, p. 264, pi. 9, figs. 5, 6.

The measurements given for this species by different authors are
quite variable. The dimensions of the zoosporangia given in our
diagnosis above are taken from Kuckuck (1894, p. 244) and our speci-
mens agree ; these were taken as representing the average typical size.
Farlow (1881, p. 87) gives the measurements of the zoosporangia as
38/x long and 19/. wide. De-Toni (1895, p. 312) gives 65-80-lOOfi
long, and 15-30,/j, wide. The material quoted from Unalaska is all
sterile. Careful comparison with mature material from the eastern
and European coasts may show that our California specimens are of
a distinct species.

The material distributed from southern California under B. ver-
rucosa in Collins, Holden and Setchell, Phycotheca Boreali-Americana,
no. LV, is very young and the rows of cells are mostly perpendicular
to the substratum instead of being curved as is the case with the
typical U. verrucosa. It seems quite possible that this plant belongs
to some other species, or possibly to another genus.

4. Ralfsia hesperia S. and G.

Thallus flattened, coriaceous, circular in outline, adhering firmly
to the substratum by the whole lower surface, at times developing
rhizoids, 3-4 cm. diam., 1-1.5 mm. thick, with concentric zonation ;
color light yellowish brown, dark on drying; paraphyses 290-360/.
long, very slightly clavate, 9-12 cells long, 8-11/. broad at the apices,
densely crowded with chromatophores above, but few below ; cells of
paraphyses 7-9 times as long as broad below, 2-3 times above ; sori
extensive, often confluent ; zoosporangia broadly clavate, 120-140/i
(up to 180/.) long, 28-34/1 broad; gametangia unknown.

Growing on rocks in the upper littoral belt. Central California.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 2.

Ralfsia hesperia is very closely related to R. verrucosa in morpho-
logical characters. The plants are, as a rule, smaller in diameter and

1925] Setchell-Gardner: Melanophyceae 499

larger in thickness, are more spongy, lighter colored and have very
decidedly larger zoosporangia and longer paraphyses than that
species. The exact southern limits of either of these species has not
been determined.

5. Ralfsia fungiformis (Gunn.) S. and G.

Thallus licheniforni, coriaceous, dark brown in color, black on
drjdng, loosely attached in the center by numerous multicellular
rhizoids, free around the margin, with free overlapping lobes seemingly
from the top of the thallus as well as the margin, circular to irregular
in outline, with concentric zones and radiating striae, 2-6 cm. diam.,
250-300/x thick ; cells forming rows curving upward and downward
from a central layer in the lobes ; zoosporangia, gametaiigia and hairs
unknown.

Growing on rocks in the lower littoral belt. Port Clarence to
Sitka, Alaska.

Setchell and Gardner, Pliyc. Cont., VII, 1924, p. 11. Fu-cus fungi-
formis Guimerus, Fl. Norv., 1772, p. 107 ; (Eder in Flora Dan., 1770,
pi. 420. Ralfsia deusta J. Agardh, Sp. Alg., 1848, p. 63; Setchell and
Gardner, Alg. N.W. Amer., 1903, p. 253 ; Saunders, Alg. Harriman
Exp., 1901, p. 424. Zonaria deusta Agardh, Syn. Alg., 1817, p. 40.
Padina deusta Postels and Ruprecht, Illus. Alg., 1840, p. 20.

The method of reproduction in this species has, apparently, never
been observed. It is a plant of the colder waters and has not been
collected and studied as much as the other species of the genus. All
of the material collected on our coast has been taken in midsummer.
In habit and structure, our Alaskan specimens agree with those from
northern New England (Phyc. Bor.-Amer., no. 419 and Phyc. Univ.,
no. 418).

11. Hapterophycus S. and G.

Thallus decumbent, dorsiventral, deeply separated into linear lobes
radiating from the center, fleshy, short, several times dichotomously
branched, concave on the ventral side, convex above, with crenate
margins, rounded sinuses and wide, blunt ends ; growth apical ; central
part of the frond consisting of large horizontal hyaline, more or
less cylindrical cells, from w^hich rows of hyaline cells curve upward
and downward terminating in the small surface cells which are filled

500 University of California Publications mi Botany [yoh. 8

with chromatophores above, hyaline below; cryptostomata with fas-
cicles of included hairs scattered promiscuously over the upper sur-
face; reproduction by zoosporangia, arising from the same basal cell
as the multicellular paraphyses; gametangia unknown.

Setchell and Gardner, in Setchell, Alg. Novae I, 1912, p. 233, pi. 25,
figs. 1-4.

•^53 "■

Hapterophycus canaliculatus S. and G.

Plate 41, fig. 54, and plate 77

Fronds up to 5 cm. wide ; color yellowish brown ; sporangia 65-75/x
long, 18-25;u. wide ; paraphyses cylindrical, composed of 4—7 cells, 6-7ju,
wide, 12-1 6/x long.

Growing on rocks in small tide pools, or on the horizontal surface
of flat rocks subjected to wave action, in the upper littoral belt, or
even above high-tide limit. San Pedro, California.

Setchell and Gardner, in Setchell, Joe. cit.

Hapterophycus, up to the present, is a monotypic genus, canali-
culatus being the only known species. It has been found only in the
vicinity of San Pedro, California, but doubtless may extend much
farther south.

In its histological structure, it resembles Ralfsia deasta, especially
in the free lobes of that species. Since the publication of the original
description, it has been discovered that on the upper surface there
are numerous, small, inconspicuous cryptostomata containing fascicles
of hairs which do not extrude beyond the surface of the frond.

12. Lithoderma Aresch.

Thallus consisting of a thin parenchymatous disk, circular or irre-
gular in outline, of indefinite expansion, attached to the substratum
by the entire under surface without rhizoids ; hairs absent ; growth
in diameter marginal, and in thickness by horizontal cell division, the
cells arranged more or less regularly in vertical rows ; reproduction by
zoosporangia and by gametangia borne in sori on different individuals,
the zoosporangia on the outer ends of the vertical rows of cells, the
gametangia either terminal on the vertical roM^s of cells or lateral on
special filaments.

Areschoug, Observ. Phyc, part 3, 1875, p. 22.

The type selected by Areschoug to represent his newly proposed
genus is Lithoderma fatiscens, which is the Ralfsia extensa of Crouan

1925] Setchell-Gardner: Melanophijceae 501

(Flor. dii. Fin., p. 166). It is credited only to the extreme northern
part of our region. All of the specimens previously reported within,
our territory are sterile and there is the greatest doubt as to their
identity, both generic and specific. Two species have been assigned to
our coast, but it is doubtful whether they are correctly referred. We
can onh' list these two species with description and comment.

Key to the Species

Thallus nearly circular in outline, up to 3 dm. diam 1. L. fatiscens (p. 501)

Thallus irregularly elongated or almost linear in outline 2. L. lignicola (p. 501)

1. Lithoderma fatiscens Aresch.

Thallus Ijroadl}' expanded, nearly circular in outline, up to 3 dm.
diam., scarcely 1 mm. thick, smooth, shining when wet ; the basal
layers of cells minute, and somewhat polyhedral, vertical filaments
12-18/x diam., 8-12 cells long; gametangia small, cylindrical or oval,
formed by transformation of terminal cells or borne on short, almost
colorless branches from the terminal cells; zoosporangia terminal,
sessile on the vertical rows of cells, ellipsoid or ovoid.

Growing on stones in the lower littoral and upper sublittoral belts.
Port Clarence, Alaska, and Iliuliuk, Unalaska,

Areschoug, Observ. Phyc. part 3, 1875, p. 22 ; Kjellman, Om
Bering. Alg., 1889, p. -49; Setchell and Gardner, Alg. N.W. Amer.,
1903, p. 253.

The plants of Kjellman from Port Clarence are sterile and those
of Setchell and Lawson from Iliuliuk are not to be found for
reexamination.

2. Lithoderma lignicola Kjellm.

Thallus forming thin, uneven, elongated, almost linear, somewhat
confluent crusts, 250/x or more thick ; the basal layer composed of
united, radiating branched rows of thick-walled cells, nearly parallel,
not flabellate; the basal layer giving rise to vertical rows of cells, 20
or more, branching occasionally, loosely united above ; cells 10-15/a
long, 8-lO/i, diam. ; character of reproductive organs unknown.

Growing on piles in the littoral belt. St. Michael, Alaska?.

Kjellman, Alg., Arc. Sea, 1883, p. 256, pi. 26, figs. 8-11 ; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 253.

This species was discovered by Kjellman ' ' in the Norwegian Polar
Sea." Specimens were taken at Talvik in the middle of September,

502 University of California Publications in Botany [Vol. 8

as described by Kjellman {loc. cit.). The specimens were all sterile.
We have seen no data as to the reproductive characters of the species
since. A sing'le collection of sterile material taken by one of us
(Setchell) at St. Michael, Alaska, in July, 1899, was referred by us
(loc. cit.) to this species. A reexamination of the material and com-
parison with material of the type collection, demonstrate that the
Alaskan plants are different, but without assisting in definitely
placing them.

FAMILY 5. ELACHISTEACEAE kjellman

Thallus pulvinate or penicillate, more or less lubricous, composed
of loosely to compactly intertwined, more or less monosiphonous fila-
ments at times agg'lutinating or anastomosing: and forming a pseudo-
par enchymatous base, often penetrating the host below and always
giving rise above to numerous free assimilating filaments, either
zoosporangia or gametangia or both and at times colorless hairs ; game-
tangia uniseriate or interruptedly pauciseriate, occasionally lateral,
single or grouped; "paraphyses" present or absent. -

Kjellman, Handbok Skand, Hafsalgfl., I, 1890, p. 41 ; Folsie, Contr.
knowl. mar. alg. Norway, I, 1890, p. 92. Elachisieae Reinke, Algenfl.
westl. Ostsee 1889a, p. 49.

The family of the Elachisteaceae, as separated and indicated by
recent writers, includes a number of pulvinate species, all small, but
some even minute, largely epiphytes, or possibly growing at times on
rocks. They are to be distinguished by their basal portions. In
Elachistea, the basal portion is solid due to the agglomeration, or
anastomosing of the filaments in this portion of the thallus. In
HalotJirix, Giraudia, Symphoricoccus, Leptonema, etc., the basal por-
tions of the filaments are not united to any considerable extent or at
all, but are intertwined and woven together to form a spongy mass.
AVe are inclined to refer to this family Cylindrocarpus micro scopicus
Crouan (cf. Kuckuck, 1899, pp. 49-55, figs. 1-5) as being a link
between the Ectocarpaceae and the Elachisteaceae but, because of its
pulvinate habit and densely interwoven filaments, approaching the
Elachisteaceae more nearly than typical Ectocarpaceae. Several
species of Ectocarpus and especially E. liemispliericus Saund. seem
to foreshadow the Elachisteaceae, but are not sufficiently compacted
or agglutinated at the base to properly be included in this family.
The method of origin of the tufts of the members of this family does
not seem to have been traced, but prostrate filaments, either free or

i^-'^] Setchell-Gardner : Melanophyceae 503

more or less united into a disk or irregular membrane probably pre-
cede the development of the compacted or agglutinated erect filaments,
thus assisting in linking the Elachisteaceae with the Ectocarpaceae
and the more pulvinate Myrionemataceae,

Key to the Genera

Basal cells of the thallus more or less firmly agglutinated; paraphyses present

13. Elachistea (p. 503)

Basal cells of the thallus loosely interwoven; paraphyses wanting

14. Gonodia (p. 505)

13. Elachistea Duby

Frond tufted or expanded, composed of a colorless basal pseudo-
parenchymatous mass of cells, which may or may not penetrate the
host, and arising from these, closely compacted "paraphyses," long
free assimilating filaments and numerous long hairs, present only in
certain species; reproduction by zoosporangia and by gametangia,
both borne among the ' ' paraphyses. ' '

Duby, Bot. Gall., 1830, p. 972.

We have adopted the original spelling of Duby who founded the
genus on the Conferva scutulata Engl. Bot., pi. 2311. The type species
of the genus, E. scutulata, was collected in 1811 by Mr. Borrer at
Brighthelmston on the coast of Sussex, England. There seem to be
two species within our territory, both occurring to the north of the
California coast. Opospermum of Rafinesque (Precis decouv. et tra.v.
somiol., 1814) seems, from the description, to be unidentifiable, at
least by no means certainly to apply to any member of Elachistea.

Key to the Species

1. Paraphyses decidedly curved, strongly moniliform, with upper cells as long or
longer than broad 1. E. fucicola (p. 503)

1. Paraphyses straight or slightly curved, barely constricted at the di.ssepiments,
with upper cells shorter than broad 2. E. lubrica (p. 504)

1. Elachistea fucicola (Velley) Aresch,

Plate 38, figs. 33-35

Thallus fasciculate, 4-7 mm. high, the basal portion composed of a
small, usually hemispherical, dense, lubricous mass of closely inter-
twined branched filaments, giving rise to numerous free paraphyses,
sporangia and assimilating filaments ; paraphyses clavate, uncinate,
moniliform, 170-200/x long, 18-24/^, diam. above, 7-lOfi diam. below ;

504 Universitij of California Publications in Botany [Vol.8

assimilating filaments briefl}^ tapering" near the base, and long and
gradual tapering above, variable in size, 20-70/a, mostly 40-50/x, diam.
not ending in a hair ; cells 0.5-2 times as long, cell wall thick ; chro-
matophores in actively growing cells a network extending throughout
the cell, later becoming- granular; zoosporangia narrowly cuneate,
rounded above, 90-llOju, long, 22-26/a diam., 200 or more spores escap-
ing at the apex in a utricle ; gametangia unknown.

GroAving on different species of Fucus in the lower littoral belt.
Sitka, Alaska, to central Oregon (Coos Bay).

Areschoug, in Linnaea, vol. 16, 1842, p. 235, pi. 8, figs. 6, 7.
Conferva fucicola Velley, Mar. PI., 1795, no. 4.

This species was first reported as being in our region by Ruprecht
in 1851 (p. 389) from Sitka. No subsequent report of its having been
collected had come to us until one of us (Gardner) collected it in 1917
at the same station.

There seems to be some question as to the exact limits of Elachistea
fucicola, but it has appeared most reasonable to us to adopt, as typical,
the plant figured by Kuetzing in the Tabulae Phycologicae (vol. 7,
pi. 95). We find plants within our territory corresponding in general
to Kuetzing 's figure and epiphytic on species of Fucus. The free
filaments are long attenuate at both the tip and the base. They vary
decidedly in diameter and in length and breadth of cells even in the
same individual, and the paraphyses are decidedly curved and de-
cidedly moniliform, with the uppermost cells usually as long as, or
longer than, broad. Our plants agree fairly well, also, with no. 417 of
the Phykotheka Universalis collected at Warnemiinde on the north
coast of Germany by Heiden. Our figure (plate 38, figs. 33-35) gives
the essential characteristics of structure. Elachistea fucicola is usually,
at least, more olivaceous than E. lubrica and all parts of the plants are
firmer and collapse less on drying than do those of E. lubrica. Thus
far we have detected E. fucicola. only on species of Fucus.

2. Elachistea lubrica Rupr.

Thallus usually forming distinctly separate tufts, 3 mm.-2 cm.
(9-11 mm. Rupr.) high, lubricous; mass of paraphyses relatively
small and loosely intertwined ; hairs absent ; erect free filaments
abruptly attenuated at the base, and gradually attenuated above the
center, 18-22/a diam. at the base, 38-48/i, in the widest part, tapering
to 9/7. or less at the apex ; cells of erect filaments cylindrical to slightly

1!^-''] SetcheU-Gardner : MeJanophyceae 505

doliiform, 2-3 times as long as the diameter at the base, 0.3-0.8 times
as long- a short distance above the base, 1-1.5 times as long in the
widest part, np to 6 times as long at the apex ; chromatophore a fine
nodular network in the young cells, separating more or less into
spheroidal masses in the older cells ; zoosporangia subovate to broadly
clavate, 75-100,(x (up to 135fi) long, 30-lOjU broad at the outer end;
gametangia unknown ; paraphyses clavate, about 225jli long, slightly
curved at times but not sensibly constricted at the dissepiments.

Growing on other species of algae, especially those of, Rhodymenia.
From Prince William Sound to Wrangell, Alaska.

Ruprecht, Tange Ochots. Meer., 1851, p. 196 (388 Orig.) ; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 248; Collins, Holden and
Setchell, Phyc. Bor.-Amer. (Exsicc), no. 828.

The type of Elachistea Juhrica Rupr. came from the Ochotsk Sea,
l)ut Ruprecht states that specimens from Russian Lapland are in
close agreement. The most typical specimens seem to grow upon
species of HnJosaccian and Bhodymenia. Specimens from Yakutat
Bay collected by Saunders (Phyc. Bor.-Amer., no. 828) and by x\lbin
Johnson as well as at Orca by Setchell, seem to agree well with speci-
mens distributed from Spitzbergen by Kjellman and from Mehavn in
Finmarken (Phyk. Univ., no. 366) collected by Foslie. The free fila-
ments taper both above and below as in E. fucicola, but the paraphyses
are less curved, only slightly, if at all, constricted at the dissepiments,
and the upper cells are inclined to be broader than high.

14. Gonodia Nieuwland

Fronds minute, tufted, composed of a mass of colorless, densely
intertwined filaments, mostly penetrating the host, giving rise to a
dense mass of erect assimilating filaments and "hairs," at least at
times; "paraphyses" wanting; reproduction by zoosporangia and
gametangia.

Nieuwland, Critical Notes IX, 1917, p. 30 ; Setchell and Gardner,
Mar. Alg. Gulf Calif., 1924, p. 722 ; MyriacHs Kuetzing, Phyc. gen..
1843, p. 330.

The distinctions between Gonodia and Elachistea are not always
readily discernible in all of the species. In the younger stages of some
of the species of Elachistea, and at times in the older stages, the basal
portion, especially if it penetrates the host, is of about the same degree
of compactness as it is in the mature stages of the species of Gonodia.

506 University of California Putlications in Botany ['^o-l. 8

Typically, however, in the mature stages of the two genera, Elachistea
has a firm, closely intertwined, agglutinated mass of filaments for the
basal portion, and Gonodia remains free and loosely intertwined. "We
are using this character as a basis of separating the two genera.

Key to the Species

1. Basal cells of the erect free filaments symmetrical 1. G. Johnstonii Cp. 506)

1. Basal cells of the erect free filaments asymmetrical.... 2. G. Marchantae (p. 506)

1. Gonodia Johnstonii S. and G.

Plants forming dense minute tufts in the conceptacles and on other
parts of the host, 160-200/x high, attached by rather deeply penetrat-
ing, sparsely branched, rhizoidal filaments ; erect fronds forked at the
surface of the host, vegetative filaments unbranched above, decidedly
clavate and blunt, at times tapering upwards above the center; cells
in widest part IO-II/a diam., 1-1.5 times as long as broad, slightly
constricted at the dissepiments ; zoosporangia broadly clavate, 65-75/a
long, 18-22ju, broad; gametangia cylindrical, 60-75/x long, 6-9/x, broad,
densely fasciculate ; both sets of reproductive organs borne on the same
plant at the surface of the host.

Growing on Sargassum insiUare. San Marcos Island, Gulf of
California. ■ .

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 722, pi. 17,
figs. 46, 47.

2. Gonodia Marchantae S. and G.

Fronds forming dense tufts with a dense mass of colorless filaments
at the base penetrating the host, the free portion about 200/x long ; fila-
ments unbranched aboA'e the host, the lower portion composed of 2-3
long, narrow cells, abruptly changing into 2-3 asymmetrical swollen
cells, then gradually attenuated upwards to blunt apices ; widest cells
18-22/x, the length of the cells in the upper part equaling the breadth ;
pseudoparenchymatous cells doliiform to subspherical ; zoosporangia
broadly clavate, 55-65jx long, 22-26/.^ broad; gametangia cylindrical,
densely fasciculate 55-65ju. long, 6-7 jjl broad ; both sets of reproductive
organs borne on the same plant at the base of the free filaments.

Growing on the fronds of Sargassum horridum. La Paz, Lower
California.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 723. pi. 17,
fig. 48.

1^25] Setchell-Gardner : Melanophyceae 507

G. Marchantae differs from G. Johnstonii in the character of the
basal penetrating portion, the former having few narrow filaments
and the latter having a dense, copious base. Two or three cells in the
lower part of the free filaments are usually asymmetrical in G. Mar-
chantae and not in G. Johnstonii. Neither species has hairs nor any
indication of having had them, a prominent character as figured bj^
Tliuret and Bornet in Etudes Phycologiques, pi. 7, figs. 2-6, for
Elachista pulvinata and mentioned by Yendo as " paraphysibus pau-
cioribus" in Myriactis Sargassi (Novae Alg. Japon., 1920, p. 3).
G. Marchantae is close to G. pulvinata in the character of the basal
penetrating portion. G. Johnstonii in general resembles G. monili-
formis but is much smaller throughout.

FAMILY 6. LEATHESIACEAE pam. nov.

Thallus thick, carnose, expanded on the substratum and solid or
irregularly globular and hollow, arising from a flattened, monostro-
matic (?) persistent or evanescent disk; inner cells large, colorless,
loosely parenchymatous of di- to trichotomous filaments, outer colored,
in anticlinal rows, generally decreasing in size from within outwards,
held together, at least loosely, by the surrounding jelly ; zoosporangia
and gametangia with loculi uniseriate or nearly so, both immersed
among the external anticlinal rows present and borne on the same or
similar plants; epiphytic or saxicolous.

Corynophlaeceae Oltmanns, Morph. und Biol, der Alg., 1922, p. 23.

As will be made apparent later, it seems most desirable to restrict
the family Chordariaceae to the genus Chordaria, as typified by
C. flagelliformis, an erect branched plant with distinct elongated
axis and lateral branches of limited growth, with subapical terminal
meristem, not trichothallic in the same degree as in the ^giraceae
and with only the unilocular zoosporangia thus far known, suggest-
ing the strong probability of a microscopic gametophyte. The
^giraceae are more or less elongated, simple or branched plants, with
strongly trichothallic apical meristems, and both zoosporangia and
gametangia borne on macroscopic plants. The Corynophlaeaceae of
Oltmanns (1922, p. 23) includes the genera Cylindrocarpus, Micro-
coryne, Strepsithalia, Corynophlaea, Myriactis, and, presumably,
Leathesia, although Oltmanns might be suspected of merging {loc cit.,
p. 26) Leathesia (Gray, 1821) in Corynophlaea (Kuetzing, 1843). We
have referred Myriactis (Gonodia.) to the Elachisteaceae because of
its close resemblance to Elachistea. Cylindrocarpus microscopicus

508 University of California Publications in Botany [Vol. 8

Crouan seems to us to be more properly included in the Ectocarpaceae,
as a further development of such forms as Ectocarpus hemisphericus
and E. chantransioides and on the way toward simpler ^^giraceae, such
as Ave conceive may be represented by Microcoryne (cf. Oltmanns, loc.
clt., pp. 25, 26). Strepsithalia is unknown to us, but on account of its
gelatinous character may possibly be considered to be a slightly
developed, or even degenerate relative of Corynoplilaea. Corynoplilaea,
if it be distinguished from Leathesia, is more generally solid, with
longer assimilating filaments which are also less swollen at the tip ;
Kylin (1907, p. 83) is of the opinion that the three genera, 3licro-
coryne, Corynoplilaea, and Leathesia are to be retained as representing
a series in complexity, leading up to the most highly developed species
of Leathesia. As indicated above, Microcoryne, on account of its elon-
gated form and greater or less differentiation of layers seems to
approach JEgira and the resemblance in structure to Mgira has been
generally noticed and commented upon. Between Leathesia and Cory-
noplilaea there seems to be no satisfactory distinction either as between
solid and hollow, or between longer and shorter cortical filaments. We
have therefore merged the two genera under Leathesia. and designated
the family as Leathesiaceae. AVe can see no resemblances sufficiently
close to unite Petrospongium with Cylindrocarpus (considering C.
microscopiciis Crouan as the type). We have therefore kept them
distinct for reasons to be given later.

The Leathesiaceae are carnose algae of flattened or globular form,
solid or hollow, arising from a primitive basal disk, with the inner cells
large and colorless, more or less agglutinated, the cortical cells in
short filaments with chromatophores, without exserted structures of
any kind except colorless hairs, with peripheral growth, and having
gametangia and zoosporangia on the same or on similar plants.

Key to the Genera

1. Zoosporangia more or less difform, usually attached laterally

,.15. Petrospongium (p. 508)

2. Zoosporangia pjTiform to ellipsoidal, attached at the base. 16. Leathesia (p. 510)

15. Petrospongium Naeg.

Thallus small, thin, flat, usually encrusting rocks, with or without
rhizoids, gelatinous, composed of 3 more or less distinct tissues, viz. ;■

(1) a basal layer, monostromatic ( ?), giving off rhizoids, whence arise

(2) loosely compacted, di- or trichotomous filaments, decumbent below,
curving to erect above, consisting of elongated, cylindrical or swollen.

1925] Setchell-Gardner : Melanaphyceae 509

colorless cells with scanty chromatophores, and (13) a coi'tieal layer
of short, branched, closely compacted, erect filaments composed of
short, colored cells forming- a more or less smooth or wrinkled surface ;
zoosporangia, embedded within the thallus, more or less cylindrical,
becoming difform, usually attached laterally ; gametangia unknown.

Naegeli, in Kuetzing. Tab. Phyc, vol. 8, 1858, p. 2, pi. 3, fig. II.

Comparison of Petrospongium Berkeleyi (Grev.) Naeg". with Cylin-
drocarpus mkroscopicus Crouan reveals such a difiPerence in habit,
structure, etc., that it seems impossible to place the two species in the
same genus. We are also of the opinion that they belong in separate
families. The figrures of Cyiindrocarpus niicroscopiciis, especially
those of Kuckuck (1899, p. 88, pi. 6, figs. 1-5) show a plant of definite
habit, growing in tufts (mueose, fide Crouan) rather tending to
elongate although only slightly so, without its individual branched
filaments being agglutinated into the definite thallus with inner
and outer distinct tissues as in the Leathesiaceae, but united into
a spongy (fide Kuckuck) mass by intertwining rhizoidal corticat-
ing filaments. We are inclined to place Cyiindrocarpus micro-
scopicus in the Ectocarpaceae, as an extremely differentiated type
of such forms as Ectocarpus hemisphericus and E. chantransioides
and looking toward the ^Egiraceae and perhaps, also, the Leathe-
siaceae, but not properly of them or, to the same degree, differen-
tiated into definite inner transparent and outer colored layers.
Petrospongium Berkeleyi (Grev.) Naeg. and P. rugosum (Okamura)
S. and G., especially the latter, have their tissues differentiated and of
agglutinated, level-topped filaments and are, in our opinion, perfectly
typical members of the Leathesiaceae.

Petrospong-ium rugosum (Okamura) S. and G.
Plate 39, figs. 42, 43

Thallus adhering more or less loosely to the substratum, circular to
irregular in outline, flat or thrown up into folds, more or less spongy
and lubricous, 2.5-5 cm. (up to 10 cm., cf. Okamura) diam., 1.5-2.5
mm. thick, of a dark, glossy, chestnut brown color ; cells of the cortical
layer 8-11/a diam., 1-2 times as long as the diameter; zoosporangia
narrowly ellipsoidal, at times decidedly difform, attached laterally a
little below the middle and near the base of the cortical filaments,
75-90/.1 long, 16-22/x broad.

Growing on rocks in the upper littoral belt exposed to surf.
Central and Southern California.

510 University of California Puhlications in Botany [Vol. 8

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 12. Cylindro-
carpus rugosa Okamura, Alg. Japon. Exsicc, no. 88, 1903, Icon, Jap.
Alg., vol. 1, 1907, p. 20, pi. 5, figs. 1-6. Petrospongium Berkeleyi
Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 232,
1896 (not of Naegeli).

This curious, dark colored, rather firm, yet carnose plant is com-
mon on rocks exposed to surf along the central Californian coast and
has been found on that of southern California as well. Its peripheral
filaments differ in size (about twice the diameter) and in branching
from those of the European species and agree closely with those of the
Japanese species.

16. Leathesia Gray

Fronds irregularly globose, solid in the juvenile stage, soon becom-
ing hollow in some species and remaining solid in others, decidedly
carnose, interior consisting of radiating, dichotomously or trichotom-
ously branched filaments composed of large, irregular, colorless cells
toward the base or center as the case may be, merging into smaller
cells toward the periphery ; cortex composed of a short series of small,
color bearing cells set in a firm mucous matrix; zoosporangia and
uniseriate gametangia known, both arising at the base of the cortical
filaments; hairs single or in fascicles.

Gray, Nat. Ar. Brit. PI., vol. 1, 1821, p. 301.

The genus Leathesia, as understood and interpreted by us, should
include the Corynophlaea and Corynephora of Kuetzing (1843, p. 331).
The chief distinction between Corynophlaea and Leaihesia as diag-
nosed, seems to be that in Corynophlaea the adult thallus is attached
by a single layer of cells from which arise all the other cells, forming
a flattened, hemispherical or spherical, solid structure depending upon
the nature of the substratum, while in Leathesia the adult thallus is
more or less spherical and usually hollow. Structurally and in mode
of development there seems to be no fundamental difference. The
basal part is practically the same in both genera in the early stages
of development (cf. Oltmanns, 1922, p. 26, but cf. also Okamura, 1907,
p. 81, pi. 18, figs. 9-14). The cells of the basal portion are more or
less modified to serve as attaching cells in both genera. It is difficult
to determine in the diminutive species of both genera whether or not
all of the specimens are solid or hollow. Kuetzing indeed transferred
the type species of his genus Corynophlaea, C. haltica, to his genus
Corynephora in Tabulae Phycologicae (1858, pi. II), which is hollow

192.5] Setchell-Gardner : Melanophyceae 511

and differs in no fundamental characters from Leathesia. In the
larger species of Leathesia the Avhole under portion sooner or later
disintegrates, leaving only the marginal cells for attaching the plant.
Until more can be learned of the life-history of and extent of variation
in these genera, it seems to us best to combine them and place them
all under the oldest name, Leathesia.

Key to the Species

1. Fronds diminutive, up to 1.25 mm. diam 1. L. nana (p. 511)

1. Fronds variable in size, up to 8 cm. diam 2

2. Paraphj'-ses terminating in a spherical or subspherical cell

3. L. amplissima (p. 513)

2. Paraphyses clavate, gradually and uniformly enlarging upward

2. L. difformis (p. 511)

1. Leathesia nana S. and G.

Plate 43, fig. 67

Thallus solid, spherical, 500-800/i, (up to 1.3 mm.) diam.; color
dark brown ; cells in the center subspherical, 40-55ju, diam. ; paraphyses
30-40/A long, composed of 3-4 cells, terminal cell enlarged, sub-
spherical, 10-11/i, diam. ; hairs 300-400/i, long, 4-5/* diam., scattered,
not in fascicles; gametangia with 4-5,u loculi, uniseriate, 3.5-4.5ju.
diam. ; zoosporangia unknown.

Growing on the leaves of Phyllospadix near the outer ends.
Monterey, California.

Setchell and Gardner, Phyc. Cont. VII, 1924, p. 3.

The host upon which this species of Leathesia was found has a
wide distribution along the west coast of the United States and careful
observation may show that this little known epiphyte is coexistent
with its host. In structure it is not far unlike the Corynephora marina
of Kuetzing (1843, p. 331 and 1858, pi. 3, fig. I) but is of smaller
dimensions and has much smaller interior cells.

2. Leathesia diflformis (L.) Aresch.

Plate 40, fig. 52 and plate 43, figs. 65, 66

Fronds gregarious, very variable in size, up to 6 cm. diam.,
spherical and solid when young, becoming hollow and more or less
lobed at maturity, hairs in small scattered groups; cortical filaments
clavate, 5-6 cells long, with enlarged, subspherical to spherical terminal
cells; color olive brown; gametangia cylindrical, 34-45/* long, 4-6;ti

512 University of California Puhlications in Botany [Vol. 8

broad, with 6-8 uniseriate loculi; zoosporangia pyriform or ovoid,
35-60ju, long, 16-22/A broad.

Growing on other algae and on rocks in the littoral belt. Ranging
from Bering Sea, Alaska, to southern California.

Areschoiig, Phyc. Scand. I, 1846, p. 154 (Repr.) ; Saunders, Alg.
Ilarriman Exp., 1901, p. 423 ; Setchell and Gardner, Alg. N.W. Amer.,
1903, pp. 249, 250; Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. 829; Tilden, Amer. Alg. (Exsicc), no. 243 (?). Tre-
meUa difformis Linnaeus Fl. Suec. (ed. two), 1755, p. 429.

Regarding the dimensions of Leathesia difformis as a whole or
especially of its parts the literature is practically silent. Areschoug
(loc. cit.) states "Thallus nunc semine Cannabidis vix major, nunc
2-5 uncias diametro aequans, ' ' but gives no definite statements regard-
ing the size of the various parts. Farlow (1881, p. 82) gives as the
diameter of the frond "from half an inch to two inches." De-Toni
(1895, p. 422) states sporangia "35 x 17." It seems to us quite prob-
able that this name has been used freely to include a number of species
somewhat similar in general appearance, but in reality distinct. We
feel that much more work should be done on our Pacific coast material
before the limits of the species can definitely be determined.

Gardner no. 3462 was collected at Tomales Bay, California, in
August and shows mature zoosporangia and gametangia on the same
individual. The gametangia have 6-8 loculi. Gardner no. 4501 was
collected at Pacific Grove, California, in December, and has only game-
tangia. These have 9-13 loculi. It may be that we are dealing with
two entities, but for the present we are grouping them under L. dif-
formis. Plants from both of these numbers are figured on plate 43.

The dimensions given in our diagnosis above are more or less a
composite but represent a fair average of the size of the structures
found in the plants which we have included in the species. We have
examined material of this species in our herbarium from Kylin, col-
lected at Strangahnfund and at Varberg, Sweden, which may be
considered to be topotypes of L. difformis. The material from both
of these localities is bearing fruit and apparently mature. The fol-
lowing measurements of their parts will indicate the relation in this
respect to our Pacific coast material. Paraphyses, composed of 3-4
cells, 38-44/x long in plants with gametangia, up to 15fx long in plants
with zoosporangia, terminal cells 14-18/x in diameter. Gametangia
composed of 4—6 uniseriate loculi, 20-25;li long, 5-6/x in diameter.
Zoosporangia are 30-50;a long, 15-22fi in diameter.

1925] Setchell-Gardner: Melanophyceae 513

3. Leathesia amplissima S. and G.

Plate 43, fig. 64

Thalliis hollow, subglobose to irregular, attached by a broad area,
the central part of which disintegrates on nearing maturity, 3-8 cm.
diam. ; color yellowish brown ; interior colorless cells relatively large,
compact and angular from compression, becoming radially arranged
and smaller toward the surface ; hairs single, scattered over the whole
free surface, ll-13ju, diam. ; paraphyses clavate, enlarging gradually
from the base to the apex, 55-65ju, long, 4-7/* diam. at the base, 7-12/a
at the apex, 4-5 cells long; chromatophores densely crow^ded; zoo-
sporangia ovoid to ellipsoidal, 38-46/* long, 22-28/x. broad ; gametangia
unknown.

Growing on rocks and over other small algae, in tide pools in the
lower littoral and upper sublittoral belts. Central California (Pacific
Grove and Monterey).

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 3.

Leathesia amplissima is not readily distinguishable in gross mor-
phological characters from L. difformis as we understand that species
at present. The histological structure, however, is essentially different
as may be seen from our descriptions and figures.

FAMILY 7. PUNCTARIAC'EAE kjellm.

Fronds from filiform and almost monosiphonous to saccate, ligulate
or broadly expanded and membranaceous, attached by a relatively
small parenchymatous disk or by rhizoidal filaments, all but the mono-
siphonous species differentiated into an inner and an outer tissue, the
inner mostly of colorless cells and the outer of color bearing cells;
reproduction both asexual, by zoosporangia projecting slightly or not
at all beyond the surface, and sexual, by gametangia projecting w^holly
or in part beyond the surface; both zoosporangia and gametangia
transformed single surface cells, either scattered or collected into
definite sori ; hairs (and paraphyses?) present in some.

Kjellman, in Gleerup, Enum. PI. Scand., 1880, p. 9 (fide Foslie,
Contr. Knowl. Mar. Alg. Norway, I, 1890, p. 98) ; Oltmanns, Morph.
und Biol, der Alg., 1922, p. 49 et seq. Pimctarieae Thuret, in Le Jolis,
Liste alg. mar. Cherb., 1863, p. 14 (in part) ; Farlow, Mar. Alg. New
Eng., 1881, p. 63.

As usually limited, the Punctariaceae include the genera Omphalo-
phyllum Rosenv., Pha.eosacci'On Farlow, Punetaria Grev., Desmo-

514 University of California Puhlications in Botany [Vol. 8

trichum Kuetz., Pogotrichum Reinke, Litosiphon Harv. and Corycus
Kjellm. We know of but one of these in our territory, viz., Punctaria.
Structurally the simplest forms of the Punetariaceae, as generally
understood, are almost monosiphonous. The more complex forms
usually included in the family are either filamentous and solid, saccate
or membranaceous. They agree in general in that the zoosporangia
and gametangia arise as transformed single surface cells, projecting
beyond the surface but slightly, if at all, but if projecting not accom-
panied by pluricellular paraphyses. They form fairly definite and
circumscribed sori.

The Punetariaceae have for their type genus Punctaria Grev. whose
type species is P. latifolia Grev. Their relation is closely with
Striariaceae. on the one hand, and the Scytosiphonaceae on the other.
This relationship is so close that it is largely a matter of convenience
to separate them. From the Striariaceae they may be separated by
the tendency of their fronds even in the simplest Desmotrichum
species, to develop a membrane rather than a cylindrical frond. In
case this distinction is made, Pogotrichum, and Litosiphon, under such
understanding, should be referred rather to the Striariaceae and to
the neighborhood of Stictyosiphon. While the species of Desmotrichum
and- Punctaria are both supposed to have hairs, yet some of our Pacific
coast species of Punctaria have no trace of hairs in adult plants. There
is an absence of hairs also in the known species of Phaeosaccion and of
Omphalophyllum, two genera usually referred to the Punetariaceae.
The two genera just mentioned also lack specialized sporangial cells in
groups and it may be found desirable to separate them and give them
a family designation of their own. Neither of these genera have
been detected as yet in our territory, so that we have no opportunities
for careful study. Corycus of Kjellman is hollow and, so far as
descriptions inform us, devoid of hairs in the adult stages. It is
flattened, even if it is hollow, and seems to belong in the Punetariaceae
on that account as well as agreeing with both Striariaceae and
Punetariaceae in having circumscribed sori which do not project
beyond the surface.

The Punetariaceae differ from the Scytosiphonaceae in having the
sori circumscribed and, commonly, of regular shape. The Scyto-
siphonaceae have extensive, indefinite and confluent sori and, thus far,
have been detected only with gametangia.

1^25] Setchcll-Gardner: Melanophyceae 515

17. Punctaria Grew

Fronds simple or more or less branched, monosiphonous, usually
complanate, membranaceous, linear to broadly expanded, attached by
a small discoid base, composed of several layers of cuboidal cells
similar in size or those of the surface layer much smaller ; reproduc-
tion by zoosporang-ia immersed in the tissue or slightly projecting or
bulging" beyond the surface, and by gametangia usually partly or
wholly immersed in the tissue; both gametangia and zoosporangia
representing transformed surface cells, single or in small groups,
scattered promiscuously on both sides of the frond ; hairs present in
some species; paraphyses wanting.

Greville, Alg, Brit., 1830, p. 52 and Syn., p. xlii. Diplostrommm
Kuetzing, Phyc. Gen., 1843, p. 298. Phycolapathum Kuetzing, Phyc.
Gen., 1843, p. 299 (pro parte maxima). Desmotrichum Kuetzing,
Phyc. Germ., 1845, p. 244. Homoeostroma J. Agardh, Anal, alg.,
Cont. Ill, 1896, p. 7. Nematophlaea J. Agardh, loc. cit., 1896, p. 12.
Ehadinocladia Schuh, Rhodora, vol. 2, 1900, p. 3, pi. 18.

There can be little question as to the idea underlying Greville 's
proposal of the genus Punctaria, if we accept his P. latifolia as the
type. The specimen examined by Greville in its living state was
collected at Sidmouth on the south coast of England by Mrs. Griffiths.
It seems sufficiently certain that Wyatt's Algae Damnonienses, no. 9,
is practically a topotype, having been collected along the same coast
and not far removed from the actual type locality, Wyatt 's specimen
answers in every detail to Greville 's description and we base our idea
of the genus Pimctaria and of the type species, Punctaria latifolia
Grev., on it. It is probably the same as the plant figured by Harvey
in Phycologia Britanniea (plate 8). The Wyatt specimen is a
Punctaria also in the sense of J. G. Agardh (1896, p. 4, pi. 1, fig. 1),
although a distinct species from any included by him. The Punctaria
of Greville, besides including P. laiifalia Grev." and P. plantaginea
(Koth) Grev., was also extended to include P. tenuissimum (Ag.)
Grev. This last species was removed later by Kuetzing to his Diplo-
stramiuni (1843, p. 298) and this genus, in turn, in spite of the refer-
ence to it as first or "type" species of Ulva plant aginif alia Lyngb.,
is identical, partly in foundation and partly in content, with Kuetz-
ing's later genus Desmotrichum (1845, p. 244). Kuetzing's genus
Diplostromium has, then, the characters of Greville 's genus Punctaria.

516 University of California Puhlications in Botany [Vol. 8

The Pimctaria tenuissima Grev. is probably identical with P. undulata
J. Ag., now generally referred to Desmotrichum.

It is much more difficult to determine the exact status of Pimctaria
plantaginea (Roth) Grev. The original is the plant figured in Dil-
lenius (1741, pi. 9, fig. 4) which, as well as the description, was
founded on a plant studied by Micheli. Smith (1810, pi. 2136) again
figured it and described it from Micheli 's material and notes which
had been communicated to him by Professor 0. Targioni-Tozetti long
after the death of Micheli. Whence the original material was derived
is uncertain, but it seems likely to have been the Adriatic Sea since
Micheli lived at Florence. In such a case, it is probable that it is not
the species which Greville referred to his Punctaria. plantaginea but
nearer to P. latifolia Grev. There seems to be little reason for doubt-
ing that the Pimctaria plantaginew as Greville understood it is also a
Punctaria^ in the sense of J. G. Agardh (loc. cit.).

J. G. Agardh has proposed a segregate from Punctaria, viz.,
Homoeo'stroma to receive the plant from Cherbourg, described and
figured as Punctaria- latifoUa by the French phycologists, Thuret and
Bornet (1878,p . 13, pi. 5). Whereas both the Punctaria latifolia and
the P. plantaginea of Greville have the external layer on each surface
of the frond distinctly smaller and more deeply colored than the
internal layers, in the P. latifolia of Thuret and Bornet (but not of
Greville) the external layers are of the same sized cells as those of the
interior and slightly, if at all, more deeply colored. In our experience,
however, there are forms difficult of placing in case Punctaria and
Homoeostroma are considered separate genera.

There remain for consideration the relations of Punctaria and
Desmotrichiim. Kuetzing (1845, p. 244) founded the latter genus on
D. balticum, a slender, yet membranaceous form with hairs borne
singly and gametangia conical and practically superficial, at least
judging from his figures (Kuetzing, 1843, pi. 20 II, figs. 2, 4, 5-8).
Reinke, however, considers Kuetzing 's D. halticum to be a mono-
siphonous species although rarely it may be two to four cells wide,
while D. undulatum (J. Ag.) Reinke is the broader species. Aside
from the slender character of the frond, the strongly projecting or
even superficial gametangia and the hairs borne singly seem to dis-
tinguish the genus. We find, however, strongly projecting, even
superficial gametangia in broad species with hairs in groups. We also
find certain delicate, but broad species with single, promptly deciduous
hairs, and we find species in which there is no indication of hairs,

1925] Setchell-Gardner : Melanophyceae 517

even traces of deciduous ones. Consequently we do not feel certain
of any constant difference among' these three genera, viz., Punctaria,
Homoeosiroma and Desmotrichum. We also do not find any definite
distinctions to separate Shuh's Rhadinocladia (1900, pp. Ill, 112)
from the simpler Desmotrichums (cf. especially Kylin, 1907, pp.
66-68), nor are we able to distinguish satisfactorily Kuetzing's Diplo-
stromium (1843, p. 298) while J. G. Agardh's Nemataphloea (1896,
p. 12), if Kuetzing's figure (1856, pi. 45, I) is to be considered typical,
seems to have nothing to distinguish it from Kuetzing's Phycolapathum
(excluding synonymy of type species, pro parte) and Pwnctaria.
Phycolapathum Kuetzing (1843, p. 299, pi. 24, II) is founded on
Greville's Punctaria I at i folia, although Laminaria debiHs Ag. is
quoted, possibly erroneously, as a synonym. The plate of Kuetzing
represents a Pwnctaria. We have, therefore adopted the name Punc-
taria for all of our species, but we have seen none of the slender
Desmotrichum type on our coast. Punctaria has received the sanction
of the International Botanical Congress (Rules, 1912, p. 76) for the
generic nama.

Key to the Species

1. Fronds with numerous marginal crenulate lobes 2

1. Fronds with entire, smooth or undulate or coarsely ruffled margins 3

2 Fronds large, up to 50 cm. long, chartaceous, composed of 6-7 layers

of cells 6. P. chartacea (p. 521)

2. Fronds small, 10 cm. long, composed of 4 layers of cells.. 4. P. lobata (p. 519)
3. Fronds broadly expanded, up to 30 cm. broad, very fragile....?. P. expansa (p. 521)

3. Fronds narrower, up to 10 cm. broad 4

4. Fronds narrowly linear, distinctly stipitate, smooth, entire 5

4. Fronds irregular, more or less broadly expanded, obscurely stipitate,

with undulate or ruffled margins 6

5. Fronds epiphytic, base broadly cuneate, apex not laciniate..l. P. hesperia (p. 517)
5. Fronds saxicolous, long stipitate, base narrowly cuneate, the apex deeply

laciniate 2. P. fissilis (p. 518)

6. Fronds with undulate, much ruffled margins .5. P. occidentalis (p. 520)
6. Fronds with smooth or shghtly undulate margins. .3. P. latifolia (p. 519)

1. Punctaria hesperia S. and G.

Plate 37, fig. 30, and plate -19, fig. 18

Fronds 1.5-2.5 cm. high, 5-10 mm. wide, solitary or in small
clusters, tapering at the base to a very short stipe and to a blunt apex,
35-50/* (up to 80/1 ) thick, composed of 4-6 layers of cells, those of the
two middle layers much larger than those of the cortical layer ; cortical
cells thin walled, 4-5 sided in surface view, 18-22/a diam. containing
numerous parietal disk-shaped or small band-shaped chromatophores,

518 University of California Piiblications in Botany [Vol. 8

medullary cells thin walled, closely joined, the length and breadth
dimensions greater than the cross-dimensions, containing a few disk-
shaped chromatophores ; zoosporangia numerous, promiscuously scat-
tered among the gametangia on both surfaces of the frond; game-
tangia very numerous, closely congested at times or scattered without
order ; projecting hairs very delicate, in small groups, early deciduous.

Growing on Phyllaspadix. Pacific Grove, California.

Setchell and Gardner, Phyc. Cont., 1924, p. 3.

This species is apparently not abundant, however the plants are
small and inconspicuous, and quite probably ephemeral. Of all of
our species, thus far discovered, this one fruits most abundantly. The
larger part of the surface cells become transformed into reproductive
cells. The type, as stated above, comes from Pacific Grove, but we
have two other collections from our territory, viz., growing on Pliyl-
lospadix at Victoria (Gardner, no. 3849&) and one from San Pedro
(Mrs. H. D. Johnston, Herb. Univ. Calif., no. 99456) on Zostet^a which
may be referred here, at least provisionally.

2. Punctaria fissilis S. and G.

Fronds frag:ile and flabby, ligulate, 12-15 cm. high. 10-16 mm.
broad, 65-70/x thick, tapering at the base to a very delicate distinct
stipe 10-14 mm. long, attached by a small parenchymatous disk, light
brown in color, more or less deeply lacerated into linear laciniae, com-
posed of four layers of cells of fairly uniform size arranged in more
or less longitudinal rows, with thin walls and very small disk-shaped
chromatophores arranged closely along the walls ; zoosporangia and
gametangia unknown; hairs present in small groups.

Port Clarence, Alaska,

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 4.

The flaccid nature of the fronds, the long, delicate stipe, their
laciniate habit, the arrangement of the cortical cells in fairly regular
longitudinal rows along with the other combination of characters,
characterize this as a distinct species. P. fissilis seems to have its
nearest relations with P. glaciale Rosenv. and P. plantaginea (Roth)
Grev. We have not been able to examine any of the material from
Yakutat Bay collected by Saunders and reported under P. plantaginea
(1901, p. 420). We have not seen any typical P. plantaginea from our
territory and suspect that possibly his P. plantaginea may be found
to belong to P. fissilis.

1^^5] SetcheJI-G arrlner : Mclanophyreae 519

3. Punctaria latifolia Grev.

Fronds gregarious, flaccid, adhering firmly to paper, 1-3 dm. high,
2-7.5 cm. wide, 70-85/^, thick, linear to oboviite, attenuated at the base ;
color yellowish to light brown ; composed of four layers of cells, the
outer layers smaller than the inner ; gametangia projecting but slightly
above the surface ; zoosporangia unknown.

Occasional in quiet sandy coves (fide Saunders).

Greville, Alg. Brit., 1830, p. 52 ; Setchell and Gardner, Alg. N.W.
Amer., 1903, p. 240. Hamoeostroma latifoUum Saunders, Alg. Harri-
man Exp., 1901, p. 420.

We have examined a few small specimens of a Punctaria collected
by Saunders at Metlacatla and on Baranoff Island, Alaska, and pub-
lished {loc. cit.) as P. latifolia, which seem to belong to this species.
We are referring them here with some doubt on account of their
much reduced size as compared with Wyatt's no. 9, of the Algae Dan-
monienses, which we are taking as fairly typical of the species. The
measurements of the cross-section of the fronds and the shapes and
sizes of the cells composing the four layers agree very well with those
of her distribution, both being 70ju, to 80/^ in diameter.

4. Punctaria lobata (Saunders) S. and G.

Fronds broadly linear, lanceolate or ovate, 10 or more cm. long, 1-5
cm. wide, narrowed below to a short stipe of a dark olive brown color,
composed of 4 layers of cells, the 2 inner layers larger than the outer
layers; edges deeply and irregularly lobed, the lobes irregularly cut
and divided ; zoosporangia scattered over the whole surface of the
plant; gametangia unknown.

Attached to Zostera marina. Prince William Sound and Sitka,
Alaska.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 12. Homoeostroma
lobattom Saunders, Alg. Harriman Exp., 1901, p. 420.

We have not seen any material belonging to this species and are
including it here without comment, having previously made the above
combination {lac. cit.).

520 University of California Publications in Botany [Vol. 8

5. Punctaria occidentalis S. and G.

Plate 35, fig. 6, and plate 80'b

Fronds moderately rigid, linear-lanceolate or oblanceolate to
broadly ellipsoidal, usually tapering at the base to a short stipe about
5 mm. long, attached by a small disk, 1.5-2 dm. high, 3-10 cm. broad,
40-180jU, thick, with undulate and coarser ruffled margins, of a light
brown color, soon changing to green on exposure, composed of 2-7
layers of cells, thickest in the median region, gradually thinner toward
the margins ; cells of medulla largest diminishing slightly toward the
surfaces, cortical cells being the smallest ; zoosporangia cuboidal to
subspherical, scattered on both sides of the frond, 30-40jU, (up to
70ju,) diam., very slightly bulging on the outer surface; gametangia
numerous, scattered promiscuously on both sides of the frond among
the zoosporangia and usually protruding beyond the surface about
one-half their length ; hairs unknown.

Growing on Zostera, in the upper sublittoral belt. Monterey,
California.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 4. HomoeostroTna
Jatifolia Saunders, Phyc. Mem., 1898, p. 159, pi. 30, figs. 4, 5. Punc-
taria latifoUa Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. 873; Tilden, Amer. Alg. (Exsicc), no. 355.

We have not been able to observe the young fronds of this species,
but the fruiting fronds have no hairs so far as we have been able to
ascertain. Rosenvinge (1910, p. 121) has found a similar condition
in his Punctaria glacialis. The surface is moderately sticky and the
plants adhere firmly to paper on drying.

As cited above, this species has been referred to P. laiifolia since
its discovery on our coast some years ago. We have not been able to
examine any of the material upon which Greville founded the species.
We have, however, examined a specimen distributed by Wyatt (no. 9)
from Torbay on the southeast coast of England near the type locality.
Since no other species have been cited from this general locality, and
Wyatt 's specimen agrees so well with Greville 's original description,
we are assuming this to be typical P. latifolia and using it as a basis
of comparison. Our plants are much larger and thicker and have
a larger number of layers of cells all of which are much more variable
in shape than those of Wyatt 's distribution. The cortical cells which
do not produce zoosporangia. and gametangia usually divide hori-

1925] Setchell-Gardner : Melanophyceae 521

zontally into two. The fructiferous cells remain undivided. The
undivided cortical cells are about the same shape and size as the
medullary cells which also vary in shape and size.'

6. Punctaria chartacea S. and G.

Plate 45, fig. 79, and plate 81

Fronds more or less finely buUate, membranaceous, chartaceous,
linear to broadly elliptical, base often cordate, margins finely fimbriate,
25-35 cm. (up to 50 cm.) long, 8-12 cm. (up to 30 cm.) broad, mature
plants 90-120ja thick, attached by a very small disk ; composed of 6-7
layers of cells, the surface cells being much smaller than the medullary
cells; cell walls thick, of light brown color, with intercellular spaces
among the medullary cells ; color dark brown, not changing on drying,
not adhering at all to paper; hairs, zoosporangia and gametangia
unknown.

Growing on Eel grass in the lower littoral and upper sublittoral
belts. Sitka. Alaska.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 4.

This species of Punctaria was found in great abundance, both
attached and floating. A large number of specimens were examined
in the last part of June and no reproductive cells could be found.
Neither were any hairs observed.

7. Punctaria expansa S. and G.

Plate 49, fig. 19

Fronds gregarious, broadly expanded, more or less circular in out-
line, 20-30 cm. broad, 100-130/i, thick, very fragile, of medium brown
color, fading to green very soon after removal from water, composed
for the most part of four layers of cells, the medullary cells being
much larger than the cortical cells; neither the medullary nor the
cortical cells arranged in any definite series ; cortical cells four to six
sided and 30-40;u, diam. in surface view ; zoosporangia and gametangia
on the same frond, the former projecting but slightly beyond the sur-
face, the latter more or less conical and projecting one-half or more of
their length beyond the surface ; hairs present but sparse.

Growing in a small quiet cove at the head of Penn's Cove, Whidbey
Island, Washington.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 5. Punctaria
latifolia Setchell and Gardner, Alg. N.W. Amer., 1903, p. 240 (in part).

522 University of California Puhlications in Botany [Vol. 8

Punctaria expansa is apparently a very rare though distinct species,
the foregoing material referred to being the only collection known.
The habit of growth' is the same as that of a group of fair sized Vivas.
It was found growing in a secluded cove in which the water rises and
falls but slightly through a small channel with the change of tide, and
is not disturbed by the action of wind. The plants are exceedingly
fragile, being scarcely tough enough to hold their own weight when
first removed from the water. They give up their brown color soon
after being removed from the water when they superficially resemble
almost perfectly an expanded Viva. The fruit, especially the zoo-
sporangia in our specimens, is decidedly sparse, likewise the hairs.

We have been able to examine a few small young specimens of
Homoeostroma undulatum collected and thus labeled by Saunders
(1901, p. 419). They are not fruiting, are densely covered with
diatoms and do not soak out readily. We are, therefore, on account
of the state of the material, unable to place it and hence are not
including it in our list of species.

FAMILY 8. ASPEEOCOCCACEAE foslie •

Fronds ligulate or saccate, simple, differentiated into two kinds of
tissue, an inner one, composed of nearly colorless cells and an outer
cortical color bearing one ; growth trichothallic at first, later inter-
calary ; hairs present, and in some species, paraphyses ; zoosporangia
on full-sized, or macroscopic, plants ; gametangia, so far as known, on
reduced, but not microscopic plants, not so definitely collected into
sori as is the case with the zoosporangia ; both zoosporangia and game-
tangia projecting from the surface and collected into sori scattered all
over the frond except at the base.

Foslie, Mar. Alg. Norway, Cont. I, 1890, p. 88. Asperoco'cceae
Zanard., Sagg. Class. Nat. Ficee, 1843, p. 10 (in part) ; Farlow, Mar.
Alg. New England., 1881, p. 88.

The Asperococcaceae, as limited to those ligulate, bullose, or nearly
spherical forms, solid or hollow, with the zoosporangia in distinct and
elevated sori, projecting and accompanied, usually at least, by distinct
paraphyses or hairs and in having the gametangia, so far as known, in
usually less' definite sori and on plants more or less reduced in size,
but still macroscopic, is closely related in its early trichothallic, later
intercalary growth to the Striariaceae, Punctariaceae, and Scyto-
siphonaceae, so that it is occasionally difficult to determine exactly

1925] Setch en-Gardner: Melanophyceae 523

where to place a particular genus. The genus Asperoeoceus, as
exemplified by A. hullosus Lamour. is distinct and definite as is also
Soramthera ulvaidea Rupr., and both of these are typical as to hollow
frond, manner of growth and type of sorus which is definite in outline,
with paraphyses and zoosporangia intermingled and with a group of
hairs more or less centrally located. The genus Halarhipis of Saunders
has also projecting sori with zoosporangia, but H. Winstonii (C. L.
Anderson) Saunders has less definite sori and is without paraphyses
but provided with hairs. The Californian species seems to belong in
Asperococcaceae because of its superficial and projecting zoosporangia,
and because of its general structure and appearance ought to be placed
near to Asperoeoceus compresses GrifT. We have seen no gametangia
on the plants of any of our species, but judging from the statements
of Buffham (1891), Sauvageau (1895), and Kuckuck (1899), the
gametophyte may be very much reduced, a tendency toward, but by
no means approximating, the microscopic gametophyte known to occur
in some of the other Melanophyceae.

Key to the Genera

1. Fronds ligulate, solid IS. Halorhipis (p. 523)

1. Fronds cylindrical or more or less spherical, hollow 2

2. Fronds cylindrical or shghtly clavate, more or less twisted

20. Myelophycus (p. 527)

2. Fronds spherical, sessile 19. Soranthera (p. 525)

18. Halorhipis Saunders

Fronds solid, ligulate, arising from a disk-shaped holdfast, differen-
tiated into two fairly distinct tissues, a medulla of several layers of
larger, nearly colorless, closely united, more or less cylindrical cells
extending lengthwise of the frond, and surrounding this a cortical or
surface tissue composed of more or less cuboidal, color bearing cells ;
zoosporangia superficial among hairs and collected into minute sori
scattered abundantly over both sides of the frond except toward the
base ; gametangia unknown.

Saunders, Phyc. Mem., 1898, p. 160.

Halorhipis was proposed by Saunders to receive a single species
found at Pebble Beach, Monterey County, California. The genus is
closely related to Haloglossum. Kuetz.

The genus Haloglossum was founded by Kuetzing (1843, p. 340)
based upon the material of Asperoeoceus eompressus Griffiths {in
Hooker, 1833, p. 278) from the southern shores of England. He

524 University of California Publications in Botany [Vol. 8

changed the name to Haloglossum GriffUhsianum which accordingly
becomes the type of the genus. The chief distinction between Halo-
glossimi and Asperococcm lies in the character of the interior of the
frond. The former is ligulate and solid and the latter is saccate. Mrs.
Griffiths expressly states {loc. cit.) "there is not the slightest tendency
to be tubular or inflated," in speaking of A. compressus. We feel
inclined to keep the two genera separate, based mainly upon this
character, although the method of reproduction in the two is similar.
The genus Halorhipis is very similar in appearance and structure to
Haloglossum, but the sori are destitute of paraphyses although the
protruding zoosporangia are aggregated about a cluster of hairs.

Halorhipis Winstonii (C. L. Anderson) Saunders

Plate 35, fig. 8

Fronds mostly aggregated into tufts, thin, membranaceous, lanceo-
late to obovate or spatulate, 8-20 cm. high, 2-5 cm. wide, about 200/i,
thick, with very slender filamentous stipes ; color dark olive green ;
zoosporangia collected into sori mostly in slight depressions, ellipsoidal,
obovate or pyriform, 30-45,ju, long, 20-30/* broad ; sori numerous,
variable in shape and size, linear to irregular in outline ; hairs in
groups in the sori ; gametangia and paraphyses unknown.

Growing on the fronds of Egregia Menziesii and on rocks. Known
only from a single locality, Pebble Beach, Carmel Bay, California.

Saunders, Phyc. Mem., 1898, p. 161, pi. 28; Collins, Holden and
Setchell, Phyc. Bor.-Amer. (Exsicc), no. 532. Punctaria Winstonii
C. L. Anderson, Some new and some old algae, 1894, pp. 358, 359.
Aspero'coccus Winstonii Svedelius, in Engler and Prantl, Natiirl.
Pflanzenfam., Nachtr. z. 1 Th., Abt. 2, 1910, p. 158 (by implication).

This plant, although known for only a little over a quarter of a
century, has had a somewhat varied history. It was first described
by C. L. Anderson (loc. cit.) who saw in it affinities with the genus
Punctaria Grev. Saunders (loc. cit.) next studied it and came to
the conclusion that a new genus, Halorhipis, should be erected for its
reception. It was distributed in the same year under that name in
the Phycotheca Boreali-Americana as no. 532. In 1910, as above
stated, Svedelius placed it in the genus Asperococcus Lamour. Our
studies, while leading us to retain the genus Haloglossum Kuetzing,
discarded by most of the later phycologists, and to place Asperococcus
compressus Griff, in that genus as a species distinct from all others.

1925] Setchell-Gardner: Melanophyceae 525

yet also lead us to keep Halorhipis Winstonii distinct because of the
lack of the pluricellular paraphyses so characteristic of both Aspero-
cocctis and HaJoglossum. It differs from a Punctaria, as we under-
stand that g'enus, in having: the thallus decidedly differentiated into
two sorts of tissues, and in having the zoosporangia superficial.

19. Soranthera Post, and Rupr.

Fronds spherical and solid in the juvenile stage, later becoming
hollow, inflated and membranaceous, sessile, attached by a rather
broad and penetrating base, composed of two kinds of tissue, an
inner one 4-5 cells deep, of large, nearly colorless cells, and a surface
layer of small, angular, color bearing cells ; reproduction by zoo-
sporangia in very distinct, conspicuous sori, scattered promiscuously
over the surface of the frond ; paraphyses pluricellular ; hyaline hairs
grouped in the center of the sori ; gametangia unknown.

Postels and Ruprecht, Illustr. Alg., 1840, p. 19.

The type species of the genus is S. ulvoidea discovered "Ad
insulam Sitcha, parasitica in fronde Rhodomelae Laricis," (fide
Postels and Ruprecht, Joe. cit.). The question of parasitism of the
genus has been investigated by Miss E. S. Barton and was discussed
by her in an article in the Journal of the Linnaean Society (1898,
p. 479). She questions the statement quoted above from Postels and
Ruprecht as to the parasitism of the species, taken in the sense that
we understand parasitism. However, her investigation shows that
the fronds are attached by a mass of rhizoidal filaments which sur-
round the host, many of which actually penetrate into the cells even
far into the interior of the host. She saw and figured what she inter-
preted as ' ' haustoria. ' ' Whether these plants and many others among
all four of the large sections of the algae which penetrate in a similar
waj', actually obtain food from the host has not been satisfactorily
determined, so far as we are aware.

Soranthera ulvoidea Post, and Rupr.

Fronds membranaceous, comparatively thin, oval, ellipsoidal or
nearly spherical, entire or irregularly lobed, up to 7 cm, diam. ; color
olive brown ; sori abundant, conspicuous, distributed fairly evenly
over the whole free surface of the frond ; zoosporangia clavate, 70-100/x
long, interspersed among numerous clavate, pluricellular paraphyses
which surpass them in length.

526 University of California Puhlications in Botany [Vol. 8

Growing on species of Rhodomela and Odonthalia, frequently in
tide pools, in the littoral belt. Common along the whole coast from
Unalaska, Alaska, to southern California.

Postels and Ruprecht, Illus. Alg., 1840, p. 19 ; Kjellman, Om
Beringh. Algfl., 1889, p. 47, pi. 7, figs. 4, 5; Saunders, Ph-yc. Mem.
1898, p. 165, pi. 29, figs. 4, 5 ; Setchell and Gardner, Alg. N.W. Amer.,
1903, p. 244. ■

Two forms have been segregated as follows :

Soranthera nlvoidea f. typica S. and G,

Plate 39, figs. 40, 41, and plate 836

Fronds nearly regular in outline, membrane thicker, oval to
spherical, up to 7 cm. diam.

Growing on species of Rhodomela and Odonthalia. From Pacific
Grove, California, northward into Bering Sea, and more common in
the southern part of the range than f . dijformis.

Setchell and Gardner, lac cit.; Kjellman, loc. cit.; Saunders, loc.
cit., Alg. Harriman Exp,, 1901, p. 422 ; Collins, Holden and Setchell,
Phyc. Bor.-Amer. (Exsicc), no. 417; Tilden, Amer. Alg. (Exsicc),
no. 245.

Soranthera ulvoidea f. difformis S. and G.

Fronds gregarious, membrane thinner, usually smaller than the
above, variously and deeply lobed.

Growing on the same hosts as the two preceding, but is more
common on the northern part of the range, from Cape Platterj^
northward.

Setchell and Gardner, Joe. cit.; Collins, Holden and Setchell, loc.
cit., no. 1130.

Forma difformis seems to prefer either brackish or muddy water or
is induced by such medium.

Soranthera ulvoidea Post, and Rupr. is a very common species
along the entire Pacific coast of North America from Pacific Grove,
California, northward. Careful search has failed to reveal the
gametangia.

1925] Setchell-Gardner: Melanaphyceae 527

20. Myelophycus Kjellm.

Fronds filiform, attached by numerous rhizoidal filaments densely
interwoven, solid throughout when young, later becoming hollow
above; growth trichothallic (?), later usually intercalary; fronds
composed of three kinds of tissue, an inner, colorless layer of a few
rows of large, rounded, isodiametric, or elongated cells, outside of
which is a layer of small prismatic, thick-walled cells giving rise to a
dense mass of erect, assimilating filaments (paraphyses ?) ; zoo-
sporangia immersed among the assimilating filaments; gametangia
unknown ; hairs present in the juvenile stage.

Kjellman, m Engler and Prantl, Natiirl. Pflanzenfam., 1893, 86
Lief., p. 202, Om Fuc. Mtjelophijcus, 1893, pp. 1-11.

The type species of the genus is M. caespitosum from the coast of
Japan. The species of our coast seems to agree in all generic require-
ments. Kjellman has followed the neuter gender of the Greek noun,
phykos, although the Latin form would indicate a grammatical
masculine. It seems best, however, to follow Kjellman as being more
in accord with the proper usage.

We place this genus in Asperococcaceae with considerable doubt.
The species may prove to be the zoosporangial forms of gametangial
plants similar to Chordaria Giinjii Yendo.

Myelophycus intestinale Saunders

Fronds solitary or loosely caespitose, 1-15 dm. high, 2-4 mm. diam.,
solid in the juvenile stage, hollow at maturity above the short, solid
stipe, cylindrical, abruptly attenuated at the tip, but later eroded,
usually twisted, unbranched, dark reddish brown in color; central
tissue composed of 2-3 layers of elongated, cylindrical, colorless cells
merging toward the outside into 3-4 irregularly arranged rows of
thick-walled, cuboidal cells, these in turn giving rise to numerous
paraphyses composed of 4-8 thick-walled cells and to zoosporangia ;
zoosporangia ellipsoidal to obovate scattered over the frond except the
stipe, 45-60ju long, 20-30/t broad.

Growing attached to rocks in the lower littoral and upper sub-
littoral belts. Shumagin Islands, Alaska, to Puget Sound, Washington.

"Myelophycus intesimalis" Saunders, Alg. Harriman Exp., 1901,
p. 420 ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 241.

528 University of California Puhlications in Botany [yo-L. 8

Myelopliycus intestinale presents a different aspect when young
from that of the older plant. This is seen particularly at the tips
which are long acuminate when young, but blunter and often worn
away in age. The younger plants seem less conspicuously^ twisted than
the older. Our plant has fewer layers of cells in the intermediate
layer and shorter cortical filaments than the M. caespitosum of Japan.

Myelophycus intestinale f. tenue S. and G.

Plate 40, fig. 50

Fronds densely caespitose, inconspicuously twisted, 1.5-2.5 cm.
high, 0.25-0.75 mm. diam. ; zoosporangia broadly ellipsoidal, 40-45/x
long, 30-35/A broad.

Growing on rocks, usually in shaded localities, and where the spray
dashes against the rocks, along high-tide level. Coos Bay, Oregon, to
central California.

"Myelophycus intestinalis f. tenuis" Setchell and Gardner, in
Gardner, New Pac. Coast Mar. Alg. I, 1917, p. 385.

FAMILY 9. STRIAEIACEAE kjellm. .

Fronds filiform, solid, more or less branched, trichothallic, mono-
siphonous at first, later usually of two sets of tissues, the inner of
larger, colorless, more or less elongated cells, the outer a single layer
of medium sized cells placed in horizontal rows and having chromato-
phores; unilocular zoosporangia and plurilocular gametangia super-
ficial or projecting beyond the surface in sori of more or less definite
circumscription and usually in transverse rows; with or without
paraphyses,

Kjellman, Handbok I, 1890, p. 53.

It seems best to follow Kjellman and keep the seemingly very
distinct genera, Stictyo siphon, Striaria and their near relatives
separate from the Asperococcaceae and the Scytosiphonaceae and
retain them among the Striariaceae. Both the gametophytes and the
sporophytes are known and are similarly macroscopic, but conditions
are otherwise in the other two families just mentioned. In the
Asperococcaceae there are dissimilarities in size, at least in some
species, between the individuals bearing gametangia and those bear-
ing zoosporangia, although neither is properly microscopic so far as
known. Among the Scytosiphonaceae, only the gametangial form is
known, suggesting that the zoosporangial form may be heteromorphic.

1925] Setchell-Gardner: Melanophj/rcae 529

whether microscopic or eliminated by some method of syncopation
may not even be surmised.

Two genera have been reported from the western coast of North
America, but there is so much doubt about one of them (Striaria,
as indicated below) that we have taken only one under serious
consideration.

21. Striaria Grev.

Greville, Algae Brit. Syn., 1830, p. XLTIT. Striaria aitentmia
(Ag. ) Grev. Greville (loc. cit.). Solenia attenuata Agardh., Syst.,
1824, p. 187.

A hand lens examination of the single scanty specimen from Orcas
Island, Washington, in the Herbarium of Trinity College, Dublin,
referred to Striaria attenuaia by Harvey (1862, p. 160) left the
impression that it may belong rather under Stictijo siphon tortiJis than
under Striaria. It was very slender and seemed to be sterile.

22. Stictyosiphon Kuetz.

Fronds attached by numerous branched rhizoids, mostly growing
in dense clusters, filiform, branched, terminating in one or more
hyaline hairs, solid in the juvenile stage, fistulose below, growth inter-
calary ; plant body consisting of parenchymatous cells, the interior of
more or less elongated, rectangular, colorless cells and the exterior of
a single cortical layer of color bearing cells ; reproduction by unilocular
zoosporangia and plurilocular gametangia produced by the trans-
formation of certain cortical cells, solitary or more or less aggregated,
borne on similar macroscopic plants.

Kuetzing, Phyc. Gen., 1843, p. 301.

The type species of the genus is S. adriaticus with the type locality
Trieste. The northern species added later seem to belong under the
same genus so that there is no necessity for retaining the genus
Phloeosporaoi Areschoug (1873, p. 164). The species of Stictyosiphon
differ principally from those of Striaria in not having paraphyses
intermingled with the reproductive organs in their sori.

Stictyosiphon tortilis (Rupr.) Reinke

Fronds filiform, long attenuate, closely aggregated, repeatedly and
alternately branched, the branches bearing minute ramuli mostly
opposite, 5-8 cm. high, often with a small, longitudinal, central cavity
in the lower portions; cortical cells with branched band-shaped chro-

530 Vniversity of California Publications in Botany [Vol. 8

matophores ; color dark olive brown ; zoosporan^ia and gametangia
scattered or aggregated into more or less transversely arranged groups.

Port Clarence, Golofin Bay and Unalaska Bay, Alaska.

Reinke, Algenfl. westl. Ostsee, 1889a., p. 55, Atlas, 1889, p. 47, pis.
31, 32; Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no.
987. Phloeospora tortilis Setchell and Gardner, Alg. N.W. Amer.,
1903, p. 245. Scytosiplion tortilis Ruprecht, Tange, 1851, p. 373.

The plants of Bering Sea appear to belong in the form cycle of
this Arctic species. We seem to remember having seen specimens
from farther south along our coast, but cannot definitely place them.

FAMILY 10. SCYTOSIPHONACEAE foslie

Fronds cylindrical and hollow, flattened and solid, or more or less
globular and hollow, main growth in length at first trichothallic, later
intercalarj% simple, of two or three tissues, one of internal hyphae in
Endarachne, but of large colorless cells in others, second, an inter-
mediate layer of large colorless cells, surrounded by an outer layer
of small, mostly cuboidal, colored cells in longer or shorter anticlinal
rows ; zoosporangia unknown ; gametangia in extended and confluent
sori of indefinite outline, palisade-like, of equal length and closely
packed together, not projecting beyond the surface ; hairs in groups ;
unicellular paraphyses ( ?) present in some species.

Foslie, List Mar. Alg. Isle of Wight, 1892, p. 13. Scytosiphoneae
Thuret, in Le Jolis, Liste des alg. mar. de Cherb., 1863, pp. 14 and 67.

Thuret seems to have been the first to have stabilized the genus
ScytosipJion and to have recognized it as the type of a group cor-
responding to a family. Foslie, so far as we may learn, first used the
family ending as now usually adopted.

The Scytosiphonaceae are close to the several preceding families as
has been noted previously. The method of growth is very similar, but
only the g-ametangia are known in the genera attributed to this family
and they are in expanded and indefinite sori which do not project
above the surface of the frond nor are they associated with pluri-
cellular paraphyses. The Punctariaceae possess the simplest members
of this series and they have both zoosporangia and gametangia, these
commonly produced on the same individual. In the Asperococcaceae,
gametangial plants are rarely found and are smaller than the zoo-
sporangial plants. In the Striariaceae both zoosporangial and game-
tangial plants are found. While in the Scytosiphonaceae, although
most of the species are common and widespread, no zoosporangial
plants have been detected.

1925] Setchell-Gardner : Melanophyceae 531

Key to the Genera

1. Fronds solid and strap-shaped 2

1. Fronds hollow and cylindrical, flattened, globose or difform 3

2. Interior composed of hyphal tissue 25. Endarachne (p. 538)

2. Interior composed of parenchymatous tissue 24. Hea (p. 535)

3. Fronds cyhndrical or flattened 23. Scytosiphon (p. 531)

3. Fronds globose or difform 4

4. Membrane entire 26. Colpomenia (p. 539)

4. Membrane reticulately perforated 27. Hydroclathrus (p. 542)

23. Scytosiphon Ag. (emend. Thuret)

Frond unbranched, solid when young, later tubular, cylindrical or
compressed, constricted at short intervals or smooth throughout, con-
sisting of two layers of cells, an inner of thick-walled, vertically
elongated, colorless cells, and an outer of small, rounded, cuboidal,
assimilating cells, the latter giving rise to the plurilocular gametangia
covering the entire surface, except at the base, and to scattered uni-
cellular paraphyses (?) and hairs when present; growth intercalary
near the base.

Agardh, Dispos. Alg. Suec, 1811, p. 24 (fide Pfeiffer) (pro parte) ;
Thuret, Rech. sur les zoospores, 1850, p. 239.

The genus Scytosiphon originally included what we now call
Dictyosiphon foenicidaceus, Chorda Filum and Scytosiphon Lomen-
taria. Thuret seems to have been the first to restrict the generic name
to the Chordaria Lonientaria Lyngbye and to point out the generic
characteristics. 0. Kuntze (1898, pp. 430 and 434) has proposed the
generic name Tuhicutis but the International Rules (Briquet, 1912,
p. 76) decided in favor of the retention of Scytosiphon.

Scytosiphon Lomentaria (Lyngb.) J. Ag.

Plate 44, figs. 72, 74

Fronds usually fasciculate, sometimes solitary and scattered, 15-30
cm. (up to 60 cm.) high, 3-6 mm. diam., cylindrical, at intervals more
or less deeply constricted, attenuated below into a small solid stipe,
attached by a small disk ; color dark brown to olive green ; plurilocular
gametangia forming a compact palisade layer over the surface of the
frond; paraphyses (?) scattered among the gametangia.

Growing on rocks in the whole of the littoral belt. Common along
the entire Pacific coast from Port Clarence, Alaska, to southern
California.

532 Unwersity of California Publications in Botany [Vol. 8

J. G. Agardh, Sp. Alg., vol. 1, 1848, p. 126 ; Thuret, Rech. sur les
zoospores, 1850, p. 239 (1851, p. 30, Repr.), pi. 29, figs. 1-4; Setchell,
Alg. Prib. Islands, 1899, p. 591 ; Saunders, Phyc. Mem., 1898, p. 163,
pi. 31, figs. 8-10, Alg. Harriman Exp., 1901, p. 421; Setchell and
Gardner, Alg. N.W. Amer., 1903, p. 243 ; Okamura, Icon. Japan. Alg.,
1908, p. 144, pi. 30. Chorda Lomentaria Lyngbye, Hydrophyt. Dan.,
1819, p. 74, pi. 18, E. ■

Scyto'siphon Lomentaria, if all the forms referred to it belong under
it, is a widespread and variable species. It varies from a centimeter
or two to twenty-five or more centimeters in height and from a milli-
meter or two up to ten millimeters in thickness. It may vary from
clearly cylindrical to decidedly flattened and it may vary from regu-
larly or irregularly and pronouncedly constricted (especially in the
larger plants) to those with no trace of constriction whatever. It
may be covered almost uniformly with gametangia or, rarely, the
gametangia may be segregated into distinct and limited sori. The
gametangia may be longer or shorter, with variations from two up to
twenty-five or more longitudinal uniseriate loculi. It has been the
custom to separate the var. complanata Rosenvinge, but this occurs in
a major and minor form, as do all the variants with the exception per-
haps of the typical form (f. typicus) which is constricted, but while
this varies much in size, it never occurs among the true minor forms
of the species. The larger and smaller forms of each series of forms
differ from one another in the size of the cells of the inner layer, and
in the length (absolute and in number of loculi in each longitudinal
series) as well as in diameter of the gametangia. In order to coordinate
the variables as mentioned above, we have tabulated them in the fol-
lowing key, rather to give a view of the combinations of characters
than that they should be considered of any definite taxonomic import-
ance, since intermediate forms are, by no means, absent.

Key to the Forms

1. Plants constricted at intervals, medium to large 1. f. typicus (p. 533)

1. Plants not constricted 2

2. Plants cylindrical 3

2. Plants complanate 5

3. Plants 6 cm. or more high; interior cells large; gametangia long

2. f. cylindricus major (p. 533)

3. Plants less than 6 cm. high; interior cells small; gametangia short 4

4. Sori covering the frond 3. f. cylindricus minor (p. 533)

4. Sori restricted, definite 4. f. cylindricus maculatus fp. 534)

5. Plants large, 6 cm. or more; inner cells large; gametangia long

5. f. complanatus major (p. 534)

5. Plants small, less than 6 cm.; inner cells small; gametangia short

6. f. complanatus minor (p. 534)

1925] Setchell-Gardner: Melanophyceae 533

The occurrence or non-occurrence of hairs is such a variable char-
acter as is also the presence or absence of the ' ' unicellular paraphyses ' '
(possibly hypertrophied gametangia ?) in all of our variants of the
species that we have not found it possible to use either structure in
any of them as a diagnostic character of much significance.

1. Scytosiphon Lomentaria f. typicus S. and G.

Plate 39, fig. 45, and plate 44, fig. 75

Plants of the largest size of the genus, both as to length and
diameter, dark brown to olive green in color, more or less regularly
and deeply constricted; paraphyses ( ?) cylindrical to slightly clavate,
not projecting beyond the gametangia, with chromatophores often
collected at the outer end ; gametangia with 10-15 loculi.

Growing on rocks in pools in the lower littoral belt. Port Clarence,
Alaska, to southern California.

Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 323&;
Tilden, Amer. Alg. (Exsicc), no. 246.

2. Scytosiphon Lomentaria f. cylindricus major

Fronds from 6-25 cm, long. 1,5-2.5 mm. diam., not constricted,
cylindrical, long-attenuated at both ends, light brown in color.

Growing on rocks in the lower littoral or in tide pools in the upper
littoral belts. Central California.

Collins, Holden and Setchell, Phyc. Bor.-Amer, (Exsicc), no.
1389. Chordaria attenuata Tilden, Amer. Alg. (Exsicc), nos. 347a,
3476 ?

The cylindrical unconstricted forms seem to be fairly well estab-
lished both as regards habitat and form. They grow in tide pools,
often even above high-tide level, and the size seems to be materially
influenced by the size and depth of such pools. In these pools where
they are frequently to be found, they are generally far removed from
the typical form which grows usually in deep tide pools in the middle
and lower littoral belts.

3. Scytosiphon Lomentaria f. cylindricus minor

Fronds less than 6 cm. high, 1-1.5 mm. diam., cylindrical, uncon-
stricted.

Growing on rocks along high-tide line in localities exposed to wave
action. Central California (San Francisco).

534 University of California Puhlications in Botany [^^ol. 8

4. Scytosiphon Lomentaria f . cylindricus maculatus

Plate 42, figs. 61, 62

Fronds 3-4 em. hig^h, cylindrical, not constricted, dark brown in
color ; g'ametangia collected into definite, larger or smaller sori.

Growing on rocks exposed to heavy surf along high-tide level.
Central California (San Francisco).

Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1235.

This form is distinguished by the presence of gametangia in groups,
giving the small fronds a warty appearance. It grows in exposed
situations in which the rocks are kept more or less moist continuously
by the spray, but for the most part the plants are not submerged in
pools.

5. Scytosiphon Lomentaria f . complanatus major

Fronds flat, 9-14 cm, high, 1-2.5 mm. wide, dark brown in color,
soon changing to green on removal from the water.

Growing on rocks in the lower littoral belt. Pebble Beach, Carmel
Bay, Monterey County, California.

6. Scytosiphon Lomentaria f . complanatus minor

Plate 44, fig. 74

Fronds 3-5 cm. high, up to 1.5 mm. wide, irregular and variable
in thickness, cylindrical below and solid or hollow, flattened above
and hollow, light brown in color ; gametangia composed of 5-7 loculi,
completely dissolving at maturity leaving the medulla clean ; medulla
composed of two layers of cells, variable in size, much longer than
broad ; hairs and paraphyses absent.

Growing on stones along high-tide level or above. Fort Point, San
Francisco, California.

This form is a short-lived winter and spring form, usually matur-
ing in February and March, so far as our experience shows. It has
been detected for many years in the above mentioned habitat, often
found growing on wood as well as on a stone wall.

1925] Setchell-Gardner: Melanophyceae 535

24. Ilea Fries

Fronds solid, or rarely M'ith occasional small cavities, membran-
aceous, considerably flaccid, simple, ligiilate, tapering at the base to a
small, solid stipe, attached by a parenchymatous disk-shaped holdfast,
or by rhizoids, differentiated into two distinct tissues, a medulla com-
posed of more than one layer of mostly large, more or less colorless
cells and a cortex of small, cuboidal, color bearing cells ; the cortical
cells on both surfaces giving rise by numerous horizontal divisions to
gametangia with uniseriate loculi, gametes escaping by the complete
dissolution of the entire wall; zoosporangia unknown; paraphyses( ?)
and hairs present in some species.

Fries, Flor. Scan., 1835, p. 321 (pro parte) ; not Ilea Fries, Syst.
Orb. Veg., part 1 (pi. homon.), 1825, p. 336; not Ilea J. G. Agardh,
Till Alg. Syst., afd. 3, 1883, p. 114. Phyllitis Kuetzing, Phyc. Gen.,
1843, p. 342 (not Phyllitis Hill). Petalonia Derbes et Solier, Sur les
org. repr. des alg., 1850, p. 265.

As to the confusion and the different views concerning the adoption
of Ilea as the name for this genus, one may consult Nordstedt (1911,
p. 265) and M. A. Howe (1914, p. 51). Peialonm is preferable from
the point of view of possible confusion, but Ilea seems to have the
right of way. The genus tends toward complanate forms of Scyto-
siphon in certain forms which, while not completely hollow, never-
theless are not completely solid. The type species is Ilea Fascia
(Muell.) Fries, whose type specimens come from somewhere on the
coast of Norway. In the genus Ilea, a condition exists very similar to
that found in Scytosiphan and in Colpcmienia, that is, of a number of
fairly striking form-types between which it is extremely difficult to
make other than very arbitrary distinctions. We are inclined to
recognize a single species. Ilea Fascia, and refer the other described
species or forms to it.

Ilea Fascia (Muell.) Fries
Plate 44, figs. 68-71, 73

Fronds attached by a small, parenchymatous disk, often fasciculate
and several disks coalescing, thin, plane or at times more or less
crisped, linear-lanceolate to oblanceolate, often rounded and more or
less eroded above, 6-12 cm. (up to 25 cm.) high, mostly 1.5-2.5 cm.
wide, but at times up to 12 cm. wide, exceedingly variable in thickness ;

536 University of California Puhlications in Botany [Vol.8

gametes escaping by the simultaneous dissolution of the entire cell
walls of the gametangia.

Growing on rocks and on PhyUospadix in the littoral belt. Occur-
ring in its various forms from Unalaska, Alaska, to southern California.

Fries, Flor. Scan., 1835, p. 321. PJiyUitis Fascia. Kuetzing, Phyc.
Gen., 1843, p. 142, pi. 24, III, figs. 1-6, in Linnaea, vol. 17, 1843, p. 97 ;
Saunders, Alg. Harriman Exp., 1901, p. 421 ; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 243; Farlow, Anderson and Eaton, Alg.
Exsicc. Amer.-Bor., no. 199. Petalonia. Fascia Kuntze, Rev. Gen. PI.,
vol. 3, 1898, p. 419 ; Howe, Mar. Alg. Peru, 1914, p. 50. Fiicus Fascia
Mueller, in Flor. Dan., 1778, pi. 768.

So far as we have been able to ascertain, typical Ilea Fascia has
neither hairs nor paraphyses. The Pliyllitis Fascia of Okamura (1901,
pi. 10) shows a group of hairs among plurilocular gametangia. Later,
however, he distributed this plant under Endarachne Binghamiae, in
his Algae Japonicae Exsiccatae, no. 86, referring to his previous
publication (Joe. cit.) as a synonym.

The size of the plants, as noted by different authors, is exceedingly
variable, as the foregoing diagnosis shows. We have included plants up
to 25 cm. long, and up to 12 cm. wide. The thickness varies with age
in the same plant, and varies in different collections of plants,
Mueller's Fu-cus Fascia {loc. cit.), as illustrated, is about 20 cm. long
and 5 mm. broad. The f. zosterifalia in size, form, and in being inter-
ruptedly and slightly hollow, approaches ScytosipJion Lomentaria f.
complanatus of medium size.

We suggest the arrangement of the forms of the species in the
following key:

1. Frond broad, stipe distinct 2

1. Frond narrow to almost filiform, almost stipeless 3

2. Tapering gradually at the base to a short stipe .... 3. f. caespitosa (p. 537)

2. Tapering abruptly at the base to a short stipe 2. f. debilis (p. 537)

3. Frond flat, solid throughout 4

3. Frond with few small internal spaces 4. f. zosterifolia (p. 537)

4. Frond flat, attached by a solid disk 1. f. typica (p. 537)

4. Frond nearly fiUform, attached by rhizoids 5. f. filiformis (p. 537)

With the exception of the last, the forms thus far described for the
genus, whether as species or varieties, seem to be represented in our
territory.

i92o] SetcheU-Gardner: Melanophyceae 537

1. Ilea Fascia f. typica S. and G.

Setchell and Gardner. Phyc. Cunt., VII, 1924, p. 12.

f. typica is founded on Fiicus Fascia, Mueller, Fl. Dan., 1778, pi.
768. The plants distributed in Collins, Ilolden and Setchell, Phyc.
Bor.-Amer. (Exsiec), no. 1131, are fairly representative of the typical
form. It extends along the coast from Puget Sound, Washington, to
southern California.

2. Ilea Fascia f. debilis S. and G.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 12.

f. dehUis is founded on Lammaria dehilis Agardh. We have a few
specimens representative of this form from San Juan Island, Wash-
ington. They measure up to twelve centimeters wide and twenty-five
centimeters long.

3. Ilea Fascia f. caespitosa S. and G

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 13.

f. caespitosa is founded on Laminaria caespitosa J. G. Agardh, Sp.
Alg., I, 1848, p. 130, and is well illustrated in Thuret and Bornet,
Etudes Phyc, 1878, pi. 4. The plants distributed in Collins, Holden
and Setchell, Phyc. Bor.-Amer. (Exsiec), no. 736 may be considered
to represent this form. It is known only in the extreme northern
portion of our region.

4. Ilea Fascia f. zosterifolia S. and G.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 13.

f. zosterifolia is founded on Phyllitiis zosterifoii-a Reinke, Algenfl.
westl. Ostsee, 1889a, p. 61. This form has been detected in but a
single locality on our coast, viz., Pebble Beach, Carmel Bay, Monterey
County, California.

5. Ilea Fascia f . filiformis S. and G.

Setchell and Gardner, Phyc Cont., VII, 1924, p. 13.

f. filiformis is founded on Phyllitis filiformis Batters, in Linn. Soc
Journ. Bot., vol. 24, 1888, p. 451, pi. 18, figs. 1-6. This form has not
been detected on our coast.

538 Unwersity of California Puhlicaticns in Botany [Vol. 8

25. Endarachne J. Ag.

Fronds plane, entire, ecostate, solid, attached by a small solid
disk, composed of three quite distinct tissues, a surface tissue of more
or less cuboidal cells holding the plastids, merging below into larger,
thick-walled, slightly parenchymatous cells which in turn merge into
the medulla, composed of very thick-walled, densely intertwined,
branched filaments extending longitudinally for the most part ; repro-
duction by plurilocular gametangia forming a palisade layer over the
whole frond except the stipe ; paraphyses unknown ; hairs in fascicles.

J. G. Agardh, Analecta Alg., Cont. 3, 1896, p. 26.

The type species of the genus is Endarachne Binghamiae collected
by Mrs. Bingham in the vicinity of Santa Barbara, California. So far
as is known at present, this is the only species of the genus.

In the genus Ilea, the central cells of the medulla are, at times,
more or less elongated, but there is no such conspicuous development
of hyphal tissues as is to be found in the present genus. Endarachne
resembles forms of Ilea Fascia so closely in general appearance that it
is necessary to examine sections to distinguish them with certainty.

Endarachne Binghamiae J. Ag.

Plate 38, figs. 37, 38, and plate 83a

Fronds usually aggregated into clusters, 10-18 cm. high, 1-3 cm.
broad, linear to broadly spatulate, at times irregular in outline, usually
eroded above at maturity, with cuneate base and distinct, small, short
stipe ; color dark brown, young plants usually turning green on drying ;
gametangia formed by horizontal division of surface cells, 44-50/i. long,
4-5/A diam., with uniseriate loculi, all enclosed by a common cuticular
layer.

Growing on rocks in the middle and upper littoral belts. Southern
California and as far as Ensenada, Lower California.

J. G. Agardh, Analecta Alg., Cont. 3, 1896, p. 26, pi. 1, fig. 5;
Saunders, Phyc. Mem., 1898, p. 162, pi. 30, figs. 6, 7 ; Collins, Holden
and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1593; Okamura, Alg.
Japon. Exsicc, no. 86. Phyllitis Fascia Okamura, Illus. Mar. Alg.
Japan, vol. 1, 1901, pi. 10.

1^--^] SetcheU-Gardner: Melanophyceae 539

26. Colpomenia Derb. and Sol.

Frond solid in the juvenile stage soon becoming hollow, rather thin
and membranaceous, whole when young, becoming irregularly, more
or less lacerated in age, consisting of two tissues, an inner tissue of
large, rounded, nearly colorless, thin-walled cells and a cortical tissue
of small, more or less cuboidal, assimilating cells; plurilocular game-
tangia in early stage of development growing around groups of hair
filaments, later spreading over the entire surface of the frond ; para-
physes (?) unicellular, clavate, scattered.

Derbes and Solier, Mem. phys. alg., 1856, p. 11.

This is the third genus among the Scytosiphonaceae to show a mul-
titude of variations without sharp distinctions and no less than five
forms, varieties, or species have been described and named. The type
of the genus is conceived of as nearly a globular plant, thin-walled and
while, at times, slightly bullose, at least neither thick-walled nor pro-
vided with conspicuously projecting, wart-like or finger-shaped lobes.
Nevertheless, such variations occur, especially on our coast, where the
variation within the genus seems to have reached its maximum. We
have placed all of these species, varieties, and forms under one species,
with a series of varieties, much as we have done in the case of Scyto-
siphon and Ilea, but these variations, being more pronounced, have
been segregated into varieties and forms rather than into forms and
subforms. Nevertheless they may be simply environmental effects or
"ecads."

Colpomenia sinuosa (Roth) Derb. and Sol.

Plate 45, figs. 82-86

Frond sessile, attached by a broad base, thin and membranaceous,
4-10 cm. diam., filled with water when young, later collapsed and
somewhat complanate, spherical to irregular in form, 0.25-0.35 mm.
thick ; color dark brown ; cortical layer of cells consisting of 1-2 rows
of cuboidal or polygonal cells, the inner layer of 2-5 rows of larger,
more or less rounded cells; plurilocular gametangia 18-22/* long, 5-8/*
broad, the loculi frequently in two rows.

Growing on rocks and on other algae in the middle and lower
littoral belts. Common along the whole Pacific coast, from Yakutat
Bay, Alaska, to southern California.

540 University of California Publications in Botany [you 8

Derbes and Solier, Mem. phys. alg., 1856, p. 11, pi. 22, figs. 18-20 ;
Saunders, Phyc. Mem., 1898, p. 164, pi. 32, figs. 7, 8, Alg. Harriman
Exp., 1901, p. 421; Setehell and Gardner, Alg. N.W. Amer., 1903,
p. 242; Collins, Holden and Setehell, Phyc. Bor.-Amer. (Exsicc),
no. 278; Tilden, Amer. Alg. (Exsicc), no. 522. Ulva sinuosa Roth,
Cat. Bot., vol. 3, 1806, p. 327, pi. 12.

The type of the species was collected at Cadiz by Mertens and
from the figures and what we know of the plants of that region is
evidently of the more regularly globular or lobed, thin-walled, more
or less smooth form of the species, although the description of Roth
might seem to apply to the more thick-walled and rugose form. The
typical form is less common on our coast but is found in certain
localities. As stated above, it seems best, after a study of all of our
west coast material, to consider all of our forms under one species
and we have arranged them according to the following key :

1. Fronds thin, membranaceous, surface nearly smooth 2

1. Fronds thick, chartaceous, surface buUose or strongly lobed 3

2. Fronds approximately globose, very thin and smooth..!, f. t3rpica (p. 540)

2. Fronds elongated or irregular, somewhat thicker and less smooth

2. f. expansa (p. 540)

2. Fronds expanded to several decimeters, thin but with short, spine-like

tubercles 3. f. expansissima (p. 541)

3. Fronds sinuose, strongly buUose and tuberculate, but never lobed

4. var. tuberculata (p. 541)

3. Fronds extending into one to several long, finger-like lobes

5. var. deformans (p. 542)

1. Colpomenia sinuosa f. typica nom. no v.

The typical form is thin, about 25-30/x thick, fairly regular in
form, usually attached at one place to a filamentous alga, and growing,
most commonly, in fairly quiet waters. The clusters of hairs are not
sunk in the tissue of the frond and the layers of colorless cells are
few (usually 2 to 3).

2, Colpomenia sinuosa f. expansa Saunders

Fronds aggregated forming expansions of indefinite size and shape,
0.35-0.45 mm. thick, inner layers of cells composed of 5-7 rows of cells.

Near Avalon Bay, Santa Catalina Island, California.

Saunders, Phyc. Mem., 1898, p. 164, pi. 32, figs. 4-6; Collins,
Holden and Setehell, Phyc. Bor.-Amer. (Exsicc), no. 825.

1^25] Se f eh ell-Gardner: Melanophyceae 541

The f. expansa is a variation of the typical form in which the frond
is more ample, less regular in shape, usually creeping over filamentous
algae and forming secondary attachments. It has a smooth surface
in the sense of not being tuberculate or conspicuously lobed. It is
somewhat thicker than the typical form, with the clusters of hairs
sunk in tissues. The layers of colorless cells are greater in number
than in the typical form (usually 5 to 7).

3. Colpomenia sinuosa f. expansissima S. and G.

Fronds 3-6 dm. diam., thin, sinuose, with minute, spine-like pro-
jections.

Floating in billowy masses in San Francisquito Bay, Lower Cali-
fornia.

Setchell and Gardner, Mar. Alg. Gulf of Calif., 192-1, p. 726.

This form is probably an extreme of f. expansa, but it has only
been found floating. It is thinner and less even in outline and surface,
but is similar in every other way. The only specimens are sterile and
show no hairs.

4. Colpomenia sinuosa f. tuberculata (Saunders) S, and G.

Fronds sessile, hollow, rigid and somewhat coriaceous, very irregu-
lar in shape and size, usually somewhat flattened, 5-10 cm. diam., 1-2
mm. thick ; surface much convoluted, wrinkled and folded, at maturity
covered with blunt warts or tubercules ; color dark brown ; cortex of
3-5 rows of cuboidal cells, the inner layer of 5-8 rows of large,
irregular, thin-walled cells; plurilocular gametangia 20-25/x long,
3-4/1, wide ; paraphyses 22/* long, 5/t wide.

Growing on rocks and on other algae, often aggregated in large,
brain-like masses, in the middle of the littoral belt. From Unalaska,
Alaska, to La Paz, Lower California.

Setchell and Gardner, Alg. N.W. Amer., 1903. p. 242 ; Howe, Phyc.
Stud. V, 1911, p. 495; Setchell and Gardner, Mar. Alg. Gulf of Calif.,
1924, p. 725. Colpomenia tuhercuJata Saunders, Phyc. Mem., 1898,
p. 164, pi. 32, figs. 1-3 ; Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. 826.

The CoJpomenm tuhercidata Saunders is thick-walled, of general
globose form, but with the distribution of growth areas interrupted in
such a way as to give rise to bullosities and tumor-like warts. Some-
times these swellings develop and fall out, leaving holes or lacunae, a
suggestion of the process in Hydroclathrus clathratus.

542 TJniversity of California Piiblicaiions in Botany [Vol. 8

5. Colpomenia sinuosa f . deformans S. and G.

Fronds rigid, erect, membranaceous, cavernous, lower part tuber-
culate in the juvenile state, later developing one or more irregularly
cylindrical, sack-like, at times flattened, projections from the upper
surface, 1-7 cm. high, surface of frond smooth, somewhat lacerated
when aged ; color dark brown.

Attached to rocks and to other algae by a broad sessile base, in the
lower littoral belt. Ranging from Cook Inlet, Alaska, to the Gulf of
California.

Setchell and Gardner, Alg. N.W. Amer., 1903, p. 242, pi. 18, figs.
13-15 ; Mar. Alg. Gulf of Calif., 1924, p. 726, pi. 19, figs. 61, 62. Scyto-
siplion huUosus Saunders, Phyc. Mem., 1898, p. 163, pi. 31, figs. 1-7,
Alg. Harriman Exp., 1901, p. 421; Tilden, Amer, Alg. (Exsicc), no.
351.

The Scytosiphon huUosus Saunders always arises from a sinuous,
lobed base which may be so small as to seem almost like a disk, but
varies to a Colpomenia sinuosa type of considerable size. We have
already discussed this elsewhere (1903, p. 243, pi. 18, figs. 13-15).
When the long finger-like lobes are reduced to one and the surface is
smooth, it is the typical form of Saunders, but more often the long
lobes are several and unequal and, at times, all are short. It seems
that certain areas grow more rapidly and this differentiation completes
the series from the forms of var. typica to the extremes of var.
deformans.

27. Hydroclathrus Bory.

Fronds spherical or irregularly ovate, hollow, similar to those of
Colpomenia, entire when young, at first becoming fenestrate, but
broken later into a well formed lattice work, the meshes being variable
in shape and size ; reproduction by plurilocular gametangia scattered
over the whole outer surface of the frond ; hairs growing in groups in
shallow depressions over the outer surface of the frond.

Bory, in Diet, class., vol. 8, 1825, p. 419.

The type and only known species of the genus is Hydroclathrus
clathratus. The completeness of the perforation of the thallus of the
single species of the genus Hydroclathrus readily separates it from the
various forms of Colpomenia, although odd plants of C. sinuosa var.
tuherculata often become considerably, but not regularly, perforated
through spotwise localization of groM^th. The origin of the holes has

1925] S etch ell-Gardner: Melanophyceae 543

not been investigated. When first visible, they are very small and it
seems likely that they originate through the death, or quiescence, of a
cell or small group of cells in the midst of a rapidly growing area, as
is the case in some other perforated algae.

Hydroclathrus clathratus (Bory) Howe

Fronds very irregular in form, in age much lacerated, 5-16 cm.
diam., sessile, attached to the substratum by a broad base, sometimes
several bases confluent, differentiated into two layers of cells ; the
apertures varying much in size and form and the frond involute along
their margins ; color 3'ellowish brown ; hairs in small tufts in the center
of expanded sori.

Growing in tide pools in the lower littoral belt. On the west coast
of Mexico near the mouth of the Gulf of California.

Howe, in Britton and Millspaugh, Bah. Flor., 1920, p. 590; Setchell
and Gardner, Mar. Alg. Gulf of Calif., 1924, p. 727. Hydradathrus
cancellatus Bory, m Diet, class., vol. 8, 1825, p. 419 ; Harvey, Phyc.
Austral., 1859, Tab. 98 ; Mitchell, in Murray, Phyc. Mem., part 2, 1893,
pp. 53-56, pi. 15, figs. 2-4. Encaelium ciathratum (Bory) Agardh,
Sp. Alg., vol. 1, part 2, 1822, p. 412. Fucus clathratus Bory, MS.,
fide Agardh, loc. cit.

The single species of Hydroclathrus is tropical and widespread in
both the Atlantic and the Pacific oceans. Our specimens, from the
Gulf of California, are old and we have been unable to detect any
gametangia.

FAMILY 11. ^GIRACEAE FAM. NOV.

Fronds erect, more or less branched, from slightly to decidedly
gelatinous ; main growth in length trichothallic, with central axes
monopodial and cortical axes sympodial, of two or three tissues, (1)
medullary of larger and longer colorless longitudinal filaments, easily
separable or compact, (2) intermediate of broad but short colorless
cells, and (3) cortical of longer or shorter distinct anticlinal filaments,
often curved and enlarged upwards ; both unilocular zoosporangia and
plurilocular gametangia known (in some species at least) and borne
on similar (macroscopic) plants.

The family which we have felt constrained to designate as ^gira-
ceae is to be separated both from the Chordariaceae (in the narrower
sense of Chordaria as limited by the type C. flagelliformis) and from

544 University of California PuhUcations in Botany [Vol. 8

the " Mesogloiaceae " (also in the narrower sense of the genus Meso-
gJoia, as limited by the type species of the genus M. vermiculata) both
because of the method of terminal growth ("trichothallic" as con-
trasted with "subapical") and because neither in Chordaria. nor
Mesogloia (as limited to conform with the type species in each case)
are plurilocular gametangia known, while they are knowm (or
reported) in the typical species of Myriodadia and Liehmamnia. There
seems to be represented here a distinct group agreeing in character-
istics of typical trichothallic apical growth and with verj^ similar mac-
roscopic gametophytes and sporophytes although more exact knowl-
edge of the types of these genera is extremely desirable, since the
various statements are perplexing and authentic material difficult (or
impossible for us) to obtain. We have made, therefore, an arrange-
ment of our genera and species based on the most certain information
available to us. It is to be borne in mind that Mesogloia (based strictly
on the type of the species) has been removed to the Chordariaceae
where it is close to, if not identical with, the genus Chordaria. Other
species than the type, referred by different authors to Mesogloia are
referred variously to Liehmannia, ^gira, Myriocladia, Myriogloia and
Chordaria.

Key to the Genera

1. Gametangia arranged unilaterally on the upper portions of the cortical filaments
28. .fflgira (p. 544)

1. Gametangia not unilateral, arising from the lower portions of the cortical fila-
ments 29. Meneghiniella (p. 548)

28. -ffigira Fries

Fronds branched, filiform, with trichothallic growth, composed of
a medulla of colorless, branched filaments firmly agglutinated, giving
rise to short, cortical, assimilating filaments and to ba.sal zoosporangia,
or to gametangia unilaterally arranged on their outer extremities or,
to both zoosporangia and gametangia.

Fries, Syst. Orbis, 1825, p. 342. Cladosiphon Kuetzing, Phyc.
Gen., 1843, p. 329. Castagnea Derbes and Solier, Sur les organes
repr. des algues, 1850, p. 269. Eudesme, J. Agardh, Till Alg. Syst.
II, 1880, p. 29.

Kuetzing founded the genus Cladosiphon (1843, p. 329) on a
Mediterranean species, C. mediterraneus, which seems definitely, as
stated below, to be either the same or most certainly closely related
to, the species on which Derbes and Solier founded their Castagnea.

1925] Setch ell-Gardner: Melanophyceae 545

J. G. Agardh (1882, p. 40) took up Cladosiphon from the point of
view of Meneghini's Liehmannia Posidoniae, a species seemingly at
present, at least, to belong to another genus (MeneghinieUa) from the
tj'pe of Kuetzing.

Derbes and Solier founded the genus Castagnea on a species they
named C. polycarpa, presumably from the neighborhood of Marseilles,
but the locality is not mentioned. This species has the gametangia
produced as outgrowths from the convex sides of the upper cells of
the curved cortical filaments and this peculiarity is made a part of
the diagnosis of the genus. Derbes and Solier state that their genus
Castagnea is readily distinguishable from the related genera Lieh-
mannia, Stilophora, and Nereia by the form of the filaments associated
with the organs of fructification in that they are cylindrical and not
terminated by large swollen cells. Castagnea polycarpa Derbes and
Solier has been variously and usually doubtfully referred by later
writers and we have found no reference to the existence of an authentic
specimen. Derbes and Solier compare its appearance to that of a
stunted Liehmannia. In their later ( ?) paper, Derbes and Solier
(Sur les organes repr. des algues, 1850, p. 269, pi. 33, figs. 12, 13)
seem inclined to consider their C. polycarpa closely related to, if not
identical with, the "Cladosiphon. fistulosum" of Kuetzing, a binomial
unknown to us, but presumably the Mesogloia fistulosa Zanard. (in
Meneghini, 1843, p. 292) and considered by Kuetzing (1849, p. 547)
to be a synonym of his Cladosiphon mediterraneus. There is certainly
suspicion here that Castagnea and Cladosiphon may have been founded
on the same type species. In the absence of that exact proof which is
difficult to obtain other than from type specimens and experience
from type localities, we feel justified in considering these two genera
as identical in foundation. We may call attention to the fact that
Bornet (1892, p. 236) has placed the Cladosiphon mediterraneus
Kuetzing under Castagnea.

The genus Eudesme was founded by J. G. Agardh (1882, p. 29)
on the Mesogloia virescens Carmichael, i.e., at least technically. There
is good reason to believe that Agardh did not restrict his ideas to type,
but had a composite idea as to the nature of his E. virescens. He
speaks of an "axis solidescens" whereas Eudesme has been applied as
typically including species whose axial filaments are readily separable
from one another (cf. Kjellman, 1893, pp. 225, 226) and as they are,
to a certain extent at least, in what may be considered to be typical
E. virescetis, viz., Alg. Danm., no. 49 ; but even here there is a definite

546 University of California Puhlications in Botany [Vol. 8

pseudoparenchymatous axis (cf. also Harvey, Phyc. Brit., pi. 82,
1847). Agardh also emphasizes in his generic diagnosis the formation
of gametangia as projecting ramelli arising from the upper cells of
the cortical filaments. The illustrations of Thuret (1850, pi. 27) show
exactly the type of cortical filaments and gametangial formation upon
which, as distinctive characters, Derbes and Solier founded their
genus. So far as we are able to discover, the three genera, Castagnea,
Cladosiphon, and Eudesme are to be united and, since Cladosiphon is
the earliest name, unless we adopt JEgira Fries of doubtful status, we
might feel compelled to use Cladosiplion as the name of the genus to
which we may assign species usually referred to Castagnea or to
Eudesme. Cladosiplion, as we understand Kuetzing's type species,
CI. niediterraneiis {Mesogloia fistidosa Menegh., Castagnea fistidosa
(Menegh.) Derb. and Sol., C. polycarpa Derb. and Sol. and Castagnea
mediterramea (Kuetz.) Bornet) is however, usually understood in the
sense of J. G. Agardh (1882, pp. 40 and 42). J. G. Agardh, on the
other hand, based his idea of Cladosiphon on Liehniannia Posidoniae
Meneghini (1843, pp. 300-304, pi. 5, fig. 1) which has its gametangia
situated very differently from those of the members of the Clado-
siphonr-Castagnea-Eiidesme complex. Meneghini 's species seems to
have the gametangia arising as transformed branches of the cortical
filaments and slender, with uniseriate loculi, not pronouncedly sili-
quaeform as in Liehniannia and not arising unilaterally from the
upper cells of the cortical filaments as in the complex we are discussing.
We have one species, from the Gulf of California, seemingly cogeneric
with Meneghini 's Liehniannia Posidoniae for which we have proposed
the generic name Meneghiniella (Phyc. Cont. VII, 1924, p. 5).

It remains to consider the oldest name possible for this Cladosiphon-
Castagnea-Eiidesme complex and that is ^gira Fries (1825, p. 342).
Fries founded his genus on the Linckia Zosterae Lyngbye (1819,
p. 194, pi. 66, fig. C) which may or may not be the Rividaria Zosterae
Weber and Mohr (1810, p. 367). The plant of Lyngbye has been
variously interpreted by J. G. Agardh (1848, p. 53) and Areschoug,
who had an authentic specimen for comparison (Areschoug, Alg.
Scand. Exsicc, ser. 1, no. 67, and 1842, pp. 228-230). The evidence
is well discussed by Kylin (1907, pp. 85-87). While there may be
reasonable doubt as to whether Lyngbye 's plant is the same as Eudesme
virescens (Carm.) J. Agardh, or Eudesme Zosterae (J. Ag.) Kylin,
it is to be placed in the Cladosiphon-Castagnea-Eudesme complex
which ought, therefore, to take the name jEgira (cf. also, M. A. Howe,
1918, p. 505).

1925] Setchell-Gardner : Melanophyceae 547

Since the foregoing was written, we have been allowed, through
the kindness of Professor C. H. Ostenfeld, to examine one of Lyngbye's
specimens of Linckia Zosterae. It is clearly very closely related to
Eudesme virescens as we understand it. We, therefore, feel justified in
assuming that JEgira is synonymous with Eudesme.

Mgira, virescens (Carm.) S. and G.

Plate 42, figs. 59, 60

Fronds very flaccid and gelatinous, 8-20 cm. (up to 45 cm.) high;
branches arising from the main frond at wide angles, nvimerous,
variable in length, usually enlarging above, blunt ; the ultimate ramuli
short, arising almost at right angles ; medullary filaments few to many
very loosely bound together, composed of cells very variable in size
even in the same filament, up to 55|U, diam., 1.5-2.5 times as long as the
diameter, constricted at the cross-walls, with numerous, small, disk-
shaped chromatophores, giving rise to longer or shorter lateral, color-
less, branched filaments bearing in turn the loose fascicles of free,
branched, assimilating filaments, zoosporangia, and long colorless hairs,
the latter terminating the colorless branches; cortical assimilating
filaments slender, branched, tapering slightly at the apices, terminal
ramuli 10-15 cells long, fascicled, arcuate, cells cylindrical below,
becoming slightly monilif orm above, 15-20ju. diam. ; hairs numerous,
long, terminating the lateral branchlets from the medullary filaments ;
zoosporangia broadly ellipsoidal, short cylindrical, obovate to rhombic-
ovate, arising near the outer ends of the colorless lateral filaments,
70-90/t (up to 120ju) long, 25-65ju. broad; gametangia short, lateral
and seriate, secund on the cortical filaments.

Growing on rocks and on eel grass in the littoral and sublittoral
belts. Shumagin Islands to Sitka, Alaska.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 11. Eudesme
virescens Saunders, Alg. Harriman Exp., 1901, p. 423 (?); Setchell
and Gardner, Alg. N.W. Araer., 1903, p. 249. Liehmannia sp.
Saunders, lac. cit., p. 424, pi. 49 ( ?). Mesagloia virescens Carmichael,
in Hooker, Engl. Fl., vol. 2, 1833, p. 387.

While we suspect that the plant referred by Saunders to Eudesme
virescens and so referred to by us in our Algae of Northwestern
America ( loc. cit. ) , may be the same species as that collected by one of
us at Sitka, we have not been able to examine a specimen to make
certain. Our plants show no gametangia but have zoosporangia and

548 University of California Publications in Botany [Vol. 8

the vegetative structure of Eudesme J. Ag. A careful comparison of
Wyatt's Mesogloia virescens, no. 49 (Alg. Danm.) with our various
specimens shows so much variation in dimension of zoosporangia and
peripheral filaments that we feel nearly safe in referring all to the
European species which we refer, for reasons given above, to JEgira.

29. Meneghiniella S. and G.

Fronds cylindrical, more or less profusely branched, flaccid and
lubricous, with growth trichothallic ; medulla composed of numerous
colorless filaments compactly coalescing, giving rise to an abundance
of short, usually arcuate, color bearing, cortical filaments, whose lower
branches are transformed into linear, plurilocular gametangia, with
nearly uniseriate loculi, and usually more or less fasciculate ; zoo-
sporangia uncertain.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 5. Cladosiphon
J. Agardh, Till Alg. Syst., II, 1882, p. 40 (not of Kuetzing).

The genus described above is founded on a plant growing in
the Gulf of California. It seems, however, that our type must be
cogeneric with the Liehmanma Posidoniae Meneghini, whose type
locality is Naples and which was selected by J. G. Agardh as typical of
Kuetzing 's genus CladosipJion. We have shown above the results of
our attempt to determine the type of JEgira, which have led us to the
conclusion that the genus, as founded by Fries, includes the later
genera published under the names of Cladosiphon, Castagnea, and
Eudesme. Our plants seem to be typically trichothallic and have
longer or shorter, slender, linear gametangia, which are, at times, very
slightly swollen at, or about, the middle. The gametangia arise near
the base of the cortical filaments and are more or less fasciculate. As
we interpret Meneghini 's figures of Liehmannia, Posidoniae (1843,
pi. 5, fig. 1), he has represented the gametangia on the upper side of
figure 6 and also in figure c. A confusion is introduced, however, in
his representation of the contracted contents of the swollen terminal
cells of the cortical filaments (figs. 6, d, e, and /) by which they are
made to seem to agree with the gametangia of Liebmannia' Levillei
J. Ag. The figures of J. G. Agardh (1882, pi. 2, figs. Sa, 3&) represent
the cortical filaments with their swollen terminal cells and their game-
tangia of his Cladosiphon zostericola of Australia which seems clearly
to be referred to Meneghiniella. The genus Meneghiniella, in our esti-
mation, included M. Brandegeei S. and G. (type), M. Posidoniae

1925] SetcheU-Gardner: Melanophyceae 549

(Menegh.) 8. and G. {Liehmannia Posidoniae Meneghini, 1843, p. 300,
pi. 5, fig. 1), M. zostericola (J. Ag.) S. and G. {Cladosiphon zostericola
J. G. Agardh, 1882, p. 43, pi. 2, figs. 3a., 36), and possibly M. erythraea
(J. Ag.) S. and G. (CladosipJwn erifthraeum J. G. Agardh, 1848,
p. 55).

Meneghiniella Brandegeei S. and G.

Plate 47, fig. 11, and plate -49, fig. 16

Fronds very slender, soft and flabby, 8-16 cm. high, up to GOO/x
diam., irregularly and alternately branched, with branches coming off
at wide angles ; medullary filaments composed of large thin-walled,
colorless cells up to lOOfx diam. and several times longer than the
diameter, firmly agglutinated, becoming much smaller toward the
periphery ; cortical assimilating filaments simple, clavate, free, not
compact, more or less arcuate, composed of 7-12 moniliform cells ;
gametangia fasciculate with uniseriate loculi for the most part, 40-50/x
long, 7-lOjti diam. ; zoosporangia and hairs unknown.

Habitat unknown. La Paz, Lower California.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 5.

The specimens on which this species, as well as the genus, are
founded are not as representative as could be desired, but the char-
acters are plain both as to the structure of the frond and the shape,
position, and the structure of the gametangia which can clearly be
determined. The tips of the filaments indicate trichothallic growth.

FAMILY 12. HETEROCHORDAEIACEAE fam. nov.

Main fronds erect, with main axis and lateral branches of limited
growth ; growth in length subapical as in the Chordariaceae ; axes of
three tissues: (1) medullary or central of elongated parallel, colorless
cells; (2) intermediate of shorter colorless cells, and (3) cortical of
short, anticlinal rows of colored cells, the distal cell being more or less
swollen ; zoosporangia and gametangia on full-sized, similar plants.

It seems something like violent procedure to separate this family
from- the Chordariaceae, but the presence of full-sized gametangial
plants in this family and the lack of any such plants among the Chor-
dariaceae (as limited by us) is certainly significant, in view of the more
recent indications as to the relative characteristics of the gametophyte
and sporophyte among the Melanophyceae.

550 University of California Puilications iyi Botany [Vol. 8

30. Heterochordaria S. and G.

Main fronds cylindrical, at first solid, later becoming hollow and at
times slightly swollen, not forked but densely clothed on all sides with
longer or shorter, subcylindrical or slightly flattened, ramuli, few to
many arising from a thin prostrate, profusely branched or lobed,
parenchymatous thallus firmly adhering to rocks; interior of fronds
composed of thick-walled, colorless, parenchymatous cells surrounded
by small color bearing surface cells in anticlinal rows ; zoosporangia
situated among numerous paraphyses and scattered hairs ; gametangia
arising through the transformation of approximately the lower two-
thirds of the cortical filaments.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 6.

The finding of both zoosporangia and gametangia on macroscopic
plants of Chordaria aiietina Rupr., and the finding only of zoo-
sporangia on the well known Chordaria flagelliformis (Muell.) Ag.
indicates the extremely strong probability of a fundamental difference
between C. abietina and C. flageUiformis as to the character of the
gametophyte. In view of our present attitude as to the bearing of
such a difference on the taxonomic position of the two plants, it seems
desirable to make C. abietina the type of a new genus and a new family
of the Ectocarpales and to place C. fiageUiformis and its allies of the
Chordariaceae (in restricted sense) in another order, the Chordariales
and near to the Dictyosiphonales to which they (the Chordariaceae as
we restrict them) seem fairly closely related.

There may be associated with this family, another genus, or even
two, viz., Ruprechtiella Yendo (Trav. Mus. Bot. de I'Acad. Imp. Sci.
St. Petersb., vol. 10, 1913, p. 117) which may be the same as Analipus
Kjellm., and the Chordaria Gunjii Yendo (Nyt. Mag. Naturvidensk.,
vol. 51, 1913, p. 280) which may simply be the gametangial form of
some species of Myelophycus.

Heterochordaria abietina (Rupr.) S. and G.

Plate 36, figs. 18, 19, and plate 91

Erect fronds, 10-30 cm. high, flaccid, gregarious, many arising
from a single, widespreading, profusely branched, parenchj^matous
horizontal thallus; main axis slightly compressed above, terete below,
thickly beset on all sides with radiating ramuli 1-5 cm. long, tapering
at both ends, cylindrical to narrowly linear, both main axis and ramuli

1925] Setchell-Gardner: Melmwphyceae 551

solid when young, becoming more or less fistulose when old ; color dark
olive brown ; erect fronds annual, base perennial ; zoosporangia ellip-
soidal to obovoid, numerous, sessile on the parenchymatous cells or at
times lateral near the base of the paraphyses, 50-60/^ long, 30-40ju
broad, paraphyses numerous, clavate, 70-85/a long, terminal cell much
enlarged, pyriform to spherical; gametangia very compact, covering
the whole surface of the ramuli, without paraphyses or hairs, pluri-
seriate, terminating in a 2-3-celled, sterile, usually clavate tip ; plants
dioecious.

Common on rocks in the middle and lower littoral belts. Ranging
from St. LawTence Island in the Bering Sea to central California
(near Point Conception).

Setchell and Gardner, Phyc. Cont., VIT, 1924, p. 6. CJiordaria
abietina Ruprecht, MS. in FarloAv, List Mar. Alg. U. S., 1876, p. 357 ;
J. Agardh, Till Alg. Syst., part 2, 1882, p. 74, pi. 3, figs. 2a, 2h;
Saunders, Alg. Harriman Exp., 1901, p. 424 ; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 251, pi. 18, figs. 16, 17; Collins, Holden
and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 281; Tilden, Amer. Alg.
(Exsicc), no. 348; Farlow, Anderson and Eaton, Alg. Exsicc. Amer.-
Bor., no. 94.

Heterochordaria ahietina (Rupr.) S. and G. is one of the very fre-
quently occurring algae of western North America from Point Con-
ception in California northwards. It has two striking peculiarities,
viz., the possession of a well developed horizontal thallus from which
the erect fronds arise and also of the gametophyte of the same size as
the sporophyte. The horizontal thallus is often a fair sized and
reasonably conspicuous structure, at times several centimeters in
diameter, spreading at the periphery and consisting of overlapping
narrow, blunt and emarginate lobes. Prom certain appearances, it
seems possible that it may be perennial. In the sporophyte, the zoo-
sporangia seem to be confined to the ramuli, and this seems to be the
case also with the gametangia of the gametophyte which have not been
seen to occur on the main axis. J. G. Agardh (1882, p. 74, pi. 3, figs.
2a, 2h) has figured and described the gametangia and Okamura (1910,
p. 122, pi. 85, figs. 14, 15) has figured and described both the zoo-
sporangia and the gametangia. We have previously (1903, pp. 251,
252, pi. 18, figs. 16, 17) figured and described the horizontal thallus.

552 University of California Puhlications in Botany [Vol. 8

FAMILY 13. CHNOOSPOEACEAE fam. nov.

Fronds elongated, solid, slender, more or less compressed, more or
less regularly dichotomous or proliferous from the margins, growth
subapical, not triehothallic, of two sets of tissues, an inner of large
cells, an outer of one or more layers of small colored cells in short
anticlinal rows ; hairs in clusters arising from the bottoms of shallow
pits ("cryptostomata") ; zoosporangia unknown; gametangia uni-
seriate, arising about the clusters of hairs and spreading over the
surface, at times into confluent sori.

The genus Chnoospora is usually ranked as of uncertain position
("incertae sedis"), but by some placed in the Encoeliaceae, a family
now abandoned and with its members distributed and separated. In
its sori, surrounding the hair clusters, it resembles closely those of
Colpomenia, but in habit and terminal growth it is very different from
that genus and its near relatives. The frond is not directly to be com-
pared with that of any brown alga at present known to us. The slender,
compressed and regularly dichotomous frond at once sets it apart. The
subapical method of terminal growth relates it more nearly to Hetero-
chordaria, but Chnoospora is known only with gametangia as are the
members of the Scytosiphonaceae. The method of subterminal growth
is like that of the genus Stilophora, but it is very different from that
genus in not having free cortical filaments. Chnoospora may be con-
sidered to be a further development of the cylindrical Scytosiphon-
aceae, with the loss of the terminal hair and the growth in length
localized. It certainly seems best, at present, to place Chnoospora in
a family of its own.

The structure and the fructification were first described by Ethel
Sara Barton (1897). Borgesen (1924, p. 264) calls attention to the
resemblance of the growth point to those of Scytothamnus australis
and Coilodesme hulligera. Both of these plants probably are to be
referred to the Chordariales since they possess only zoosporangia of a
distinct unilocular type and it is to be suspected that they have
microscopic gametophytes.

31. Chnoospora J. Ag,

Fronds cylindrico-compressed, solid, ecostate, profusely branched,
attached by a solid holdfast ; branching dichotomous, f astigiate ; fronds
differentiated into two tissues, the center of larger, colorless prismatic
cells, elongated longitudinally, surrounded by tw^o layers of small cells

1925] Setchell-Gardner : Melanophyceae 553

containing the endochrome ; reproduction by phirilociilar gametangia
surrounding depressions containing tufts of extruding hairs, and, by
extension at times, nearly covering the fronds.

J. G. Agardh, Nya Alg., 1847, p. 7.

The type species of the genus is C. pacifica from St. Augustin,
Mexico, listed later by J. G. Agardh as C. fastigiata var. pacifica.
Chnoospora is the only genus we feel inclined to refer to the family
Chnoosporaceae, for the reasons given above in discussing the family.

Chnoospora pacifica J. Ag.

Fronds densely caespitose, many arising from a solid disk-shaped
holdfast, of the same diameter throughout except at the slightly atten-
uated apices, 2.5-7.5 cm. high, 2 mm. wide, 1 mm. thick ; branches
forming acute angles.

J. G. Agardh, Nya Alg., 1847, p. 7 ; Setchell and Gardner, Mar.
Alg. Gulf Calif., 1924, p. 728. Chnoospora fastigiata var. pacifica
J. G. Agardh, Sp. Alg., 1848, pp. 171, 172. Barton, On the fruit of
Chnoospora fastigiata, 1898a;, p. 507. Sargassum piluliferuni Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 537&.

The material upon which this species, as well as the genus, was
founded was collected by Professor F. M. Liebmann at St. Augustin,
in the southern portion of the state of Oaxaca, on the Pacific coast of
Mexico. This is a tropical locality and the Chnoospora was accom-
panied by several other marine algae described also in the same paper.
J. G. Agardh also described a species of Chnoospora, at the same time,
from La Guayra on the Caribbean Sea which he named C. atlantica.
In 1848, he united these two species as varieties under a new name,
C. fastigiata. The name C. pacifica has the rights of priority and we
have retained it. It seems widespread in the Pacific and Indian oceans,
although possibly appearing in a number of forms. It seems likely
that C. pannosa J. G. Agardh, from Hawaii, is simply a pannose
growth form of C. pacifica. C. atlantica J. Ag. from the Caribbean has
the fronds dilated under the axils and may be a distinct species.
C. implexa Hering is doubtful and may very possibly prove to be of
quite a different genus, but there are forms of our own coast species
approaching it in habit.

554 University of California Puhlications in Botany [Vol. 8

Order 3. DESMARESTIALES ord. nov.

Frond erect, filamentous or membranaceous, cylindrical to slightly
compressed, ligulate or even broadly membranaceous ; growth terminal,
trichothallic ; frond clothed with densely crowded exserted, simple or
branched, colored, monosiphonous filaments (hairs) when young or
in the growing portion, these filaments early or tardily deciduous as
age advances ; tissues several ; reproduction by zoosporangia ; game-
tangia unknown, probably borne on a microscopic gametophyte.

The posse.ssion of longer or shorter, exserted, colored, monosiphon-
ous filaments, especially on young plants or younger portions of peren-
nial plants, has always been considered as characteristic of Desmarestia
and the related genus Arthrodadia. ' It also is characteristic of the
later described and very nearly related genus Phaeiirus Skottsb.
There have been several other algae with this permanent or partially
transient covering of colored monosiphonous filaments, viz. : Mesogloia
Andersanii Farlow, Myriocladia Sciurus Harv., M. Chorda J. Ag.,
M. grandis Howe, M. callitriclia Rosenv., M. Kuromo Yendo, and pos-
sibly M. capensis J. Ag. and Mesogloia natalensis Kuetz. The first two
seem to have been investigated by Kuckuck, who did not live to publish
his results, but some of his notes and sketches Avere used by Oltmanns
(1922, pp. 19-22). From these we learn that Kuckuck had assigned
Mesogloia Andersonii Farlow (at least in part) and Myriodadia
Sciurus Harv. to a new genus " Myriogloea" and had proposed for
these a distinct family, presumably named Myriogloiaceae. Reserving
the discussion of the different species until later, we may state that it
seems best to adopt Myriogloiaeeae, as of Kuckuck and assign it to our
newly established order Desmarestiales, since the terminal growth is
most typically trichothallic exserted hairs are prominent, mono-
siphonous and colored, and the reproduction is typically onty by
zoosporangia.

Key to the Families

1. Cortical filaments free 14. Myriogloiaceae (p. 555)

1. Cortical filaments coalescent into a tissue 15. Desmarestiaoeae (p. 558)

1925] Setchelh-Gardner : Melanophyceae 555

FAMILY 14. MYRIOGLOIACEAE kuckuck

Fronds filamentous, erect, cylindrical, solid or later tubulose,
central strands of larg-e colorless cells, peripheral of two sorts, free,
short, cortical filaments with rounded terminal cell and exserted mono-
siphonous filaments, with basal meristematic cells and outer cells pro-
vided with phaeoplasts and, consequently colored ; colorless hairs
absent ; terminal growth conspicuously trichothallic ; zoosporangia
arising at the base of the cortical filaments, unilocular ; gametangia
unknown.

Kuckuck, in Oltmanns, Morph. und Biol, der Algen, ed. 2, vol. 2,
1922, p. 19 (by implication). Myriogloeeae Oltmanns, loc. cit.

The family thus far is credited with a single genus, Myriogloia,
although it is far from certain that Mesoglaia Andersonii Farlow is
cogeneric with Myriodadia Scmrus Harv., as a glance at Kuckuck 's
figures (loc. cit., p. 22, fig. 312, 1 and 2) will indicate. The very
robust plants assigned to Myriodadia Scinrus Harv., M. Chorda J.
Ag., and 31. gramdis Howe may, or may not, belong to the same genus
as Mesogloia Andersonii Farlow and Myriodadia callitricha Rosenv.
We feel reasonably certain in referring the two last species to Myrio-
gloia, which is absolutely distinct from Myriodadia J. Ag., as typified
by M. Lovenii J. Ag., designated type of the genus, at least if one
credits Kuckuck 's sketches (cf. Oltmanns, 1922, p. 37, fig. 328 and
p. 38, fig. 329) as having been drawn from correctly determined
specimens. Influenced by these sketches and the various descriptions
(notably that of Kylin, 1907, pp. 88-90, fig. 21), we are inclined to
refer Myriodadia J. Ag., as limited to M. Lovenii, to the Myrio-
gloiaceae as here established, as a second genus, distinguished by hav-
ing its axis made up of a single longitudinal and persistent cellrow
(monosiphonous axis), whereas in Myriogloia this condition is soon
lost, the frond becoming polysiphonous or hollow^ below, and in the
thicker species {Myriodadia Sciurus, M. Chorda, and M. grandis) the
medullary portion is probably polysiphonous from the first (cf.
Kuckuck 's figure 312, 2, on p. 22 of Oltmanns, 1922) and remains
solid. So far as our opportunities are concerned, there is insufficient
material to follow out a detailed investigation and we cannot proceed
farther than the above suggestions. Oltmanns, however, keeps Myrio-
dadia (1922, p. 34) distinct and establishes a special tribe, Myrio-
cladieae, for it under the family Spermatoehnaceae. Spermatodinus,
however, has a distinct apical cell.

556 University of California Publications in Botany [Vol. 8

32. Myriogloia Kuck.

Fronds cylindrical, more or less profusely branched, flaccid and
lubricous, with distinct trichothallic growth; medulla composed of
numerous colorless, equally large filaments slightly coalescent, giving
rise to numerous, narrow, long descending filaments and to an abun-
dance of short, straight or uncinate, cortical, colored filaments and
long, exserted, hair-like, assimilating filaments with cells densely con-
gested with chromatophores ; reproduction in macroscopic plants by
zoosporangia borne among the cortical filaments.

Kuckuck, MS (fide, Oltmanns, Morph. und Biol, der Algen, 1922,
p. 19 et seq.).

We have already explained the situation in regard to this genus
and its position. It is closely related to Myriocladia but lacks the
distinct and persistent monosiphon.ous axial filament. Its relation to
the southern hemisphere Pacific species of Myriocladia is to be con-
sidered, with the strong possibility that, upon careful study, they
may be segregated generically. Myriogloia, in restricted sense, prob-
ably includes three species: M. Andersonii (Farlow) Kuckuck {Meso-
gloia Andersonii Farlow, Report U. S. Fish Comm. 1876, p. 715), M.
callitricha (Rosenvinge) S. and G. {Myriocladia callitricha Rosen-
vinge, Gronl. Havalger, 1893, p. 855), and M. capensis (J. Ag.) S.
and G. {Myriocladia capensis J. G. Agardh, Spec. Alg. vol. I, 1848,
p. 54). If the Mesogloia natalensis Kuetzing is to be united with the
last, the gametangia (?) depicted in Kuetzing 's figure (Tab. Phyc,
vol. 8, pi. 10, fig. II, e) , if they really occur in the plant described and
figured, call into question our disposition of this entire family.

Myriogloia Andersonii (Farlow) Kuck,

Plate 47, figs. 8-10, plate 48, figs. 12-14, plate 49, fig. 17, and plate 76
Frond very gelatinous, solitary or gregarious, cylindrical, 8-30
cm. (up to 50 cm. high), 1-3 mm. diam., profusely branched; branches
mostlj' alternate, long and filiform, very slightly attenuated at the
extremities, when young beset on all sides with relatively long, mono-
siphonous, conspicuous filaments with band-shaped chromatophores,
disappearing when older ; color dark olive brown ; cortical filaments
moniliform, mostly uncinate, 6-9 celled, upper cells enlarged; uni-
locular zoosporangia developing at the base of the cortical filaments,
uniformly distributed over the frond, pyriform, 36-45/a long, 18-21/^
broad.

1925] SetcheU-Gardner: Melanspliyceae 557

Growing on rocks in the lower littoral and upper sublittoral belts,
or in tide pools higher up. The present known range is from Sitka,
Alaska, to San Diego, California.

Kuckuck, loc. cit. Mesogloia Andersonii Farlow, New Alg. U. S.,
1889, p. 9, pi. 87, fig. 2 ; Setchell and Gardner, Alg. N.W. Amer., 1903,
p. 250 ; Farlow, Anderson and Eaton, Alg. Exsicc. Amer.-Bor., no.
163; Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no.
925. Chordaria flageUiformis Tilden, Amer. Alg. ( Exsicc. ) , no. 349.

Much variation in the size of plants of MyriogJ&ia Andersonii has
been noted in collections from different localities. Tilden 's plant
(loc. cit.) is only about seven centimeters long and quite slender. No.
163 of Farlow, Anderson and Eaton (loc. cit.) is a young plant with
abundance of hair filaments and is about twenty-five centimeters long.
No. 925 of Collins, Holden and Setchell (loc. cit.) is an older plant,
more robust throughout, about twenty centimeters long. Plants from
Point Arena, California, in the Herbarium of the University of Cali-
fornia, measure about fifty centimeters long. Likewise variations in
the size of the mature zoosporangia are very noticeable, some collec-
tions having them twice as long as those in others. The specimens
distributed by Farlow, no. 163, 1881, are entirely clothed with brown
exserted filaments. Farlow, however, in his description (1889, p. 9)
states that it was supposed that all the specimens distributed were
fertile and speaking of projecting filaments says : "In my specimens
preserved in alcohol, they are not to be seen." We have been able
to examine the alcoholic specimens referred to and find some,
although scanty, exserted filaments. We find specimens of all condi-
tions from being densely clothed with hairs to those with very few,
but none in which they are entirely absent. We have tried to find out
whether these various forms may be separated by any other characters
than frequence or scarcity of exserted filaments, but can find none
applying satisfactorily. We have, therefore, placed all of our Pacific
coast material under the one species. Judging from what Oltmanns
states from the imperfect notes in his possession, Kuckuck has two
entities from the Mesogloia Andersonii Farlow, viz., MyriogJoia Ander-
sonii, as stated above, and CJiordarva Andersonii (cf. Oltmanns, 1922,
pp. 20 and 23, fig. 313, 4). Of the latter, we have no further knowl-
edge, although Oltmanns says "Chordaria Andersonii Kuckuck mscr.
{Mesogloia Andersonii Farlow) . ' ' There is nothing in Farlow 's mate-
rial, either dried or in alcohol, which corresponds in methods of growth
to Kuckuck 's figure.

558 University of California Piiblications in Botany [Vol. 8

FAMILY 15. DESMAE.ESTIACEAE kjellm.

Fronds erect, sparsely branched in some species, profusely branched
in others, from cylindrical through compressed to broadly membran-
aceous, solid, arising- from a simple monosiphonous branched filament
of trichothallic growth in length, soon becoming complex through the
downward growth of closely applied branching corticating filaments
which results in the production of several distinct tissues; central
monosiphonous and branching filament remaining distinct, although,
at times, becoming obscured through thylose growth filling the cavities
of the large cells; tissues outside the central monosiphonous axis,
various, usually large celled and more or less regular, the outer being
smaller and forming a distinct outer tissue of one, or few, layers of
cells ; zoosporangia known in only a few species scattered or in small
patches, arising by direct transformation of the outer cells; plants
mostly undergoing change of color (to verdigris green) and odor
(''sharp") soon after being removed from the water and exercising
a bleaching effect on other algae with which they come into contact.

Kjellman, Enum. PL Scand., 1880, p. 10, Alg. Arctic Sea, 1883,
p. 261.

The Desmarestiaceae form a very distinct group and one which is
readily separable from that of the Myriogloiaceae. It represents a
very distinct type of frond development, at least so far as the final
product is concerned. There are two centers of distribution, viz.,
North Atlantic and North Pacific as contrasted with the Antarctic and
Subantarctic regions, and it is doubtful whether there are any species
common to the two centers.

There are, at present, referred to the Desmarestiaceae three genera,
Desmarestia, the type genus, Arthracladia, by some referred to a
separate family, and the Phaeurus of Skottsberg. The first is from
both Arctic and Anarctic centers, the second is North Atlantic
(European and American) while Phaeurus is from Fuegia. While
the Myriogloiaceae are seemingly the more primitive and related to
such forms as ^giraeeae, the Desmarestiaceae are more complex, at
least, with Phaeurus as the more simple and the ligulate Desmarestias
as the more complex.

19--^] Setchell-Gardner: Melanophyceae 559

33. Desmarestia Lamour.

Fronds attachod by a solid, parenchymatous disk, exceedingly
variable in size, with a longer or shorter, usually cylindrical stipe,
more or less profusely branched, wholly or in part terete or slightly
flattened or decidedly complanate and ligulate to broadly expanded
and membranaceous, with or without a midrib ; branching distichous
and either wholly opposite or alternate, or both opposite and alternate ;
fronds composed of a single fundamental axial filament of cylindrical
cells and more or less profusely branched, colored and projecting
beyond the surface at the apices and along the margin and in the
juvenile stage, early deciduous, with trichothallic growth, sur-
rounded in the frond by one or more layers of large colorless cells
enclosed by a cortical tissue of small assimilating cells ; reproduction
b}' zoosporangia composed of slightly transformed surface cells,
inconspicuous.

Lamouroux, Essai, 1813, p. 23.

The genus Desmarestia may be divided into three groups, some-
times reckoned as distinct genera, or at least it may be divided into
two genera. The cylindrical or at most only slightly flattened species
with mostly opposite branching, constitute Dichloria, while the flat-
tened forms are retained under the name of Desmarestia, or the sub-
genus Eudesmarestia, with branching mostly alternate. The Dichloria
group has the hairs scattered as in Phaeiirus, but unlike that
genus, soon loses them. In the Eudesmarestia group, the hairs are
branched and are the very tips of incipient' branches, some, or even
many, of which do not develop into permanent structures. In
Phaeiirus the hairs are scattered, but are more permanent and per-
sistent for a longer time than in the Dichloria group. In Arthrocladia,
the hairs are in whorls, branched, persistent, and possibly represent
the tips of incipient branches. One well-known characteristic of the
species of Desmarestia may well be emphasized and that is their habit
of turning verdigris green on drying. This color is associated also with
their power to bleach other algae in contact with them. The change
of color is followed, sooner or later, by decomposition. This happens
quickly in the more slender species of the Vw'ides-iieciion and fairly
early in the members of the JFZ^er&aceae-section, but affects only the
younger and more delicate portions of the members of the Aculeatae-
section.

560 University of California Piiblications in Botany [Vol. 8

The genus Desmarestia is well represented on the Pacific coast of
North America, extending in its various species from the Bering Sea
on the north to Santa Catalina Island off the coast of southern Cali-
fornia on the south, so far as is known of its distribution at present.
The species of both the ligulate and the filiform groups manifest a
degree of variability wliich has up to date made it exceedingly difficult
to determine specific limits. It is evident from the relatively good
supply of herbarium material at our disposal, that more than mere
superficial resemblances must be resorted to in order to make a satis-
factory disposition of the group. Knowledge of the entire life-history
is highly essential. Little is known of reproduction. The character
of the hairs, present only in the early stages of growth — absent in
most of our herbarium specimens — is, so far as our experience shows,
of marked constancy and would doubtless be of decided assistance in
classification. The histological structure, very difficult to restore after
the material has once been thoroughly dried, certainly will prove of
decided value in establishing specific entities Avhen studied in fresh
material.

We have made use of all of these different phases as far as the
material and time at our disposal permits and have come to the follow-
ing conclusions regarding the number, arrangement, and distribution
of the Pacific coast species.

Key to the Species

1. Fronds filiform and either terete or compressed 2

1. Fronds ligulate or membranaceous (Herbaceae) 6

2. Fronds terete or in part subterete, branching mostly opposite (Virides).... 3
2. Fronds compressed or subterete below, branching mostly alternate

(Aculeatae) 5

3. Fronds profusely branched; tylloses few or wanting 4

3. Fronds moderately branched; tylloses filling central tube to near tip

3. D. farcta fp. 562)

4. Branches long, attenuated, successive orders gradually reduced in

size 1. D. media (p. 561)

4. Branches slightly attenuated, blunt, successive orders abruptly changing

to smaller size 2. D. pacifica fp. 562)

5. Fronds up to 15 dm. high and 3 mm. wide, with aculeate margins, moderately

branched 4. D. latifrons fp. 563)

5. Fronds up to 7 dm. high and 1.5 mm. wide, vdih smooth margins and profusely

branched 5. D. intermedia (p. 564)

6. Fronds unbranched or with a few short branches at the base

10. D. foliacea fp. 569)

6. Fronds regularly branched 7

7. Fronds membranaceous, up to 1 m. wide 9. D. latissima (p. 568)

7. Fronds ligulate, up to 1 dm. wide 8

8. Fronds more than 3 m long with prominent aculeae along the margins

8. D. munda fp. .567)

8. Fronds less than 3 m. long, smooth or ^\^th fine aculeae along the margins.. 9

1925] Setchell-Gardner : Melanophyceae 561

9. Fronds up to 8 dm. high, 6-8 mm. wide, without distinct lateral veins, stipe

cylindrical, 3-4 mm. diam 6. D. ligulata (p. 566)

9. Fronds up to 2.5 m. high and 2 cm. wide, with distinct lateral veins, stipe

flattened and short 7. D. herbacea (p. 566)

Section 1. Virides
Desmarestia, subgen. Dichloria (Grev.) DeToni.

Fronds cylindrical or only very slightly compressed ; branching
mainly opposite.

DeToni, Syll. Alg., vol. 3, 1895, p. 456. Dichloria Greville, Alg.
Brit., 1830, p. 39.

1. Desmarestia media (Ag.) Grev.

Fronds cylindricaj, considerably cartilaginous, branching very
profusely and opposite throughout ; color dark brown, nearly black on
drying; branches of 5-6 orders, each successive order gradually
reduced in size, composed of an axial row of relatively large cells sur-
rounded by 2-3 layers of large, more or less irregular, elongated cells
surrounded in turn by a single palisade layer of cortical assimilating
cells, the terminal ramuli long-attenuate and acute.

Growing on rocks in the upper sublittoral belt. Unalaska, Alaska.

Greville, Alg. Brit., 1830, p. xl; Pease, P. S. M. S. Publ., vol. 1,
1917, p. 386. Desmarestia aculeata f. media Setchell and Gardner,
Alg. N.W. Amer.. 1903, p. 246; Collins, Holden and Setchell, Phyc.
Bor.-Amer. (Exsicc), no. 1036. Sporochnus medius Agardh, Sp. Alg.,
A'ol. 1, 1820, p. 153, Icones Alg. Ined., 1821, pi. 16. Desmarestia viridis
Saunders, Alg. Harriman Exped. 1901, p. 422 ( ?).

Desmarestia media var. tenuis S. and G.

Fronds cylindrical and with opposite branching throughout, very
profusely branched, 3-4 dm. high, stipe 2-3 mm. diam., ultimate
ramuli very slender and flaccid, 200-250/x diam. ; axial filament of the
fronds very large, 100-125ju, diam., surrounded by a single layer of
large, colorless cells between the axial filament and the small cortical
cells.

Growing on stones, etc., in the upper sublittoral belt. Alaska
(Juneau) to Puget Sound, Washington.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 7. '

562 University of California Publications in Botany [Vol. 8

There has been considerable confusion as to this species which was
described by C. A. Agardh from specimens said to have been collected
at Unalaska by Chamisso. Our specimens were collected at Unalaska
and in general seem to answer definitely to the description and speci-
mens of Agardh. The fronds are nearly exactly cylindrical through-
out, the branching is almost entirely opposite and the branches
decrease in size gradually for each successive order of ramifications.
In habit, branching, and in structure, D. media is clearly related to
D. viridis, but is a more ample and a coarser plant. It is clearly
distinct in habit, branching and structure from any of the actdeata-
group. Cross-sections do not show the dense external palisade-layer
of cells of the D. aculeata^ and its allies. The variety tenuis is typically
a much more slender plant, but is, nevertheless, coarser and more rigid
than any of the forms of D. viridis. The variety seems to be more
southern in its range than the type of the species.

2. Desmarestia pacifica S. and G.

Fronds slightly compressed below, terete above, branching opposite
throughout, each succeeding order of branches much reduced in size ;
subterminal branches 5-8 cm. long, tapering upwards but very slightly,
clothed with many, mostly opposite, blunt, ultimate ramuli, 2-5 mm.
long ; axial filament 40-50ft diam., surrounded by two layers of larger
cylindrical to subcylindrical cells; cortical layer not definitely
delimited.

Santa Catalina Island, southern California.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 6.

We have only a single and imperfect specimen upon which to base
this new species, but, while it closely resembles Desmarestia media,
each successive order, the branches of which do not taper appreciably
upwards, gives the plant a characteristic seemingly very distinct.

3. Desmarestia farcta S. and G.

Fronds cylindrical throughout, at least 5 dm. high ; branches long,
opposite throughout, not attenuated, blunt, of 3-4 orders, 0.5-1 mm.
diam. ; stipe and holdfast unknown ; primary cylinder composed of
cylindrical cells 20-25,/x diam. in the center of a distinct "stele" of
3-4 layers of larger cylindrical cells 35-45/x diam., these surrounded
by 2-3 layers of large colorless cells irregular in form, 100-125/x, diam.,

1925] Setchell-Gardner : 3Iehinophyceae 563

and surrounded in turn by the cortex of colored, radially elongated
cells ; reproduction and hairs unknown.

Cast ashore. Argyle, San Juan County, Washington.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 7.

We have but two portions of what may be a single ample plant of
this sjiecies upon which to base this description. The character of the
holdfast and of the lower parts of the frond cannot be chronicled at
this time, but the characters of the parts which we have are so pro-
nounced as to leave little doubt that the species has hitherto been
overlooked and is undescribed. One of the incomplete specimens,
probably a portion of the same plant, is in the collection of the Marine
Biological Station at Friday Harbor, Washington, and we are able
to examine it through the courtesy of Professor T. C. Frye. The
densely "stuifed" condition of the cells of the central monosiphonous
axis begins within two or three cells of the tip of each axis through
the growth of tylloses, a condition not noted by us in any other of the
specimens available to us.

Section 2. Aculeatae
Desmarestia, subgen. Eudesmarestia DeToni.

Fronds compressed to slightly flattened, never strictly flattened-
foliaceous, f oliaceous, or cylindrical ; branching alternate, occasionally
opposite below.

DeToni, Syll. Alg., vol. 3, 1895, p. 457 (in part).

4. Desmarestia latifrons (Rupr.) Kuetz.
Plate 90

Fronds of a firm, rigid consistency, dark brown in color, moderately
branched, flattened throughout except the base of the stipe, several
main lateral branches often exceeding in length the central axis,
midrib manifest only in the lower older parts, 8-15 dm. high, 2-3 mm.
wide ; branches of 3-4 orders, those of the fourth order f oliaceous
above, reduced to small aculeae below or entirely absent ; hairs oppo-
site, branched, arising from every cell of the central axial filament,
closely congested along the entire margin of the corticated frond and
of the ramuli except at their stipitate bases.

Growing on rocks in the lower littoral and the upper sublittoral
belts. Central Oregon (Coos Bay) to central California (Point Sur).

564 University of California PuhJications in Botany [Vol. 8

Kiietzing, Tab. Phyc, vol. 9, 1859, p. 40, pi. 95 ; Collins, Holden
and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 279; Farlow, Anderson
and Eaton, A\g. Exsicc. Amer-Bor., no. 121. Spinularia latifrons
Kuprecht, Tange, 1851, p. 375.

The type locality of D. latifrons is, according to Ruprecht (loc.
cit.)., Fort Ross, California. We have not seen any specimens from
the type locality, although we have both collected algae in that vicinity.
The species, as at present known, does not seem to be very abundant,
but is scattered sporadically from Coos Bay, Oregon, to Point Sur,
Monterey County, California.

As far as our experience goes, it is one of the most stable, least
variable species of the genus, and can readily be told from all other
known species. The hairs are relatively long, very abundant and
remain on until the plants are nearly maximum growth. These are
replaced in part by numerous prominent teeth or aculeae at maturity.
The fronds are decidedly rigid and of a dark color.

Desmarestia latifrons resembles Desmarestia intermedia more than
any other of the Acnleatae, but is definitely flattened rather than com-
pressed, and is not at all either membranaceous or foliaceous. It
seems to be a very distinct species of the central Californian coast
(extending over into the southern Oregon coast at Coos Bay), but is
scarce even in its own territory. The species is related to D. actdeata
(L.) Lamour., but more closely to D. intermedia P. and R., but is
broader and more flattened than either.

5. Desmarestia intermedia P. & R.

Fronds subcoriaceous, up to 6 dm. high and 1.5 mm. wdde, dark
brown in color, almost black on drying, very profusely branched, sub-
opposite and fasciculate below, strictly alternate above ; stipe and
lower parts cylindrical to subcylindrical ; ramuli decidedly compressed
and lanceolate, densely clothed in the juvenile stage with relatively
long hairs with alternate or opposite branching ; frequently, especially
in the lower parts and in the older plants, two or more branches arising
at the same node and on the same side of the frond.

Growing on rocks in the upper sublittoral belt. Bering Sea to
Puget Sound.

Posters and Ruprecht, lUus. Alg., 1840, p. 13, pi. 26 (Excl. Syn-
onymy) ; Desmarestia aculeata Setchell and Gardner, Alg. N.W. Amer.,
1903, p. 246 (in part); Tilden, Amer. Alg. (Exsicc), no. 352.
Possibly, Saunders, Alg. Harriman Exp., 1901, p. 422.

1925J Setchell-Gardner: Melanophyceae 565

Desmarestia intermedia has usually passed for D. aculeata, true
representatives of which do not seem to occur on our coast. D. aculeata
is characterized, at least in typical forms, by having its shorter and
even longer branches margined by short branches of limited growth,
asuming the form of spines. D. intermedia, however seldom has what
appear to be genuine aculeae, i.e., short rigid spine-like branches of
very limited growth such as are characteristic of D. aculeae and D.
latifrons, usually there being little, if any, appearance of this sort. One
characteristic of D. intermedia is the fasciculate branching below. The
groups of branches springing from one point are subopposite, i.e., arise
near one another, but one bunch is inserted higher on the axis whence
both arise and not exactly on the opposite side. The plants we have
assigned to this species vary from fairly broad (type) to medium (var.
fuscescens P. and R., Joe. cit.) and on to the most slender and almost
terete forms (var. teretifolia P. and R., loc. cit.).

Postels and Ruprecht {loc. cit.) seem to feel certain that the nomen
nudum of Mertens, Fucus pseudoaculeatus, refers to their plant, hav-
ing probably examined a specimen, but they consider the other three
quotations, viz., Sporochnus medius Ag., Trinitaria canfervoides Bory,
and Desmarestia media Grev., as dubious. The first and the last refer,
in our opinion, to what we call Desmarestia media. The middle plant
is presumably another species, having been assigned to a plant of
the southern hemisphere.

Section 3. Herbaceae

Desmarestia, subgenus Eudesmarestia, De-Toni {loc. cit.), In part.

Fronds ligulate to broadly membranaceous or foliaceous ; branch-
ing abundant to very sparse or none, opposite or subopposite.

De-Toni, Syll. Alg., vol. 3, 1895, p. 457.

There is a very considerable variation in length, width, and branch-
ing in our west American plants of the Herbaceae and there are very
considerable difficulties in the way of classifying them either as dis-
tinct species or as varieties. Miss Vinnie Pease (1917 and 1920) has
made an effort in this direction and we have considered her account
and the opinions she has expressed therein, very carefully. As a
result of our study of the literature and of the specimens, we have
decided to mark off the more striking plants of our coast as more or
less distinct species.

566 University of California Publications in Botany [Vol. 8

6. Desmarestia ligulata (Lightf.) Lamoiir.
Plate 87

Fronds plumose, up to 8 dm. high (in our territory), 6-8 mm. wido,
firmlj" attached hy a relatively large disk ; primary axis closely beset
with opposite, primary branches throughout, exceedingly variable in
size and complexity of branching, some of which extend beyond its
apex ; ramuli all opposite, closely crowded, of 4-5 orders, the ultimate
reduced to tine teeth or aculeae, all very decidedly flattened ; midrib
barely discernible in the primary axis and main laterals ; hairs very
abundant, with short, very acute, opposite branches, giving rise along
the margins of the ultimate branches to numerous small acute ramuli,
many of which disappear at maturity.

Growing on rocks in the upper sublittoral and lower littoral belts
among and in the vicinity of the San Juan group of islands,
Washington.

Lamouroux, Essai, 1813, p. 25 ; Harvey, Phyc. Brit., vol. 1, 1847,
pi. 115 ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 247 ; Pease,
P.S.M.S. Publ., vol. 1, 1917, p. 388, and vol. 2, 1920, p. 314; Wyatt, Alg.
Danmon., no. 55 ; Le Jolis, Alg. mar. Cherb., no. 268 ; Hauck and
Kichter, Phyk. Univ., no. 420. Fucus Ugulatus Lightf oot. Flora Scotia,
vol. 2, 1777, p. 946, pi. 29.

There may be still a question as to whether typical Desmarestia
ligidata occurs on our coast. Pease (1920, p. 316) has argued that it
does and we have adopted her opinion. There does occur on the island
of San Juan and apparently in abundance a narrow, branched plant
with short filiform stipe without prominent midvein, and lacking
lateral veins, which seems to correspond to Lightf oot 's figure of Fucus
Ugulatus and description and with British plants. The type locality
is on the Scottish coast and is probably on the Firth of Forth,

7. Desmarestia herbacea (Turner) Lamour.

Plate 88
Fronds ligulate, 1.5-2.5 m. long, 1-2 cm. wide, the rhachis and
the primary branches with a distinct, conspicuous midrib ; stipe very
short, almost immediately flattened; branches of 3-4 orders, each
order much reduced but exceedingly variable in size, the primary
branches often reaching the size of the rhachis, but interspersed with
branches only a few centimeters long ; all of the branches, though

1925] Setchell-Gardner : Melmiophyceae 567

relatively wide, tapering at the base to a small, distinct, almost cylin-
drical connecting stipitate portion, and rounding at the apices; the
branches exceedingly abundant in the juvenile stage, and all densely
clothed at the apices and along the margins with acute hairs, the main
terminal hairs with profuse opposite branching.

Growing in tide pools in the lower littoral and upper sublittoral
belt. Kodiak Island, Alaska, to southern California (La Jolla).

Lamouroux, Essai, 1813, p. 25 ; Kuetzing, Tab. Phyc, vol. 9, 1869,
pi. 100 ; Pease, P.S.M.S. Publ., vol. 1, 1917, p. 389 ; Ibid., vol. 2, 1920,
p. 340, pi. 54, fig. 3, pi. 60, figs. 1-5. Desmia herbacea Postels and
Ruprecht, Illus. Alg., 1840, p. 13. Desmarestia ligulata var. herbaceo.
Collins, Holden and Setchell, Phyc. Bjor.-Amer. (Exsicc), no. 19b.
Desmarestia ligtdata Farlow, Anderson and Eaton, Alg. Exsicc. Amer.-
Bor., no. 166. Fncus herbaceus Turner, Hist. Fuc, 1809, pp. 77, 78,
pi. 99.

The exact locality from which the material forming the type of the
species of D. herbacea was collected is not known. Turner {loc. cit.),
says, "North-west coast of America. Mr. Menzies. " There is much
variation in the size of the fronds as they occur along our coast.
Turner says, "two feet or more long." We have seen specimens two
and a half meters long. Some specimens, apparently mature, from
La Jolla and from Pacific Grove, California, in our herbarium are
almost as narrow as the specimens from San Juan County, Washington,
which we have referred to D. ligulata. Other specimens approach very
nearly to the narrow forms of I), niimda as we have limited that species.
However, the typical and abundant specimens are quite distinct from
either of these species. They are midway between the two species as
regards the number and character of the branches. Typical plants
also show distinct, often heavy, lateral veins, at least on all the broader
axes.

8. Desmarestia munda S. and G.

Plate 89
Fronds attached by a firm parenchymatous disk, ligulate, relatively
rigid and coriaceous, up to 8 m. long, 4—10 cm. wide, glossy yellowish
brown in color, sparingly branched at maturity ; midrib prominent in
the stipe and lower parts, becoming very inconspicuous or appearing
only as a mere nerve above ; branches usually of 2 but in part 3 orders,
the largest primary branches arising near the base at times as long as
the central rhachis, and even wider in part, tapering rather abruptly

568 University of California Publications in Botany [Vol. 8

to a small cylindrical connection at the base, acuminate or rounded
above, the margins of all bearing very prominent, rather distant spine-
like projections with more or less rounded angles above ; stipe flattened
almost to the base.

Growing principally on rocks, just below extreme low water level
down to ten fathoms or more in depth. From Puget Sound, Wash-
ington, and probably considerably farther north, to southern Cali-
fornia (San Pedro).

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 7. Desmarestia
ligulaia var. Jierhacea Setchell and Gardner, Alg. N.W. Amer., 1903,
p. 247; Tilden, Amer. Alg. (Exsicc), no. 244; Collins, Holden and
Setchell, Phyc. Bor.-Amer. (Exsicc), no. LXXIX A (not B). Des-
marestia latissima Pease, P.S.B.S. Publ., vol. 2, 1920, p. 319 (in part).

Desmarestia munda is a long and moderately broad plant, with
indistinct midvein, usually faint above, and lateral veins branching
and disappearing before reaching the margins. The texture of the
living plant is firm and its surface is smooth, shining, and usualty
free from epiphytes and epizoa. It is for this reason that we have
chosen the specific designation. Pease (1920, p. 319) has published a
Desmarestia latissima as of Setchell and Gardner and states that the
type is in the Herbarium of the University of Minnesota. We are
not responsible for the type, which we have not yet seen, but from the
description in Pease, conclude that it includes D. munda as we estab-
lish it, as well as a plant to which we did assign the provisional name
D. latissima. The latter is the plant to which we limit the name, D.
latissima: In our opinion, our D. latissirma is sufficiently distinct from
D. munda to justify keeping them separate.

9. Desmarestia latissima S. & G.

Fronds membranaceous, broadly expanded, 3-4 m. long, up to 12
dm. wide, sparsely branched, exceedingly fragile; branches similar to
the main axis, of one order only, or possibly at times a few small
secondary branches, abruptly tapering at the base to a very slender
connection with the main frond, rounded at the outer end, midrib and
lateral veins inconspicuous.

Growing in quiet water in bays and on mud flats, in the upper
sublittoral belt. San Juan County, Washington.

Setchell and Gardner, in Pease, P.S.B.S. Publ., vol. 2, 1920, p. 319
(in part).

1925] Setch ell-Gardner: Mehmophi/ceae 569

This is by far the largest species of Desmarestia yet discovered,
and seems, as far as known to us, to inhabit relatively quiet water, at
times merely resting on mud flats where the water is rarely disturbed
except as it rises and falls with the tidal movements. One of its dis-
tinguishing characters is its very large size. The most noticeable
feature about it, however, is its exceedingly fragile nature. It is very
crisp and brittle, wholly unable even to bear more than a fraction of
its own weight without being ruptured. It is further characterized by
the paucity of branches, as a rule bearing only a few primary laterals.
This plant fully merits the name Desmarestia latissima, as already
given by us, and is a plant of entirely different color, texture, habit,
and place of growth from our D. munda.

10. Desmarestia foliacea Pease

Fronds membranaceous, unbranched, attached by a small disk,
tapering below to a very delicate cylindrical stipe, up to 1 m. long,
15 cm. wide, margins irregularly sinuate with occasional sinuate teeth,
midrib evident, with opposite branches which break up into fine
veinlets.

Growing in the upper sublittoral belt. San Juan County, Wash-
ington.

Pease, P.S.B.S. Publ., vol. 2, 1920, p. 322, pi. 58, figs. 5-10, pi. 61,
figs. 1-5. Desmarestia tabacoides Pease, P.S.B.S. Publ., vol. 1, 1917,
pi. 84, figs. 2-7.

We are not acquainted with this plant and know it chiefly through
the above mentioned publications. There is a plant in the Herbarium
of the University of California (Gardner, no. 2321) dredged in ten
fathoms off Canoe Island, San Juan County, Washington, which may
possibly belong here. The plant was attached to a clam shell by a
relatively small, flat disk, has a small cylindrical stipe, a blade about
four meters long and three decimeters wide. There are, however, a
few small, short branches arising near the base, differing in this
respect from the type material as described by Miss Pease.

Through the kindness of Miss Pease, we have received, since writing
the foregoing, two sheets of specimens of Desmarestia foliacea from
Professor Josephine Tilden, of the University of Minnesota. The
species seems closely related to D. tabacoides Okam., but differs in size,
shape, lobing and venation.

570 University of California Publications in Botany [Vol. 8

Order 4. CHORDARIALES ord. nov.

Fronds erect, cylindrical, branched or unbranched, arising from
a discoid holdfast or from an expanded and lobed horizontal thallus,
with terminal, but subapical, growth, of three sets of tissues, central
core of vertical rows of elongated cells (often disappearing leaving
the erect filaments hollow), an intermediate layer of cells slightly
longer than broad or thick, and an outer layer of longer or shorter
assimilating filaments, more or less distinct or combined into a cortical
tissue, near, or at, whose bases the zoosporangia are formed ; zoo-
sporangia immersed among the assimilating filaments ; colorless hairs
generally present ; gametangia and gametophytes unknown, the later
presumably microscopic.

In pursuance of the idea stated elsewhere, that those groups of
Melanophyceae closely related in general vegetative structure and par-
ticularly with the same method of growth in length in which no member
is known to occur with gametangia, are to be suspected of possessing
microscopic gametophytes, we have established the order Chordariales
with the family Chordariaceae as the typical family, which includes
Chordaria, typified in Chordaria flag ellif ormis (Muell.) Ag., as the
type genus. The method of growth is that designated as subapical, as
distinguivshed from trichothallic (e.g.,Ectocarpales and Desmarestiales)
on the one hand and truly apical (e.g., Sphacelariales, Dictyo-
siphonales, etc.) on the other. The meristematic cells are situated in
short apical filaments similar to the external assimilatory filaments,
but are not terminal in those filaments, being situated two or three, or
occasionally more, cells below the terminal cell. In other respects the
structure approaches that of the ^giraceae and the Myriogloiaceae
where, however, the growth is trichothallic and both macroscopic
sporophytes and macroscopic gametophytes are knoM^n, From Hetero-
chordariaceae, the families of this order differ in lacking, so far as
known, a macroscopic gametophyte. While the family Chordariaceae
can be assigned to the order Chordariales, as the type family, yet the
close relationship of the genera Coilodesme and Scytothamnus lead
us to add a family for the first (Coilodesmaceae) and one for the second
(Scytothamnaceae), although we have no members belonging to the
latter within our territory.

Key to the Families

1. Assimilating filaments distinct 16. Chordariaceae (p. 570)

1. Assimilating filaments united into a cortical tissue ...17. Coilodesmaceae (p. 577)

1925] SetcheU-Gardner: Mclanophyceae 571

FAMILY 16. CHORDARIACEAE reichexh. (in part)

Fronds erect, cylindrical, branched or simple, arising from discoid
holdfasts or from lobed and expanded horizontal thalli, composed of
three tissues or layers, an inner, of elongated, usually slender fila-
ments (often disappearing and leaving a hollow center), an inter-
mediate, of large, rounded, or somewhat elongated cells, and an outer
cortical layer, of short, anticlinal, assimilating filaments, distinct from
one another, but enclosed in an enveloping jelly ; zoosporangia
immersed among the cortical filaments ; gametangia and micro-
scopic (?) gametophyte unknown.

Reichenbach, Conspectus Regn. Veg., 1828, p. 25 (in part; fide
Pfeiffer, Nomen. Botan., vol. 1, 1873, p. 732, et auct. var.) ; Harvey,
Ner. Bor. Amer., I, 1852, p. 121 (in part). Chordarieae Agardh,
Syst. Alg., 1824, p. xxxvi (in part).

The family of the Chordariaceae, while actually very distinct from
all others of the Melanophyceae except that of the Heterochordariaceae
because of its subterminal growth and frond structure, has been the
recipient of several families in which the growth is clearly tricho-
thallic. We may assume with reason, we think, that the subapical
method is intermediate between the apical and the trichothallic yet
distinct from each of them. The outer, cortical, layer of short, but
distinct, anticlinal filaments distinguish this familj^ from both the
Coilodesmaceae and the Scytothamnaceae.

Key to the Genera

1. Fronds simple (in our species) or very sparingly branched 2

1. Fronds, typically, much branched 34. Chordaria (p. 571)

2. Fronds composed of a flattened, sterile part and an erect, fertile part

35. Analipus (p. 575)

2. Fronds not composed of two parts 36. Gobia (p. 576)

34. Chordaria Ag. (emend Grev.)

Fronds cylindrical, filiform, branched, more or less rigid and
cartilaginous to slightly lubricous, solid or at times becoming fistulous
in age, composed of two tissues, a medulla and a cortex ; growth sub-
apical ; medulla composed of relatively large, colorless, longitudinally
arranged, firmly agglutinated filaments, with numerous small fila-
ments, especially abundant in the center of the frond, interspersed ;
cortical assimilating filaments short, firmly agglutinated ; reproduc-
tion in macroscopic plants by zoosporangia borne among the cortical
filaments.

572 University of California Publications in Botany [Vol.8

Agardh, Syn. Alg. Scand., 1817, p. xii and p. 12, Urn. mid.;
Greville, Alg. Brit., 1830, p. 44.

The name Chordaria was first proposed by Link in 1809 to include

a series of different plants, none of which, however, belong to this

entity as now constituted. C. Agardh (1817, p. xii) modified the

limits of the genus to include C. rotunda, C. divaricata, C. flagelli-

formis, C. cabrera, and C. Filum and three other doubtful species. Of

these C. flagelliformis is the only northern species of the series now

generally included under the genus, but some authors also include

C. divaricata. Greville (1830, p. 44, pi. 7) was the first definitely to

establish the present limits of the genus by pointing out a type, viz.,

C. flagelliformis.

Key to the Species

1. Fronds flagelliform, cylindrical throughout 2

1. Fronds irregular in diameter, flattened at the branching ...3. C. dissessa (p. 574)
2. Fronds relatively robust, up to 9 dm. high, usually with many long,

slender, primary branches 1. C. flagelliformis (p. 572)

2. Fronds slender, up to 3.5 dm. high, primary branches patent, scattered
along the whole main axis up to 4 cm. long 2. C. gracilis (p. 573)

1. Chordaria flagelliformis (Muell.) Ag.

Fronds attached by a small disk, solitary or in groups, lubricous,
solid, 2-5 dm. (up to 9 dm.) long, branching close to the base into
many branches of nearly equal length, or the main axis percurrent
giving rise to numerous alternate branches, shorter above ; branches
usually of 3 orders; peripheral assimilating filaments 6-8 cells long,
with terminal cell nearly spherical, and with numerous hyaline hairs
interspersed ; color dark olive brown, nearly black ; zoosporangia ovoid
to pyriform, arising at the base of the cortical filaments.

Growing on rocks in the upper sublittoral belt.

Agardh, Syn. Alg. Scand., 1817, p. 12 and p. xii, Fucus flagelli-
formis Mueller, Flor. Dan., 1771, pi. 650,

This species in its various forms seems to be exclusively boreal,
though extending much farther south on the Atlantic shore than on
the Pacific shore of North America. It is very common on the
American and European shores of the North Atlantic. It has been
reported as far north as the Spitzbergen Islands and in the Baltic
Sea. The following forms have been reported within our territory.

1925] SetcheU-Gardner: MeJanophxjceae 573

Chordaria flagelliformis f. typica Kjellm.

Plate 39, fig. 44

Fronds much branched, primary branch arising near the base,
usually longer than the others, main axis nearly percurrent.

Growing on rocks in the upper sublittoral belt. Bering Sea (St,
Lawrence Island) to Sitka, Alaska.

Kjellman, Om Spets. Thall. II, 1877a;, p. 28 ; Reinke, Atlas, 1889,
pi. 39, figs. 1-7; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 250;
Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1037.

Chordaria flagelliformis f. ramuscvilifera Kjellm.

Fronds cylindrical, up to 30 cm. long, slender, with scattered, short,
simple branches 1-1.5 cm. long, arising nearly perpendicular to the
main axis; cortical filaments 5-6 cells long, distinctly clavate, terminal
cell moderately swollen ; zoosporangia elliptical, 55-60/a long, 25-30/x
wide.

Growing in the sublittoral belt. St. Lawrence Island, Alaska.

Kjellman, Om Spets. Thall. II, 1877a, pp. 29, 30, pi. 1, figs. 10-12 ;
Setchell and Gardner, Alg. N.W. Amer., 1903, p. 251.

Chordaria flagelliformis f. chordaeformis Kjellm.

Fronds simple or sparsely branched ; cortical filaments nearly
cylindrical with terminal cell slightly swollen ; zoosporangia 60-70ju.
long, 25-30ja broad.

Kjellman, Om Spets. Thall. II, 1877a, pp. 28, 29, pi. 1, figs. 13-15 ;
Setchell and Gardner, Alg. N.W. Amer., 1903, p. 251.

This form has been reported by Kjellman in the upper sublittoral
belt at Konyam Bay, Siberia, and may be expected to occur in our
waters.

2. Chordaria gracilis S. and G.

Plate 84
Fronds 2.5-3.5 dm. high, with a single main percurrent axis
400-500/x diam. arising from a disk ; primary branches numerous,
short, scattered along the entire length of the 'main axis and alter-
nately arranged on all sides, up to 4 cm. long, standing almost at right
angles to the main axis; secondary branches 3-6 mm. long, standing

574 University of California Publications in Botany [^'ou 8

at wide angles with the primary branches ; color light brown ; zoo-
sporangia unknown ; cortical filaments short, composed of 2-3 cells
with a subspherical terminal cell.

Growing in abundance on stones. West shore of Amaknak Island,
Unalaska, Alaska.

Setchell and Gardner, Phyc. Cont., 1924, p. 8.

The slender percurrent axis bearing numerous short, almost equal,
lateral branches over most of its length with short branchlets and all
axes patent to one another, give the plants of the single collection
available a very distinctive appearance. The plants are young and
sterile, but the structure of the frond, the terminal subapical growth
and the discoid holdfast indicate its affinities with Chordaria. In the
Algae of Northwest America (1903, p. 248) we placed this plant
under Dictyosiphon foenicidaceivs (Huds.) Grev., an untenable posi-
tion, since the subterminal growth and other details of structure plainly
show its different nature.

3. Chordaria dissessa S. and G.

Plate 41, fig. 57, and plate 75

Fronds attached by a small disk, 15-25 cm. high, up to 2 mm. thick,
moderately and alternately branched, usually wider at the branching,
but swollen in some parts and contracted in others, older parts at
times fistulous, olive brown in color, composed of a medulla of longi-
tudinal filaments giving rise to a cortex of short assimilating filaments ;
filaments of the medulla composed of large, colorless, closely compact,
thick-walled cells up to 120/^ diam. in the central part of the frond,
many times longer than the diameter, becoming much shorter and
narrower toward the periphery ; cortical erect filaments cylindrical,
simple or branched, not closely compact, 4-6/x diam., composed of 2-4
cells, terminal cell swollen and globular ; hairs unknown ; branches of
each different order reduced in size, terminal ramuli long-attenuate
and acute, angles mostly wide and rounded ; zoosporangia obovoid to
ellipsoidal, 36-42/^ long, 28-32jli broad ; gametangia unknown.

Growing on eel grass in the middle and lower littoral belts. East
Sound, Orcas Island, Washington.

Setchell and Gardner, Phyc. Cont., 1924, p. 8. Castagnea divari-
cata Setchell and Gardner, Alg. N.W. Amer., 1903, p. 249 (not C.
divaricata (Ag.) J. Ag.).

The resemblance between Chordaria dissessa and C. divaricata
(Ag.) Grev. is superficially close, but the former has its main axes less

1925] Setch ell-Gardner : Melanoplnjceae 575

regularly cylindrical, is less profusely branched, seems less gelatinous,
with fewer or no hairs and has the assimilating cortical filaments
shorter and of fewer cells, being 2-4 celled, while in C. divaricata they
are 4-6 even more celled. The aspect of C. dissessa is that of a coarser,
less abundantly branched, plant than is that of C. divaricata.

35. Analipus Kjellm.

Thallus composed of an expanded, more or less deeply lobed,
pseudoparenchymatoiLs, sterile, basal portion attached to the sub-
stratum by numerous, unicellular or multicellular, rhizoidal filaments,
and giving rise above to simple, more or less cylindrical, solid or
fistulous, fertile fronds; fertile fronds composed of a dense mass of
pseudoparenchymatous cells with a few chromatophores giving rise
on the surface to short, densely congested, cortical, assimilating fila-
ments and to numerous zoosporangia ; gametangia unknown.

Kjellman, Om Beringh. Alg., 1889, p. 48.

The type and only known species of the genus is A. fiisiformis
from Bering Island.

^t5

Analipus fusiformis Kjellm.

Fertile fronds 2-6 cm. high, 1-3 mm. diam., one to several arising
from the upper side of the same horizontal base; the sterile base pro-
fusely and deeply lobed, terminal lobes rounded and blunt, attached
by short, somewhat branched, monosiphonous hairs; cortical filaments
of fertile fronds 3-5 cells long, clavate, with enlarged, rounded,
terminal cell ; zoosporangia arising from the base of the paraphyses,
globose to ellipsoidal, 35ju. long, 25/a broad (fide Kjellman), up to 65/x
long.

Growing on rocks. St. Paul Island, Bering Sea, Alaska.

Kjellman, Om Beringh. Alg., 1889, p. 49, pi. 7, figs. 6-12 ; Setchell,
Alg. Prib. Isl., 1859, p. 591 ; Setchell and Gardner, Alg. N.W. Amer.,
1903, p. 252.

The material reported here seems to be almost identical with that
found at Bering Island, the type locality, and reported by Kjellman.
The zoosporangia are somewhat larger than the dimension given by
Kjellman, some being as long as 65/x.

Analipus seems to belong to the Chordariaceae as we have limited
it, and we have been able to assure ourselves that the method of growth
in length is subapical and, consequently, that the agreement with

576 University of California Puhlications in Botany [Vol. 8

Chordaria in family characteristics is close. In case Ruprechtiella
filiformis (Rnpr.) Yendo (Trav. Mus, I'Acad. Imp. Sci. St. Petersb.,
vol. 10, 1913, p. 118) proves to be identical, generically at least, with
Analipus fusiformis Kjellm., Analipus must be removed to the Hetero-
chordariaceae.

36. Gobia Reinke

Frond filiform, more or less lubricous, simple or branched, hollow
except in the very young stage, growth apical or subapical, composed
of an inner tissue of loosely anastomosing colorless filaments giving
rise to short, compact, vertical, assimilating filaments, to zoosporangia
and to scattered colorless hairs; gametangia unknown.

Reinke, Algenfl. westl. Ostsee, 1889a:, p. 65.

The type of the genus, and the only other species known except the
following, is Gohia haltica, based upon material of Cladosiphon
halticus Gobi (Brauntange finn. Meerbus., 1874, p. 12).

The growth in length of the species of Gohia is said to be from an
apical cell at first, but this soon ceases and intercalary growth sets in
(cf. Reinke, 1889a, pp. 65, 66). If we are correct in referring the
Mesogloi<a simplex Saunders to Reinke 's genus Gohia, with which it
seems to agree in details of structure, we are probably also correct in
interpreting the growth of the genus as being subapical in that the
apical cell in very young plants and in very young branch initials very
early cuts off a subapical cell which takes on the principal meristematic
function. The same thing happens in Chordaria and presumably also
in Myelophycus and in Analipus. This is very different from the
situation in Dictyosiphan, where the terminal cell retains its meris-
tematic function and activity as long as active growth in length con-
tinues. Gohia, from this point of view, differs from the other genera
of the Chordariaceae in the loosely placed anastomosing filaments of
its inner tissues.

Gobia simplex (Saunders) S. and G.

Plate 42, fig. 58, and plate 786
Fronds olive brown, 5-12 cm. high, 2-3.5 mm. diam., simple, cylin-
drical or in part decidedly clavate, often arcuate, very blunt, solid in
the juvenile stage, soon becoming hollow except at the very base of the
apex, attached by a small disk ; central filaments loose, colorless, ana-
stomosing freely, 25-35ju, diam. ; cortical filaments straight, perpendi-
cular to the main axis, the terminal segments composed of 2-4 cells,

1925] 8 etch ell-Gardner: MeJanophyceae 577

terminal cell pyriform, 30-35ju, long, 18-22/x diam., with large ehroma-
tophores; zoosporangia obovoid to ellipsoidal, 40-50|U. long, SO-SSjit
broad ; hairs sparse, 4.5-5.5/* diam.

Growing on Heterochordaria ahietina (Rupr.) S. and G. Extend-
ing from Cook Inlet, Alaska, to Vancouver Island.

Setchell and Gardner, Phyc. Cont., 1894, p. 12. Mesoghia simplex
Saunders, Alg. Harriman Exp., 1901, p. 423, pi. 50, figs. 3, 4 ; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 250.

The measurements given in the diagnosis apply to the collection
from Sitka. Saunders (loc. cit.) gives 25-35/* long and 15-25ju, broad
for the dimensions of the zoosporangia. Those in the collection of
Polley and Butler in the Herbarium of the University of California
(no. 99329) from Vancouver Island, have zoosporangia up to 80/* long.
Otherwise they agree well with the Sitka plants. Small immature
specimens of this plant were found growing on Heterochordaria
ahietina distributed by Miss J. E. Tilden (Amer. Alg., no. 348 sub
Chordaria ahietina) .

Oobia simplex differs from Gohiu. haltica in being simple and in its
tendency to be somewhat enlarged above. Gol)ia simplex has always
been found growing on Heterochordaria ahietina S. and G.

FAMILY 17. COILODESMACEAE fam. nov.

Fronds membranoiLS, tubular or ligulate, growth subterminal at
least in early stages, possible terminal at the beginning, composed of
at least two tissues, inner composed of more or less elongated cells,
becoming smaller outwards and passing into short anticlinal filaments
firmly agglutinated into a cortical assimilating tissue ; zoosporangia
immersed in the cortical tissue but extending to, or near to, the sur-
face ; gametangia unknown and gametophyte presumably microscopic.

Coilodesmeae Kjellman, in Engler und Prantl, Die natiirl.
Pflanzen fam., 1 Th., 2 Abt., 1893, p. 200.

Stroemfelt (1886, p. 47) assigned his new genus CcUodesme to the
Chordariaceae, a.s then understood, and as an intermediate type
between the Chordarieae and the Scytosiphoneae. In vegetative struc-
ture, Coilodesme is more highly differentiated than Chordaria, in that
its cortical filaments have lost their distinctness and are combined
into a definite tissue in which the zoosporangia are immersed. The
method of growth has not been carefully or completely determined
but seems (cf. Oltmanns, 1922, p. 62, fig. 354, 2) to be subterminal.

578 Universitij of California Publications in Botany [^"ol. 8

No species of Coilodesme has as yet been seen with gametangia and
this suggests the extreme possibility of a microscopic gametophyte.
This combination of characters leads us to suggest the new familj^
Coilodesmaceae of the order Chordariales.

Key to the Genera

1. Fronds solid, ligulate 38. Phaeostrophion (p. 585)

1. Fronds hollow, complanate or saccate 37. Coilodesme (p. 578)

37. Coilodesme Stroem.

Fronds saccate, complanate from the beginning or more or less
flattened in age, linear to ovate or obovate in outline, rounded above,
tapering more or less at the base into a longer or shorter solid stipe,
with small disk-shaped holdfast or with penetrating rhizoids epiphytic,
parasitic or saxicolous, composed of two tissues, the inner consisting of
large, irregular, thick-walled cells, with few chromatophores, the
outer cortical portion consisting of a few to many layers of smaller
color-bearing assimilating cells arranged more or less in radial rows ;
hairs absent ; unilocular zoosporangia developed within the cortex,
scattered over the whole surface of the frond except the stipe and
extreme basal portion ; paraphyses absent ; gametangia unknown.

Stroemfelt, Meeresalg. Isl., 1886a, p. 173.

The type species of the genus is C. hulligera, of Icelandic origin.

The species of Coilodesme seem to divide themselves fairly defi-
nitely into two groups, the one saxicolous, the other epiphytic. Of
the epiphytic plants in the collections available to us, all are from the
western coast of North America or the northeastern coast of Asia and
seem to differ from one another essentially according to their host
plant, only one host plant having more than one species associated
with it. This is not surprising, since Rathbone (Journ. Linn. Soc,
1904, p. 674, pi. 24, figs. 5, 6) has shown the actual penetration of the
host and our own investigations have demonstrated an intimate meta-
bolic relation. The parasitism of all the epiphytic species of our coast
is also striking in that the hosts are all members of the related genera,
Halidrys, Cystoseira, and Cystophyllum, each of which seems to have
one or more species of Coilodesme peculiar to it. Okamura (Icones
Jap. Algae, vol. IV, no. 3, 1918, p. 55, pi. 144, figs. 10-13) figures
and describes a Coilodesme Cystoseirae growing on Cystophyllu-m
liaJiodatense at Akkeslie, Japan. From the data presented, we are
inclined to question the reference of this plant to C. Cystoseirae as
interpreted by us, but careful comparison is necessary to determine

1925] Setchell-Gardner: Melanophyceae 579

exactly the nature of Ruprecht's Asperococcus Cxjstoseirae and its
relation to the Japanese plant on the one hand and the Alaskan plant
on the other. Ruprecht states that the type of A. Cystoseirae is para-
sitic on Cystoseira Lcpidium but not on C. thyrsigera.

Key to the Species

1. Fronds complanate from the beginning, rigid, membrane up to 375/z thick,

epiphytic, stipe short 6. C. rigida (p. 584)

1. Fronds inflated at first, collapsing, especially when ruptured, membrane less

than250/i thick 2

2. Fronds ample, up to more than 1 m. long, and 1 dm. broad, epiphytic,

stipe short 1. C. californica (p. 579)

2. Fronds smaller in all proportions 3

3. Fronds narrowly cylindrical, up to 8 mm. diam., and 8 dm. long, epiphytic,

stipe short 5. C. Cystoseirae (p. 583)

3. Fronds more or less irregular, subcylindrical 4

4. Fronds finely corrugated, subsessile, epiphytic, stipe short

3. C. corrugata (p. 582)

4. Fronds undulate, decidedly stipitate 5

5. Fronds saccate, slightly undulate, saxicolous, stipe slender, elongated

2. C. bulligera (p. 581)

5. Fronds flattened, decidedly bullose, with ruffled margins, saxicolous, stipe

slender, elongated 4. C. polygnampta (p. 582)

5. Fronds moderately bullose, epiphytic, stipe slender 7. C. sitchensis (p. 584)

1. Coilodesme californica (Rupr.) Kjellm.

Plate 46, figs. 1, 2, and plate 86

Fronds flabby and fragile, smooth when young, more or less
wrinkled, or bullate, and ruffled along the margins at maturity, inflated
in the juvenile stage, soon collapsing because of ruptures, usually at
the apex, chiefly cylindrical, rounded above, tapering rather abruptly
at the base to a very short stipe 1-2 mm. long, variable in size from a
few decimeters to over a meter long, up to 12 cm. wide, 175 to 215ju,
thick, composed of an inner tissue of 4-5 layers of large, colorless cells
intermixed with smaller cells, and a cortical tissue of cells in anti-
clinal rows, 2-4 cells in a row ; zoosporangia very irregular in form
and size, 40-45/* long; color light bro^ai, changing to greenish on
exposure.

Epiphytic on Cystoseira and Cystophyllum. Puget Sound region
to southern California (San Pedro).

Kjellman, Undersok. Slagtet Adenocystis, 1889a, pp. 4-8, figs. 1-8
(excl. specimen) ; Saunders, Phyc. Mem., 1898, p. 160, pi. 29, figs. 1-3,
Alg. Harriman Exp. 1901, p. 422; Setchell and Gardner, Alg. N.W.
Amer., 1903, p. 241 ; Collins, Mar. Alg. Vancouver Isl, 1913, p. 106 ;

580 University of California Puhlications in Botany [Vol. 8

Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), Fasc. A,
no, I; Tilden, Amer. Alg. (Exsicc.) no. 354.

Adenocystiis {Lessonii var. ?) calif ornica Ruprecht, Tange Och.,
1851, p. 291. Coilodesme amplissima, Setchell, Algae novae I, 1912,
p. 232.

The question as to the identity of the Adenocystis (Lessonii var.?)
calif ornica of Ruprecht (loc. cit.) rests upon an imperfect description
on the part of Ruprecht. The more definite statements of Kjellman
and his illustrations seem to indicate that the plant intended was, at
least, not ample, but in fact short and with a comparatively long
slender stipe, dark brown, and an epiphyte. This was the inference
of Setchell (1912, pp. 231, 232) although the herbarium specimens
selected under that impression did not thoroughly correspond. Yendo
(1913, pp. 279, 280) has stated that Kjellman 's Bering Island plant
(loc. cit.) does not agree with Ruprecht 's type, which does agree with
Tilden 's plant (no. 354, American Algae) from Vancouver Island
which Setchell (loc. cit.) had referred to his Coilodesme wmplissvtna.
Yendo makes no statements as to characters or differences and his
opinion is so little in agreement with the figures quoted by Ruprecht
(viz., in Voy. Coquille, Bot. Atlas, 1826, pi. 11, fig. 2) that we hesitate
to prefer his judgment to that of Kjellman. The important character
separating the Coilodesme calif ornica of Kjellman and the C. amplis-
sima of Setchell is the stipe, comparatively long and slender in Kjell-
man 's plant and practically wanting in Setchell 's species. Ruprecht 's
plant, however, was epiphytic on Cystoseira osmundacea, as is C.
amplissima, while Kjellman 's plant is said to be epiphytic on larger
algae. C. amplissima is widespread along the coast from Yakutat Bay,
Alaska, to San Pedro, California. From the ma2;e of uncertainty, it
seems best to adopt Yendo 's statement as to the identity of Ruprecht 's
plant and to regard C. amplissima as a synonym under C. calif ornica.

Coilodesme calif ornica, as identified by Yendo, is an ample saccate
species, the largest of the genus, with delicate walls, usually light
yellowish brown in color, with extremely short stipe and parasitic on
Cystoseira osmundacea and Cystophyllum. geminaium. It extends
from the Puget Sound region and possibly even from Yakutat Bay,
Alaska (Saunders, 1898, p. 422), although the more northern refer-
ences are subject to suspicion as to possible confusion with our C.
sitchensis. Its southern limit is at San Pedro, California, so far as
shown by available specimens. It is the only species as yet detected on
the coasts of central and northern California where is situated Fort
Ross, the designated type locality of the species.

1925] Setchell-Gardner : Melanophyceae 581

2. Coilodesme bulligera Stroem.

Plate 45, figs. 77, 78

Fronds usually fasciculate, 5-12 cm. high, 15-22 mm. wide, 75-85/>i
thick, moderately firm, acuminate or rounded above, gradually nar-
rowed below into a delicate stipe about a decimeter long, composed
of 1-2 layers of large, thin-walled, colorless cells within and a cortical
tissue of three layers of cells in vertical filaments or anticlinal rows;
color dark brown, plants adhering firmly to paper on drying.

Growing on rocks in the lower littoral belt, Shumagin Islands,
Alaska, to Coos Bay, Oregon.

Stroemfelt, Meeresalg. IsL, 1886a., p. 173, Om Algenveg. vid Isl.
Kuster, 1886, p. 48, pi. 2, figs. 9-12; Saunders, Alg. Harriman Exp.,
1901. p. 422; Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc),
no. 923a; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 240 (in
part).

The saxicolous species of Coilodesme are found in the Arctic Ocean
where it joins the North Atlantic (Iceland and Greenland) and in the
northern portion of our own territory. Coos Bay, Oregon, is as far
south as any has been found. The specimens collected by Gardner
at Coos Bay agree well in habit and structure with some collected
at Dixon Harbor, Alaska (somewhat northwest from Cape Spencer)
by G. B. Rigg. The habit of both sets of plants is close to those col-
lected and referred to C. californica by Kjellman at Bering Island
(cf. Kjellman, 1889a, figs. 1-3) but his were epiphytic and, conse-
quently, to be suspected as being of some different species. The plants
already mentioned agree also with the plant distributed by DeAlton
Saunders from '* Alaska" under 923rt of the Phycotheca Boreali-
Americana (at least in two copies in our possession) and with certain
specimens collected by Gardner at Whidbey Island (no. 201). These
plants all agree fairly well with the Stroemfelt figure of Coilodesme
l)ulligera in being gradually and insensibly attenuated into a distinct
slender, comparatively long stipe and in having the cortical layer of
anticlinal rows of cells of three to four each and each terminating
outwardly in a larger, dark brown, rounded cell. We are, consequently,
referring all these to Stroemfelt 's species, but with some uncertainty.
It is to be noted that 923^ of the Phyc. Bor.-Amer. is referred by us
to our C. pohjgnampta.

582 University of California Publications in Botany [Vol. 8

3. Coilodesme corrugata S. and G.

Plate 46, figs. 3, 4

Fronds very fragile and flaccid, very closely wrinkled all over the
surface, 3-7 cm. long, 8-14 mm. broad, 40-55//, thick, attached to the
host by (penetrating?) rhizoidal filaments, rounded above, attenuated
below into a short delicate stipe ; color light brown ; internal tissue
composed of 2 layers of large, irregular, colorless cells and the cortical
tissue composed of 2-3 layers of small, angular cells; zoosporangia
irregular in form, usually broader than long.

Epiphytic upon Cystoseira neglecta S. and G. In the upper sub-
littoral belt. Santa Catalina Island, southern California.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 8.

This is the smallest and most delicate species of the genus, and
seems related to C. amplissima in general outline and structure. These
characters, together with the delicate wrinkling and its occurrence on
a distinct and not common host, lead us to keep it distinct. Appar-
ently it is quite limited in its distribution.

4. Coilodesme polygnampta S. and G.

Plate 82

Fronds usually densely fasciculate, 1-4 dm. high, 1-5 cm. broad,
moderately firm, complanate, at least at maturity, with definitely
undulate and crisped margins, attached to rocks by a small, paren-
chymatous, disk-shaped holdfast, rounded and blunt above, tapering
more or less abruptly at the base into a definite, small, solid, cylin-
drical stipe, dark reddish brown in color ; histologically the interior
tissue is composed of about 2-3 layers of colorless cells, the cortical
tissue is composed of about 3 layers of small, assimilating cells in
anticlinal rows.

Growing on rocks in the lower littoral and upper sublittoral belts.
Bering Sea.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 9. Coilodesme
hiilligera Setchell and Gardner, Alg. N.W. Amer., 1903, p. 240 (in
part) ; Collins, Holden and Setchell, Phyc. Bor-Amer. (Exsicc),
no. 923&.

The plants referred from our Pacific Coast to Coilodesme hiilligera
Stroemf., do not all seem, on further study and careful comparison,
to be the same as those of Iceland and Greenland. Our specimens

1925] Setchell-Gardner: Melanophrjceae 583

from Bering: Sea are more ample, broader, taperinfj more abruptly
into the slender stipe (at least in older specimens) than is represented
in the figure of Stroemfelt (1886, pi. 2, fi(j. 0) or as compared Avith a
si:)ecimen from Greenland distributed by Kosenvinge. In Stroem-
felt's figure and in Rosenvinge's specimen, the cortex is made up of
closely packed anticlinal rows of 3-4 cells each and each row ends
outwardly in a somewhat larger, dark brown, rounded cell. In the
Bering Sea plant, the anticlinal rows are not recognizable, or are at
most of only two cells, the outer deep colored cell usually sitting imme-
diately upon the larger transparent outer cells of the inner layer. The
typical form of our plant which we consider a distinct species is repre-
sented by no. 9236 of the Phycotheca Boreali-Americana, and the tj'pe
locality is Amaknak Island in the Bay of Unalaska.

5. Coilodesme Cystoseirae (Rupr.) S. and G.

Fronds flaccid, irregularly cylindrical, blunt at the apices, taper-
ing abruptly into a short stipe about 2 mm. long, 1-4 dm. (up to 8 dm.)
high, 3-8 mm. diam., broad, usually inflated, composed of 2 layers of
large, thin-walled, colorless cells on the interior and 2 layers of small,
cuboidal, cortical cells; zoosporangia ovoid, 15-20/i. long, 11-14/x wide.

Epiphytic on CystophyUum geminatum. Kukak Bay and Yakutat
Bay, Alaska.

Setchell and Gardner, Alg. N.W. Amer., 1903, p. 241; Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1128. Coilo-
desme linearis Saunders, Alg. Harriman Exp., 1901, p. 421, pi. 48 ;
Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 824.
Asperococcus Cystoseirae Ruprecht, Tange Och., 1851, p. 370 ; not
Eyicoelium Cystoseirae Kuetzing, Tab. Phyc, vol. 9, 1859, pi. 6, fig.
IV, e-h.

This species of Coilodesme may readily be distinguished from all
other known species of the genus by its relatively narrow and long
cylindrical fronds, gradually attenuated at the base to a short stipe.
According to Saunders, it replaces the broader plant which he referred
to C. californica (probably our C. sitchensis) on the coast of Alaska
to the westward of Yakutat Bay. The dimensions given by Ruprecht
for his plants are far under those of the plants referred by us to his
species, but his plants were probably very young, similar to the very
young and slender individuals included by Saunders {loc. cit.) in the
group figured by him. Ruprecht 's plants, also, showed no reproductive
bodies.

584 University of California Publications in Botany [Vol.8

6. Coilodesma rigida S. and G.

Plate 45, fig. 76, and plate 79Z>

Fronds rigid, more or less coriaceous, complanate from the begin-
ning, and smooth, thickened along the margins, attenuated toward
the base into a thick, short stipe, broadened and rounded above, 5-10
cm. (up to 25 cm.) high, 1-2.5 cm. wide, 300-375/* thick, light brown
in color ; inner tissue composed of large, colorless, thick-walled cells
intermingled with numerous smaller filaments extending irregularly
in all directions, but toward the surface becoming smaller, perpendi-
cular to the surface and dividing di-trichotomously, terminating in
anticlinal rows of small, cortical cells ; zoosporangia irregular in form,
several times longer than broad.

Epiphytic on Halidrys dioica Gardner in the upper sublittoral
belt. Redondo to San Diego, southern California.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 9.

Coilodesme rigida is one of the smaller species of the genus. It is
very firmly attached to the host by numerous deeply penetrating
rhizoidal filaments. It may readily be disting-uished by its rigid,
smooth, thick frond, usually thickest along the margin at maturity,
and by its complanate form from the beginning. It is of a decidedly
dark brown color, complanate from its early stages and, apparently,
confined to HaUdrys dioica as a host plant. The plant is smooth even
on the margins which, in the adult plant, are decidedly thickened,

7. Coilodesme sitchensis S. and G.

Fronds thin and flaccid, inflated, moderately undulate, 15-25 cm.
high, 3-6 cm. wide, cylindrical to slightly clavate, tapering abruptly
at the base into a slender short stipe ; color dark reddish brown, espe-
cially so on drying, internal tissue of 2 layers of large cells, cortex of
2-3 layers of smaller cells, radially elongated, both tissues bearing
chromatophores.

Growing on CystophyUum geminatum (Ag.) J. Ag. Sitka, Alaska.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 9.

Cailodesme sitchensis occurs on the same host as C. linearis and
C. calif arnica. From the former it is to be distinguished by its broader
and more bullose habit and from both by its longer and more definite
stipe. It is also darker in color than C. calif ornica and rather more
bullose and undulated on the margins after collapsing. It may be
this species which was reported by Saunders on "CystophyUum
Lepidium" in localities north of the Puget Sound region.

i^--5] 8etcheH-Gardner: Melanophyceae 585

38. Phaeostrophion S. and G.

Fronds ligulate, solid, polystromatic, more or less stipitate, attached
by a small disk ; central tissue composed of elongated cells, irregular
in shape and size, merging on each side into smaller parenchymatous
cells which in turn merge into palisade-like cortical cells ; reproduction
by zoosporangia on both surfaces of the frond immersed in the cortex
extending to but not projecting beyond, the surface ; gametangia and
hairs unknown.

Setchell and Gardner, Phyc. Cont., VTI, 1924, p. 10.

The genus Phaeostrophion resembles \QTy closely the genus Enda-
rachne in size, general shape and consistency of the frond, but is quite
different from that genus in its histological characters and in its
method of reproduction. The development of the frond of Phaeostro-
phion has not been determined since only old battered specimens are
available. The outer ends seem to be the oldest, judging from the
more or less battered condition in which the specimens were found.
The tissues are almost parenchymatous throughout, but there is a
slight tendency in the center of the old fronds to a thick-walled fila-
mentous condition, in this respect resembling the central tissue of
Eudarachiie, in which the whole central tissue is composed of narrow
intertwined filaments.

In Endarachne the small surface cells divide tangentially to build
up the solid palisade layer of gametangia, but in Phaeostrophion cer-
tain cells of the inner layers of the cortex are transformed directly
into zoosporangia. In this last character Phaeostrophion resembles
very closely the genus Coilodesme which, however, is always hollow.
In its parenchymatous character, and in having larger cells in the
center and smaller toward and on the surface, it closely approximates
to the condition prevailing in the fronds of Ilea Fries.

The structure of the frond in connection with the method of
reproduction would seem to be sufficient to relate Phaeostrophion to
the members of the Coilodesmaceae, in which family we propose to
place it.

586 University of California Puhlications in Botany [Vol. 8

Phaeostrophion irregulare S. and G.

Plate 38, fig. 36, plate 50, fig. 8, and plate 85

Fronds linear to linear-spatulate, frequently much distorted and
irregularly notched or lobed ; 15-25' cm. (up to 40 cm.) high, 1.5-4 cm.
wide ; base long and gradually attenuate, becoming very slender at the
small disk-shaped holdfast ; zoosporangia numerous, elongate-polygonal
to regularly ellipsoidal, 38-44ja long, 26-34/a broad ; gametangia and
hairs unknown.

Growing on rocks in pools in the middle of the littoral belt. Mouth
of Coos Bay, Oregon, and Bolinas Bay, California.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 10.

The plants collected at Coos Bay were all much smaller than those
found growing at Bolinas. It was supposed at the time of collecting
that the former were rather rigid specimens of Ilea Fries and no micro-
scopical examination was made. Later they were soaked out and the
structure found to be practically the same as that of the plants from
Bolinas. Although collected nearly two months earlier, the Coos Bay
plants have zoosporangia which seem to be as nearly mature as those
of the plants from Bolinas. In neither collection have any zoospores
been seen, but the protoplast seems on the verge of transformation.
The zoosporangia are very numerous and are scattered quite uniformly
over the whole of both surfaces of the frond except the stipitate
portion.

Order 5. DICTYOSIPHONALES ord. nov.

Fronds cylindrical, filiform, repeatedly branched, solid or fistulose,
attached by a solid parenchymatous disk, composed of two, or in some
cases three, tissues, an inner of elongated slender or stouter cells, and
an outer of shorter, nearly iso-diametric or somewhat flattened cells ;
growth from a distinct apical cell ; hairs scattered singly over the
entire surface of the frond, soon deciduous; zoosporangia unilocular,
embedded in the cortex, borne on macroscopic plants ; gametangia
plurilocular on microscopic filamentous gametophytes.

The species of the genus Diictyosiphon (including Coil one ma) are
to be distinguished by the possession of an apical cell which persists
as long as growth in length continues, when it seems to be replaced,
in some instances at least, by a hair (cf. Kuckuck, im Oltmanns, 1922,
pp. 60, 61). The species of Dmtyosiphon, also, were never known
except as to the macroscopic plant with zoosporangia until Sauvageau

i92o] SetchcU-Gardner: Melanophyceae 587

(1917, pp. 829-831) annoimced tlie discovery of a reduced, practically
microscopic gametophyte, filamentous like a dwarfed Ectacarpiis with
plurilocular gametangia giving rise to motile gametes Avhich either
fuse or develop parthenogenetically. Such a gametophyte is inter-
mediate between the reduced, yet still macroscopic, gametophyte in
some of the Asperococcaceae, and the very much modified gameto-
phyte of the Laminariaceae. Besides the Dictyosiphonaceae, it may
be desirable also to include in the Dictyosiphonales, the family of the
Spermatochnaceae, at least as limited to species with growth from a
single apical cell, in which ease our diagnosis of the order must
necessarily be modified.

FAMILY 18. DICTYOSIPHONACEAE detoni
Characters of the order

De-Toni, Syst. Uebers. Fucoid., 1891, p. 179 ; Syll. Alg., vol. 3, 1895,
p. 4-18 ; Kjellman, m Engler and Prantl, Die natiirl. Pflanzenfam.,
1 Th., 2 Abt., 1893, p. 212. Dictyosiphoneae Kuetzing, Sp. Alg., 1849,
p. 484; Thuret, in Le Jolis, Liste alg. mar. Cherb., 1863, p. 72 (all
in part).

We have restricted the Dictyosiphonaceae to the species of Dictyo-
siphon in the broader sense as including Coilanema Aresch. The genera
Gohia and Scytothamnus seem to us to be more properly assigned to
the Chordariales, since the growth in length is subapical rather than
strictly terminal. The resemblances between the Dictyosiphonaceae
and the Spermatochnaceae consist in having growth in length from
an apical cell and producing only zoosporangia on macroscopic plants.
The genus Dictyosiphon is readily recognized and delimited, but with
the species, it is otherwise. We have restudied the specimens from
our coast and arranged them as best we may.

39. Dictyosiphon Grev.

Fronds attached by a small parenchymatous disk, terete, filiform,
with several orders of branches arising alternately from all sides,
solid or at times fistulose, growing from an apical cell, composed of
two tissues, a central core of longitudinally elongated, colorless cells,
surrounded by a tissue of small, more or less polygonal, color bearing
cells ; reproduction by zoosporangia embedded in the cortical tissue ;
colorless hairs abundant.

Greville, Alg. Brit., 1830, p. 55.

588 University of California Publications in Botany [Vol. 8

The type and only species mentioned by Greville is D. foenicu-
laceus. The type locality is "Mona Island" (Isle of Man) (cf.
Dillenius, 1741, p. 16).

Key to the Species

1. Fronds relatively small and delicate, with opposite branching. . 1. D. tenuis (p. 588)

1. Fronds more robust, chiefly with alternate branching 2

2. Fronds with slender, attenuated branches 3

2. Fronds with fewer, shorter, thicker, blunt branches

4. D. hippuroides (p. 589)

3. Hispid with slender, short ultimate branchlets 2. D. hispidus (p. 588)

3. Not as above 3. D. foeniculaceus (p. 589)

1. Dictyosiphon tenuis S. and G.

Fronds very fragile and flaccid throughout, 5-10 cm. long, with
main axis percurrent and 225-250/x diam. ; branches opposite, of 3-4
orders, flagellif orm ; ultimate ramuli blunt, 40-50ju, diam. ; sporangia
scattered, completely submerged in the cortical tissue.

Golofin Bay, Alaska.

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 10. Dictyosiphon
foeniculaceus f. americanus Setchell and Gardner, Alg. N.W. Amer.,
1903, p. 248 (not of Collins).

We have but a few specimens of this species upon which to base
the diagnosis. They were collected by Mr. E. C. McGregor in 1900.
The species is based principally upon the delicate character of the
fronds, the main axis being only about a quarter of a millimeter in
diameter, and upon the prevailing opposite branching, alternate
branching prevailing in the other species. Our plant is much more
slender than the plants referred by Collins to his f. americanus of
D. foeniculaceus which also lacks the regularly opposite branching. .

2. Dictyosiphon hispidus Kjellm.

Fronds delicate and flaccid, freely branched on all sides into several
relatively long, primary, flagelliform branches, these in turn irregu-
larly branched, all clothed with numerous, short, subulate or cylin-
drical, ultimate ramuli, tubular below, solid above ; olive brown in
color, darker on drying.

Growing on Chordaria in the lower littoral belt. Orca, Alaska.

Kjellman. Algenveg. Murm. Meer., 1877, p. 47, Spets. Thall. II,
1877a., p. 39, pi. 2, fig. 1. Dictyosiphon foeniculaceus Setchell and
Gardner, Alg. N.W. Amer., 1903, p. 248 (in part).

1925] Setchell-Gardner : Melanophyeeae 589

A single collection from Orca in the Prince William Sound region
of Alaska is all we have of this species and this was earlier (1903,
p. 248) referred by us to D. f&eniculaceus. The abundance of short
branchlets over the plant, however, give it the hispid appearance of
Kjellman's species. Certain of the specimens we refer to D. foeni-
cnlaceus approach it, but do not seem by any means, to equal it in
hispidity.

3. Dictyosiphon foeniculaceus (Huds.) Grev.

Plate 40, figs. 47-49

Fronds solitary or occasionally several arising from a confluent
disk-shaped holdfast, terete, repeatedly much branched, beginning
close to the base, 15 cm. to 4 dm. high; branches either alternate or
occasionally opposite, the different orders much reduced in size, the
ultimate being very fine, short and subulate, all gradually attenuated ;
when 3'oung the ultimate branches covered with a dense growth of fine
hairs; superficial cells small, rounded or angular, irregularly disposed
or at times in long, longitudinal rows ; zoosporangia spherical or
ellipsoidal, distributed at random over the surface of the frond.

Growing on stones and on other plants, in the middle and lower
littoral belts. From Bering Sea, Alaska, to Puget Sound, Washington.

Greville, Alg. Brit., 1830, p. 56, pi. 8, figs. 1-4; Saunders, Alg.
Ilarriman Exp., 1901, p. 422 ; Setchell and Gardner, Alg. N.W. Amer.,
1903, p. 248 (in part) ; Harvey, Phyc. Brit., pi. 326. Conferva
foeniculacea Hudson, Fl. Angl., 1762, p. 479.

This species has more slender as well as coarser forms. It is larger
and more robust than Dictyosiphon tenuis and lacks the small slender
ultimate branchlets arranged along the branches which give character
to D. hispidus. All three of these species differ from D. hippwrokles
in having more tapering branches which decrease in size with each
order of branching.

4. Dictyosiphon hippuroides (Lyngb.) Kuetz.

Frond solid, or occasionally slightly fistulose below, filiform, rigid,
moderately branched, 15-60 cm. high, up to 1.25 mm. diam. ; branches
of different orders very similar, flagelliform, tapering but slightly ;
surface cells in the lower part of the frond arranged in horizontal
series, but irregular above ; color dark brown.

Growing on rocks in the lower littoral belt. Apparently rare in
our territory. Reported from St. Paul Island and from near Iliuliuk,
Unalaska, Alaska.

590 University of California Publications in Botany [Vol. 8

Kuetzing-, Tab. Phyc, vol. 6, 1856, p. 19, pi. 52, II ; Setchell, Alg.
Prib. Is., 1899, p. 591 ; Setchell and Gardner, Alg. N.W. Amer., 1903,
p. 248. Dictyosiphon Chordaria f. gelatinosa Setchell and Gardner,
Alg, N.W. Amer., 1903, p. 248 (not Stroemfelt). Scytosiplion hip-
piiroides Lyngbye, Hydr. Dan., 1819, p. 63, pi. 14 B.

D. hippuroides resembles D. foenicid.aceiis very closely, and is often
confused with it. It is a coarser plant generally, is less profusely
branched, each order of branches approximating the same size, and is
less tapering. Collins (Rhodora, 1900, p. 164) states that this species
frequently reaches a meter in length on the Atlantic coast. The mate-
rial which we have referred to this species is scanty and of small size.
Some of our plants may possibly be referred to the var. fragilis
(Harv.) Kjellman.

Order 6. LAMINARIALES oltmanns

Fronds of large size, solid or hollow, simple or branched, cylindrical
to flattened, usually with three distinct regions, viz., (1) a holdfast
varying from discoid to clusters of simple or branched hapteres, (2)
a stipe cylindrical or more or less flattened and simple to dicho-
tomously or irregularly branched, and (3) a flattened blade or blades,
of usually three sets of tissues, inner of colorless, elongated, usually
intertwined, hyphal cells, some with enlarged ends ("trumpet hy-
phae"), intermediate of somewhat vertically elongated or nearly
isodiametric cells with scattered chloroplasts (phaeoplasts) passing
over into an outer layer of small cells usually deeply colored ; mucilage
glands, or canals, present in some species; zoosporangia on macroscopic
sporophylls, borne in extended sori, accompanied by closely packed
unicellular paraphyses having, usually, terminal hyaline appendages;
growth in length from a meristematic tissue intercalated between blade
and stipe, or, in Chordaceae, just above the holdfast ; gametophyte
microscopic, usually very much reduced, confervoid, bearing uni-
locular gametangia ; sperms motile, biciliate ; eggs non motile, fertilized
and germinating in position.

Oltmanns, Morph, und Biol, der Algen (ed. 2), vol. 2, 1922, p. 121.

The discoveries by Sauvageau (1915) and by Kylin (1916, 1918)
of the gametophytes of several members of this group inaugurated a
change in the whole attitude toward the classification of the Brown
Algae and led to interpretations of certain peculiarities of the occur-
rence of the reproductive bodies in certain groups. Following these

I9i'>| Setchell-Gardner : MelanopJiyceae 591

indications to their seeminprly logical outcome has led us to arrange the
Brown Algae as we have, trusting to be justified, at least in large
measure, when our knowledge of development within the group shall
have been made more exact. The order of the Laminariales, however,
is clearly delimited by the intercalary region of growth in the complex
plants belonging to it associated with the intermingling of the uni-
locular zoosporangia with unicellular paraphyses in extensive elevated
sori. The question as to the division of the order into families is not
so clear as that of its cleavage from other orders. Two families have
been recognized and there is great temptation to increase the number.
There certainly exists a certain tendency to group genera into some
sort of larger divisions, but the difficulty lies in finding sharp lines of
cleavage. The best distinctions lie according to the differentiations
of the frond arising at the transition place between the blade and the
frond and the relation to this and the fertile areas. The alternatives
seem to be these : either to retain all the genera under one large and
varied family (possibly excepting the Chordaceae) divided into num-
erous tribes, or to divide them between numerous families of unequal
value and of often indistinct cleavage. We have separated the Lami-
nariaceae into five families, one of which Haligeniaceae (including
Haligenm and Saccorhiza) is not represented in our territory.

Key to the Families

1. Stipe absent or not differentiated, paraphyses without hyaline appendages

19. Chordaceae (p. 591)

1 . Stipe distinct, at least when young, paraphyses with hyaline appendages 2

2. Transition place neither giving rise to splitting nor to outgrowths

20. Laminariaceae (p. 593)

2. Transition place not as above 3

3. Splitting arising at transition place or within its influence

21. Lessoniaceae (p. 621)

3. Outgrowths arising at transition place or within its influence

22. Alariaceae (p. 633)

FAMILY 19. CHORDACEAE dumort.

Fronds cylindrical, hollow, interrupted by diaphragms, with a
discoid holdfast, growth in length basal (situated just above the hold-
fast), innermost cells elongated, supporting hyphal filaments united
to form the diaphragms ; outermost layer a single row of slightly elon-
gated cells with colorless hairs and with or without exserted colored
filaments ; sori extended, paraphyses destitute of colorless appendages ;
gametophyte microscopic as in the order.

592 University of California Publications in Botany [Vol. 8

Dumortier, Comm. Bot., 1822, p. 72 (Me Pfeiffer) ; Kylin, Stud,
iiber Algenfl. der Schwed. Westkiiste, 1907, p. 97.

The single genus Chorda', clearly a member of the Laminariales,
but differing from all the other genera of the order in its hollow
cylindrical frond, is taken to represent a distinct family.

40. Chorda Stackh.

Fronds arising from a small disk-shaped holdfast, unbranched,
cylindrical, solid below, hollow above, the cavity separated by cross-
partitions into many chambers, color olive brown, growth intercalary
near the base ; median tissue of the hollow portion composed of longi-
tudinally elongated cells, hexagonal in cross-section, firmly united,
lined with delicate, loose filaments which unite to form the cross-parti-
tions at intervals; the outside of the median tissue is clothed with
unicellular paraphyses, hairs and zoosporangia ; reproduction asexual
by unilocular zoosporangia.

Stackhouse, Physiol. Observ., m Ner. Brit., 1797, p. xvi.

The type of the genus is the Fuous Filum of Linnaeus (Sp. PL,
1753, p. 1162). The type locality is unknown.

Chorda Filum (L.) Lamour.

Fronds gregarious, several at times arising from the same confluent
base, cylindrical, sometimes spirally twisted, slightly tapering at the
base and at the apex, from 3 dm. to 4 m. long, 2-5 mm. diam.,
lubricous, when young clothed with a dense growth of hyaline or
yellowish colored hairs ; paraphyses unicellular, densely crowded,
clavate, slightly longer than the zoosporangia and nearly obscuring
them above ; unilocular zoosporangia oblong to ellipsoidal, 30-50;u,
long, 10-15/i, wide; plurilocular gametangia 35-40/i- wide.

Growing on stones in the upper sublittoral belt. From Port
Clarence, Alaska, to the Straits of Juan de Fuca.

Lamouroux, Essai, 1813, p. 26 ; Saunders, Alg. Harriman Exp.,
1901, p. 424; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 254;
Reinke, Atlas, 1889, p. 35, pis. 26, 27, 28. Fiiciis Filum Linnaeus,
Sp. Plant., 1753, p. 1162.

There is little to be said under Chorda, except to note that it is a
plant of the colder waters and of which we have seen specimens from
our coast collected at Port Clarence, Safety Harbor, near Cape Nome,

19-'5] SetcheU-Gardner: MeJanoi^hyceae 593

St. Michaels, and Golofin Bay on the Alaskan coast. Tt is credited by
Postels and Rnprecht (1840, p. 19) to Sitka and Unalaska, by Saunders
(1901, p. 424) to Popof Island and Prince William Sound, and by
MacMillan (1902, p. 219) to the Straits of Juan de Fuca. The last
locality is not verified by any specimens, so far as we know. Such
specimens as we have seen do not differ from those of the North
Atlantic.

FAMILY 20. LAMINAKIACEAE reichenb.

Fronds simple, composed of holdfast, stipe and blade, at least in
the earlier stages, never with true dichotomous branching of the stipe,
adventitious or false branching arising in a few cases from the blade,
composed of the usual three tissues; mucilage glands and passages
present or absent ; blades destitute of cryptostomata (or bunches of
hairs) ; sori extended, always on the blade, paraphyses with hyaline
appendages.

Reichenbach, Conspect. Reg. Veg., 1828, p. 29 (fide Pfeiffer).

We have adopted the family of the Laminariaceae in restricted
sense as including those members of the Laminariales having the prin-
cipal meristematic region situated at the summit of a more or less well
developed stipe, at the base of a flattened blade, and initiating neither
splitting nor outgrowths, of or from, either the stipe or the blade. In
the simplest genera (e.g., Laminaria) the transition place is perfectly
plane, but in some genera folds (e.g., Pleurophycus and Cymathaere),
in others ribs (e.g., Costaria and Agarum) are developed, while in
some genera (Costaria, Agarum, and TludassiophyUum) bullosities and
resulting perforations are the final results of its activity. In certain
species the center of the blade wears away to the transition place which
extends and growing on, produces two false branches each with half a
blade attached (Hedophyllum subsessile and Thalassiophyllum). It
has seemed best to group the various genera into tribes according to
these various methods of behavior.

Key to the Tribes

1. Blades without perforations, longitudinal ribs or folds, occasionally bullose.... 2

1. Blades with perforations, longitudinal ribs or folds 3

2. Stipe persistent 1. Laminarieae fp. 594)

2. Stipe early disappearing 4. Hedophylleae (p. 616)

.3. Blades with longitudinal folds 2. Cymathaereae (p. 606)

3. Blades with longitudinal ribs or perforations 3. Agareae (p. 609)

594 University of California Fuhlications in Botany [Vol.8

TRIBE 1. LAMINARIEAE bory (lim. mut.)

Fronds with well developed stipes and flattened blades; hapteres
sometimes elongated and producing new fronds ; blades plane, undi-
vided or splitting from above downwards, at times bullose, but without
perforations, longitudinal ribs or folds.

Bory de Saint Vincent, in Diet. Class., vol. 1, 1822, and vol. 9, 1826,
p. 187 (as family) ; De-Toni, Syll. Alg., vol. 3, 1895, p. 317 (in part) ;
Setchell, Kelps of the U. S. and Alaska, 1912ff, p. 146.

The tribe of the Laminarieae contains the single large genus,
Laminaria. It may be looked upon as containing the simplest, or,
certainly, the least modified of the members of the family and as the
type whence divergences have arisen or taken their departure.

41. Laminaria Lamour. (in part)

Fronds differentiated into three distinct parts, a basal holdfast,
composed either of a solid disk of more or less branched hapteres, a
stipe of greater or less length, and a blade ; both the stipe and the
blade may or may not have mucilage ducts; blade simple or more or
less deeply lacerated into few to many segments, plane or with intra-
marginal bullae and marginal ruffles ; growth intercalary at the base of
the blade; reproduction asexual, by zoospores borne in unilocular
zoosporangia among unicellular paraphyses in extensive sori nearly
covering both surfaces of the blade of the macroscopic plant, and
sexual, oogamous, on microscopic plants.

Lamouroux, Essai, 1813, p. 20.

The Laminaria of Lamouroux {loc. cit.) included plants now refer-
red to several genera and well distributed through the order Lamin-
ariales. The first three species, however, are of the genus Laminaria
as gradually restricted by later authors. The Lamouroux name is
antedated by the Laminarius of Roussel (1806), by the Saccharina
of Stackhouse (1809) and the PJiycodendron of Olafsen and Povelsen
(1772). Laminaria has, however, received the sanction of the Inter-
national Botanical Congress of Brussels (1910, see Briquet, 1912,
p. 76). While most of the species of Laminaria are readily separated
into those whose blade is digitate (i.e., split deeply into segments) and
those whose blade is entire, there are a few species seemingly present-
ing both conditions, e.g., L. cuneifolia of our coasts.

1925] SetcheU-Gardner: Melanophyceae 595

Key to the Species

1. Blade entire or slightly split at the outer end 2

1. Blade split more or less deeply into several narrow segments 8

2. Fronds, except when young, arising from creeping "rhizomes" 3

2. Fronds without creeping "rhizomes'' 4

3. Mucilage ducts absent from stipe 3. L. longipes (p. 597)

3. Mucilage ducts present in both stipe and blade 4. L. Sinclairii (p. 598)

4. Holdfast a solid parenchymatous disk 6. L. personata (p. 599)

4. Holdfast of branched hapteres 5

5. Blade densely bullate all over 5. L. Farlowii (p. 599)

5. Bullae in two rows within the margin or scattered 6

6. Stipe long, complanate, except at base, very dark brown

2. L. complanata (p. 596)

6. Stipe short, terete, color yellowish brown 7

7. Mucilage ducts in the blade only 1. L. saccharina (p. 595)

7. Mucilage ducts in both blade and stipe 7. L. cuneifolia (p. 600)

8. Holdfast a sohd parenchymatous disk 8. L. ephemera (p. 603)

8. Holdfast composed of stout branched hapteres 9

9. Fronds plane, stipe long, up to 1 m 10

9. Fronds more or less bullate, stipe relativelj- short 7. L. cuneifolia (p. 600)

10. Stipe cylindrical to near apex 11

10. Stipe compres.sed from just above holdfast 11. L. platymeris (p. 605)

11. Mucilage ducts of stipe in a circle just beneath the cortex, stipe slender

9. L. dentigera (p. 604)

11. Mucilage ducts of stipe deep seated, stipe rigid 10. L. Andersonii (p. 605)

1. Laminaria saccharina (L.) Lamour.

Holdfast of numerous, branching, more or less rigid hapteres often
extending for 5-8 cm. up the stipe which is very variable in length,
from 5-50 cm., cylindrical, flattening above into the blade, 6-9 mm.
diam., without mucilage ducts; blade undivided, usually coriaceous or
membranaceous when growing in quiet w^aters, plane, undulate, or
with two distinct rows of large bullae along the margin, ovate to ovate-
lanceolate, often with cuneate base, with mucilage ducts ; color rich
brown.

Growing on rocks in the upper sublittoral belt. Reported from
various localities from the Alaskan peninsula to Coos Bay, central
coast of Oregon.

Lamouroux, Essai, 1813, p. 22; Setchell and Gardner, Alg. N.W.
Amer., 1903, p. 261 ; Setchell, Kelps of the U. S. and Alaska, 1912rt,
p. 149. Fucus saccharimis Linnaeus, Sp. PL, 1753, p. 1161.

All attempts to separate this widespread species of the cooler waters
of the northern hemisphere have been unsatisfactory. The type has
been assumed to be the plant with mucilage ducts in the blade, but
lacking them in the stipe. This seems to be a condition with all the

596 University of California Puhlications in Botany [Vol. 8

plants we have referred here. We have not met with any forms
referrable to L. AgardliU, i.e., entirely destitute of mucilage duets.

On our coast at least two forms of this species may easily be
distinguished :

Laminaria saccharina f . linearis J. Ag.

Holdfast composed of an abundance of relatively small branched
hapteres extending along the stipe up to 8 cm. above the base, stipe
slender and short, blade with cuneate base, 12-18 cm. broad, 2.5-3.5 m.,
or more, long, with two distinct rows of bullae extending the entire
length of the blade just within the margin.

Growing on rocks in localities subjected to the action of the water,
in the upper sublittoral belt. From Unga, Alaska, to Puget Sound,
Washington.

J. Agardh, De Lamin., 1867, p. 12 ; Saunders, Alg. Harriman Exp.,
1901, p. 429, ''forma b."; Setchell and Gardner, Alg. N.W. Amer.,
1903, p. 261 ; Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc),
no. LXXXVIII.

Laminaria saccharina f. membranacea J. Ag.

Hapteres strong, stipe longer than in the preceding, frond ample,
membranaceous, often of delicate consistency, not holding together
when lifted from the water, more or less undulate or bullate ; large
fruiting specimens at Sitka, Alaska, are stiff and rigid, not readily
torn.

Growing on rocks, logs, floats, etc., in the upper sublittoral belt.
From Alaska southward to Coos Bay, Oregon.

J. Agardh, De Lamin., 1867, p. 13; Setchell and Gardner, Alg.
N.W. Amer., 1903, p. 261 ; Saunders, Alg. Harriman Exp., 1901, p. 429,
"forma a." Laminaria saccharina Tilden, Amer. Alg. (Exsicc),
no. 240.

2. Laminara complanata (S. and G.) Setchell

Plate 57
Holdfast of few, rigid, slightly branched hapteres ; stipe 3-5 dm.
long, terete below, much flattened above, varying from 8-12 mm. diam.
below, to 20-25 mm. wide and 2-4 mm. thick above, Mdthout mucilage
duets; blade 80-100 cm. long, 40-50 cm. wide just above the base, in
many specimens short and nearly circular in outline, amply ruffled,

1925] Setchell-Gardner: Melanophyceae 597

with base decidedly cordate and with mucila<?e duets large and
abundant, just under the surface layer of cells; color dark.

Growing on piles and on rocks in the upper sublittoral and lower
littoral belts. Observed in but a single locality, Friday Harbor, San
Juan Island, Washington.

Setchell, Kelps of U. S, and Alaska, 1912fl, p. 149. Laminaria
saccharina f. complanata Setchell and Gardner, Alg. N.W. Amer.,
1903, p. 262 ; Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc),
no. LXXXVII.

Laminaria complanata, known as yet only from the type locality,
seems very distinct from other members of the sac char ina-^TO\\Y>, on
account of its rather long and flattened stipe. It does not seem, so
far as the material available is concerned, to merge into true L.
saccharina found in the same locality, even growing on the same piles
with it.

3. Laminaria longipes Bory

Holdfast at first composed of a few hapteres arising from the
basal portion of the stipe, these in turn, becoming rhizome-like and
giving off lateral secondary hapteres and lateral erect fronds ; stipes
terete, smooth below, compressed at the summit, 8-15 cm. long, 4-6
mm. thick, without mucilage ducts ; blade plane, undivided, acuminate
or rounded at the apex, 10-20 cm, long, 2-A: cm. wide, with large
mucilage ducts within the cortex.

Growing on rocks in the Bering Sea (Agattu, Kyska and St. Paul
Islands).

Bory, in Diet. Class., vol. 9, 1826, p. 189 ; Kjellman, Om Beringhaf.
Algfl., 1889, p. 43 ; Setchell, Alg. Prib. Isl., 1899, pp. 591, 592, pi. 95,
Kelps of the U. S. and Alaska, 1912a, p. 150; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 260. Lessonia repens Ruprecht, Tange Och.,
1851, pp. 232, 350. Laminaria Ruprechtiana Le Jolis, Examen, p. 71,
or 590. (For explanation of the dates of this publication see Setchell,
1891, p. 214.) Anthrothamnus? l&ngipes J. Agardh, De Lamin., 1867,
p. 26.

We have followed J. G. Agardh in assigning the northern plant
with distinct rhizome-like basal portion to the Laminaria Jongipes
Bory, although we have not seen the type specimen. It closely
resembles the more southern L. Sincladrii, but lacks the mucilage
glands of the stipe so well developed in the latter species.

598 University of California Publications in Botany [Vol. 8

4. Laminaria Sinclairii (Harv.) Farlow

Holdfast at first composed of small, branching- hapteres, soon send-
ing out numerous extensively creeping- rhizomes from which fronds
subsequently arise ; stipe "^dth small, deep-seated mucilage ducts, cylin-
drical, 2-3 mm. diam., 2-3 dm. long; blade linear, undivided or
slightly lacerated at the free end, plane, glossy, 1.5-3 cm. wide, 4—7 dm.
long, with mucilage ducts ; color very dark brown, black on drying.

Growing on rocks in the lower littoral and upper sublittoral belts.
Abundant from the southern end of Vancouver Island to Pecho, San
Luis Obispo County, California.

Farlow, in Farlow, Anderson and Eaton, Alg. Exsicc. Amer.-Bor.,
no. 118 (nomen nudum) ; Setchell, Notes on Kelps, 1896, p. 44; Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. VII. Lessonia
Sinclavrii Harvey, in Hooker, Flora Antarc, vol. 2, 1846, p. 460.
Laminaria saacharina Harvey, in Hooker and Arnott, Bot. Beechey,
1841, p. 407. Hafgygia Sinclairii Areschoug, Obs. Phyc, IV, 1883,
p. 6.

The combination Laminaria Sinclairii was first used by Farlow
as a nomen nudum, in connection with the plants collected at Santa
Cruz, California, by Dr. C. L. Anderson and distributed in Farlow,
Anderson and Eaton's Algae Exsiccatae Americae-Borealis, fascicle
3, no. 118, in 1878. The specific name was first used by Harvey in
Hooker's Flora Antarctica {lo'C. cit.) in honor of Dr. Sinclair, who
first collected the species and sent it to Sir W. J. Hooker in whose
herbarium Harvey first saw it. Harvey states on page 87, part 1, of
the Nereis Boreali-Americana "The Lessonia Sinclairii, from Cali-
fornia, mentioned by Dr. Hooker, Fl. Antarct., vol. 2, p. 460 must
for the present remain undescribed. ... is the Laminaria saccharina
of Harvey in Hook, and Arn. Bot. Beechey, p. 407." It M-as listed
by Anderson (1891, p. 220) as ^'Laminaria Sinclairii Farlow and
Eaton" and De-Toni (1895, p. 343) adopted this combination. Ares-
choug (1883) gave the first diagnosis of the species.

Laminaria Sinclairii may readily be distinguished from all other
species on our coast, except L. longipes Bory, by its relatively long
and narrow blade and slender stipe, and by its creeping rhizomes. It
is to be distinguished from L. longipes of the Bering and Ochotsk
seas, which it most closely resembles, by the absence of mucilage ducts
in the stipe of that species.

1925] Setchell-Gardner: Melanophyceae 599

5. Laminaria Farlowii Setchell
Plate 56o

Holdfast of strong, compact, branching liapteres; stipe short,
terete, flattening suddenly into the blade, 4-7 cm. long, 4-6 mm, diam.,
without mucilage ducts; blade thick, coriaceous, abundantly wrinkled
or pitted with rather deep depressions, more or less in longitudinal
rows over the whole blade on both sides with scanty large mucilage
ducts ; color very dark brown.

Growing in the lower littoral and very upper sublittoral belts.
Central California to southern California.

Setchell, in Anderson, List of Calif. Mar. Alg., 1891, p. 220 (nomen
nudum), Trans. Conn. Acad., 1893, vol. 9, p. 355 (description), Regen-
eration among kelps, 1905, pi. 16, fig. 17 ; Collins, Holden and Setchell,
Phyc. Bor.-Amer. (Exsicc), no. XXXI.

Laminaria Farlowii is not a very abundant or commonly collected
species, although in its general region, it is scarce rather than rare.
The peculiar corrugated appearance of the blade is difficult to describe,
but is a constant and a striking characteristic.

6. Laminaria personata S. and G.

Plate 61

Fronds attached by an ample parenchymatous discoid base, yellow-
ish brown in color, 4-6 dm. high ; blade plane, broadly cmieate beloM%
12-20 cm. wide, mucilage ducts in the outer cortex, relatively sparse ;
stipe 8-12 cm. long, 4—5 mm. diam., terete below, flattening into the
blade, without mucilage ducts.

Growing on rocks in the upper sublittoral belt. Alaska [Yakutat
Bay, Kukak Bay and Popof Island (Saunders), Sitka (Gardner)].

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 10. Laminaria
solidungula Saunders, Alg. Harriman Exp., 1901, p. 429 ? ; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 260 (?) ; Setchell, Kelps of
U. S. and Alaska, 1912a, p. 150 ( ?) (none of these of J. G. Agardh).

This species of Laminaria is the only one we have seen from
Alaskan waters which has a disk-shaped holdfast, and in this respect
it resembles L. ephemera Setchell, a species extending southward
from Puget Sound to central California (Carmel Bay). Only a few
specimens were observed at Sitka, and Saunders {Joe. cit., p. 429)
remarks on the occurrence of his L. salidungula: "Occasional in the

600 University of California Puhlications in Botany [Vol. 8

sublittoral zone" and he also remarks (loc. cit.) that in his plants
"the stipe is thick and abundantly supplied with mucilage canals as
are the broad flattened rhizoids. " The statements, "thick stipe,"
"broad flattened rhizoids," and "mucilage canals" are indications
that he must have located and described some other plant. We have
been unable to locate a complete specimen of Saunder's plant. The
plants from Sitka, upon which we founded L. personata, have no
mucilage ducts in the stipe, another indication that they are different
from those collected by Saunders and are, therefore, different from
Agardh's L. solidungida.

We have searched carefully for L. solidtmgula in our territory
since J. G. Agardh calls attention to Ruprecht's statement (1851, p.
351) that he found a young abnormal specimen of L. latifolia Ag. (a
form of L. saccliarina) with a scutate holdfast among his Ochotsk
specimens. Kjellman does not mention it among his Bering Sea plants,
but Saunders {loc. cit.) credits it to Yakutat Bay, Kukak Bay, and
Popof Island, Alaska. Possibly both Ruprecht 's and Saunders ' plants
may prove to be the same as our Sitkan specimen. Our Sitkan plant
probably belongs to the yS'a.cc/iar ma-group, although the blade is some-
times broad and deeply split (as in the case of the older plants in our
illustration, plate 61).

7. Laminaria cuneifolia J. Ag.

Plate 59a, h, and plate 60

Holdfast of a few, stout, branched hapteres ; stipe usually short and
flexuose, terete at the base, flattening above into the blade, 6-10 cm.
long, 3-4 mm. diam., with moderately large mucilage ducts in a circle
near the surface ; blade entire or with a few lacerations at apex, usually
cuneate at the base, but at times even cordate, becoming linear, very
variable in size, about 6-9 dm. long, 7-12 cm. wide, coriaceous, with a
row of prominent, transverse bullae within each margin, in some only
at the base, in others extending to the apex, with abundant large
mucilage ducts ; color of the whole plant a very dark brown or nearly
black.

Growing in a narrow belt along low-tide line. St. Lawrence Island,
Alaska, to Puget Sound, Washington.

J. G. Agardh, De Lamin., 1867, p. 10. Laminaria hullata Kjellman,
Om Beringh. Algfl., 1889, p. 46, pi. 2, figs. 5-9 ; Saunders, Alg. Harri-

1925] Setchell-Gardner: Melanophyceae 601

man Exp., 1901, p. 428 ; Setchell and Gardner, Alg. N.W. Amer., 1901,
p. 257; Setchell, Kelps of IT. S. and Alaska, 1912o, p. 151.

There are two series of Laminarias in our northern waters present
in great variety of form and seemingly overlapping forms which are
puzzling as to their proper taxonomic treatment. The one series has
passed under the names of forms of Laminaria bullata Kjellman and
the other under those of forms of L. Bongardimia P. and R. The
L. hullata type (cf. Kjellman, 1889, pi. 2, fig. 5) has a simple individual
blade and a series of pronounced bullae within each border. From this
simple type, forms with a great variety of lengths of stipe, breadth of
blade, more pronounced or less pronounced bullae (even almost, or
quite absent) are found. The compression of the stipe also varies
greatly in degree. The wider blades become broadly oblong to even
cordate at the base and split deeply into a few broad segments. In
several of its states it is to be distinguished from very similar forms
of the plants, usually grouped under L. Bongardiana, by its some-
■\vhat less flattened stipe and the presence, or often only indication, of
bullae. Both are species possibly better reckoned in the digitate
section, yet L. Bongardiana (see below under L. platynieris) is more
properly digitate while L. cuneifolm is perhaps more typically of the
saccharina-gvo\\\). We hesitate somewhat in referring the L. hullata
Kjellman series of forms to L. cuneifolia J, Ag., but J. G. Agardh cer-
tainly included the simpler forms of Kjellman 's species under his.
Agardh makes no statement as to type, but probably regards the
Ochotsk 8ea plants from that point of view. The American plants
placed by Agardh under his L. cuneifolia, viz., one from Esquimault,
is the same as the L. hullata f. simplex S. and G. We can see no way
of distinguishing between the two species from the descriptions and
consequently unite them, placing the forms we have described under
L. cuneifolia. The L. cuneifolia of Greenland, however, seems to be a
different species and is closely related to L. groenlandica Rosenvinge.

We are uncertain as to the nature of the Alaskan plant referred
by Saunders (1901, p. 429) to L. cuneifolia; since he says that the
blade is "thin, papyraceous, light olive green and very brittle in
drying, quite regularly wavy on the margin" and have not included
it in our synonymy.

602 University of California Puhlications in Botany [Vol. 8

Laminaria cuneifolia f. angusta S. and G. comb. nov.

Holdfast of distinct branched hapteres; stipe short, 1-2 cm. long,
terete or slightly flattened above, with mucilage ducts closely crowded
in a circle just beneath the periphery, elongated radially in section
view, surrounded by conspicuous secreting cells ; blade undivided,
15-45 cm. long, 3-5 cm. wide, cuneate at the base, usually more or less
falcate, with a well defined row of bullae within each margin ; mucilage
ducts in the blade large, about one-third of the way between the sur-
face and the distinctly marked off, wide medulla; color very dark
brown, turning black on drying.

Growing abundantly on rocks in the upper sublittoral belt.
Reported at Cape Flattery and along the w^est coast of Whidbey
Island, Washington, and doubtless at other localities in the lower
Puget Sound region and farther north.

Setchell and Gardner, Alg. N.W. Amer., 1903, p. 257; Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. LXXXV (both
siib L. hullata).

Laminaria cuneifolia f. subsimplex S. and G. comb. nov.

Hapteres well developed; stipe short and stout, 4-8 cm. long, 4-6
mm. diam., terete below, somewhat compressed above, with mucilage
ducts forming a circle just beneath the periphery; blade thick and
somewhat coriaceous, 50-150 cm. long, 10-15 cm. wide, plane in some
specimens, particularly the younger ones, with a distinct row of
bullae extending for some distance above the base within each margin,
entire or split from a half to a third of the distance to the base into
two, or at most a few, broad segments, with large mucilage ducts
situated about halfway between the surface and the distinctly marked
off medulla ; color very dark in old specimens.

On rocks in the upper sublittoral belt. Common in the lower Puget
Sound region.

Setchell and Gardner, Alg. N.W. Amer., 1903, p. 257 (sul) L.
hullata). Laminaria hidlata Collins, Holden and Setchell, Phyc. Bor.-
Amer. (Exsicc), no. XXIX.

l92o] Setchell-Gardner: Melanophyceac 603

Laminaria cuneifolia f. amplissima S. and G, comb, no v.

Stipe 2-4 cm. long:, soon much flattened into the blade, with
mucilage ducts in a discontinuous circle just Avithin the periphery ;
blade up to nearly 3 m. long-, 1 m. wide, but usually smaller, broadly
cuneate to cordate at the base, rounded above, split into few segments,
the rows of bullae large and fairly distinct, particularly at the base
when the plants are young, at maturity the whole blade strongly
buUate ; mucilage ducts conspicuous and plentiful, associated with
groups of secreting cells one-half to one-third the way between the
surface and the distinct medulla; color very dark brown, black when
dry.

Growing on piles, floats, and rocks in the upper sublittoral belt in
quiet water. Sitka, Alaska, to Puget Sound and Cape Flattery,
Washington.

Setchell and Gardner, Alg. N.W. Amer., 1903, p. 258; Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. LXXXIV (both
suh L. hiUlata).

8. Laminaria ephemera Setchell

Plate 58

Holdfast strictly discoid, firmly attached to rocks; stipe terete,
slender, 2-4 mm. diam., 6-10 cm. long, devoid of mucilage ducts ;
blade broadly rounded to narrowly cuneate at the base, linear, entire
or split more or less deeply into a few linear lobes or segments, 5-8
cm. wide, 3-4 dm. long, about 1 mm. thick, without mucilage ducts ;
sori arranged in several longitudinal bands ; color light brown ; annual ;
fruiting in May.

Growing in the upper sublittoral belt. Known from only four
localities. Cape Flattery and Tacoma, Washington, Port Renfrew on
the southwest coast of Vancouver Island, British Columbia, and
Carmel Bay, California.

Setchell, Notes on Algae I, 1901, p. 121, Kelps of U. S. and Alaska,
1912a, p. 150. Eenfrewia parvula Griggs, Postelsia, 1906, pp. 247-
274, pis. 16-19; Tilden, Amer. Alg. (Exsicc), no. 609.

This is one of the few seemingly strictly annual species of the genus
Laminario> and the only one on our coast. It shares this distinction
with comparatively few species either in the family or in the entire

604 University of California Publications in Botany [Vol.8

order. In the family Laminariaceae, the only suspected annuals of
our coast are Costaria costaia, Cymathaere triplicata. and Agarum
fimhriatum. The annual character and the discoid holdfast clearly
distinguish this species.

9. Laminaria dentigera Kjellm.

Holdfast composed of a dense mass of rigid hapteres ; stipe 30-35
cm. long, smooth, flexuose, thickest at the base, slightly attenuated
upward, somewhat compressed at the apex, mucilage ducts large,
densely crowded in a circle just beneath the surface ; blade sub-
lanceolate, simple cuneate at the base, thick, smooth, dark brown or
almost black, shining, dissected nearly to the base with narrow or
w^ide linear lobes with lacerated margins.

Growing in the sublittoral belt. Aleutian Islands to the Bering
Strait.

Kjellman, Om Beringh. Algtl., 1889, p. 45, pi. 2, figs. 10-14;
Setchell, Kelps of U. S. and Alaska, 1912a:, p. 151. Laminaria denti-
gera f. hrevipes Setchell and Gardner, Alg. N.W. Amer., 1903, p. 259,

Laminaria dentigera, while seemingly distinct in appearance from
any of the truly digitate species, is somewhat difficult to diagnose with
certainty. The rather stout, nearly cylindrical, or often more or less
compressed, stipe, the sharply cuneate base of the blade and the
usually numerous and narrow segments of the blade with lacerate
margins as if irregularly dentate, distinguish the typical plants. As
is frequent among the Laminariaceae, there are long-stiped and short-
stiped forms with other variations tending to obscure the limits of the
species. The form, previously described by us as var. hrevipes
(Setchell and Gardner, 1903, p. 259) is probably close to Kjellman 's
type while var. longipes S. and G. {loc. cit.) is in our present opinion,
to be separated on account of its decidedly flattened stipe. We now
refer the latter to L. platymeris de la Pyl. In spite of these, however,
Laminaria dentigera seems clearly distinct from the forms usually
reckoned under L. Bongardiana P. and R. (L. pla.tymeri^ De la Pyl.)
with the normal form of which, according to Kjellman, Areschoug
confused it. It is to be distinguished from L. Andersonii Farlow in
having the circle of mucilage glands of the stipe situated among the
outer tissues. The type of L. dentigera came from Bering Island, on
the Asiatic side of the Bering Sea.

1925] SetcheU-Gardner : Melanophyceae 605

10. Laminaria Andersonii Farlow

Fronds perennial ; holdfast spreading out at the very base of the
stipe, composed of numerous strong, overlapping branched hapteres ;
stipe large and very strong and elastic, terete, tapering upward and
flattening into the blade, 1-1.5 cm. diam., 5-8 dm. long, smooth and
glossy, with concentric layers and numerous deep-seated mucilage
ducts ; blade coriaceous, thick, smooth, divided often nearly to the
base into several narrow segments, 15-25 cm. wide, 5-8 dm. (up to
10 dm.) long; color rich dark brown.

Growing in the upper sublittoral belt. Sitka, Alaska, to central
California.

Farlow, in Anderson, List Calif. Mar. Alg., 1891, p. 220 ; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 255 ; Setchell, Regeneration
among kelps, 1905, p. 145, pi. 17, figs. 19-25, Kelps of IT. S. and Alaska,
1912a, p. 151 ; Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. IV. Hafgygia ''Andersoni" (Farlow) Areschoug,
Observ. Phyc, IV, 1883, p. 3.

This seems to be a perfectly distinct species, with typically digitate
blade, stout, cylindrical, erect and rigid stipe, the latter having deep-
seated mucilage ducts. Its color, when living, is light brown.

11. Laminaria platymeris De la Pyl.

Holdfast forming a more or less globose mass of strong, irregularly
branched hapteres ; stipe thick, long or short, terete at the base, begin-
ning to flatten a short distance above the base, often becoming widely
complanate at the apex ; blade plane, from cuneate to deeply cordate
at the base, free from transverse bullae, more or less lacerated into
segments of different lengths and widths, mucilage ducts in both stipe
and blade.

Ranging from Bering Sea to the Strait of Juan de Fuca.

De la Pylaie, Flore de Terre-Neuve, 1829, p. 52; Setchell, Kelps
of the U. S. and Alaska, 1912rt, p. 151. Laminaria pl^tytoha De la
Pylaie, Ann. Sci. Nat., vol. 4, 1825, p. 178, pi. 9, fig. I {nomen
nudum?). Laminaria Bongardiana< Postels and Ruprecht, Illus. Alg.,
1840, p. 10, pis. 13, 14 (in part); Kjellman, Om Beringh. Algfl.,
1889, pp. 43, 44 (excl. syn. Hafgygia Bongardiana Areschoug, et
varr.). Laminaria taeniata Postels and Ruprecht, Illus. Alg., 1840, p.

606 University of California Publications in Botany [Vol. 8

10, pi. 38, fig. /. Laminaria nigripes J. G. Agardh, Spetsb. Alg., Till.
1868, p. 29. Laminaria atrofulva J. G. Agardh, Groenl. Alg., 1872,
p. 16. Laminaria fissilis J. G. Agardh, De Lamin., 1867, p. 18.

After trying in vain to find definite lines of cleavage between the
Arctic species, L. nigripes J. Ag., and L. atrofulva and the North
Atlantic L. platymeris De la Pyl., all three digitate species, with
mucilage ducts in the flattened stipes, Setchell (1912a, p. 151) decided
to unite them. In a similar fashion he found it impossible to separate
satisfactorily Laminaria fissilis J. Ag. and L. Bongardiana P. and R.
of the North Pacific. He found also that the two groups of North
Atlantic (Arctic) and North Pacific had no definite characteristics to
segregate one from the other. He consequently, combined them all
under the oldest tenable name, viz., L. platynieris De la Pyl.

Each of the components of this general group shows form varia-
tions as to length of stipe, thickness of stipe, degree of flattening or
compression of stipe, and length, breadth, shape of base, and degree
of splitting of blade. The following sets of form names may illustrate
this variation : f . taeniata, f . ohlonga, f . elliptica, and f . reniformis as
to shape and base of blade; f. hrevipes and f. longipes as to varying
length of stipe ; f . complanata, f. compressa, and f. suhteres, as to com-
pression or flattening of the stipe. Most forms may be better designated
by combinations of the foregoing names than by any single one of them.

TRIBE 2. CYMATHAEREAE s. and g.

Members of the Laminariaceae (in sensu restrict) having the
blades provided with one or more longitudinal folds.

Key to the Genera

1. Fold single, broad, median 42. Pleurophycus (p. 606)

1. Folds several, narrow 43. Cymathaere (p. 608)

42. Pleurophycus Setchell and Saunders

Holdfast of numerous branching hapteres ; stipe simple ; blade long,
undivided, with a single median, longitudinal, rather broad, shallow
fold and with broad, at times ruffled, margins ; sori narrow on both
surfaces of the median fold ; mucilage duets absent from both bladder
and stipe.

Setchell and Saunders MS., in Setchell, Notes on Algae I, 1901,
p. 123, in Saunders, Alg. Harriman Exp., 1901, p. 427 (description).

1^25] Setchell-Gardner : Melanophyceae 607

Pleurophycus is a monotypic genus as at present known, and was
discovered in 1898 on the west coast of Whidbey Island, Washington,
where it grows in abundance in the upper sublittoral belt. A little
later (1899) Saunders found it cast ashore at Yakutat Bay, Alaska,
At Neah Bay, Washington, and on the west coast of Vancouver Island,
it grows in abundance. Being so large and having such distinct char-
acter, it is rather remarkable that it escaped the notice of collectors
for so long. Technically it is nearest in structure to Cymatliaere from
which it differs much in general appearance.

Pleurophycus Gardner! Setchell and Saunders

Plate 80a

Holdfast of numerous whorls of rigid, branched hapteres; stipe
30-50 cm. long, solid, in mature specimens terete at base, gradually
flattened above into the relatively thin, meristematic, transition region ;
blade of flabby, elastic consistency, not perforated, undivided, but
more or less eroded at the outer end, with single, median, longitudinal,
broad, shallow fold, with broad, undulate margins, and with delicate
wrinkles along either side of the fold, rounded or cuneate at the base,
margin entire; blade 6-9 dm. long, 12-20 cm. (up to 40 cm.) broad;
fold 2-4 cm. (up to 15 cm.) broad; color dark olive green.

Growing on rocks in the upper sublittoral belt. Yakutat Bay,
Alaska, to Coos Bay, Oregon.

Setchell and Saunders, MS., in Setchell,' Notes on Algae I, 1901,
p. 123, in Saunders, Alg. Harriman Exp., 1901, p. 427, pi. 52 (descrip-
tion) ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 264; Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. XC ; Tilden,
Amer. Alg. (Exsicc), no. 346.

So far as is known at present, this member of the Laminariaceae,
seemingly very closely related to Laminaria., fruits but once and then
dies, quite different in this respect from many species of Laminaria
which are perennial and regenerate the blade several times. Its
present known southern limit of distribution is the mouth of Coos
Bay, on the coast of Oregon. It possibly extends many miles farther
south, even to the coast of California.

60S University of California Puhlications in Botany [Vol. 8

43, Cymathaere J. Ag.

Holdfast discoid or with inconspicuous, rudimentary, simple liap-
teres ; stipe short, flattened, persistent ; blade long, narrow, wider at the
base and tapering upward, longitudinally grooved, or loosely folded,
folds deep at the base becoming nearly plane above ; mucilage ducts
abundant and large in both stipe and blade ; sori broad, on both sur-
faces, at the base of the blade, paraphyses unicellular with well
developed hyaline appendage (not as in Griggs, Ohio Naturalist, vol. 1,
1907, p. 92, as to absence of either mucilage ducts or hyaline tips to
the paraphyses).

J. G. Agardh, De Lamin., 1867, p. 29.

The genus Cymathaere resembles Pleurophycus in having longi-
tudinal grooves or folds. It is also peculiar in having a discoid hold-
fast, although this is not a primitive structure as Griggs (1907) sup-
poses. Griggs is, also, in error in denying mucilage ducts and hyaline
tips to the paraphyses on C. tripUcata, the only known species.

Cymathaere triplicata (Post, and Rupr.) J. Ag.

Holdfast discoid, 6-15 mm. broad ; stipe short and stout, 5-25 cm.
long, cylindrical below, flattened above ; blade undivided, narrowly
linear, tapering upward, with acuminate apex and rounded or cuneate,
rarely cordate base, 1.5-4 m. long, 8-18 cm. wide, thick and coriaceous
at the base, thinner above, possessing three characteristic longitudinal
folds extending throughout the length of the blade ; sori broad, occupy-
ing the basal portion only ; paraphyses linear with hyaline appendages ;
color yellowish brown.

Growing on rocks in the upper sublittoral belt. From Bering Sea
to Puget Sound.

J. G, Agardh, De Lamin,, 1867, p. 30 ; Saunders, Alg, Harriman
Exp., 1901, p. 430; Setchell and Gardner, Alg. N.W. Amer., 1903,
p. 264; Griggs, Cymathere, 1907, pp. 89-96, pi. 7; Collins, Holden and
Setchell, Phyc. Bor.-Amer. (Exsicc), no. XXXIII a, 1) ; Tilden, Amer.
Alg. (Exsicc), no. 343; Laminaria tripUcata Postels and Ruprecht,
Illus. Alg., 1840, p. 10, pi. 10.

The type and only species of Cymathaere resembles in appearance,
color, texture, etc., Costaria costaia more nearly than any other alga.
In pressed specimens, the longitudinal folds are often obscured or even

1925] SetcheU-Gardner : Melanophyceae 609

obliterated. Such specimens are parti}" responsible for the report of
Laminarkt dermatodea on our northwest coast. Griggs (1907) looks
upon this species as of a more simple type than can be adopted by other
students of the plant.

TRIBE 3. AGAREAE kuetz. (lim. mut.)

Members of the Laminariaceae having blades, either longitudinally
ribbed or perforated, or both.

Kuetzing, Phyc. Gen., 1843, p. 347; Setchell, Kelps of the U. S.
and Alaska, 1912a, p. 153.

Key to the Genera

1. Adult fronds simple; blades bilaterally symmetrical 2

1. Adult fronds branched; blades not bilaterally symmetrical

45. Thalassiophyllum (p. 613)

2. Rib single; perforations numerous and constant 46. Agarum (p. 614)

2. Ribs several; perforations frequent, but never constant

44. Costaria (p. 609)

44. Costaria Grev.

Holdfast of branched hapteres ; stipe variable in length usually
flattening above ; blade with longitudinal percurrent ribs, each pro-
jecting on one side only and alternating on the two surfaces, bullate
and not rarely perforate, mucilage ducts absent from both stipe and
blade ; sori broad, distributed over most of the blade.

Greville, Alg. Brit., 1830, p. xxxix.

The genus Costaria was founded by Greville {he. cit.) to receive
a single species, C. Turneri Grev., dedicated to Dawson Turner, who
had previously (1819, p. 72, pi. 226) described and figured it as Fucus
castatus. Greville also quotes as a synonym the Laminaria costata of
Agardh (1820, p. 109 and 1824, p. 269).

The members of this genus, as at present known, are confined to
the waters of the Pacific Ocean, and largely on the North American
coast. They are closely related to the species of Agarum, but differ
in number, breadth, and structure of ribs, as well as in constancy of
perforations.

610 University of California Publications in Botany [Vol. 8

Costaria costata (Turn.) Saunders

Plate 56b, and plate 79a

Holdfast of extensive, widespreading, dichotomously branched
hapteres; stipe very variable in length and thickness, from slender
forms 3 mm. diam. and 5 cm. long, up to forms 1.5 cm. diam. at the
base, and 65 cm. long, with numerous, fine, parallel, longitudinal
grooves, flattened at the top ; blade also variable in size and form,
from narrowly lanceolate with cuneate base, to ovate with marked
cordate base, 1-3 dm. wide, 5-25 dm. long, coarsely bullate, with five,
prominent, parallel, longitudinal ribs alternately projecting only on
one side of the blade, leaving a depression on the opposite side ; color
dark brown, soon changing to green on drying.

Growing on rocks, wood, and other large algae in the lower littoral
and upper sublittoral belts. Common along the whole Pacific coast
from the Bering Sea to San Pedro, California.

Saunders, m Bot. Gaz., 1895, vol. 20, p. 57 ; Setchell, Kelps of
U. S. and Alaska, 1912ai, p. 154. Costaria turneri Greville, Alg. Brit.,
1830, p. xxxix; Postels and Euprecht, Illus. Alg. 1840, p. 12, pi. 24;
Saunders, Alg. Harriman Exp., 1901, p. 431 ; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 265 ; Collins, Mar. Alg. Vancouver Island,
1913, p. 109 ; Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), nos. 631 and XXXV. Costaria Turneri var. pertusa Collins,
Holden and Setchell, loc. cit., no. XXXIV. Costaria Mertensii J. G.
Agardh, Sp. Alg., vol. 1, 1848, p. 140; Tilden, Amer. Alg. (Exsicc),
no. 238. Fucus costatus Turner, Fuci, 1819, pi. 226.

Much diversity of opinion has existed among phycologists as to
the number of distinct species of Costaria that should be recognized.
Fortunately but few species have been proposed. The earlier descrip-
tions of these, however, have been inadequate, probably on account of
insufficient data, to enable one to mark their limits with much degree
of certainty. Four species and one variety were recognized by De-Toni
(1895, p. 361) as follows: C. Turneri Grev. and its var. pertusa Harv.,
C. Mertensii J. Ag., C. quadrinervia Rupr. and C. reticulata? Saunders,
all of which have been reported within our territory. The last men-
tioned is now known to be a young stage of Dictyoneuron calif ornicum
Rupr. C. quadrinervia has not again been reported, so far as we know,
since its discovery, and we do not feel that the description sufficiently
distinguishes it to warrant retaining it as a distinct species.

1925] Setchell-Gardner: Melanophyceae 611

Our i)roblem lies in the distribution of the bulk of our plants
between '*C Twrnerb" and '^C. MertensH," one or both, and in the
proper combination to use by which to designate them. Castaria
Turneri was first proposed by Greville in 1830, when he established
the genus, in the following language: "Fucus membranacea, Integra,
linearis, multi-costata. Fructus ignotus. " No additional diagnosis
was given for the species. The diagnosis is wholly inadequate to dis-
tinguish the species, were it not for the fact that he cites Fucus costatus
Turner (Hist. Fuc, 1819, pi. 226), and Laminaria costata Agardh
(Sj'st., 1824, p. 269), who in turn cites Turner's plant as a synonym.
Turner proposed F. costatus in 1819, thus antedating Agardh 's L.
costata^ first used in 1820 (Sp., vol. 1, p. 109). Turner's description
and figure were based upon material collected "On the west coast of
South America" by Menzies, and is so complete as to leave no doubt
that his plant was a true species of Castaria as now recognized. As
diagnosed, his species was "a foot and a half or more long, nearly
linear, about two inches wide." He states that he had but a single
specimen and that was pressed very flat, so he relied largely upon the
description and figure of Menzies for his information. We are thus
left very much in doubt as to whether Menzies' knowledge was of a
normal medium-sized plant, or, as it seems to us more likely, of a young
or a dwarf specimen. We have, on the California coast, plants which
answer very well to the description of '^ F. cast aid}" except as to size.
They are wider and longer, three to six inches wide and three to six
feet long, although Saunders (1895, p. 57) reports fruiting plants
from Pacific Grove, California, which are two inches wide and two

f

feet long. We feel fairly certain luitil we can obtain further knowl-
edge of South American plants, that the citation of South America is
erroneous and that our long, narrow forms are of the type species.
Observing the Vienna Code, therefore, they should be listed as
Castaria costata (Turner) Saunders.

Costaria Mertensvi of J. G. Agardh (1848, p. 140) as diagnosed
differs in no essential particulars from the diagnosis of C. Turneri
Postels and Ruprecht (1840, p. 12, pi. 24) which Agardh quotes as a
synonym. Whether or not C. Turneri Post, and Rupr. is the same as
Fucus costatus Turner, which Postels and Ruprecht quote as a
synonym, C. Mertensii J. Ag. has no legitimate right to supplant C.
Turneri Post, and Rupr., and we are thus disregarding it in our
account.

612 University of California Publications in Botany [Vol. 8

We have made no attempt to correlate anatomical differences with
the gross morphological characters of the various forms of the genus
which we have studied. Such studies as we have made of a rather
extensive array of material from different habitats and localities,
have all been confined to the general shape and size of the plant as a
whole, and in this we have found a great range of variation. As to the
hapteres, Fucus costatus Turner is said to have numerous, slender,
unbranched hapteres. We have come across no forms except those
in the juvenile stage that have this character. Ours are all many
times dichotomously branched in unmutilated specimens. The thick-
ness of the hapteres seems to depend upon the habitat. Those growing
in quiet waters are usually longer and more slender than those growing
in exposed situations. Regarding the stipe, it is to be noted that there
are extreme variations. In Turner's description, the stipe is only an
inch long and ''of the size of a crow's quill." On two occasions,
seven years apart, one of us (Gardner) has studied colonies of living
plants growing in quiet water at Oak Bay, Victoria, British Columbia,
and at Roche Harbor, San Juan Island, Washington, whose rigid
stipes were up to six and a half decimeters long and ten centimeters
wide at the flattened top. In all cases, they are covered with finer
or coarser longitudinal grooves. Between these two extremes there are
innumerable gradations in length and thickness. The blades, as we
have studied them, range in width from about five to seven centimeters
in C. Turneri var. pertiisa (cf. Collins, Holden and Setchell, lac. dt.,
no. XXXIV) to forms mentioned above at Oak Bay and Roche Harbor,
with blades up to sixty-five centimeters wide and only one to two times
as long. Some of the forms of medium width, growing in the Puget
Sound region, are up to three meters long. The bullations are finer
or coarser, deeper or shallower, but not in any way correlated with
other characters. The bases are narrowly to broadly cuneate, passing
over into slightly cordate and finally auriculate-cordate. The con-
sistency of the blade seems to be correlated with the exposure. They
are more or less thick and rigid in the locations exposed to the heaviest
surf, and thinner and flabby in quiet water. With our present knowl-
edge, we do not feel able to recognize more than one exceedingly
variable species, and we are not able even to state the limits of forms.

1925] Setchell-Gardner: Mehinophyceae 613

45. Thalassiophyllum Post, and Riipr.

Holdfast of stout, diehotomously branched hapteres ; primitive
stipe short, not elongating but thickening, soon buried among the stout
hapteres ; blade soon eroded to the base and developing two lateral
scrolls which unroll as they develop from thickened margins and form
fan-shaped, spirally twisted, closely and regularly perforated, partial
blades ; numerous secondary stipes or branches, arising along the
thickened margins, bearing small blades ; no mucilage ducts in either
the stipe or blade ; sori in irregular dark brown areas ; plants perennial.

Postels and Ruprecht, Illus. Alg., 1840, p. 11.

There are two peculiarities of the genus ThalassiopliijUum, viz.,
the one-sided fan-shaped blades unrolling from a one-sided scroll,
borne on M'hat seems to be a branched, heavy, solid stipe. From the
development (cf. Setchell, 1905a, p. 123, pi. 13, figs. 6-13) it appears
that the early stages resemble those of Agarnm in being bilaterally
symmetrical, with basal margins inrolled. By the disintegration of
the central portion of the blade, the basal margins only are left with
their scrolls. One margin ceases to develop, as does the short primi-
tive stipe. The other basal margin thickens, elongates, and becomes
stipe-like, bearing its one-sided blade. The branches (false) arise
(cf. Rosenthal, 1890, Flora, vol. 73, pp. 140, 141, pi. 8, figs. 33, 34) by
adventitious outgrowths from the developing scroll-like base of the
blade. The branching of ThalassiopliyUum is a pseudo-branching and
of a pseudo stipe not to be confounded with that of the Lessoniaceae
or in Egregia of the Alariaceae.

Thalassiophyllum Clathrus (Gmel.) Post, and Rupr.

Plate 72-

Plants 12-15 dm. high, rigid ; holdfast and stipe as in the genus ;
blade thick and coriaceous, with the numerous, large elliptical per-
forations with roughened margins radially arranged ; color dark
brown to deep olive green, much darker toward the basal portions.

Growing on rocks in the sublittoral belt. Bering Sea to the Straits
of Juan de Fuca (the last mentioned locality fide MacMillan, 1902,
p. 219).

Postels and Ruprecht, Illus. Alg., 1840, p. 11, pis. 18-19; Setchell,
Alg. Prib. Isl, 1899, p. 592, Post-emb. Stages, 1905a, p. 123, pi. 13,

614 University of California Piihlications in Botany [Vol. 8

figs. 6-13, Kelps of U. S. and Alaska, 1912a, pp. 155, 156; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 266 ; Collins, Holden and
Setchell, Phyc. Bor.-Amer. (Exsice.), no. XXXVI. Fucus Clatlims
Gmelin, Hist. Fuc, 1768, p. 211, pi. 33.

The juvenile blade disintegrates in the center, back to the stipe,
after which two meristems are established at the base and along the
margin of the remaining two partial blades. For a time these continue
to develop together but sooner or later one takes the ascendancy and
becomes erect while the other usually degenerates sooner or later. As
the blade continues to grow, it rolls and thickens along the margin,
leaving behind a false stipe, which branches here and there, the
branches developing numerous small blades.

46. Agarum (Bory) Post, and Rupr.

Holdfast composed of small branched hapteres ; stipe compara-
tively small, cylindrical or flattened, at times fimbriated ; blade rela-
tively thin, with narrow to broad percurrent midrib and perforated
alae; sori broad.

Bory, in Diet, class., vol. 9, 1826, p. 193.

The genus Agarum was founded by Bory (Joe. cit.). He enum-
erated five species, as follows: A. quinquecostatum, A. cribrosum,
A. escidentum, A. Belisei, and A. Pylaii, all but one of which have
since been distributed between two other genera, the first one to
Costaria, the last three to Alari<a, thus indicating that he had a much
broader generic concept than can be accepted by later algologists.
Thus Agarum cribrosum, though imperfectly described specifically by
Bory, is the type of the genus as reconstituted by Postels and Ruprecht
(1840, p. 11). They enumerated five species, as follows: A. Gmelini,
A. Turneri, A. pertusum, A. platyneurum, and A. hrassicae forme, all
of which were reported within our territory with the exception of
A. Tiirneri.

Key to the Species

1. Stipe cylindrical, not fimbriate; blade with two basal scrolls

1. A. cribrosum (p. 615)

1. Stipe flattened, with fimbriate margins; blade with rounded or cordate base ....

2. A. fimbriatum (p. 616)

19-5] Setchell-Gardner : Melanophyceae 615

1. Agarum cribrosum Bory
Plate 63

Holdfast consisting of numerous branched hapteres, slender and
diffuse in young specimens, thicker and more compact in older exposed
specimens ; stipe cylindrical, 3-30 cm. long, 5-8 mm. diam. ; blade
with median percurrent midrib 1.5-3 cm. wide, rather rigid, mem-
branaceous, plane, 5-9 dm. long, 2-5 dm. wide ; mature specimens
broadly ovate, cordate at base, margin smooth, unfolding at the base
from a scroll on either side of the midrib, profusely perforated ; per-
forations smooth, circular to very irregular in outline and up to 2 cm.
in diam. ; color dark brown, changing to olive green on drying ; sori
generally distributed over the blade.

Growing on rocks in the sublittoral belt. Apparently very gen-
erally distributed from the Bering Sea to Sitka, Alaska, and recently
reported as far south as San Juan Island, Washington.

Bory, in Diet, class., vol. 9, 1826, p. 193 (in part) ; Setchell, Kelps
of the U. S. and Alaska, 1912a, p. 155; Muenscher, Key to Phaeo-
phyceae, 1917, p. 260, fig. 12.

Fucus crihrosus Mertens, in Linnaea, vol. 4, 1829, p. 52. Agarum
Gmelini Post, and Rupr., Illus. Alg., 1840, p. 11, pis. 20, 21; Saunders,
Alg. Harriman Exp., 1901, p. 430, pi. 61. Agarum Turneri Post, and
Rupr., Illus., 1840, p. 12, pi. 22 ; Saunders, Alg. Harriman Exp.,
1901, p. 431. Agarum pertusum Post, and Rupr., Illus., 1840, p. 12,
pi, 23. Fucus Agarum Turner, Fuci, vol. 2, 1809, p. 10, pi. 75.

The earliest name for this plant was Fucus Agarum Turner, but
since the generic name Agarum was limited by Postels and Ruprecht
to this and related species, this specific name became untenable and
they chose Turneri. By the laws of priority, however, the earlier
specific name crihrosum must be considered as the one to be adopted.
Postels and Ruprecht attempted to distinguish five species of which
three had broad flattened midribs and two had narrow prominent mid-
ribs, but our experience, which includes a study of a great variety of
Agarums from both the Atlantic and the Pacific coasts of North
America, leads us to feel that there is no distinct cleavage as to con-
stancy^ of broader or narrower midrib and as to coarser or finer per-
forations, all gradations appearing on both coasts. The unrolling of
the blade from two small cone-like scrolls distinguishes Agarum
crihrosum as a true Agarum.

616 University of California Puhlications in Botany [Vol. 8

2. Agarum fimbriatum Harv.

Plate 71

Holdfast of branched hapteres, profuse and slender in specimens
growing' on wood in sheltered localities, larger and denser in old
specimens growing in situations exposed to the action of the w^ater ;
stipe flat, 2-6 cm. long, 4-7 mm. wide, beset with numerous, often
branched, fimbriae on the margins, particularly near the blade; blade
thin and bullate, nearly circular to narrowly elliptical in outline, 2-8
dm. long, 1.5-2.5 dm. wide, base rounded or slightly cordate, margin
crisp and fimbriate, midrib fairly broad, complanate, perforations
few and irregular in outline ; color light brown.

Growing in the sublittoral belt from 60 fathoms up to low water
mark on stones, other algae, or on piles, woodwork, etc. Abundant
in the quiet water of the lower Puget Sound region, particularly Island
County, Washington. Rare on the California coast but found floating
as far south as San Pedro.

Harvey, Coll. Alg. N.W. Amer., 1862, p. 166 ; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 266; Setchell, Kelps of U. S. and Alaska,
1912fl, p. 155 ; Muenscher, Key to Phaeophyceae, 1917, p. 262, fig. 13 ;
Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. XCI;
Tilden, Amer. Alg. (Exsicc), no. 345,

We have been able to examine hundreds of specimens of this species
from Puget Sound and to note its characters. The lack of distinct
scrolls at the base of the blade and the thin extremely flattened stipe
giving ofif, more or less abundantly fimbriate, haptere-like outgrowths,
are characters distinguishing this species from Agaruni crihrosiim
most satisfactorily. Outside of a few floating specimens, possibly
from deep water, found in southern California, A. fimhriatum is nar-
rowly confined to the region of the inner Straits of Juan de Fuca and
to the shores of Island County, Washington.

The type locality is Esquimalt, near Victoria, British Columbia,
where it was dredged in 4-10 fathoms of water.

TRIBE 4. HEDOPHYLLEAE setchell

Stipes present only in very young plants, later disappearing or
remaining short and obscure; blade plane (or irregularly bullate),
soon becoming sessile and usually more or less decumbent, thickened
and emitting hapteres either scattered or in whorls ; meridional region

1^--^] Setchell-Gardner: Melanophyceae 617

usually wearing away, leaving the thickened (and sometimes auri-
culate) basal margins separated from one another, to grow on into
simple, or, in turn, dichotomously falsely branched fronds.

Members of the Laminariaceae with plane blades and with false
branching of pseudostipes formed from the thickened lower margins
of the fronds.

Setchell, Kelps of the U. 8. and Alaska, 1912a, p. 151.

Key to the Genera

1. Pseudoblades without auricles 47. Hedophyllum (p. 617)

1. Pseudoblades with auricles 48. Arthrothamnus (p. 619)

47. Hedophyllum Setchell

Holdfast of branched hapteres; stipe distinct at fir.st, later dis-
appearing in some species; blade short, broad and plane, at first often
becoming bullate, not auriculate at the base, later giving rise to
branched hapteres from the lower margin, either scattered or whorled ;
sori basal, irregular in outline.

Setchell, Notes on Algae, I, 1901. p. 121.

Key to the Species

1. Stipe very short, soon vanishing; blade sessile, often much bullate

1. H. sessile (p. 617)

1. Stipe longer, flat, persistent, blade subsessile 2. H. subsessile (p. 618)

1. Hedophyllum sessile (Ag.) Setchell

Plate 73

Stipe of young plant very short and much flattened, soon entirely
disappearing ; blade at first ovate, entire, .soon splitting deeply even to
the base, and becoming decidedly cucullate, in age becoming ample,
much plicated and absolutely attached, sessile, not greatly thickened
at the base, giving ri.se to hapteres along the sessile margin, 30-50 cm.
(up to 15 dm.) long, and 8 dm. wide; surface of blade either per-
fectly smooth or Avith irregular bullate swellings scattered over the
whole blade or only the basal portion, with numerous mucilage ducts
surrounded by small secreting cells ; .sorus extensive, basal, irregular
in outline.

Growing on rocks in the middle and lower littoral and in the
upper sublittoral belts. Ranging from Yakutat Bay, Alaska, to Point
Sur, California.

618 University of California Publications in Botany [Vol. 8

Setchell, in Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. VIII, Notes on Algae, I, Zoe, 1901, p. 121; Saunders,
Alg. Harriman Exp., 1901, p. 429, pi. 51 ; Setchell and Gardner, Alg.
N.W. Amer., 1903, p. 262 ; Setchell, Post-emb. Stages, 1905, pp. 119-
123, pi. 12, Kelps of the U. S. and Alaska, 1912a, p. 152. Laminaria
sessilis Agardh, Syst. Alg., 1824, p. 270; Tilden, Amer. Alg. (Exsicc),
no. 344. Laminaria apoda Harvey, Notice of a collection of algae, 1862,
p. 167.

Hedophyllum sessile is the type of the genus. It is the Laminaria
sessilis of Agardh (lo'C. cit.), said to have come from "Mari Australi. "
The type specimen of L. sessilis is in the herbarium of J. G. Agardh
under number 1730. Also in the same herbarium under number 1731
is a plant marked "Laminaria apoda Harv., Esquimalt. " This was
probably contributed by Harvey,

Both of these specimens have been carefully examined by one of
us (Setchell) and found to be our plant, H. sessile. It was next col-
lected by Dr. David Lyall in the "Fuca Straits" and later classified
as Laminaria apoda by Harvey {loc. cit.). There are six specimens
in the herbarium of Harvey at Dublin, probably of this collection,
which have been examined by one of us (Setchell) and found to be
good H. sessile.

The size of the plant and the amount of bullations depend largely
upon the habitat in which they grow. On exposed coasts, e.g., at
Fort Ross, California, they are short, scarcely over three decimeters
long and practically the whole blade is much bullated. At Neah Bay,
Washington, in a sheltered location, where they grow in great pro-
fusion at and below extreme low water level, the blade frequently
becomes fifteen decimeters long and eight decimeters wide, folding
back and forth as it widens and becoming attached at the base. The
blades may be entirely smooth or only the lower parts bullate.

2. Hedophyllum subsessile (Aresch.) Setchell

Young fronds provided with a normal holdfast, attached by a few
branched hapteres ; stipe short, stout, somewhat flattened ; blade of the
young plant at first narrow and simple, later becoming broader than
long and divided, with a cordate base and marginal rows of bullae ;
at maturity the base becomes deeply cordate or even reniform, the
basal margins are much thickened and give rise to hapteres which
attach the thickened bases to the substratum ; the central portion of

192o] SetcheU-Gardner : Melanophyceae 619

the blade dies away clear to the base, leaving two separated, partial
blades, each at the extremity of a thickened basal margin which
resembles very closely a running rootstock; blade 60-75 cm. long,
segments very variable in width, somewhat rolled at the base, but not
auriculate ; mucilage ducts present in both blade and stipe.

Growing on rocks in exposed localities in the middle littoral belt.
Known in Bering Sea and possibly extends for some distance south
along the coast of Alaska.

Setchell, in Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. XXVII, Notes on Algae, I, Zoe, 1901, p. 122; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 263, pi. 20; Saunders, Alg.
Harriman Exp., 1901, p. 430. Arthrothamnns Bongardianus J. G.
Agardh, De Lam., 1867, p. 28 (in part). Hafgygia Bongardiana f.
subsessilis Areschoug, Obs. Phyc, part 4, 1883, p. 5.

This very curious plant has been described and illustrated by us
in our Algae of Northwestern America. It is intermediate between
the simpler decumbent Hedophylhini sessile and such a decumbent
species as Arthrothamnus hifidus (Gmel.) Ruprecht. HedopJiyllum
spirale Yendo (1903) is an erect species, even more closely related to
the erect species of Arthrothamnus, difficult to place with precision.
Yendo 's later account (Notes on algae new to Japan II, 1914,
pp. 269-273, with fig.) seems to us to have confused several different
plants as well as distinctly different views on relationships. This is
particularly true as regards Postels and Ruprecht 's Laminaria Bon-
gardiana^ and is due, we think, to the fact that Yendo had access to
so little material for comparison. The plates of Postels and Ruprecht
(1840, pis. 13, 14) show clearly a digitate Laminaria, even plate 14
being quite possible of this interpretation. Very possibly plants of
Hedophylhini suhsessile or of H. spirale were mixed in the material,
yet the conception of the species is largely that of a digitate Laminaria
and one that we have referred under L. platynieris. The Hedophyllum
suhsessile of Setchell is very different from H. spiral of Yendo, the
latter approximating Arthrothamnus.

48. Arthrothamnus Rupr.

Original true stipe short, simple, flattened, attached by short stout
hapteres; original blade at first simple, smooth, flat, soon thickening
along the lower margin next to the stipe, and becoming more or less
cucullate ; the two thickened margins become the meristems, the
original blade disintegrates to the top of the stipe, the thickening

620 University of California Puhlications in Botany [Vol.8

elongating margins become secondary stipes, develop spirally and each
produces a secondary blade at its summit, the process perfecting dicho-
tomous branching; the secondary stipes become more or less decum-
bent and produce numerous hapteres along the lower margin ; the
blades produce fruit, disintegrate and bifurcate as before, the process
being repeated many times till a comj)lex organism is built up ; sori
unknown.

Ruprecht, Bemerkungen, 1848, p. 67 (11).

The type of the genus is A. kurilensis Rupr. from the island of
"Urup" in the Kurile group. At the same time that Ruprecht pro-
posed the genus he also proposed a second species, viz., A. radicans,
which is the Fucus bifidus of Gmelin (1768, p. 201, pi. 29, fig. 2)
which Ruprecht later 1851, p. 350) so recognized. The observations
on the development of ArthrotJiamnus by Yendo (1903) have thrown
much light on the systematic position of this previously little known
genus. It is obvious that it has close resemblances in development to
Thalassiopliyllum and Hedophyllum in the method of formation of
the secondary stipe and to the members of the sub-family Lessonieae
in the splitting of the transition region. Yendo 's later discussion
of Arthrothaninus (1914) is less clear than the earlier, but serves to
emphasize certain points.

Arthrothamnus bifidus (Gmel.) J. Ag.

Secondary or false stipes spirally twisted, marked with scars of
successive eroded blades, decumbent, attached by numerous hapteres,
ascending at the apices, developing into a scroll bearing the terminal
blades ; mucilage glands in the blades just beneath the cortex ; blade
membranaceous, linear, relatively long and acuminate.

Aleutian Islands, Alaska.

J. G. Agardh, De Lam., 1867, p. 28 ; Setchell and Gardner, Alg.
N. W. Amer., 1903, p. 267 ; Setchell, Kelps of the U. S. and Alaska,
1912a, p. 152. Arthrothamnus radicons Ruprecht, Algenst., 1848,
p. 68. Laminaria bifida Agardh, Sp., vol. 1, 1820, p. 122; Postels and
Ruprecht, Illus. Alg., 1840, p. 10, pi. 15. Fucus bifidus Gmelin,
Hist. Fuc, 1768, p. 201, pi. 29, fig. 2.

We are including this rather singular species of algae in our
account on the authority of Areschoug (1884, part 5, p. 14), who
reports it from the Aleutian Islands. Being quite abundant just across
on the Asiatic coast, it may justly be expected to occur somewhere
among the islands of the Alaskan peninsula.

1925] Setchell-Gardner: Melanophyceae 621

FAMILY 21. LESSONIACEAE fam. nov.

Fronds more or less compound, composed of holdfast, branching
stipe and few to numerous blades; structure as in Laminariaceae;
mucilage ducts present or absent ; blades destitute of cryptostomata
or tufts of hairs; paraphyses unicellular, with hyaline appendages.

We have included under the family Lesson iaceae all those members
of the Laminariales in which a distinct division of stipe and frond
occurs by a splitting (rather than disintegration, e.g., schizogenetic
rather than lysigenetic) process in the principal region of growth, i.e.,
at the transition place. With one exception (Lessoniopsis), there are
no outgrowths arising at the transition place as in the Alariaceae.

Key to the Tribes

1. Sori on distinct sporophylls arising as outgrowths (alarioid) at the transition-
place 3. Lessoniopseae (p. 631)

1. Sori on the ordinary blades 2

2. Stipe regularly dichotomous 1. Lessonieae (p. 621)

2. Stipe scorpioid sympodial 2. Macrocysteae (p. 626)

TRIBE 1. LESSONIEAE setchell

Members of the family of the Lessoniaceae with regularly dicho-
tomous stipes, but without specialized sporophylls.

Setchell, Kelps of the U. S. and Alaska, 1912a, p. 157.

Key to the Genera

1. Stipe flattened, solid, blade reticulately ribbed 49. Dictyoneurum (p. 621)

1 . Stipe swollen, hollow; blade plane or longitudinally grooved 2

2. Main stipe swollen at summit into a pneumatocyst

50. Nereocystis (p. 623)

2. Main stipe cylindrical to the apex 51. Postelsia (p. 624)

49. Dictyoneurum Rupr.

Original stipe very short, attached by branched hapteres, at first
erect, later becoming decumbent, much flattened and thin, developing
hapteres along the margins and later dying and disappearing as the
anterior portion advances, meristem at the advancing end of the flat-
tened prostrate stipe becoming erect and splitting dichotomously,
finally dividing the blade longitudinally into two equal parts; original
blade and often the mature later blades with distinct "midrib" and
reticulate alae, later the "midrib" disappearing more or less and the
blade becoming reticulated with coarse ribs ; sori on both sides of
blade, irregular, broad.

622 University of California Publications in Botany [Vol. 8

Ruprecht, Neue Pflanzen, 1852, p. 80, pi. 7.

Dictyoneurufii is one of several monotypic genera confined to the
west coast of North America. It was discovered by Wosnessenski in
the vicinity of Fort Ross, California, and specimens were contributed
by him to Ruprecht. The reticulate character of the blade marks it
as distinct from all other genera. In its method of the dichotomous
splitting of the blade, beginning at the transition meristematic region
between the stipe and the blade, it closely resembles Lessonia.

Dictyoneumm californicum Rupr.
Plate 70

Holdfast and stipe as in the genus ; transition or growing region at
the juncture of the stipe and blade ; no mucilage ducts ; blade rigid
and somewhat brittle, margins smooth or with scattered short spine-
like projections, variable in width with age, up to 18 cm. wide, 4-6 dm.
long, increasing in number by dichotomous splitting in the transition
point, constantly eroding at the outer free end ; the stipe portion of
the transition region constantly becoming prostrate after longitudinal
splitting, developing hapteres along the margin, and by wearing away
at the rear end the plants are multiplied vegetatively ; color yellowish
brown; perennial.

Growing on rocks in the lower littoral and upper sublittoral belts.
From Vancouver Island to Pecho, San Luis Obispo County, California.

Ruprecht, Neue Pflanzen, 1852, p. 80, pi. 7; Setchell, Notes on
Kelps, 1896, pp. 46-49, pi. 1 ; Setchell and Gardner, Alg. N.W. Amer.,
1903, p. 267 ; Farlow, Anderson and Eaton, Alg. Exsicc. Amer.-Bor.,
no. 115; Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc),
no. XI; Tilden, Amer. Alg. (Exsicc), no. 519. Costaria reticulata
Saunders, Bot. Gaz., vol. 20, 1895, p. 58, pi. 7.

Costaria reticulata Saunders (loc. cit.) was founded upon material
collected at Pacific Grove, California. His excellent illustration of
the species leaves no doubt that it is the Dictyoneurum of Ruprecht.
Saunders' illustration, unlike that of Ruprecht, is of a broad mature
blade, probably of the first or original blade and stipe of a young
plant, while the large illustrations of Ruprecht {loc. cit.) are of older
plants whose blades have recently split and which have no ' ' midribs. ' '
Superficially, in the wrinkled bullate character, the blade of Dic-
tyoneurum resembles that of a Costaria, but the tissue composing the
so-called "nerves" of the former is different and distinct from the

1025] Setchell-Gardner : Melanophyceae 623

other tissues of the blade, whereas in Costaria the tissues are uniform
among the bullae, between the five longitudinal costae. The so-called
"midrib" represents only a very slight modification of the blade. The
histological characters are the same as the remainder of the blade.
The part between two parallel longitudinal nerves is usually smooth
and free from cross nerves and is only slightly thickened at the
margins. The stipe in Dictyoneuriim is erect only in the juvenile
stage, soon becoming prostrate and attached by secondary hapteres,
only the short terminal portion remaining erect. That of Costaria
always remains erect. Dictyoneurum is perennial, lasting as long as
it can maintain attachment, and new blades are constantly being
formed. Costaria never fruits more than once.

50. Nereocystis Post, and Rupr.

Holdfast of a dense, interwoven mass of strong branched hapteres ;
stipe much elongated, slender, solid and cylindrical from the base
through the most of its length, enlarging above into a thick hollow
tube, contracting suddenly at the outer end just below the large,
terminal, spherical pneumatocyst, which bears at its summit a row of
short dichotoraous branches, on the ends of which are the long, narrow,
thin blades which bear the sori.

Postels and Ruprecht, Illus. Alg., 1840, p. 9, pi. 39, figs. 24-30.

Nereocystis differs so greatly in aspect from any species of Lessania
and even more so from the single species of Dictyoneurwm that the
resemblances are more in need of emphasis, in order to justify associa-
tion in the same tribe of the same family. In Nereocystis the first
segment of stipe is so long, so swollen, and the pneumatocyst so large,
while the branches (above the pneumatocyst) are so abbreviated and
condensed, that the Lessonia relationship is obscured, but becomes
plain on careful analysis. While Nereocystis differs so greatly in
aspect from other members of the Lessonieae, it resembles in the modi-
fication of the stipe, at least, Pelagophycus, of the tribe Macrocysteae,
so nearly that one must realize the significance of the unilateral branch-
ing of the stipe in the latter to feel justified in separating the two.
Postelsia>, however, is closely related to Nereocystis, the difference
between the main types of the type species of the two genera, apart
from dimensions, consisting largely in the development of a globular
pneumatocyst in Nereocystis, and the lack of such a terminal globular
inflation on the main stipe of Postelsia. The blades are much more
differentiated in the latter genus.

624 University of California Publications in Botany [Vol.8

Nereocystis Luetkeana (Mert.) Post, and Rupr.

Stipe very strong and elastic, 20-25 m. long, 1-1.5 cm. thick below,
the hollow portion, or pneumatocyst, 3-4 m. long, 15-17 cm. broad;
sporophjdls 25-40 in number, 3-4.5 m. long, 6-15 cm. broad ; sori in
large, irregular, promiscuously distributed areas on both sides of the
sporophylls.

Growing on rocks and epiphytic in water 5-25 m. in depth. Extend-
ing from the Shumagin Islands, Alaska, to Santa Barbara, California.

Postels and Ruprecht, Illus. Alg., 1840, p. 9, pis. 8, 9 ; Setchell
and Gardner, Alg. N.W. Amer., 1903, p. 268 ; Setchell, Kelps of the
U. S. and Alaska, 1912cf:, p. 158 ; Farlow, Anderson and Eaton, Alg.
Exsicc. Amer.-Bor., no. 116 ; Collins, Holden and Setchell, Phyc. Bor.-
Amer. (Exsicc), no. X; Tilden, Amer. Alg. (Exsicc), no. 237.
Nereocystis priapus (Gmel.) Saunders, Alg. Harriman Exp., 1901,
p. 431. Fucus Lu.etkea.nus Mertens fil, in Linnaea, 1829, p. 48.

The "Bull Kelp," as it is frequently called, is an annual plant,
grows in beds of greater or less extent, rising from even 10-12 fathoms
of water, particularly extensive in the Puget Sound region and is an
available source of potash salts and other substances. It is one of our
largest and most striking seaweeds. We are extremely disinclined to
adopt the binomial Nereocystis Priapus, although by the process of
exclusion its application to our plant may seem to be extremely
plausible. The Ulva Priapus Gmelin (1786, p. 231, pi. 31, fig. 2) was
neither well described nor convincingly illustrated. Its locality is in
doubt. Probably it is either Nereocystis or Pelagophycus, and more
probably the former than the latter. We have neither seen nor heard
anything of the type specimen.

51. Postelsia Rupr,

Holdfast of stout branched hapteres ; stipe large, cylindrical and
hollow, bearing a group of pendent blades on short, solid dichotomous
branches at its summit.

Ruprecht, Neue Pflanzen, 1852, p. 19 (75).

Postelsia is a unique, monotypic genus of algae, confined as at
present known to the temperate waters of the west coast of North
America. Its relation to Nereocystis has just been discussed. The
date of publication depends upon the date of issue of the separate
papers which have 1852 on the title page, thus seeming to antedate
Areschoug's genus Virginia.

1925] SetcheU-Gardner: MeUtnophijceac 625

Postelsia palmaeformis Rupr.

Plate 69

Holdfast of stout, blunt, branched hapteres; stipe stout, smooth
and g^lossy, elastic though rigid, cylindrical, tapering slightly upward,
hollow, 4-6 dm. high, 1.5-3 cm. diam., bearing at its tip numerous
solid, cylindrical, dichotomous branches ; blades terminating the small
branches, falcate, 100-150 on a plant, 15-24 cm. long, produced by
longitudinal splitting in the transition region at their bases, with
deep, parallel, longitudinal grooves on either side, in which the sori
are developed ; color a rich olive brown.

Growing only on rocks exposed to the heavy action of the waves.
Extending from the southern end of Vancouver Island to Lion Rock,
San Luis Obispo, California.

Ruprecht, Neue Pflanzen, 1852, p. 19 (75), pis. 6 and 8 ; MacMillan,
Kelps of Juan de Fuca, 1902, pp. 213 and 217 ; Setchell and Gardner,
Alg. N.W. Amer, 1903, p. 268 ; Setchell, Kelps of U. S. and Alaska,
1912a, p. 158; Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. XXXVIII and no. 131; Tilden, Amer. Alg., no. 341;
Farlow, Anderson and Eaton, Alg. Exsicc. Amer.-Bor., no. 113. Vir-
ginm Palma-Maris Areschoug, Oefvers. Kongl. Vet.-Akad. Forhandl.,
1853, p. 147, Flora, 1855, p. 652. Postelsia calif ornica Guignard, App.
Mucif. Lam., 1892, p. 41.

This plant, in many ways remarkable, was first brought to the
attention of phycologists by Wosnessenski, who found it growing on the
exposed shore of a small island at the entrance to Bodega Bay, Cali-
fornia. It was known to the Indians of that region as Kakgunu-
chale, according to Ruprecht. It is everywhere known today as the
Sea Palm, owing to the close resemblance of more or less extensive
clusters of these singularly beautiful plants to miniature groves of
palms as seen in the distance along ocean shores.

It has accustomed itself to growing only on rocks exposed to the
very heaviest action of the waves. It mainly inhabits the middle of the
littoral belt, but in certain localities in which the waves are accustomed
to beating high, they may grow at or even above the general high-tide
level. They are annual plants.

Postelsia palmaeformis was first published by Ruprecht {loc. dt.),
who gave a very excellent illustration of the three specimens collected
by Wosnessenski. This paper was published separately in 1852 and
afterwards in Memoirs Imp. Acad. St. Petersburg in 1855 (ser. 6,

626 University of California PuNications in Botany [Vol. 8

vol. 7, p. 75, pi. 6). Pfeiffer (Norn. Bot., vol. 2, p. 823) quotes this as
the original publication, thus considering Virginia Palma^Maris of
Areschoug {loc cit.) as antedating Ruprecht's Postelsia palmaeformis.
The Postelsia calif ornica of Guignard {loc. cit.) undoubtedly refers to
Postelsia palmaeformis, since he lists several other kelps of the
California coast along with this species.

TRIBE 2. MACROCYSTEAE kuetzing (lim. mut.)

Plants of the family Lessoniaceae having, in large part, unilateral
splitting, thus producing a scorpioid sympodial stipe.

Kuetzing, Phyc. Gen., 1843, p. 348 ; Setchell, Kelps of the U. S. and
Alaska, 1912rt,, p. 158.

The Macrocysteae have regularly dichotomous splitting in the first
few divisions, but after that the splitting is unilateral.

Key to the Genera

1. Stipes solid 52. Macrocystis (p. 626)

1. Main stipe hollow 53. Pelagophycus (p. 629)

52. Macrocystis Ag.

Holdfast of mature plants consisting of a large entangled mass of
dichotomously branched hapteres or of creeping flattened rhizomes
giving off lateral hapteres as well as erect fronds ; stipe in the juvenile
plant forking 1-3 times dichotomously, forming several main stipes,
later the branching is unilateral ; blade at first splitting equally, but
soon the terminal bladderless falcate blade splitting unequally ; meri-
stematic transition region at the juncture of the stipe and falcate
blade ; the mature blades undivided bearing a bladder at their base and
the sori on both sides; perennial.

Agardh, Sp. I, 1820, p. 46.

The species of Macrocystis are not well marked off from one another
but the genus, with its unilateral splitting and its long cylindrical
stipe, unbranched above the basal dichotomies, and its large and com-
plicated holdfasts or overlapping massive "rhizomes," is distinct
from any other.

Concerning the position of the sori in members of this genus,
Skottsberg (1907, pp. 104-108) and Howe (Mar. Alg. Peru, 1914,
pp. 62, 63, 65, 66) have reviewed the literature and the facts most
thoroughly. The sori, in our species, seem confined to the basal leaves,
and specimens are not commonly collected. The sori do not seem to

1925] Setchell-Gardner : Melanophyceae 627

be confined to grooves in the leaves (cf. Hoffman, 1911, p. 155) as
Smith and Whitting (1895, p. 84, pi. 20) found them, as did also
Skottsberg {Joe. cit.).

The sporelings of Macrocystis have been observed by R. P. Brandt
(1923, pp. 4-7, fig. 2), who did not succeed, however, in clearly elu-
cidating the complete life-history of the gametophyte. Brandt also
deals with the polymorphy, especially as regards the shape of the
bladders, due to more quiet or rougher waters.

Key to the Species

1. Fronds arising from masses of branched hapteres 1. M. pyrifera (p. 627)

1. Fronds arising from stout creeping rhizomes 2. M. integrifolia (p. 628)

1. Macrocystis pyrifera (L.) Ag.

Plates 64 and 65

Holdfast very large, often becoming nearly a meter in diameter in
the typical deep sea form ; stipe at first dichotomously branched, estab-
lishing 2-8 growing points ; the main stipes become 30-50 m. long, bear-
ing at their summits the young differentiating blades and along the
greater part of their length, at regular intervals, the mature lateral
blades, each with a pyriform bladder at its base, which is in turn sup-
ported by a short cylindrical stipe ; blades rigid, coarsely rugose, 3-5
dm. long, 5-9 cm. wide, with spinulose margins.

Growing on rocks usually in 20-30 m. of water. It extends from
Sitka, Alaska, to Magdalena Bay in Lower California, but is a plant
of the outer coa.sts, exposed to wave action.

Agardh, Sp. I, 1820, p. 47 ; Setchell, Kelps of the U. S. and Alaska,
1912a, p. 158; Postels and Ruprecht, Illus. Alg., 1840, p. 9, pi. 6;
Saunders, Alg. Harriman Exp., 1901, p. 431, pi. 60 ; Setchell and
Gardner, Alg. N.W. Amer., 1903, p. 270 ; Collins, Holden and Setchell,
Phyc. Bor.-Amer. (Exsicc), no. XXXTX; Tilden, Amer. Alg.
(Exsicc), no. 518. Fucus pyriferus Linnaeus, Mantissa, II, p. 311;
Turner, Hist. Fuc, vol. 2, 1809, p. 103, pi. 110.

We have not attempted to segregate varieties or forms of this
species, since we have not been able to determine any exact lines of
cleavage. The varying shape of the bladders from globular to
extremely elongated, the varying breadth, length, dentation, wrinkling,
etc., of the leaves seem possibly matters of age and character of habitat.
There are also some indications of the possibility that this species is
only the deep water condition of the next. The exact northern and

628 University of California Publications in Botany [Vol. 8

southern limits of this species have not been satisfactorily determined.
W. C. Crandall (Fert. res. U. S., 1912, p. 210) has seen ''beds as far
south as Cedros Island," off the coast of Mexico, and states that
"according to local fishermen these beds extend to Magdalena Bay."
The northern limit, as growing near Sitka, rests especially upon the
report of Edward C. Johnston (Fert. res. U. S., 1912, p. 215).

Our Macrocystis of the north Pacific seems to grow in warmer water
than that of the Antarctic and south Pacific. While the southern
hemisphere form grows in the comparatively warmer waters of Peru
it also grows about Cape Horn in waters of extreme coolness. In our
Alaska waters, it seems to stop short at Sitka and does not go farther
west and north into the coldest waters of our region, Sitka being
decidedly under the isothere of 10° C. but just about on the isocryme
of 0° C. The coldest locality for Macrocystis seems to be South
Georgia, between the isocryme of somewhat below 0° C. and the
isothere of 5° C,

2. Macrocystis integrifolia Bory
Plate 62

Fronds relatively slender, composed of a flattened, almost ligulate,
prostrate, creeping, rhizome-like portion, 2-3.5 cm. wide, profusely
and for the most part dichotomously branched, attached closely and
firmly to rocks by numerous, strong hapteres arising along its two
margins and giving rise to numerous free erect portions, relatively
slender, 5-8 m. long, bearing the vesicles with blades at their distal
ends ; vesicles subspherical to broadly pyrif orm ; blades denticulate
along the whole margin, 2.5-3.5 dm. long, 3.5-5 cm. wide, acuminate
at both ends.

Growing on rocks at slightly below extreme low water mark.
Central California.

Bory, in Diet, class, d'hist. nat., vol. 10, 1826, p. 9.

At several localities along the coast of central California, notably
at Monterey Bay and Carmel Bay, for many years we have observed
quantities of a species of Macrocystis attached to boulders and rock
ledges at and just below low-tide level which, according to our judg-
ment, is entirely distinct from the abundant and widely distributed
M. pyrif era (Turner) Ag. We have thought that possibly it might
represent only stranded individuals of that species. The fundamental
difference between the two species as represented on our coast is the

1^25] Setchell-Gardner : Melanophyceae 629

character of the attaching portion — "rhizome" and hapteres. In the
deep sea species, there is no rhizome and the mass of hapteres developed
often becomes several decimeters in thickness and up to a meter in
diameter, while in the shore species the conspicuous "rhizome" is flat
and adheres closely to the rock, dying and decaying at the rear as it
advances and spreads out in all directions. We have had an excellent
opportunity to study what might be considered a natural experiment
upon the effect of changing the deep sea species to the habitat of the
shore species at San Pedro, where a government breakwater was
extended out from the shore a long distance in the vicinity of a large
"kelp bed" of Maicracystis pyrifera. Thousands of plants attach
themselves to the rocks along low-tide level and persist until they are
torn loose. In not a single instance has the nature of the holdfast
been changed. It thus seems perfectly definite that the shore species
is a distinct entity, as well as the deep sea species. We have never
seen any specimens of either species which would seem to represent
transition stages between the two.

53, Pelagophycus Aresch.

Holdfast of several whorls of strong, dichotomously branched
hapteres diminishing in size toward their termini ; stipe solid at the
base, hollow above, constricted at the summit just below the large
spherical bladder, again becoming smaller and solid for a few centi-
meters beyond the bladder at its summit, dividing once dichotomously,
each branch again dividing unilaterally 4-6 times and each branch
bearing a single large terminal blade.

Areschoug, in Botaniska Notiser, 1881, p. 49 ; Observ. Phycol.,
part 5, 1884, p. 6.

Pelagophyciis is one of the large, conspicuous, monotypic kelp
genera confined to a relatively small area on the Pacific coast of North
America. Its resemblance to Nereocystis is striking so far as the main
stipe up to the first dichotomy is concerned. Above the first, or occa-
sionally, the second dichotomy, however, the unilateral splitting and
sympodial character of the stipe clearly places the genus with the
Macrocysteae.

630 University of California Publications in Botany [Vol.8

Pelag-ophycus porra (Leman) Setchell

Holdfast composed of whorls of dichotomously and later fasieu-
lately branched hapteres, the whole mass about 1.5 dm. diam. ; stipe
solid, terete and slender below, increasing slightly in diameter upward
for 6-7 m., merging abruptly into a swollen, hollow portion, the apo-
physis, about 80 cm. long, which is deeply constricted just below the
terminal bladder or pneumatocyst ; pneumatocyst spherical, 12-20 cm.
diam., filled with gas, giving rise on its summit to 2 solid branches,
1.3-1.6 m. long, decidedly flattened, each bearing about 5 short lateral
branches on the distal side ; the main and the lateral branches terminate
in blades which are 4-5.5 m. long, 20-45 cm. wide, with cuneate base,
borders loosely ruffled, more or less beset with small, spine-like pro-
jections, coarsely rugose, easily torn; sori scattered in irregularly
shaped areas.

Growing in open waters, 10-15 fathoms, mostly attached to rocks.
From the vicinity of Point Conception, California, to some unknown
locality on the coast of Lower California (Mexico).

Setchell, Nereocystis and Pelagophycus, in Bot. Gazette, vol. 45,
1908, pp. 129-134, The Elk Kelp, in Erythea, vol. 4, 1896a, pp. 179-
184, pi. 7, Kelps of the U. S. and Alaska, 1912a, p. 159 ; Collins, Holden
and Setchell, Phyc. Bor.-Amer. (Exsicc), no. CIX. Nereocystis
gigantea Aresehoug, in Bot. Not., 1876, p. 71. Pelagophycus giganteus
Areschoug, in Bot. Not., 1881, p. 49. Laminaria porra Leman, in
Diet. Sci. Nat., vol. 25, 1822, p. 189.

Pelagophycus porra is quite limited in its distribution as noted
above. It has been known to the civilized world in a general way
since the days of the early Spanish navigators to the west coast of
North America. Although it is a plant which is normally anchored
relatively near the shore, it may become detached and, coming into
an ocean current, may float for many miles out to sea, remaining alive
for many months. The early navigators, on approaching the shores
of Mexico and California, were constantly on the lookout for this
floating seaweed, the presence of which was looked upon by them as
an unfailing sign of their near approach to land. Porra was one of
the names by which it was known, but the term also applied to both
Macrocystis and Nereocystis as well (cf. Setchell, 1908, for a complete
account of the early literature bearing on this alga). At present it is
generally known as Elk-kelp and is one of the kelps made use of in
large quantities during the recent war as a source of potash, etc.

1925] Setchell-Gardner: Melanophyceae 631

Leman {Jac. cit.) was the first to publish a critical diagnosis of
the species under the combination, Laminaria porra. Nothing was
done with it in a botanical way until 1876, when Areschoug redescribed
it from material collected by Dr. G. Eisen, at Santa Catalina, Cali-
fornia, placing it with Nereocystis as N. gigmiiea. Later (1881) he
created the genus PeJagophycus to receive the same species where it
remained until the combination employed here was made (cf. Setchell,
Joe. cit.).

TRIBE 3. LESSONIOPSEAE setchell

Members of the Lessoniaceae having specialized sporophylls arising
as outgrowths on the outer margins of the transition place where
splitting is about to occur,

Setchell, Kelps of the U. S. and Alaska, 1912ff, p. 160.

The tribe of the Lessoniopseae might perhaps be placed with equal
propriety either under Lessoniaceae or under Alariaceae, since the
sole genus, monotypic, has the characters of each of these families.
The plant, however, has the habit of a Lessonia and this influences us
strongly to place it nearer to Lessonia than to Alaria.

54. Lessoniopsis Eeinke

Frond differentiated into holdfast, stipe, blade, and sporophylls;
holdfast consists of a dense mass of short, thick, dichotomously
branched hapteres; stipe short, cartilaginous, very dense and rigid,
arborescent, more or less deeply furrowed, merging into hapteres below,
irregular in outline, profusely dichotomously branched above, branch-
ing taking place by splitting in the meristematic region, terminating
in long narrowly linear blades with distinct percurrent midrib and
stipitate base ; the meristematic transition region at the base of each
blade giving rise to sporophylls in pairs, 1-3 pairs each season.

Reinke, Studien zur Entwick. Lam., 1903, pp. 25-28.

Lessoniopsis is one of the many monotypic genera of kelps thus
far known exclusively on the west coast of North America. It was
established by Reinke to receive Lessonia Uttoralis Farlow and
Setchell. Although among the relatively recent discoveries, its morph-
ology, anatomy and life-history, except as to the embryonal stages,
have been quite thoroughly made known.

In its method of development of the sporophylls, its affinities are
close to Pterijgophora. It resembles Lessonia in its method of dicho-
tomous branching of the blade and the consequent production of new
blades.

632 University of California PuMications in Botany [Vol. 8

Lessoniopsis littoralis (Farlow and Setchell) Keinke

Plates 67 and 68

Frond up to 2 m. high ; stipe up to 2 dm. thick at the base, very
dense and cartilaginous; blade 7-12 mm. wide, up to 8 dm. long, with
percurrent midrib 1-2 mm. wide, up to 800 in number on a single
plant, splitting longitudinally at the transition region ; sporophylls
always wider than the blade, but generally considerably shorter,
broadly ovate below, somewhat narrowed and rounded at the apex,
arising in pairs on the edges of the flattened transition region ; sori on
both sides covering most of the sporophyll; fruiting in summer and
then eroding from the apex; color dark olive green; mucilage ducts
absent from both stipe and blade.

Growing on large boulders and rock ledges exposed to the action
of the heaviest surf. Banging from Sitka, Alaska, to Point Carmel,
California.

Reinke, Studien zur Entwick. Lam., 1903, pp. 25-28, fig. 8 ; Griggs,
Sporophylls of Lessoniopsis, Ohio Nat., 1909a, vol. 9, no. 4, p. 437,
Juvenile kelps, 1909, p. 9 ; MacMillan, Observations on Lessonia, Bot.
Gaz., vol. 30, 1900, p. 318; Setchell and Gardner, Alg. N.W. Amer.,
1903, pp. 267, 268. Lessonia littoralis Farlow and Setchell, in Tilden,
Amer. Alg. (Exsicc), no. 342; Collins, Holden and Setchell, Phyc.
Bor.-Amer. (Exsicc), no. XXXVII. Lessonia fuscescensil) Farlow,
Mar. Alg. U. S., 1875, p. 355 (not of Bory). Lessonia nigrescens
Farlow, Mar. Alg. U. S., 1876, p. 707 (?) ; Setchell, Class, and Geog.
Dist. Lam., 1893, p. 357 (neither L. nigrescens Bory).

This species was first brought to the attention of phycologists by
Mr. E. Hall who sent specimens of it which he found on the coast of
Oregon to Farlow, who (1875, p. 355) doubtfully referred it to Les-
sonia fuscescens. Still later, Farlow collected living material near
Monterey, California, and recognizing it as a new species of Lessonia
he gave it the manuscript name, L. littoralis. Tilden distributed the
species from Vancouver Island, in 1900, as no. 342 in American Algae
and gave the first diagnosis, accrediting the name to Farlow and
Setchell. A complete account of the plant was given in the same year
by MacMillan. Reinke (loc. cit.) was the first to recognize the funda-
mental distinction between this species and those of the genus Lessonia
proper, viz., that all of the terminal blades with midribs are sterile
and that the other blades which are free from midribs are in reality
the sporophylls. The sporophylls are lateral in origin, and in this
respect are like those of the genus Pterygophora, and continue to

1925] Setchell-Gardner : Melanophyceae 63o

develop in the transition region in pairs year after year as the meri-
stem moves forward until the plant is dislodged and dies. Griggs
(1909) was the first to properly describe the origin of the sporophylls.

The branching is strictly dichotomous and is accomplished by the
splitting of the blades longitudinally through the midrib, the splitting
beginning at the transition region and progressing to the outer end.
A new ala develops on each half blade as the splitting proceeds.

Lessmiiopsis littoral is and Postelsia paJmaeformis are the most
typical examples of cumatophyte species of our algal flora. No surf
seems to be too rigorous to hinder their optimum development. The
former usually occupies a belt lower down than the latter. Apparently
it is unable to withstand the desiccation incident to long exposure to
the air.

FAMILY 22. ALARIACEAE fam. nov.

Fronds simple or irregularly branched, with terminal blades and
lateral outgrowths, the latter arising at the transition places ; terminal
blade with or without a midrib or central thickened area, plane or
rugose, with or without cryptostomata, or tufts of hairs ; otherwise as
in the order.

The family of the Alariaceae is intended to include all the genera
whose species possess sporophylls, either kinetic or potential, arising
as outgrowths of either stipe or blade and arising at the transition
place, with the exception of the genus Lessoniopsis, described above.

Key to the Tribes

1. Mature outgrowths confined to the stipe 1. Alarieae (p. 633)

1. Mature outgrowths not as above 2

2. Mature outgrowths confined to the blade 2. Ecklonieae (p. 645)

2. Mature outgrowths on both stipe and blade 3. Egregieae (p. 647)

TRIBE 1. ALAEIEAE setchell

Members of the Alariaceae with sporophylls only on the stipe.

Setchell, Kelps of the U. S. and Alaska, 1912^, p. 160.

The Alarieae are confined to the northern hemisphere, occurring

in the Arctic, the north Atlantic, and the north Pacific oceans. The

outgrowths at the transition place are arranged on the stipe and are

usually highly specialized sporophylls, bearing sori, maturing and

disintegrating.

Key to the Genera

1. Midrib distinct; sporophylls definitely limited in growth 56. Alaria (p. 635)

1. Midrib indistinct; sporophylls more or less indefinite in growth

55. Pterygophora Cp. 634)

634 University of California PuNications in Botany [Vol.8

55. Pterygophora Rupr,

Holdfast of stout, branched hapteres; stipe simple, solid, more or
less woody, containing mucilage ducts ; blade terminal, linear, without
distinct midrib but with central portion thickened ; sporophylls lateral
on both sides of the upper part of the stipe at the transition region,
long, and of continued growth ; sori on both sides of sporophylls and
on the terminal blade ; perennial, the stipe increasing in length and
in diameter through several years.

Ruprecht, Bemerkungen, 1848, pp. 8 (64) and 14 (70) (nomen),
Neue Pflanzen, 1852, p. 17 (73) (description).

Pterygophora was first proposed by Ruprecht in 1848, as above
mentioned, in connection with a study of the structure of the stipes
of some of the large Melanophyceae. A diagnosis of the whole plant
was deferred until 1852. No other species than the one then proposed
as the type, viz., P. calif ornica, has been discovered since, and this one
seems to be in a state of specific equilibrium, as it varies but slightly
throughout its entire range of distribution. It was discovered by
Wosnessenski in the vicinity of Fort Ross, California, in 1840. It is
less specialized than Alaria.

Pterygophora californica Rupr.

Plate 74

Stipe terete below, flattened and somewhat constricted above at the
transition region, with distinct concentric rings shown in cross-section,
8-12 dm. (up to 2 m.) long, 2.5-4 cm. (up to 7 cm.) diam., giving rise
successively to elongated, blade-like sporophylls, pinnately arranged
in the transition region ; blade with mucilage ducts and with a slightly
thickened median, longitudinal band, forming a false midrib, linear,
6-9 dm. long, 6-10 cm. wide, fruiting annually then dying back and
regenerating ; sporophylls of the same general shape as the blade, but
smaller, stipitate, without median thickening, 12-18 maturing in a
season, then disintegrating, leaving only a scar on the stipe, new ones
meanwhile developing above ; sori occupying the basal half or more of
each sporophyll on both sides.

Growing on rocks and on other large algae in the upper sublittoral
belt, largely on exposed coasts. From the southern end of Vancouver
Island to Lower California.

1925] Setchell-Gardner: Melanophyceae 635

Riipreeht, Neue Pflanzen, 1852, p. 17 (73), pi. 5; Setchell and
Gardner, Alg. N.W. Amer., 1903, p. 271 ; MacMillan, Observations on
Pterygophora, 1902rt, p. 723, pis. 57-62; Muenscher, Key to the
Phaeophyceae, 1917, p. 280, fig. 45; Frye, The Age of Pterygophora
calif arnica, 1918, pp. 65-71, pi. 17 ; Farlow, Anderson and Eaton, Alg.
Exsicc. Amer.-Bor., no. 114 ; Collins, Holden and Setchell, Phyc. Bor.-
Amer. (Exsicc.), no. CVIII ; Tilden, Amer. Alg. (Exsicc), no. 520.

Pterygophora calif o^'nica is a perennial. No accurate data,
obtained from experimental evidence, has yet appeared to indicate
the age of this sturdy species of algae. Frye (loc. cit.) has sum-
marized the opinions of various writers on the subject and has given
the results of his own observations on plants in the vicinity of Blakeley
Island and at Cape Flattery, Washington. His results are based upon
the number of scars on the stipe, produced by the disintegration of
sporophylls, and upon the number of rings in the stipe. Thirteen
years is the oldest estimated age of any plant which he has discovered.

56. Alaria Grev.

Holdfast comparatively small, of more or less slender branched
hapteres from near the base of the stipe, forming a turbinate compact
mass; stipe comparatively small and usually short, solid, unbranched,
with or w^ithout mucilage ducts; blade terminal, considerably elon-
gated, entire, thin, Avith a pronounced longitudinal, percurrent, cen-
trally located midrib, usually showing tufts of hairs (cryptostomata) ;
sporophylls from both sides of the stipe, of limited growth, developing
in the transition region ; sori usually covering nearly the whole of
both surfaces of the sporophylls ; perennial.

Greville, Alg. Brit. Syn., 1830, p. xxxix and p. 25.

Greville remarks {loc. cit.) concerning the establishment of the
genus as follows : ' ' The individuals which constitute this genus I have
removed from an assemblage published by Bory de St. Vincent under
the name of Agarum.'' The individuals to which Greville refers, as
enumerated in the Synopsis, were Agarum esciilentum, A. Delisei and
A. Pylaii. As the type of his new genus he selected the first which is
the FiicAis esculentus of Turner (1809, pi. 117), this being the only
species of the three mentioned by Bory which occurs within the British
waters, the other inhabiting "Terre-Neuve." The Vienna Congress
has conserved the generic name Alaria Grev. (1830) as against Musae-
/o/m Stackhouse (1809) and Orgyia ^tackhouse (1816).

636 University of California Puhlications in Botany [Vol. 8

No genus of the Laminariaceae, not even excepting the genus
Lamvnaria, is so confusing as to specific segregation within its limits
as Alaria. Ruprecht, J. G. Agardh, Kjellman, Setchell, and Lendo
have struggled with it and assisted both in clearing some species and
confusing others. The arrangement of our species by Setchell (1912a)
and the monograph of the genus by Yendo (1919) are followed in their
essentials in our account. We cannot, however, follow Yendo in all
his distinctions since our experience leads us to assign somewhat
different degrees of importance to certain of the characters which he
stresses. We are not able to apply the distinction into "Holosoria"
and "Metasoria" with such precision even as Yendo (1919, pp. 24-26)
indicates as possible.

Key to the Species

1. Midrib solid throughout 2

1. Midrib fistulose at intervals 9. A. fistulosa (p. 644)

2. Sporophylls short 3

2. Sporophylls long 7

3. Sporophylls narrow 4

3. Sporophylls broad 5

4. Blade long and broad above 2. A. praelonga (p. 637)

4. Blade short and narrow 1. A. nana (p. 636)

5. Stipe long, very much flattened 3. A. tenuifolia (p. 638)

5. Stipe short, cylindrical 6

6. Midrib broad 4. A. marginata (p. 640)

6. Midrib narrow 5. A. Pylaii (p. 641)

7. Sporophylls narrow 8

7. Sporophylls broad 8. A. valida (p. 643)

8. Cross-section of midrib elliptical 6. A. dolichorhachis (p. 642)

8. Cross-section of midrib oblong 7. A. lanceolata (p. 642)

1. Alaria nana Schrader

Plants anchored by firm strong hapteres; stipe 4.5-7 cm. long,
robust, 5-8 mm. diam., terete; rhachis 2-4 cm. long, slightly com-
pressed, passing gradually into the midrib ; blade usually widest near
the base tapering rather abruptly below and gradually above, with
some specimens nearly linear, 40-60 cm, long, 3-8 cm. wide ; midrib
prominent, 4—6 mm, wide, nearly rectangular in cross-section ; sporo-
phylls 25-50 in number, linear to slightly elliptical, 6-12 cm. long,
8-15 mm, wide, rounded at the outer end, tapering rather abruptly at
the base to a distinct short stipe, sori covering both entire surfaces.

Growing on rocks in the upper part of the littoral belt, in localities
exposed to the action of the heavy surf, in company with Postelsia
palmaeformis and Lessoniopsis littoraHis, Port Renfrew, Vancouver
Island, British Columbia,

if^2o] Setchell-Gardner: Melanophyceae 637

Schrader, Observations on Alariu nana, 1903, p. 157, pis. 23-27 ;
Yendo, Monogr., 1919, p. 118, pi. 13, figs. 1-3. Alaria praeJonga f.
nana, Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no.
1292. Alaria marginata f. nana Collins, Mar. Alg. Vancouver Isl.,
1913, p. 110.

The above locality is the only one in which this species is certainly
known to occur. There are plants, however, growing on the central
California coast in similar habitats which seem to be closely related to,
if not identical with, the Port Renfrew plant. Specimens have been
distributed from Golden Gate, San Francisco, in Collins, Ilolden and
Setchell, Phyc. Bor.-Amer., no. 1292, under A. praelonga f. nana.
Yendo (1919, p. 118, et seq.) considers this a distinct species on
account of the shape of the sporophylls. We are still of the opinion,
in which Kjellman agreed, that it is very closely related to A.
praelonga.

2. Alaria praelonga Kjellm.

Stipe 5-8 cm. long, terete ; rhachis 3-5 cm. long, larger than the
stipe, somewhat flattened, passing almost directly into the blade,
muriculate; blade up to 4 m. long, 7-10 cm. wide, with narrowly
cuneate base, plane, smooth, shining, rigid, dark brown color ; costa
more or less prominent, narrow, in cross-section slightly enlarged at
the ends or elliptic-linear, passing abruptly into the blade ; sporophylls
numerous, fasciculate, distinctly petiolate, oblong-lanceolate to linear-
lanceolate, 8-10 cm. long, 1-2 cm. wide, plane, rigid ; sori covering
nearly the entire surface.

Growing in the lower littoral belt. From St. Paul Island, Bering
Sea, Alaska.

Kjellman, Om Beringh. Algfl., 1889, p. 38, Tab. 4, figs. 1-4.

Alaria praelonga Kjellman is a much misunderstood species and
seemingly needlessly so. Relying upon Kjellman 's opinion (in lift.),
we have classified under this name such diverse species as A. marginata
Post, and Rupr. and A. lanceolata Kjellm. The type, both as repre-
sented by the type specimen and by Kjellman 's figure, is a plant with
a short stipe, short narrow sporophylls, a long, moderately broad
blade, very long and narrowly attenuated at the base. The figure of
Yendo (1919, pi. 4) seems to have broader as well as longer sporo-
phylls than the type, with the sporophylls broader at the base. We
have only a single imperfect plant which, in the light of Kjellman 's
figures and type specimen, we venture to refer finally to this species.

638 University of California Publications in Botany [Vol. 8

It was collected on St. Paul Island of the Pribiloff Group. Yendo has
(1919, p. 88) with justice said that '*it is to be questioned as to how
Setchell comprehended A. praelonga Kjellm. " Setchell, finally,
has restricted his view to a plant closely resembling^ Kjellman's figure
and type specimen, both young plants.

3. Alaria tenuifolia Setchell

Stipe 16-60 cm. in length, cylindrical or nearly so only at the very
base, much flattened above, slender and flexible ; rhachis more or less
elongated, flattened, with the sporophylls at first remote in young
plants, later crowded ; blade 10-15 dm. long, 10-35 cm. wide, broadly
to narrowly cuneate at the base, very thin, collapsing when withdrawn
from the water, with plentiful cryptostomata of small size ; midrib
narrow to fairly broad, oblong in cross-section ; sporophylls from
narrowly to broadly lanceolate and cuneate at the base, varying to
broadly ovate or oblong and distinctly cordate at the base, sessile or
with more or less pronounced stipes.

Not uncommon, from the Bering Sea to Puget Sound, Washington.

Setchell, in Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. XLV, Kelps of the U. S. and Alaska, 1912a, p. 162,
Critical notes on the Laminariaceae, 1908^, p. 12 ; Setchell and Gard-
ner, Alg. N.W. Amer., 1903, p. 272. Alaria Pylaii Yendo, Monogr.,
1919, p. 97, pi. 7 (at least in last part, but not Agarum Pylaii Bory.).

Two well-marked forms have been noted as follows:

Alaria tenuifolia f. typica Setchell

Stipe of moderate length, 10-15 cm. long; blade comparatively
narrow, 8-15 cm. in width and cuneate at the base ; midrib not
noticeably broad; sporophylls narrowly to broadly lanceolate with
cuneate base and short stalks.

On rocks and stone at low water mark. Amaknak Island, Bay of
Unalaska, Alaska,

Setchell, in Setchell and Gardner, Alg. of N.W. Amer., 1903, pp.
273-274, pi. 22.

192.5] Setchell-Gardner: MelanopJnjceae 639

Alaria tenuifolia f. amplior S. and G.

Stipe varying from 10-90 cm. in length, usually stouter than in
the last and usually as much flattened ; blade ample, 20-35 cm. wide,
10-15 dm. long, base generally, distinctly cordate ; sporophylls broadly
ovate to oblong and cordate, each with a conspicuous stalk, frequently
very large, 45 cm. long, 25 cm. wide, sorus more or less orbicular and
covering only the basal third or fourth.

Attached to piles and boulders near low water mark. Esquimalt,
B. C, and Roche Harbor, San Juan Island, Washington.

Setchell and Gardner, Alg. of N.W. Amer., 1903, p. 274.

Yendo (1919, p. 98) has referred this species to A. Pylaii Grev.
and he may be right although J. G. Agardh's plant, so far as his
adult type specimen (cf. Yendo, 1919, pi. 8, fig. 1) is concerned,
resembles our f. longipes more nearly than f. typica, which is the
type of A. tenuifolia. J. G. Agardh's Alaria Pylaii has been the
basis of all discussion of this species and is assumed by Yendo to be
the true A. Pylaii. It is not the Agarum Pylaii Bory, however, nor in
any wise resembling it, but is possibly the Laminaria remotifolia De la
Pyl. (L. Despreauxii of J. G. Agardh's herbarium is a different and
distinct species. AV. A. S.) We have for some time been leaning toward
the opinion that the f. longipes with the truncate base to the blade
might be distinct from the typical form of A. tenidfolia<. It is cer-
tainly a very distinct form (or perhaps rather variety) if this char-
acteristic is constant (as it is in all our specimens). It is important
to note that J. G. Agardh's plant (no. 2088 of his herbarium), repre-
senting his and the prevalent idea of Alaria Pylaii, is labeled "Al.
Despreauxii Bory. L. Pylaii de la Pyl, viz., Bory (musaefolia v.
remotifolia Del,), Groenland, Sukkertoppen, S. Berggren," and that
this is the plant figured by Yendo (1919, pi. 8, fig. 1). A plant in
J. G. Agardh's herbarium (no. 2090), assigned also by him to Alaria
Pylaii, but very different from that just discussed, is labeled ''Lami-
naria musaefolia Bory, Terre Neuve, Mr. Despreaux, 1828. ded,
Guillermin." This plant is entirely different from the Alaria Pylaii
J. Ag., as understood, as well as entirely different from the Agarum
Pylaii Bory and probably also different from the Laminaria musaefolia
De la Pyl.

These various considerations lead us to retain the name Alaria
tenuifolia for our plant and to retain both forms, typica. and longipes,
under it for the present at least. What we consider to be the true
Alaria Pylaii (Bory) Grev. will be discussed below.

640 University of California Publications in Botany [Vol. 8

4. Alaria marginata Post, and Rupr.
Plate 66

Stipe moderately short, 2-5 cm. long, 3-5 mm. diam., nearly terete ;
rliachis flattened, up to two times as thick as the stipe, merging grad-
ually into the midrib ; blade 2.5-3 m. long, 15-20 cm. wide, nearly
linear, somewhat tapering above with abruptly cuneate base, produc-
ing numerous cryptostomata with abundant exserted hairs ; midrib
variable in width, from 5 mm. in some individuals up to 22 mm. in
others, merging abruptly into the blade ; sporophylls ovate, lanceolate
or elliptical, rounded above, 10-20 em. long, 2-3 cm. wide, coriaceous,
rigid, usually plane, 24-40 in number, sori in some individuals
covering both sides completely except a narrow margin, in others only
the basal half or third.

Growing on exposed rocks or in sheltered coves in the middle and
lower littoral belts. Fairly abundant along the central California
coast, possibly also in Puget Sound.

Postels and Ruprecht, Illus. Alg., 1840, p. 11 ; Setchell, Kelps of
the U. S. and Alaska, 1912a, p. 162, Notes on Kelps, 1896, p. 41, Critical
Notes on Laminariaceae, 1908a, p. 9 ; Yendo, Monogr., 1919, p. 93, pi. 6,
figs. 1-4. A. curtipes Saunders, A New Species of Alaria, Minn. Bot.
Studies, 1901a, p. 561, pi. 33; Tilden, American Algae (Exsicc), no.
521 (at least in part). Alaria. cordata Tilden, Amer. Alg. (Exsicc),
no. 241 {fide Yendo). A. lanceolatm ( ?) Collins, Holden and Setchell,
Phyc. Bor.-Amer. (Exsicc), 1901, no. XLIV (not of Kjellman).

The first notice of this species of Alaria appeared in 1840, in the
above mentioned publication of Postels and Ruprecht. Later Ruprecht
(Tange, 1851, p. 355) referred to A. marginata, as being rare, and
known to him only from Fort Ross on the California coast. One
of us (Setchell) has examined a good specimen of Alaria in the Herb.
Acad. Sci. Petrograd, labeled A. marginata "Unalaska — Wosnes-
senski," which is like our California species which we have placed
under A. marginata Post, and Rupr. No later discovery of this species
has been made in the original locality, Unalaska, although careful
search has been made for it by one of us (Setchell), and it seems quite
probable that the Wosnessenski plant may have come from Fort Ross,
California. Hence we take this plant as the type of A. marginuta
Post, and Rupr. and Fort Ross as the type locality, and place all of
the California plants, except A. nana Schrader, under that name at

1925] SetcheU-Gardner : Melanophyceae 641

present. At first we were inclined to place the broad Californian plant
under A. praelonga Kjellman, being led thereto partially by the
opinion of Kjellman, who stated in a letter that one of our plants
greatly resembled his Alariai praelonga but differed in the shape of
the blade. The blade in our plants is, at times, much nearer in shape
to that of A. praelonga than are the sporophylls.

Alaria marginata Post, and Rupr., as shown by the type, has a
short stipe, crowded with broad but rather short sporophylls, and a
long broad blade, long attenuate at the base. The midrib seems to vary
from narrow to very broad.

5. Alaria Pylaii (Bory) Grev.

Stipe 2-4: cm. long, terete ; rhaehis flattened, about twice the
diameter of the stipe, passing directly into the midrib ; blade short
and broad, with broadly cuneate or truncate-cordate base, 2.5-3.5 dm.
long, 1-2 dm. wide, thin, membranaceous ; midrib 5-8 mm. wide ; sporo-
phylls 16-24 in number, 8-10 cm. long, 2-4 cm. wide, obovate, with
slender pedicels.

Growing on stones in the upper sublittoral belt, in the vicinity of
Prince William Sound and Kadiak Islands, Alaska.

Greville, Alg. Brit. Syn., 1830, p. xxxix ; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 272; Setchell, Kelps of the U. S. and
Alaska, 1912fl., p. 162. Agarum Pylaii Bory, in Diet, class, d'hist. nat.,
vol. 9, 1826, p. 194. (Neither Alaria Pylaii J. Ag. nor A. Pylaii Yendo.)

The type locality of this species is Newfoundland and the type
specimen is in the Herbarium of the Museum of Paris, and is labeled,
Laminaria Pylaii. It was collected by De la Pylaie in 1817. The
specimen is evidently an old one, more or less battered, and has some-
what broader sporophylls than the measurements given above, other-
wise our specimens from Orca agree fairly well with the type and
until more data can be obtained from field study we are placing our
plants somewhat doubtfully under this species.

The type of Agarum Pylaii Bory in Herb. Bory shows a plant with
a short stipe, thickly placed, short but broad sporophylls, and a
broad blade, broad to even cordate at the base. Our specimens agree
sufficiently to be considered a geographical variety of this species.

642 TJniversiiy of California Publications in Botany [Vol. 8

6. Alaria dolichorhachis Kjellm.

Stipe 2-7 cm. long, terete ; rhachis 8-20 cm. long, merging abruptly
into the costa ; blade short and narrow, 3-7 dm. up to 1 m. long, 4-7
cm. wide, often splitting to the midrib into narrow segments, even
wearing completely away, lanceolate, with narrowly cuneate base,
undulate ; midrib little prominent, 6-10 mm. wide, elliptical in cross-
section ; sporophylls 30-50 in adult plants, narrowly linear to spatu-
late, more or less undulate, or spirally twisted, 14—25 cm. long, 6-9
mm. wide.

Growing on rocks in the upper sublittoral belt. Known only from
a single locality on our coast. Collected by Charles H. Townsend in
the service of the U. S. Fish Commission at Agattu Island, Alaska,
1894.

Kjellman, Algae Arctic Sea, 1883, pp. 217-220, pis. 20, 21, 25, figs.
11-18; Setchell and Gardner, Alg. N.W. Amer., 1913, p. 272; Setchell,
Kelps of the U. S. and Alaska, 1912a, p. 162. Alaria crispa Kjell-
man, Om Beringh. Algfl., 1889, p. 37, pi. 3, figs. 5-7 ; Yendo, Monogr.,
1919, p. 89, pi. 5. Alaria taeniata Setchell, Kelps of the U. S. and
Alaska, 1912a, p. 162 (not Kjellm.) (not A. dolicho-rhachis Collins,
Holden and Setchell, Phyc. Bor.-Amer., 1901, no. XLI).

We have only one collection (of several plants, however), which
have the exceedingly long and narrow sporophylls demanded by
Kjellman 's figure (loc. cit.). Kjellman 's type, however, does not
show these as does his figure and looks more like Kjellman 's A. crispa.
We are retaining our plants under A. dolichorhachis, to which species
Kjellman thought them closely related yet from which he thought
they ought to be separated.

7. Alaria lanceolata Kjellm.

Stipe short, 3-6 cm. long, subterete; rhachis short, compressed,
thicker than the stipe ; blade lanceolate, short and narrow, up to 1.5 m.
long, 10 cm. wide, decurrent, somewhat undulate and plicate, often
much worn away ; midrib 4-8 mm. wide, oblong in cross-section ;
sporophylls 30-40, linear or linear-lanceolate, cuneate at base, rounded
or spatulate apex, short stipitate ; sori covering the basal portion.

Growing on rocks in the littoral and sublittoral belts. The type
locality is Bering Island in the Bering Sea. Kjellman reports it

1925] Setchell-Gardner: Melanophyceae 643

growing in the sublittoral belt, but one of us (Setchell) has observed
it growing in abundance high up in the littoral belt at Amaknak
Island, Alaska.

Kjellman, Om Beringh. Algfl., 1889, p. 39, pi. 5, fig. 1-3 ; Saunders,
Alg. Harriman Exp., 1901, p. 426, pi. 53(?) ; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 275 ; Setchell, Kelps of the U. S. and Alaska,
1912a, p. 162; Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. XLIV. Alaria laticosta Saunders, Alg. Harriman Exp.,
1901, p. 42.5, pi. 55 (not of Kjellm.). Alaria doUchorhachis Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. XLT.

The Alaria' lanceolata of Kjellman, judging from his illustration
and a photograph of his type, seems to be a plant with short stipe,
crowded long and narrow sporophylls, with the blade fairly broad
and narrowed to a moderately broad cuneate base. On the authority
of Kjellman, we remove to this species the specimens distributed as
A. dalichorhachis under no. XLI of the Phycotheca Boreali- Americana.
The only difference noted by Kjellman in the case of these plants is
that the blade is not so dark as in his type specimens. We also feel
inclined to refer here the plant figured by Saunders {loc. cit.) as
Alaria laticosta, w-hich has, however, a much broader midrib. Yendo
refers Saunders' plant to A. niacroptera (Rupr.) Yendo, but the
figures of that species as given by Yendo show a plant with different
sporophylls (too broad) and wath the base of the blade long and
narrowly attenuated. The Alaria:. lanceolata Saunders {loc. cit.) has
shorter sporophylls than is characteristic for this species.

8. Alaria valida Kjellman and Setchell

Stipe short, 1-5 cm. long, cylindrical, at first slender, but at length
stout and showing rings of growth, reaching a diameter of 13 mm.,
showing also a well developed cork layer ; rhachis at first short, becom-
ing very much elongated, up to 15-20 cm., somewhat flattened, obtuse
on the edges, stout like the stipe ; blade up to 3-4 m. long, 15-25 cm.
wide, tapering very gradually to the base, midrib 16-22 mm. wide,
little prominent and abruptly narrowing toward each end in cross-
section ; sporophylls linear lanceolate, 15-30 cm. long, 3-6 cm. wide,
with margins undulate and all except the narrow margins and a small
portion of the tip occupied by the sorus.

In the sublittoral belt. From Unga, Alaska, to Puget Sound,
Washington.

644 University of California Puhlications in Botany [Vol. 8

Kjellman and Setchell, in Setchell and Gardner, Alg., N. W. Amer.,
1903, p. 278, pi. 21. Alaria grandifolia J. Ag., Collins, Holden and
Setchell, Phyc. Bor.-Amer. (Exsicc.), no. CV (not J. Ag.).

Alaria valida f. longipes S. & G.

Stipe long, 8-20 cm. in length, not including any of the rhachis;
otherwise as in the type.

With the typical form on the west coast of Whidbey Island,
Washington.

Setchell and Gardner, Alg. N.W. Amer., 1903, p. 279.

Alaria valida seems to be a distinct species, as was suggested by
Kjellman. This is also the opinion of Yendo (1919, p. 118). At one
time Setchell (1908(ri, p. 11) was inclined to unite it with A. grandifolia
J. Ag., but that species has the blade broadly cuneate, almost oblong
at the base. Yendo (1919, pp. 116, 118) states that Tilden's no. 521
(a specimen collected by De Alton Saunders on the shores of Monterey
Bay and presumably determined by him) is Alaria valida ICjellm. and
Setch. The specimen in our copy is Alaria marginata, as we under-
stand it, and we have not seen anything referable to A. valida on the
California coast.

9. Alaria fistulosa Post, and Rupr.

Holdfast of numerous, irregularly branched hapteres ; stipe 20-25
cm. long, 8-12 mm. diam., without mucilage ducts, terete below, much
flattened in the transition region, where the numerous sporophylls are
developed, passing imperceptibly into the blade, perennial; blade up
to 25 m. long and up to 9 dm. broad, cuneate at the base, with well
defined midrib 2-3 cm. broad, which is hollow and inflated at irregular
intervals, the inflations being distinctly separate; mucilage ducts
abundant; color very dark olive brown; cryptostomata absent.

Growing often in great profusion on rock ledges and boulders in
the upper sublittoral belt. From Augustine Bay, Dall Island, to
the Bering Sea in our waters and extending to the Kurile Islands and
Japan on the Asiatic coast.

Postels and Ruprecht, lUus. Alg., 1840, p. 11, pi. 16; Saunders,
Alg. Harriman Exp., 1901, p. 246, pi. 57 ; Setchell and Gardner, Alg.
N.W. Amer., 1903, p. 275 ; Setchell, Kelps of the U. S. and Alaska,
1912a, p. 163 ; Kibbe, Structure of Alaria fistulosa, 1915, pp. 43-57, pis.
7-9 ; Yendo, Monogr. Alaria, 1919, p. 76, pi. 1.

Two forms have been segregated :

1925] Setchell-Gardner: Melanopliyceae 645

Alaria fistulosa f. stenophylla Setchell

Blade narrow, 10-25 m. long, usually not over 30 cm. wide ; midrib
narrow ; sporophylls usually short and more or less decidedly obovate.

Growing in belts just off shore and in isolated groups in quiet
waters, 5-10 fathoms. Ranging from the latitude of the Pribilof
Islands in the Bering Sea to southeastern Alaska.

Setchell, in Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. XLIII; Setchell and Gardner, Alg. N.W, Amer., 1903,
p. 276.

Alaria fistulosa f. platyphylla Setchell

Blade usually 6-9 dm. wide, up to 25 m. long, with broad, much
inflated midrib ; sporophylls numerous, up to 200, long and narrow,
30-50 cm. long, 3-6 em. wide.

Growing in deep water, usually found floating or cast ashore. Same
range as preceding.

Setchell, in Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. XLII ; Setchell and Gardner, Alg. N.W. Amer., 1903,
p. 277.

The fistulose midrib distinguishes this large, or at least long species,
which varies much in breadth of blade and size and number of sporo-
phylls. The two forms described above are fairly easy to segregate
but are probably ecological. This kelp grows in deep water and forms
beds in fairly deep water, its northern area comparable with that of
Nereocystis, with which it may be associated.

TRIBE 2. ECKLONIEAE setchell

Members of the Alariaceae having the outgrowths from the transi-
tion place maturing on the blade ; simple, or once or twice furcate
through wearing away of the central portion of the blade.

Setchell, Kelps of the U. S. and Alaska, 1912a, p. 163.

The genus Ecklania is not represented on our coast, but Eisenia,
whose early stages are like those of Ecklayiia, occurs on the coast of
southern California and in Japan. EckJonia is represented in the
Southern Hemisphere (Australia, New Zealand, Cape Good Hope,
and even up to the Canary Islands, as well as in Japan).

646 JJniversiiij of California Publications in Botany [Vol. 8

57. Eisenia Aresch.

Holdfast of diehotomously branched hapteres ; stipe elongating
and persistent, bifurcate above, the two false branches being the
thickened lower margins of the original and subsequently eroded blade.
A small partial blade persists at the outer extremity of each false stipe
throughout the life of the plant, giving rise to numerous sporophylls
along the lower outer margin ; perennial.

Areschoug, in Bot. Not., 1876, no. 3, p. 69, Observ. Phycol., part 5,
1884, p. 7.

Eisenia is a distinct genus of two known species ; one, E. arhorea,
the type of the species, inhabiting the waters of southern California,
and the other, E. hicyclis (Kjellm.) Setchell, inhabiting the Japanese
waters.

Eisenia arborea Aresch.

Hapteres arising from very close to the base of the stipe, much
branched, the terminal branchlets fine and contorted ; stipe nearlj^
terete at the base, much flattened above, up to 1 m. long, tough and
rigid, containing mucilage ducts ; blade in young plants entire, some-
what ovate, soon giving rise to short outgrowths, the pinnules, in the
transition region, later eroding from the apex to the transition or
meristematic region, which now becomes divided longitudinally estab-
lishing two meristems; these two now moving forward, giving rise to
the two twisted stipe-like portions, each bearing a short terminal
thickened blade, in turn bearing numerous pinnules or sporophylls
along the lower outer margins ; a third meristem remaining active at
the summit of the true stipe and elongating ; each blade bearing 30-50
sporophylls ; the sori in large irregularly shaped areas.

Growing on rocks in the upper sublittoral and lower littoral belts.
Southern California, south of Redondo.

Areschoug, in Bot. Not., 1878, no. 3, p. 69, Observ. Phycol., part 5,
1884, p. 7; Setchell, Kelps of the U. S. and Alaska, 1912a;, p. 164;
Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. IX.
Ecklonia radiata Areschoug, 1884, p. 13, in part (not Fucus radiatus
Turner).

The plants reported by Saunders from Puget Sound, Washington,
and Wrangell, Alaska, proved to be large specimens of Laminaria
Andersonia with blades worn away. Areschoug reports it "in sinu
propre San Francisco" on the authority of Dr. G. Eisen, in honor of

192-5] SetcJiell-Gardner : Melanophyceae 647

whom the genns was named, but the type locality (fide Dr. Eisen) is
Santa Catalina Island. Areschoug also reports it from Santa Cruz,
California, on the authority of Dr. C. L. Anderson, who has told us
that his plant was found floating. We have no knowledge which seems
to us to be authentic of its occurrence, growing in position, north of
Redondo, California. We have no data as to its southern limit of
distribution, but we presume that it extends to some distance south
along the coast of Lower California.

TRIBE 3. EGREGIEAE setchell

Members of the Alariaceae having irregular branching and out-
growth of sporophylls on both stipe and blade.

Setchell, Kelps of the U. S. and Alaska, 1912a, p. 164.

There is a single genus of this tribe, with two species, both con-
fined to the western coast of North America.

58. Eg^egia Aresch.

Holdfast a very densely compact mass of repeatedly branched hap-
teres; stipe irregularly branched near the base, terete below, each
branch soon becoming ligulate forming a rhachis and bearing out-
growths on each margin nearly throughout its length ; some outgrowths
metamorphosing into conspicuous stipitate bladders, some becoming
ligulate and sterile, and some remaining small and bearing sori of
zoosporangia on both surfaces.

Areschoug, in Bot. Not., 1878, no. 3, p. 66, Observ. Phycol., part 5,
1884, p. 3.

The genus Egregia was established by Areschoug (loc. cit.) to
receive the Fucus Menziesii of Turner. Menzies collected it at
''Nootka, Trinidad and Monterey."

IVEY TO THE SPECIES

1. Sterile, lateral outgrowths linear spatulate 1. E. Menziesii (p. 647)

1. Sterile, lateral outgrowths ligulate • 2. E. laevigata (p. 648)

Egregia Menziesii (Turn.) Aresch.

Compact mass of hapteres arising close to the base of the stipe,
by repeated branching, often growing up around the base of the
stipe; stipe smooth, nearly terete at the base for 8-15 cm., branched
several times, soon becoming ligulate, forming a rhachis ; rhachis

648 TJfiiversity of California Publications in Botany [Vol. 8

5-8 m. long, 2.5-3.5 em. broad, densely covered on both sides with short
blunt tubercles, and along each margin with outgrowths bearing the
ellipsoidal bladders and smooth ovate or spatulate sporophylls among
the sterile leaflike structures ; blade in the young specimens coarsely
rugose, conspicuous, but soon lost.

Growing in the upper sublittoral and lower littoral belts. From
the southern end of Vancouver Island to Point Conception, California.

Areschoug, in Bot. Not., 1878, no. 3, p. 66 ; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 271; Setchell, Kelps of the U. S. and
Alaska, 1912a,, p. 164; Muenscher, Key to the Phaeophyceae, 1917,
p. 274; Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), nos.
1741 and XCII ; Tilden, Amer. Alg. (Exsicc), no. 236. Macrocystis
Menziesii Agardh, Sp. Alg., 1820, p. 49. Macrocystis obtusa Harvey,
Botany Beechey's Voyage, 1833, p. 163 (cf. Harvey, 1852, p. 62).
Phyllospora Menziesii Ruprecht, Neue Pflanzen, 1852, p. 70, pi. 4.
Fucus Menziesii Turner, Hist. Fuc, 1808, p. 58, pi. 27.

This plant is often very long, with several branches arising mostly
near the base as outgrowths which, unlike those of the sporophyll type,
have lateral blades, transition places, and lateral outgrowths in turn,
developing like the main stipe and rhachis. The bladders are frequent
but develop in no exact order.

Egregia laevigata Setchell

Holdfast and stipe much as in E. Menziesii, but the true stipe and
the rhachis smooth on both sides, the marginal outgrowths very
variable in different individuals and at different ages of the same
individual, some profusely dissected and filiform, even capillary, others
large, linear and entire, up to 15 cm. long; the bladders varying in
shape from spherical to very narrowly ellipsoidal.

Growing on rocks in the lower littoral and upper sublittoral belts.
Extending from the vicinity of Point Conception, California, to some
unknown locality on the west coast of Lower California, at least as
far down as Ensenada.

Setchell, Notes on Kelps, 1896, p. 44, Kelps of the U. S. and Alaska,
1912a, p. 164; Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), nos. 420 and XII; Tilden, Amer. Alg. (Exsicc), no. 340.
Egregia menziesii Farlow, Anderson and Eaton, Alg. Exsicc. Amer.-
Bor., no. 111.

1925] Setchell-Gardner : MeJanophyceae 649

Egregia laevigata f. borealis Setchell

Form with entire sporophylls and with blade and ligulate portions
roughened slightly witli pointed papillae.

Growing in the upper sublittoral belt. Carmel Bay, California.

Setchell, in Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. XL.

This plant, which in its well developed form with capillary dis-
sected leaflets, is known as the "Feather Boa," is usually readily to
be distinguished from its sister species. Toward its northern limit,
it is not common, occurring as f. horealis along with E. Menziesii and
presenting forms with few to fairly numerous papillate roughenings,
appearing as if of possible hybrid origin.

SERIES 2. APLANOSPOREAE ser. nov.

Fronds of moderate size, filamentous and monosiphonous (Chori-
stocarpaceae) or polysiphonous (Tilopteridaceae) or complanate
(Dictyotaeeae) ; reproduction by both sexual and asexual methods;
sexual reproduction isogamous ? or heterogamous ; non-sexual reproduc-
tion brought about by the formation of aplanospores, one or more,
usually four, in a sporangium and, in some genera, by biciliate zoo-
spores; alternation of generations known in some species (e.g., Dictyota
dichotoma in which reduction takes place at the time of aplanospore
formation).

The Aplanosporeae consists of two orders, viz., the Tilopteridales
and the Dictyotales. The series is characterized by certain of the
members of both orders bearing non-motile, non-sexual spores which
may be single (Tilopteridales) although possessing (in some cases at
least ) four nuclei at maturity or may be formed in fours from a single
mother cell (Dictyotales). The Tilopteridales resemble the Ectocar-
pales in vegetative structure and may have unilocular zoosporangia
giving rise to biciliated zoospores. There are no representatives of
this order on our coasts.

650 TJniversity of Calif ornia Publications in Botany \yo-i.. 8

Order 7. DICTYOTALES kjellm.

Fronds of moderate size, complanate, attached by a more or less
extensively stupose base, with the hairs often extending for some dis-
tance up the fronds; reproduction both asexual and sexual; the non-
sexual spores (aplanospores) usually borne in groups of four, but
rarely more, in transformed surface cells projecting singly or in
groups beyond the surface; the gametes heterogamous, the female,
borne singly in a gametangium, large and non-motile, the male, borne
many in a plurilocular gametangium, small and motile by a single ( ?)
lateral flagellum ; the male and female gametangia usually in dense
sori, and always projecting beyond the surface; hairs numerous, in
groups; paraphyses present in some forms.

Kjellman, in Engler und Prantl., Die natiirl. Pflanzenfam., 1 Teil,
2 Abt., 1896, p. 291.

The order Dictyotales is a distinct, well marked group, but with
uncertain affinities. The possession of a brown pigment in addition to
chlorophyll is a prominent character of the Melanophyceae. The
development of aplanospores in groups of four (tetraspores) is a
character common in the Rhodophyceae. The male gamete with but
a single cilium is not present elsewhere in either group. The brown
pigment and heterogamous method of reproduction seem sufficient to
relate them to the Melanophyceae and probably closely to the Fucales,
where we are placing the group. It has no zoosporangia or zoospores
represented in any of its members.

FAMILY 23. DICTYOTACEAE harvey (lim. mut.)

Characters of the order which contains only the single family.

Marvey, Ner. Bor.-Amer., vol. 1, 1852, p. 99.

Harvey included all the complex Melanophyceae ''whose spores are
superficial and disposed in definite lines or sori. ' ' The Dictyotaceous
genera as now understood were included, but also some others such as
Punctaria, Soranthera, Stilophora, Dictyosiphon, Asperococcus, and
HydroclatJiriis. Harvey, himself, realized that in external habit and
even in internal structure, the plants he referred to his order, as he
termed it, exhibited "considerable variety." He also stated that "the
fructification exhibits considerable diversity of aspect. ' '

1925] Setchell-Gardner: MelanopJiyceae 651

Key to the Genera

1. Growth in length by division of an apical cell 59. Dictyota (p. 651)

1. Growth in length by the division of many marginal cells 2

2. P>ond with a distinct midrib 60. Neurocarpus (p. 655)

2. Frond without a midrib 3

3. Reproductive organs on both sides of the frond 5

3. Reproductive organs on only one side of the frond 4

4. The ternunal margin inrolled 64. Padina (p. 661)

4. The terminal margin not inrolled 62. Chlanidophora (p. 658)

5. Sori usually in more or less distinct concentric lines partially embedded in. the

frond, even at maturity 61. Taonia (p. 656)

5. Sori seldom concentric, entirely superficial at maturity 63. Zonaria (p. 659)

59. Dictyota Lamour.

Frond plane, membranaceous, ecostate, dichotomoiis below, some-
what irregularly cleft and flabellate above, arising from a stupose base,
and consisting of two layers of cells, an inner layer of large, colorless,
longitudinally elongated, prismatic cells, and a cortex of small, asvsimi-
lating cells, arranged longitudinally in rows ; reproduction sexual, by
antheridia developed in small groups producing antherozoids with a
single terminal cilium, and by oogonia in groups, each oogonium pro-
ducing a single egg which is extruded and fertilized in the water ; and
asexual by aplanospores, 4 in a sporangium, cruciately divided,
developed from the surface cells, scattered, sparse.

Lamouroux, Nouv. Bull. Soc. Philom., vol. 1, 1809 (May), p. 331,
and in Desv., Jour, de Bot., vol. 2, 1809a, p. 38.

The older genus Dictyota was distinct among the Dictyotaceae
whose fronds, arising from a single apical cell, were flattened or com-
pressed and more or less dichotomously branched. The latest divisions
of J. G. Agardh, while attractive and representing certain tendencies
toward increase in complexity, yet present certain difficulties in the
line of cleavage. Dilophus, for example, presents species in typical
form with two or four layers of cells throughout the older portions of
the frond, but what is to be done with those species which are two-
layered only on the extreme margins, those which are two-layered
more widely on the margins, and those which are only one-layered
occasionally? All of our species are of the first or second types and
we feel better satisfied to refer them all to the genus Dictyota. Con-
cerning the other segregate of J. G. Agardh (1894) from the Dictyota
of earlier accounts, we are in no position to speak.

652 University of California Publications in Botany [Vol. 8

IVEY TO THE SpECIES

1. Margin of the frond profusely toothed and crenulate 5. D. crenulata (p. 655)

1. Margin of the frond smooth 2

2. Fronds large, 1.5-2.5 dm. long, 250-500/i thick, axils rounded 5

2. Fronds smaller, up to 15 cm. long, axils more or less acute 3

3. Fronds 7-9 cm. long, 135-160^ thick, axils wide 4. D. Vivesii (p. 654)

3. Fronds 8-15 cm. long, 80-135/i thick, axils narrow, acute 4

4. Oogonia single 6. D. hesperia (p. 655)

4. Oogonia collected into sori 3. D. Johnstonii (p. 653)

5. Fronds dark brown, coriaceous, tips rounded 1. D. Binghamiae (p. 652)

5. Fronds pale yellowish brown, deUcate, tips truncate 2. D. flabellata (p. 652)

1. Dictyota Binghamiae J. Ag.

Plate 34, figs. 1, 2

Fronds 10-18 cm. (up to 25 cm.) high, with stupose base of fine
brown hairs, closely appressed, extending for some distance up the
frond, coriaceous below, up to 500ja thick, thin and membranaceous
above, branches strict, angles rounded, terminal lobes rounded ; color
dark below, brown above.

Growing in pools in the middle and lower littoral belts. Southern
California and northern Lower California (Ensenada).

J. G. Agardh, Anal. Alg. Cont. I, 1894, p. 72 ; Collins, Holden and
Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1392. Dictyota Kunthii
Farlow, Kept. U. S. Fish Comm., for 1875, 1876, p. 705; Farlow,
Anderson and Eaton, Alg. Exsicc. Amer.-Bor., no. 93. Glossophora
Kunthii Collins, Holden and Setchell, loc. cit., no. 85.

This is a large and broad, coarse species with even margins and
rounded axils. In color it is usually darker brown than the next
species, somewhat thicker and more coriaceous, more truly dicho-
tomous above, and with the upper lobes longer and more rounded.

2. Dictyota flabellata (Collins) S. and G.

Plate 34, fig. 3; plate 35, fig. 7, and plate 36, figs. 13-17

Fronds plane, membranaceous, slightly stupose at the base, dicho-
tomous, or in mature plants certain segments growing faster than
others, presenting the appearance of being a flexuose rhachis with alter-
nate branching, 1-2 dm. high ; segments cuneate, widening upward to
each forking, terminal lobes and sinuses rounded; width variable, 2
mm. to 2 cm. wide ; color yellowish brown, darker toward the base ;
medulla consisting of a single layer of nearly cuboidal, colorless cells.

1925] Setchell-Gardner: Melanophyceae 653

except along the thickened margins of the old fronds where it is
double ; cortex of a single layer of colored cells, except in older, thick-
ened margins where there are two layers, arranged in longitudinal
parallel series ; oogonia, antheridia, and tetrasporangia all growing on
different individuals ; oogonia scattered in small sori ; antheridia and
tetrasporangia in oval or oblong sori ; reproductive cells on both
surfaces of the frond.

Growing on boulders in the lower littoral belt. Southern Cali-
fornia.

Setchell and Gardner, Phyc. Cont. VII, 1924, p. 12. Dilophus
flahellatus Collins, in Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. 834. Glossoplwra Kunthii Tilden, Amer. Alg. (Exsicc),
no. 334. Dictyota dichotoma Tilden, Amer. Alg., no. 335 (not Lamour.).

This seems to be a distinct species, thus far confined to southern
California, but with unattached specimens collected at Monterey, Cali-
fornia, and Tracyton, Washington, probably sporadic or transported
to the latter. The species has the least doubling of the marginal cells
of any in our territory and is closely related to D. Binghamiae. It is
thinner, lighter colored, more pseudodichotomous, with the uppermost
almost pinnately arranged lobes short and bent. It is not difficult to
separate the two species when in company, either living or in her-
barium specimens, but we do find it difficult to express definitely the
exact points of distinction between them.

3. Dictyota Johnstonii S. and G.

Fronds 12-16 cm. high, 5-8 mm. wide, 125-135/x thick, composed
of a single layer of large medullary cells surrounded by a single sur-
face layer of small cells except in the lower part, along the margin the
medullary and surface cells becoming doubled, slightly stupose at the
base, pinnate, dichotomously branched, angles acute to somewhat
rounded, branches strict, margins smooth, color dark brown, black on
drying ; oogonia aggregated into elliptical or elongated areas, 115-125/a
long, 80-90/i, wide ; tetrasporangia and antheridia unknown.

Growing on rocks in the upper sublittoral belt. San Marcos Island,
Gulf of California.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 730, pi. 18,
figs. 54-56, and plate 39.

Dictyota Johnstonii appears to belong to the subgenus or section
of the genus, Strigocarpus J. Agardh (Anal. Algol. Cont. I, 1894,

654 University of Calif ornia Publications in Botany [Vol.8

p. 73) and related to D. pi^matifida Kuetzing (Tab. Phyc, vol. 9, 1859,
p. 16, pi. 39, fig. 1), to Z). Pappeana. Kuetzing (loc cit,, pi. 38, fig. 2),
and to D. liturata Kuetzing (loc. cit., fig. 1). The cross-section in
the central and lower parts of the frond has a structure similar to
that shown by Okamura (1913, p. 33, pi. 109, figs. 3, 7) for D.
mnrginata. In Z>. Johnstonii the margins are thickened by divisions
of the cells of both the medulla and the surface, while in D. marginata
increase in thickness is brought about by division of the medullary
cells only.

4. Dictyota Vivesii Howe

Fronds densely caespitose, stupose at the base, 7-9 cm. high, 135-
160/A thick (240ja at the base), collapsed and thin on drying, somewhat
regularly 3-6 times dichotomous below, the branches then rather
closely 3 or 4 times subflabellately or subpinnately dichotomous,
sinuses mostly rather acute, margins entire or slightly undulate ; main
segments oblong or obcuneate, 3-8 mm. broad, diminishing in length
and width upward, terminal segments 1-2 mm. wide; cortical cells
nearly uniform in size, 19-65ju, long, 11-27^ wide, interior cells much
larger and very thin-walled ; aplanospores forming small, scattered,
inconspicuous sori.

Collected by Senor G. V. Vives at La Paz, Lower California, in
February, 1911.

Howe, Phyc. Stud. V, 1911, p. 497, pi. 27.

Dictyota Vivesii seems to be a near relative to D. Bartayresiana
Lamour. from the West Indies. According to Howe (1911, p, 498) :
"It is more caespitose in habit of growth than D. Bartayresiana, more
stupose at the base, less regularly dichotomous towards the apices,
rather broader in its broadest parts and more conspicuously dwindling
in width as the ultimate segments are approached, the axils (the
upper at least) are more acute and the segments less patent or divari-
cate, the apices are less acute, and both the cortical and the interior
cells are for the most part narrower and the cortical cells overlaying
the septa and lumina of the interior cells show scarcely any of that
differentiation in form and translucency that led J. Agardh to
describe B. Bartayresiana as 'fenestrate.' "

1925] SetcheJl-Gardner : MeJanophyceae 655

5. Dictyota crenulata J. Ag.

Frond attached by a stupose base, decompositely dichotomous, with
wide sinuses; segments linear, margins crenulate-dentate, narrow
below, tongue-shaped above, areolae rectangular ; sori sparingly scat-
tered over the whole surface of the segments.

Keported from St. August in, on the west coast of Mexico, and from
La Paz, on the peninsula of Lower California.

J. Agardh, Nya Alg., 1847, p. 7 ; Setchell and Gardner, Mar. Alg.
Gulf Calif., 1924, p. 730, pi. 18, figs. 50, 51. Dictyota Bartayresiana
var. dcuticulata Kuetzing, Tab. Phyc, 1859, p. 8 (as to synonym only).

The type locality for Dictyota crenulata is St. Augustin, Mexico.
We referred a plant to this species {loc. cit.) collected at La Paz, which,
although sparse, seems to belong to J. G. Agardh 's species from
St. Augustin,

'»"

6. Dictyota hesperia S. and G.

Fronds linear, repeatedly branched, 8-10 cm. high, 2-4 mm. wide,
80-120jU, thick, more or less finely stupose at the base, dichotomously,
or at times subdistichously branched, antheridia and oogonia distri-
buted over both surfaces on the same frond, oogonia single and
antheridia in small, circular groups ; tetrasporangia single or in small,
irregular groups.

Growing on rocks in the lower littoral and upper sublittoral belts.
San Marcos Island, Gulf of California.

Setchell and Gardner, Mar. Alg. Gulf. Calif., 1924, p. 731, pi. 18,
figs. 52, 53.

Dictyota hesperia' seems to belong to the subgenus or section of
the group designated as Pleiadophora by J. G. Agardh (1894, p. 69)
with close af^nity with D. sandvicensis Sond.

60. Neurocarpus Web. and Mohr

Fronds smooth, fairly firm, with distinct, percurrent midrib,
repeatedly dichotomo-flabellate, attached by a stupose base, composed
of two distinct tissues, a medulla of cuboidal to angular cells, many
layers deep at the midrib, diminishing to few at the margin, and a
single layer of small, color bearing cells on the surface; aplanospores
arranged in sori along: either side of the midrib ; oogonia and antheridia
scattered.

656 University of California Puhlications in Botany [Vol.8

Weber and Mohr, B,eitrage zur Naturkunde, vol. 1, 1805, p. 300;
Howe, Mar. Alg. Peru, 1914, p. 69, Dictyopteris Lamouroux, Observ.
Phys., 1809, p. 332. Halyseris Agardh, Sp. Alg., vol. 1, 1820, p. 141.

We have adopted the opinion of Howe as regards the proper
designation of this genus which is readily recognized among the
genera of our coast by its percurrent midrib, although our species has
somewhat the superficial aspect of a Zonaria.

Neurocarpus zonarioides (Farlow) Howe

Plate 34, fig. 4; plate 35, fig. 11; plate 36, fig. 21; plate 38, fig. 39, and plate 95

Fronds arising from a stupose base, 8-24 cm. high, densely and
irregularly dichotomous, ultimate dichotomies short, with prominent,
percurrent midrib, tomentose below, and laminae without lateral veins,
much incised and lacerate above, soon disappearing below ; color when
young, yellowish brown, dingy brown or almost black on drying ; ulti-
mate segments often subflabellate, short, somewhat divergent, with
obtuse or slightly retuse tips; aplanosporangia numerous, arranged in
sori parallel to the midrib on both sides of the frond, 75-100/x diam.

Growing on rocks in the lower littoral and upper sublittoral belts.
Known only from southern California and northern Lower California
(Ensenada).

M. A. Howe, Mar. Alg. Peru, 1914, p. 69; Setchell and Gardner,
Mar. Alg. Gulf of Calif., 1924, p. 728. Dictyopteris zonarioides
Farlow, in Erythea, vol. 7, 1899, p. 73 ; Collins, Holden and Setchell,
Phyc. Bor.-Amer. (Exsicc), no. 581. Haliseris polypodioides
McClatchie, Seedless plants of southern Calif., 1897, p. 354.

Farlow and Howe have made plain the characteristics of this
species. Its nearest rlative seems to be N. Cokeri Howe of the Peru-
^dan coast. For exact information, Howe's comparison {loc. cit.)
should be consulted.

61. Taenia J. Ag.

Fronds plane, ecostate, attached by a more or less stupose base,
flabellate, the cuneate segments more or less deeply divided and dis-
sected into narrow laciniae, composed of two distinct tissues, a medulla
of several layers (usually 4) of larger colorless cells, surrounded by a
single layer of cuboidal, color bearing cells, arranged mostly in longi-
tudinal series; aplanospores scattered or arranged vaguely in concen-
tric zones, usually abundant; oogonia and antheridia on different
individuals.

l92o] SetcheU-Gardner: MeJauophyccae 657

J. G. Agardh, Sp. Alg., vol. 1, 1848, p. 101.

The genus was founded by J. G. Agardh on Ulva Atomaria Wood-
ward (Linn. Trans., vol. 3, 1797, p. 53). V. Atamaria, was discovered
by Mr. Lilly Wigg on the beach at Yarmouth, England. Since its
"discovery, it has been associated with at least the following genera :
Dictyota, three different species, Zonuria Atomaria, Padina, two
species, Stypopodium Atomaria and Ulva serrata. While the genus
Taonia- is to be distinguished from Zonaria. by its usually more delicate
tissues, its absolute lack of a midrib {Zonaria^ being subcostate), and
by a greater tendency to have its reproductive bodies in concentric
lines, nevertheless there are difficulties in exactly diagnosing it. In all
the species of Zonaria, the reproductive bodies are, at least at maturity,
superficial, while in the species of Taonia., they are partially embedded
in the frond.

Taonia Lennebackerae Farlow

Plate 35, figs. 9, 10, and plate 96

Fronds arising from a slightly stupose base, 1-2 dm. (up to 3 dm.)
high, variable in width ; segments mostly broadly cuneate from near
the base of the fronds, unevenly divided at the outer end into several
lobes, often deeply lacerated ; color when young olive, dark brown in
older specimens ; cells of the interior 3-4 layers, angles considerably
rounded, slightly elongated longitudinally ; cells of the cortex mostly
arranged in longitudinal rows, 1.5-3 times as long as broad ; reproduc-
tive cells mostly scattered promiscuously and profusely on both sides
of the frond, at times vaguely in concentric lines.

Growing on rocks in the upper subittoral belt. Southern Cali-
fornia.

Farlow, in Farlow, Anderson and Eaton, Alg. Exsicc. Amer.-Bor.,
no. 160 (nomen midum) ; J. G. Agardh, Anal. Alg. Cont. I, 1894,
p. 30 (description) ; Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. 327; Tilden, Amer. Alg. (Exsicc), no. 337.

The type locality of this species is Santa Barbara, California. The
type specimens were contributed by Miss Lennebacker, of Santa
Barbara. The California species seems more delicate than the Euro-
pean, with never a sign of a stupose base and with the fruit dots
usually scattered, although at times in undulate and indistinct concen-
tric lines. According to J. G. Agardh, the cortical cells are shorter
than in other species of the genus (1.5-3 times longer as contrasted
with 3-4 times longer in surface view). Our plant is certainly less

658 University of California Publications in Botany [Vol. S

characteristic than the European T. Atomaria (Good, et Wood.) J.
Agardh, as to concentric lines of sori while, in habit, it seems to differ
decidedly from T. australasica (Kuetz.) J. Agardh, if we may rely on
Kuetzing's figure (1859, pi. 48, fig. 2, quoted by J. G. Agardh with
a query).

62. Chlanidophora J. Ag.

Fronds multifid decompound, terminal segments plane, flabellately
dichotomous, subcostate below, lateral laciniae acuminate often ending
in a terminal cell ; external cells rectangular in surface view regularly
placed, those of each surface (in transverse section) adherent in turn
and corresponding in position, forming a distromatic frond (there
being no internal cells) ; fertile cells superficial, sparse, obovate
globose ; antheridia situated on the same individuals, forming oblong-
linear superficial sori, parallel to the length of the fronds, of a double
series of cells, finally quaternate.

J. G. Agardh, Anal. Alg. Cont. I, 1894, p. 16 {Chlamdote in Key,
loc. cit., p. 6). Chlanidote De-Toni, Syll. Alg., vol. 3, 1895, p. 238.

We have paraphrased the generic description of J. G. Agardh, but
give below under our species the characters of our plant, which may
possibly be of a distinct genus. We have no specimens of Chlanido-
phora microphyUa (Harv.) J. Agardh for examination and, conse-
quently, must rely on the descriptions and on Harvey's (Ner. Austr.,
vol. 4, 1862, pi. 195) and Kuetzing's (Tab. Phyc, vol. 9, 1859, pi. 69,
fig. Ill, a^d) figures. In structure, the Australian plant is very close
to ours, yet ours seems to be more an aggregation of somewhat loosely
cohering filaments, while the type of the genus seems possibly more
tightly cohering and parenchymatous. We have altogether too imper-
fect a knowledge of the reproductive bodies in the case of either
species for a close comparison. Harvey 's plant was sterile. Kuetzing 's
plant was probably sterile. The description and figures of J. G.
Agardh are confusing.

Chlanidophora abyssicola S. and G.

Plate 50, figs. 1-7

Fronds 2-3 em. high, composed at maturity of numerous broadly
flabellate segments ; false stipe slender, in part polystromatic, com-
posed of the original narrow blade covered on both sides by numerous,
slender, closely appressed, multicellular hairs spreading out at the

19--5] SetcheJI-fjarchier: Mclanophijceae 659

base on the substratum, serving as attaching organs ; cells dividing
horizontally 1-3 cells back of the margin, making the main body of
the frond distromatic ; marginal layer of 1-3 cells monostromatic ; cells,
in surface view, quadrangular, 16-24ju, long, 10-12ju broad, marginal
cells 30-50ja long, containing numerous, small, spherical chromato-
phores ; sporangia pyriform to ellipsoidal, 32-3Sjji, long, 28-32/^ broad ;
aplanospores (?) 4-5ju, diani., numerous; paraphyses clavate, 4-7 cells
long.

Growing attached to shells of mollusks in 10-15 fathoms. Griffin
Bay, San Juan Island, Washington.

Setchell and Gardner, Phyc. Cont., A^T, 1924, p. 11.

63. Zonaria Ag. (lim. mut.)

Fronds in part decumbent or wholly erect, decidedly stupose at
the base, ecostate at first, the margins soon wearing away in the lower
part, thus becoming stipitate and subcostate by thickening, flabellately
divided ; growth in length by means of numerous cells at the ter-
minal edge; fronds composed of two tissues, a medullary layer of
several cells, and a single cortical layer, arranged more or less in longi-
tudinal rows, containing many chromatophores ; antheridia and aplano-
spores unknoAvn ; oogonia on one or both sides of the thallus, borne in
small sori, among paraphyses.

C. A. Agardh, Syn. Alg. Scand., 1817, p. xx (lim. mut.) ; J. G.
Agardh, in Linnaea, vol. 15, 1841, p. 444, Bidrag till Alg. Syst., I,
1873, p. 45 ; Anal. Alg. Cont. I, 1894, p. 12.

We have taken the genus Zonaria in the broader sense and includ-
ing certain of the segregated genera of J. G. Agardh, such as his
Gymnosorus, Homoeostrichus, and Stypopodium. This is the sense in
which Howe has used it in the Bahama Algae (1920, p. 594). While
Homoeostrichus has certain very distinct species, there seems to be
less distinct cleavage for certain others. Our single species seems
clearly of Zonaria' in this broader sense.

Nieuwland (1917, pp. 51, 52) has proposed the name ViUa.nia
instead of Zonaria (J. G. Agardh, 1872, p. 45) because of a most con-
vincingly evident misprint of "Zonaria" for "Zornia." Such an
attempted application even of the principle : ' ' once a synonym always
a synonym" seems a " reductio ad absurd urn " and especially when
applied by one seemingly in absolute ignorance of the genus affected,
other than as an abstract entity. Zonaria, however, dates back to

660 University of California Publications in Botany [Vol. 8

1801 (Draparnaud, Discours sur les moeurs des plantes, fide Steudel)
and was apparently applied to a Diseomyeete. Roussel also used
the name for a fungus in 1806 (fide Steudel). C. A. Agardh (1817,
p. XX ) first used the name for a genus of algae of far wider extent
than that in which it is used at present. J. G. Agardh has variously
applied the name, his final and rather too narrow limitation having
been published in 1894.

Zonaria Farlowii S. and G..

Plate 34, fig. 5; plate 36, fig. 20; plate 43, fig. 63, and plate 97

Fronds 8-12 cm. long, profusely and more or less flabellately
branched, terminal lobes flabellate, alae at times split into numerous,
narrow and pointed segments, lower part forming a much thickened
stipe, becoming decidedly stupose ; marginal growing cells very large,
densely filled with cell contents ; medulla composed of 6-9 layers of
cells parallelopiped in shape, having scattered chromatophores ; aplano-
spores borne in sori, irregular in shape and size, scattered promiscu-
ously on both sides of the frond, formed under the cuticle among
numerous, multicellular para.physes, growth of aplanospores and para-
physes finally rupturing the cuticle allowing the escape of the spores ;
paraphyses clavate, composed of 5-7 cells ; hairs borne in small, inde-
pendent groups or arranged in transverse bands.

Growing on rocks in the upper sublittoral belt, and in pools in the
lower littoral belt. Southern California (Santa Barbara to San
Diego).

Setchell and Gardner, Phyc. Cont., VII, 1924, p. 11. Zonaria
Tournefortid Farlow, in Farlow, Anderson and Eaton, Alg. Exsicc.
Amer.-Bor., 1878, no. 91 ; Collins, Holden and Setchell, Phyc. Bor.-
Amer. (Exsicc), no. 86; Tilden, Amer. Alg. (Exsicc), no. 336 (not
Fucus Tournefortii Lamouroux). Zonaria flava Harvey, Ner. Bor,-
Amer., part 3, Suppl., 1858, p. 123 (not Fucus fiavws Clem.).

The Zonaria abundant on the southern coast of California is near
to Z. Tournefortii (Lamour.) Farlow (i.e., Fucus Tournefortii
Lamour.) or Z. flava (Clem.) Agardh of Europe, but that is a coarser
plant than ours with larger cells, and ( judging from Kuetzing 's figures,
1859, pi. 65, I, fig. &) the sori lack paraphyses as do those of Z. Turner-
iana J. Agardh of Australia (fide specim. auth.). It is a more slender
species than Z. sowo-h's (Lamour.) Howe {Stypopodium lohatum (Ag.)
Kuetzing). Z. Sinclairii (Harv.) J. Agardh is narrow and of Homoeo-

1925] Setchell-Gardner : Melariophyceae 661

strichus structure. Z. Biesingiana J. Agardh is more prostrate and
coarser. The Japanese plant illustrated under this name by Okamura
(1907, pi. 4, figs. 1-10) has paraphyses with peculiar swollen terminal
cells. Zonaria interrupta (Lamour.) J. Agardh, from the Cape of
Good Hope, has very narrow divisions and has paraphyses with short,
swollen cells above, as in ours, but the terminal segment of each divi-
sion of the frond is peculiarly truncate and enlarged above (cf.
Phycopteris cuneata Kuetz., Tab. Phyc, vol. 9, 1859, pi. 67, fig. II).
The structure of the frond is also very different, as may be seen from
Kuetzing's figure quoted above.

61. Padina Adans.

Fronds plane, ecostate, flabellate, entire or branched, differentiated
into two kinds of tissues, a single surface layer on either side consist-
ing of color bearing rectangular cells, and a central tissue, medulla, of
several layers of somewhat elongated cells with few chromatophores ;
growth in length by divisions of many marginal cells ; margin scrolled,
or inrolled ; reproduction sexual, heterogamous, and asexual by aplano-
spores ; the reproductive organs at times developed in transverse zones
on the upper parts of the fronds and the antheridia in longitudinal
sori, on the same frond as the oogonia.

Adanson, Fam. II, 1763, p. 13 (fide De-Toni).

The " Peacock 's-Tail" algae are well-known tropical species whose
specific limits require careful study and consideration. The genus is
readily distinguished by its involved meristematic margin and by its
having the sori only on one side (the upper) of the frond.

Padina Durvillaei Bory
Plate 93

Fronds 10-22 cm. high, membranaceous to decidedly coriaceous,
highly stupose for considerable distance above the base, repeatedly
and irregularly branched by splitting of the blade more or less deeply,
the lobes laciniate to reniform-flabellate ; thickness of frond varying
from 18 cells, at the base, to 1 cell, at the broad inrolled margin, for
the most part 8-12 cells thick ; the flabellate portion of the frond
marked distinctly by numerous, narrow, concentric bands of hairs;
color dark brown, dark green to almost black on drying; sori of
oogonia and aplanosporangia distributed unevenly over both sides of

662 University of California Puhlications in Botany [Vol. 8

the frond with a slight tendency to concentric zonation; antheridia
unknown ; oogonia 140-200jli long, 70-90/x broad, wall 10-12/x thick ;
aplanosporangia about the same size, but with much thinner walls.

Growing in the lower littoral and upper sublittoral belts. Widely
distributed in the Gulf of California, and as far north as Magdalena
Bay, Lower California.

Bory, Diet, class, hist, nat., vol. 12, 1827, p. 591 ; Voy. Coquille,
Bot. Crypt., 1828, p. 147, Atlas, 1826, pi. 21, tig. 1 ; Howe, Phyc. Stud.
V, 1911, p. 497; Setchell and Gardner, Mar. Alg. Gulf Calif., 1924,
p. 729.

There may be other species represented on our coast, but all of the
specimens in our hands seem to belong to the same coarse, thick, dark
colored species which passes under the foregoing name.

Series 3. CYCLOSPOEEAE aresch.

Thallus at maturity never unicellular nor monosiphonous, simple
or branched, varying in size from a few centimeters to several meters
long, usually saxicolous but rarely epiphytic or floating, composed of
highly differentiated and complex tissues, always solid, provided in
part with specialized cavities, the vesicles, tilled with gas which serves
to buoy the plant; multiplication in a few (e.g., Sargassum) by frag-
mentation ; growth terminal ; reproduction sexual only, the oogonia, or
unilocular female gametangia, and antheridia, or unilocular male
gametangia, being located within the thallus, or frond, in specially
developed cavities (the conceptacles) scattered over the whole .surface
of the frond or limited to specialized terminal or subterminal parts
(the receptacles) ; the female gametes non-motile, the male gametes
motile- by two laterally placed cilia of unequal length and possessing a
small, red ' ' eye spot ' ' ; fertilization is effected after both gametes have
escaped into the water ; branched paraphyses associated with the
reproductive organs and often extruding through the osteole of the
coneeptacle.

Areschoug, Phyc. Scand., 1846, p. 28 Repr.

The members which now constitute this series are diverse and
heterogeneous, both as to structure and form. Scarcely any two
authors have agreed from the beginning up to the present time as to
their grouping into orders, families, etc. The various genera, now
regularly assigned to the series, are, in general, in close agreement
regarding the method of reproduction. The gametangia (oogonia
and antheridia) are both unilocular and are borne on the same or on

1^--'^] SetclieU-Gardner: Mehmophijceae 668

separate individuals in specialized cavities among the so-called para-
physes. The reproduction is heterogamous and fertilization takes
place after the escape of the gametes. The female gametes (eggs) are
non-motile, and the male gametes (antherozoids) are motile by two
laterally placed cilia of unequal length. The inequality in size of the
gametes is very marked, the female having been estimated in some
cases to be thirty thousand times as large as the male.

Areschoug {he. cit.) used the name as an order under the sub-
class Fucaceae, as he interpreted that group, and included four
genera, viz., Halidrijs Lyngb., Halicoccus Aresch., Fiicus L., and
Himanthaha as occurring in the region covered bv his account.

Order 8. FUCALES oltmanns

Fronds extremely variable in size, from a few centimeters to
several meters long, but usually slender, complanate, cylindrical, sub-
cylindrical or tumid and constricted at regular intervals, branched,
the branches either pinnate in two ranks in one plane (Fucaceae) or
arising on all sides of the main axis (Sargassaceae) ; attached by a
well developed, solid, usually disk-shaped holdfast, perennial, fruiting
annually and then the specialized fruiting parts disintegrating ; recep-
tacles limited to the terminal or subterminal parts of the fronds, the
oogonia producing 1, 2, 4, or 8 non-motile gametes (eggs).

Oltmanns, Morph. u. Biol, der Algen, vol. 2, 1922, p. 186.

Key to the Families

1. Frond differentiated into axial and lateral members 25. Sargassaceae (p. 704)

1. Fronds flattened without differentiation into axil and lateral members

24. Fucaceae (p. 663)

FAMILY 24. FUCACEAE lamour. (lim. mut.)

Members of the order Fucales with flattened fronds not differen-
tiated into axial and lateral members.
Lamour., Essai, 1813, p. 8, in part.

Key to the Geneka

1. Fronds with distinct percurrent midrib 2

1. Fronds without midrib 3

2. Oogonium producing 8 viable gametes (eggs) 65. Fucus (p. 664)

2. Oogonium producing 1 viable gamete (egg) 68. Hesperophycus (p. 703)

3. Oogonium producing 2 viable gametes (eggs) (Xl. Pelvetia (p. 700)

3. Oogonium producing 1 viable gamete (egg) 67. Pelvetiopsis (p. 702)

664 University of California Publications in Botany [Vol. 8

65. Fucus (L.) Dec'ne and Thuret

Fronds attached by a solid disk-shaped holdfast, complanate, with
more or less distinct percurrent midrib and alae of variable widths,
branching beginning 2-5 cm. from the base, dichotomous or subsecund,
stipe formed by thickening of the midrib and wearing away of the
alae; reproduction sexual, by antheridia and oogonia borne among
hyaline, more or less branched, sterile filaments, the paraphyses, within
cavities, the coneeptaeles, limited to the terminal, metamorphosed, more
or less swollen portions of the branches, the receptacles; oogonium
producing eight oospheres, or eggs, which escape together in a trans-
lucent utricle which soon disintegrates and frees the eggs ; antheridium
produces 64 antherozoids, each with a ' ' red eye spot ' ' and two laterally
affixed cilia of unequal length ; fertilization effected after the eggs
escape from the membrane ; plants synoicous or dioicous.

Decaisne and Thuret, Rech. sur Antherid., 1845, p. 13 ; Linnaeus,
Gen. Plant., 1737, p. 326 {Urn. mut.).

For a discussion of the genus Fucus, see Gardner (1922).

Key to the Species

1. Fronds with abundant caecostomata 1. F. furcatus (p. 664)

1. Fronds with few or no caecostomata 2

2. Cryptostomata absent 5. F. nltens (p. 699)

2. Cryptostomata present, usually abundant 3

3. Fronds membranaceous 2. F. membranaceus (p. 673)

3. Fronds coriaceous 4

4. Cryptostomata few, fronds usually narrow 3. F. edentatus (p. 678)

4. Cryptostomata more or less abundant, fronds wider

4. F. evanescens (p. 681)

1. Fucus furcatus Ag.

Fronds usually rigid, often arborescent, more or less cartilaginous,
for the most part decidedly mucilaginous, regularly dichotomous, olive
green to yellowish ; segments usually relatively long, slightly cuneate
to linear, in some cases decidedly crisped, midrib prominent and per-
current and often yellowish, caecostomata usually abundant ; recep-
tacles for the most part complanate, sometimes tumid, often decidedly
yellowish.

Growing in the middle and lower littoral belts. From Sitka,
Alaska, to Oil Port, San Luis Obispo County, California.

Agardh, Sp. Alg., 1820, p. 97; Icon. Ined., fasc. 2, 1821, pi. 14,
Syst., 1824, p. 279 ; J. Agardh, Sp. Alg., 1848, vol. 1, p. 209. Gardner,
Genus Fucus, 1922, p. 16, pi. 1, fig. 1 (copy of original of Agardh).

1925] Setchell-Gardner : MelanopJiyceae 665

Key to the Forms

1. Caecostomata very numerous, 250-450 per square centimeter 2

1. Caecostomata less than 250 per square centimeter 4

2. Fronds cartilaginous, much crisped and twisted 5. f. contortus (p. 668)

2. Fronds tough, coriaceous, plane or nearly so 3

3. Fronds much branched, 10-18 mm. wide, 20-30 cm. high. ...1. f. tjrpicus (p. 665)

3. Fronds loose, 10-18 mm. wide, 30-50 cm. high 3. f. elongatus (p. 667)

3. Fronds much branched, 15-30 mm. wide, up to 90 cm. high

2. f. luxurians (p. 666)

4. Receptacles cornute, wholly or in part 5

4. Receptacles not cornute 6

5. Segments linear, 5-6 mm. wide 10. f. cornutus fp. 671)

5. Segments cuneate, 18-24 mm. wide 9. f. nigricans fp. 671)

6. Young receptacles decidedly reflexed 7. f. reflexus (p. 669)

6. Young receptacles not reflexed 7

7. Fronds 8-12 cm. high 11. f. abbreviatus fp. 672)

7. Fronds over 12 cm. high 8

8. Fronds 4-12 mm. wide 9

8. Fronds over 12 mm. wide 10

9. Fronds 4-7 mm. wide, receptacles acute 13. f. angustus fp. 673)

9. Fronds 8-12 mm. wide, receptacles blunt 12. f. linearis fp. 672)

10. Segments increasing in width upward 8. f. latifrons fp. 670)

10. Segments diminishing in width upward 11

11. Receptacles covering 3-4 terminal segments, linear 4. f. rigidus fp. 668)

11. Receptacles covering 1-2 terminal segments, very variable in shape

6. f. variabilis fp. 669)

1. Fucus furcatus f. tjrpicus Gardner

Fronds somewhat caulescent, moderately cartilaginous, 20-30 cm.
high, regularly dichotomous, dark olive green, black on drying, seg-
ments plane, linear to cuneate, 10-18 mm, wide, midrib distinct, per-
current, caecostomata 250-300 per sq. cm., small, inconspicuous,
cryptostomata absent or very sparse ; receptacles complanate, mostly
broadly linear, 4-6 cm. long, mostly bifid, apices acute ; conceptacles
very numerous and relatively small.

Growing on boulders and rock ledges in the middle of the littoral
belt. From the Strait of Juan de Fuca to central California.

Gardner, Genus Fticus, 1922, p. 16, pis. 2, 3. Fucus evanescens f,
typicus, Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc),
no. 1338. Fucus inflatus f. edentatus, Setchell and Gardner, Alg.
N.W. Amer., 1903, p. 280.

The Fucus furcatus of Tilden's American Algae, no. 234, as to
the specimen distributed in Professor Setchell 's copy, is not t.ypical of
this species as found on the California coast. The specimen is only a
small portion of a plant. It is mature, considerably worn and bat-
tered, and has comparatively few caecostomata. The distribution is

666 University of California PuMications in Botany [Vol.8

probably to be referred to f. typieus, but it is desirable to consult other
specimens of the distribution before deciding definitely.

This form is abundant on the California coast, but less common
toward the north, being supplanted by numerous other forms. It
reaches its optimum growth in localities where the surf is only mod-
erately active. In such situations it attains its greatest height, not
infrequently specimens attaining a height of 4.5 dm., but when on
boulders exposed to a heavy surf, it remains much shorter, is more
arborescent, the alae wearing away up to the last segment. The
illustration on Gardner's plate 3 is of the former state, and that on
plate 2 is from an exposed situation at Fort Ross, California, pre-
sumably the type locality for the species.

3. Fucus furcatus f . luxurians Gardner

Plate 99

Fronds cartilaginous, distinctly caulescent, the alae wearing away,
the much thickened midrib becoming the flattened stipe, regularly
dichotomous, 4-5 dm. (up to 9 dm.) high, dark olive green to olive
brown, midrib and receptacles yellowish, holdfast large and firm;
segments varying from broadly cuneate to nearly linear, frequently
splitting obliquely to the midrib, 1.5-3 cm. wide, terminal often wider
and rounded, caecostomata very numerous, 300-350 per sq. cm.,
cryptostomata scattered, bearing fascicles of long exserted paraphyses.
Receptacles definitely delimited, 4-6 cm. long, complanate or decidedly
tumid, bi- tri-furcate, apices blunt or acuminate ; conceptacles very
numerous.

Growing on boulders and rock ledges in the middle of the littoral
belt. Northern and central California.

Gardner, Genus Fuctis, 1922, p. 22, pi. 10.

Plate 99 represents a plant of this form in mature fruit, producing
receptacles for the first time. The receptacles in this specimen are
all complanate, a condition which prevails very commonly, but in
certain localities the receptacles are tumid and mucilaginous.

Forma luxurians is closely related to forma typieus. The two forms
often grow intermingled, but may readily be distinguished by differ-
ences in size and color, forma luxurians being the more robust, and of
a lighter brownish or yellowish color in the upper parts, particularly
the receptacles. The southern limit of the form, as of all forms of

if--i] Setchell-Gardner : Melanopliijceae 667

Fucus, so far as is at present known, is Oil Port, San Lnis Obispo
Countj% California. The northern limit has not yet been determined,
but it probably extends as far north as Cape Flattery, Washington,
where there is an intermingling of several forms whose limits have not
yet been determined.

o.

Fucus furcatus f. elongatus Gardner

Fronds sparsely branched, distinctly arborescent, somewdiat f oliace-
ous, 3-5 dm. high, regularly dichotomous, angles wide and rounded,
dark olive green, dark olive brown on drying ; segments long, 10-18
mm. wide, euneate, terminal segment often ovate, rounded, growing
point slightly depressed, midrib narrow but distinct, frequently yel-
lowish above, alae wearing away below, remaining on 4-5 segments
from the apices, caecostomata variable but usually abundant, up to
450 per sq. cm., cryptostomata few, scattered, bearing fascicles of
long, exserted paraphyses ; receptacles dark brown to yellowish, either
complanate or much inflated, distinctly delimited, simple, emarginate
or bifid, blunt or acute, 3-5 cm. long; conceptacles not prominent or
abundant.

Growing on boulders in the lower littoral belt in localities exposed
to the surf. South end of San Juan Island, Washington, and Sunset
Beach, near the mouth of Coos Bay, Oregon.

Gardner, Genus Fucus, 1922, p. 21, pi. 9.

This form is intermediate, in several of its characters, between
F. furcatus f. fypieus and F. furcaUis f. luxurians. On drying it is
smoother than either of the above mentioned forms, the caecostomata
not standing out so prominently.

Gardner, no. 1973 (Herb. TTniv. Calif., no. 132743, and in Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. CX, sub F.
evanescens f. macrocephalus Kjellm.) is typical of forma elongatus in
color, character of branching, having mostly wide and usually rounded
angles, character of segments and receptacles, but varies from the type
in having very few caecostomata. It is certainly quite different from
Kjellman's F. evanescens f. macroceplialus in all of its important
characters.

668 University of California Piihlications in Botany [Vol. 8

4. Fucus furcatus f. rigidus Gardner
Plate 100

Fronds robust, rigid, cartilaginous, 30-45 cm. high, dichotomous or
subsecund, dark olive green, black on drying, midrib and receptacles
olive green below, olive brown and yellowish above, angles mostly
rounded; segments linear or slightly cuneate, strict, 1.5-3 cm. wide,
apices truncate, midrib prominent, caecostomata, 60-100 per sq. cm. ;
receptacles not definitely delimited, complanate, linear, apices acumin-
ate, bi- tri-furcate, 10-20 mm. wide, 5-8 cm. long; conceptacles large,
covering a cuneate area extending down the midrib from the receptacle.

Growing on boulders in the lower littoral belt. Port Townsend
and the west coast of Whidbey Island, Washington.

Gardner, Genus Fucus, 1922, p. 24, pi. 13.

According to our conception of this form, its phylogenetic position
is between forma latifrons and forma luxurians. From the former
it differs in having a more rigid cartilaginous structure, in having
narrower and more strict segments, and in having longer and narrower
receptacles. From the latter it differs in having more strict and more
nearly linear segments and longer and narrower receptacles.

5. Fucus furcatus f . contortus Gardner

Fronds 20-30 cm. high, caulescent below, foliaceous above, robust,
cartilaginous, much contorted, dichotomous or subdichotomous, olive
green below, yellowish above, dark olive brown on drying ; segments
cuneate below, margins crisped, linear above, reduced in width above
each forking, 2-2.5 cm. wide, apices rounded, growing depression
slight, midrib prominent, caecostomata 250-300 per sq. cm., minutely
papillate on drying; receptacles complanate, 4-7 cm. long, not
distinctly delimited.

Growing in quiet water on boulders in the lower littoral belt. Near
Bellingham, Washington.

Gardner, Genus Fucus, 1922, p. 25, pi. 15.

This form seems to be quite distinct from all others. Its color, its
rigid, cartilaginous consistency, its relatively wide fronds, its large
number of caecostomata, its contorted and crisped habit of growth
are characters not found combined in any other known form. The
plants were found growing in great profusion, in good vegetative
condition in midsummer. The fruiting season apparently is in the
winter, as only a few well developed receptacles could be found.

19-5] SetcheJI-Gardner: Melanophyceae 669

6. Fucus furcatus f. variabilis Gardner

Fronds caulescent, foliaceous above, 25-40 cm. high, subcartilagin-
ous, dichotoraous or in part subseciind, dark olive green to olive brown,
black on drying, midrib and receptacles yellowish ; segments linear
to moderately cuneate, 2-3.5 cm. wide, usually reduced in width above
each forking, midrib well developed, percurrent, caecostomata vary-
ing from few up to 75 per sq. cm. ; receptacles very variable in shape
and size, definitely delimited, swollen and mucilaginous or complanate,
simple and blunt or bi- tri-furcate and acuminate.

Growing on rock ledges in quiet bays in the middle of the littoral
belt. Sitka, Alaska.

Gardner, Genus Fucus, 1922, p. 26, pis. 16, 17.

A close relationship seems to exist between this form of F. furcatus
and F. evanescens f. magnificus. The color, caecostomata characters,
and the rigid, more or less cartilaginous consistency of the fronds are
characters that belong to the furcatus group. The width of segments
and particularly the fruiting habit (see under F. evanescens f.
magnificus) are very similar to those of that form. The caecostomata
vary much in number. In some specimens they are practically absent,
while in others there may be seventy-five or perhaps more to the square
centimeter. They are mostly large, deep-seated, and inconspicuous
on drying, except by transmitted light. Specimens with but a few
caecostomata usually have a few small cryptostomata.

7. Fucus furcatus f. refiexus Gardner

Fronds foliaceous, subcartilaginous, 30-45 cm. high, regularly
dichotomous, dark olive green to dark olive brown, angles mostly very
broad and rounded ; segments cuneate below, terminal often obovoid,
8-25 mm. wide, apices rounded, growing point only slightly depressed,
midrib narrow but distinct, slightly evanescent above, caecostomata
very variable in number, 50-150 per sq. cm. ; receptacles distinctly
delimited, substipitate, complanate, ellipsoidal, entire or bifid, wide-
spreading; conceptacles small and numerous.

Growing on boulders in the lower littoral belt. Point Defiance,
Tacoma, Washington.

Gardner, Genus Fucus, 1922, p. 23, pis, 11, Ha, and 12.

The consistency of this form is somewhat like that of the evanescens
group, the tissues being more delicate and soft than those of the typical

670 Vniversity of California Puhlications in Botany [Vol. 8

furcatus forms. It is placed in the furcaius group on account of the
color, dark olive green to dark olive brown, and the presence of
caecostomata in abundance.

Many of the terminal segments are very broadly ovate, almost
stipitate, on account of the narrowing of the bases, producing broad
rounded angles between the segments. The growing points differ from
those of all other forms observed on our coast, the apices of the seg-
ments being rounded and smooth, but with slight inconspicuous slits
to indicate the position of the growing point.

The receptacles, particularly when young, have the pronounced
and distinguishing character of spreading very widely from each other
at their bases, in some instances standing at right angles, or even
more, to each other. Many segments have the character of reducing
their width at each forking, the terminal segments forming the stipe-
like portion bearing the much wider receptacles.

8. Fucus furcatus f. latifrons Gardner

Plate 101

Fronds decidedly foliaceous, contorted, comparatively thin, 25-35
cm. long, dark olive brown below, light brown above, black on drying,
dichotomous, angles mostly acute; segments cuneate, terminal seg-
ments mostly rounded, 2-3.5 cm. wide, midrib prominent below,
decidedly reduced above, caecostomata 40-80 per sq. cm. ; receptacles
not definitely delimited, complanate, bi- tri-furcate, mostly acute ; con-
ceptacles large, frequently extending down the midrib, covering a
cuneate area of the segment.

Growing on rocks in the middle and lower littoral belts. Channel
Rocks, near Fort Ward, west of Seattle, Washington.

Gardner, Genus Fucus, 1922, p. 25, pi. 14. Fucus evanescens f.
pergrandis, Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc),
no. CXI.

In width of fronds, this form of F. furcatus overlaps or merges
into forms of F. evanescens, e.g., forma magnificus. It is here placed
under furcaius on account of the dark olive green color, more or less
cartilaginous consistency, and particularly on account of the abun-
dance of caecostomata.

It is closely related to F. furcatus f. rigidus, but the fronds are
more foliaceous, the receptacles are wider and shorter and the
caecostomata are less abundant.

19--'^l Setchcll-Ganlner: MelanopJniceae 671

9. Fucus furcatus f. nigricans Gardner

Fronds comparatively fragile, thin membranaceous, 35-45 cm.
high, regularly dichotomous, dark olive green, black on drying, angles
broad and slijyhtly rounded ; segments cuneate below, linear above,
reduced to a midrib 4-5 segments back of the apices, widest 3-4 back
of the apices, reduced in width at each forking, widest 18-24 mm.,
midrib narrow but distinct, percurrent, alae thin, caecostomata 70-80
per sq. cm., small, obscure ; receptacles 2.5-3.5 cm. long, complanate or
inflated, single or bifid, definitel}' delimited, apices mostly acute and
mostly reflexed.

Growing on boulders and rock ledges in the middle of the littoral
belt. Cattle Point, south end of San Juan Island, Washington.

Gardner, Genus Fucus, 1922, p. 21, pi. 8.

In the cornute habit, forma nigricans is closely akin to forma
cornutus, but dififers from it in the following characters; thinner
fronds which are more cartilaginous, darker color, black on drying, in
having widely divergent segments and in having an abundance of
caecostomata.

10. Fucus furcatus f. cornutus Gardner

Fronds subcoriaceous, 20-30 cm. high, regularly dichotomous, dark
olive green to dark olive brown, black on drying ; segments divergent,
linear, or very slightly cuneate in the narrower specimens, varying to
decidedly cuneate in the wider specimens, 5-8 mm. wide, apices acu-
minate, truncate, midrib well developed, prominent, very slightly
evanescent above, alae narrow, caecostomata very sparse, 10-20 per
sq. cm., small ; receptacles distinctly delimited, very variable, com-
planate or inflated, usually much wider than the segments, simple or
bifurcate. 2-4 cm. long, blunt to acuminate, apices mostly laterally
reflexed ; conceptacles not numerous, inconspicuous.

Growing on rocks in the middle and lower littoral belts. From
Yakutat Bay, Alaska, to Victoria, British Columbia.

Gardner, Genus Fucus, 1922, p. 20, pi. 7. Fucus evanesccns f.
cornutus Saunders, Alg. Harriman Exp., 1901, p. 432, pi. 62, fig. 2;
Collins, Mar. Alg. Vancouver Isl., 1913, p. Ill ; Collins, Ilolden and
Setchell, Phyc. Bor.-Amer. (Exsicc), no. 927.

The narrow fronds with prominent midribs and the very dark color
make this form close to certain narrow forms of edentatus, but the

672 University of California Publications in Botany [^'o^. 8

absence of cryptostomata and the small caecostomata relate it to the
furcatus group. It has the least number of caecostomata of all the
forms Avith which we are familiar, certain specimens being nearly
free from them. In this character the form approaches very closely
to the edentatu^ group.

11. Fucus furcatus f. abbreviatus Gardner

Plate 98

Fronds cartilaginous, rigid, rough, 8-12 cm. high, regularly
dichotomous, varying from olive green to olive brown or light yellow-
ish ; segments cuneate to linear, 8-16 mm. wide, apices truncate, mid-
rib narrow^ but distinct, caecostomata comparatively large, prominent
on drying, 60-80 per sq. cm. ; receptacles distinctly delimited, much
wider than the segments, 2-3.5 cm. long, complanate or swollen with
mucilage, 1-2 furcate, apices acuminate ; conceptacles numerous and
very prominent.

Growing on boulders in the middle littoral belt. Islands of San
Juan County and vicinity, Washington.

Gardner, Genus Fucus, 1922, p. 19, pi. 6.

This form is fairly abundant in the moderately quiet waters among
the islands of San Juan County, Washington, and extends into the
upper Sound region. It varies considerably in size, both in length and
in width of the fronds. Specimens 3 to 4 cm. in height have been taken
in full fruit, but these were depauperate forms, in the upper limit on
the belt in which they grow. The striking features of this form are
the short bunchy habit, the abundance of well developed receptacles
appearing simultaneously, and the numerous large caecostomata
making the fronds rough, particularly prominent on drying.

12. Fucus furcatus f. linearis Gardner

Fronds somewhat caulescent, cartilaginous, 30-35 cm. high, pro-
fusely branching, dichotomous, dark olive green, black on drying;
segments strict, linear, 8-12 mm. wade, caecostomata 40-70 per sq. cm.,
inconspicuous on drying ; receptacles very numerous, definitely
delimited, wider than the fronds, entire or bifid, mostly ovate, 1.5-2 em.
long; conceptacles numerous, large.

Growing in quiet water on boulders in the lower littoral belt.
Tracyton, Kitsap County, Washington.

Gardner, Genus Fucus, 1922, p. 19, pi. 5.

19--^] Setchell-Gardner: Melanophyceae 673

This form is remarkable for the abundance of receptacles it pro-
duces on a single plant. Plate 5 of Gardner was made from a portion
of a plant that had approximately five hundred and fifty receptacles.
It is also rather unusual that so many of the segments should fruit at
the same time as is the case in this form. Not a single segment remained
sterile in the above mentioned plant. This form does not seem to be
abundant in the Puget Sound region.

13. Pucus furcatus f . angnstus Gardner

Fronds narrow, caulescent, rigid, subcartilaginous, 20-35 cm. high,
dichotomous, dark olive green, black on drying; segments linear to
slightly cuneate, 4-7 mm. wide, apices truncate, midrib distinct and
perciirrent, caecostomata 125-175 per sq. cm. ; receptacles mostly com-
planate, occasionally inflated, definitely delimited, deeply bifid, 4-7
cm. long, apices acute or acuminate; conceptacles numerous, closely
placed.

Growing in abundance on rock ledges in the upper third of the
littoral belt. San Juan County, Washington, and the central coast
of Oregon.

Gardner, Genus Fucus, 1922, p. 18, pi. 4,

This form was found at Sunset Beach, near the mouth of Coos Bay,
Oregon, and at Cattle Point, at the south end of San Juan Island,
Washington. This latter locality is particularly rich in forms of
Fucus. There is an intermingling of the waters flowing among the
islands to the north and east, which have a varied Fucus flora, with
the waters from the Strait of Juan de Fuca and the Straits of Georgia,
Avashing the shores of land to the west and north, which have a some-
what different Fucus flora. The shore of the south side of Cattle
Point is exposed to a swiftfly flowing current and to the action of
swells from the Pacific Ocean, through the Straits of Juan de Fuca.
Around to the north side of the point the shores are affected but little
by the action of the surf and tides.

2. Fucus membranaceus Gardner

Fronds variable in size from a few cm. to 4 dm. high, decidedly
membranaceous, dark olive brown, to decidedly yellowish above; seg-
ments linear to slightly cuneate, strict, alae membranaceous, midrib
relatively small but distinct and percurrent, cryptostomata scattered.

674 University of California Publications in Botany [Vol. 8

mostly in the terminal and subterminal segments, relatively small but
papillate on drying ; receptacles mostly complanate, in part inflated,
bifid and acuminate.

Growing in the littoral belt. From the Bering Sea to Puget Sound,
"Washington.

Gardner, Genus Fucus, 1922, p. 32.

Key to the Forms

1. Cryptostomata sparse and inconspicuous 3. f. limitatus (p. 674)

1. Cryptostomata up to 60 per square centimeter 2

2. Fronds 8-14 cm. high, 5-9 mm. wide, receptacles wider and blunt

4. f. abbreviatus (p. 676)

2. Fronds over 14 cm. high 3

3. Fronds 12-20 cm. high, 4-9 mm. wide, receptacles sharply acuminate

6. f. aciiminatus (p. 677)

3. Fronds over 10 mm. wide 4

4. Fronds up to 6 cm. wide, cryptostomata 15-20 per square centimeter...

2. f. latissimus (p. 675)

4. Fronds less than 3 cm. wide 5

5. Cryptostomata 20-25 per square centimeter, receptacles acuminate or acute...

1. f. typicus (p. 674)

5. Cryptostomata 30-40 per square centimeter, receptacles blunt

5. f. obtusus (p. 676n

1. Fucus membranaceus f. typicus Gardner

Fronds caulescent, 28-40 cm. high, flaccid, membranaceous, dicho-
tomous, alae wearing away below, leaving the relatively small, thick-
ened, cylindrical midrib, light yellowish brown ; segments narrowly
cuneate to linear, 10-18 mm. wide, growing point in a slight, crescent
shaped depression, midrib moderately developed, alae thin, cryptosto-
mata small, 20-25 per sq. cm., conspicuous on drying, mostly in the
terminal and subterminal segments; receptacles relatively large, dis-
tinctly delimited, mostly inflated, bifid, apices long attenuate ; con-
ceptacles moderately abundant, relatively small, but conspicuous
because of their dark color.

Growing on rock ledges and boulders in the middle of the littoral
belt. Sitka, Alaska.

Gardner, Genus Fucus, 1922, p. 34, pis. 30, 31.

This form was sparsely interspersed in groups among the multi-
plicity of forms growing in the vicinity of Sitka. It may readily be
recognized by its light yellowish color, its mostly inflated receptacles,
and by its moderately narrow, long, membranaceous fronds.

1^25] Setchell-Gardner: Melanopliyceae 675

2. Fucus membranaceus f. latissimus Gardner

Plate 105

Fronds foliaceous, membranaceous, crisped, dichotomous, 22-25
em. high, dark olive green below, changing to yellowish brown above,
holdfast comparatively small, stipe flattened ; segments slightly ovate
above, narrower and cuneate below, subterminal 4-6 cm. wide, midrib
relatively narrow, percurrent, alae thin, membranaceous, frequently
inflated with gas, margins undulate, apices round and smooth, grow-
ing point inconspicuous, eryptostomata 15-20 per sq. cm., producing
abundant protruding paraphyses, prominent on drying ; receptacles
sparse, complanate, not definitely delimited, widest at the base, bi-
tri-furcate.

Growing on boulders in the extreme lower littoral belt. Kadiak
Island and Sitka, Alaska.

Gardner, Genus Fucus, 1922, p. 35, pis. 33, 34,

In June, 1910, Gardner first observed this form growing in the
harbor at Sitka, but at that time was unable to obtain good fruiting
specimens. On his second visit to the same place in July, 1917, he
located a large bed containing thousands of plants growing along the
extreme low-tide level, and was able to procure a number of good
fruiting specimens, although the summer season does not seem to be its
best fruiting season. This form may be readily distinguished from all
others by its width, being the widest of all forms, its membranaceous
alae, its very dark lower portion, and usually yellowish upper portion,
and by its much crisped habit of growth. Some of the plants of this
species collected by G. B. Rigg at Kadiak Island are the widest speci-
mens of Fucus that have been reported, some of the fronds measuring
over seven centimeters in width.

3. Fucus membranaceus f . limitatus Gardner

Fronds distinctly membranaceous, 22-32 cm. high, dichotomous,
light yellowish brown, dark brown on drying, stipe and holdfast
fragile ; segments strict, mostly cuneate, usually wider at each suc-
cessive forking, 12-20 mm wide, terminal lobes rounded, growing
point slightly depressed, midrib not prominent, alae thin, erypto-
stomata very sparse and inconspicuous; receptacles mostly inflated,
bifid, apices acuminate, usually divergent, very variable in size, up to
5 cm. long and 2 cm. wide ; conceptacles scattered and not prominent.

676 University of California Publications in Botany [Vol.8

Growing on rocks, logs, etc., in sheltered localities in the upper
third of the littoral belt. Point Defiance, Tacoma, Washington.

Gardner, Genus Fucns, p. 35, pi. 32.

It is not easy to delineate the distinctive characters of this form
associating it with the memhranaceus group, rather than with the
evanescens group. In many specimens the midrib is quite indistinct.
The cryptostomata characters are quite as much of the one group
as of the other. It is provisionally placed with the former group
largely on account of the predominance of inflated receptacles, on the
membranaceous character of the fronds and on the color.

Two forms of Fucus were found growing at Point Defiance, repre-
senting an interesting case in distribution. Forma limitatus occupied
for the most part the extreme upper portion of the littoral belt, and
grew on logs and boulders. F. furcatus f. reflextis grew on boulders
and occupied the lower third, with an occasional plant of either form
invading the central part.

4. Fucus membranaceus f. abbreviatus Gardner

Fronds short, membranaceous, flabelliform, profusely branched,
8-14 cm. high, dichotomous or in part secund, yellowish brown; seg-
ments strict, narrowly cuneate to sublinear, 5-9 mm. wide, apices
truncate midrib moderately prominent, only slightly evanescent, alae
thin, cryptostomata few in the lower portions of the plant, 20-40
per sq. cm. in the terminal segments, small but conspicuous on drying ;
receptacles bifid, mostly swollen wdth mucilage, apices acuminate or
in part blunt, somewhat divergent, sharply delimited ; conceptacles
numerous and prominent.

Growing on rock ledges exposed to heavy surf, in the upper limit
of the littoral belt. Sitka, Alaska.

Gardner, Genus Fucus, 1922, p. 32, pi. 27.

Forma abbreviatus grew on the same islet on which the forma
acuminaius was found growing, but mostly higher up, some specimens
were even above mean high-tide level. It differs from forma acu-
minaius principally in having shorter fronds, relatively much broader
and shorter receptacles, fewer cryptostomata, and it is less mucilagin-
ous. It seems closely akin to J. Agardh's Fucus bursigerus from
Spitzbergen, but it is more robust, and much more profusely branched
than his description and figure call for (cf. J. Agardh, Spets. Alg.
Till., 1868, pp. 41, 42, pi. 3). We have seen the specimen of Kjellman's

is^.")] Setchell-Gardner : Melanophyceae 677

F. evanescens f. hiirsigera, also collected at Spitzbergen. The plants
of Agardh and of Kjellman are undoubtedly of the same lineage, and
they are probably allied more closely to the evanescens lineage than
to any other. The Sitka plants under consideration, on the whole, and
particularly on account of size and consistency, seem closer to the
memhranaceus group than to the evanescens group. The two groups
certainly overlap through these two forms.

5. Fucus membranaceus f. obtusus Gardner

Fronds slender, more or less caulescent, membranaceous, 30-40 cm.
high, dichotomous, olive green to dark brown, very dark on drying ;
segments strict, linear to slightly cuneate, relatively long, 10-16 mm.
wide, midrib percurrent, alae thin, cryptostomata and caecostomata
nearly equal in number, 30-40 per sq. cm., small, papillate on drying ;
receptacles definitely delimited, much inflated, mostly single, in part
bifid, 2.5-4 cm. long, 1-1.5 cm. wide, blunt, yellowish brown; con-
ceptacles not numerous, comparatively small, not protruding.

Growing on rocks in the middle and lower littoral belts. Sitka,
Alaska,

Gardner, Genus Fucus, 1922, p. 34, pi. 29.

This form has characters linking together forma acuminatus and
forma typicus of this species. It has the arborescent habit common to
both forms, the width of the fronds is intermediate, and the receptacles
are mostly simple and blunt, unlike either form.

6. Fucus membranaceus f. acuminatus Gardner

Fronds caulescent, 12-20 cm. high, profusely branched, dicho-
tomous to subsecund, dark brown to yellowish brown ; segments very
slightly cuneate to linear, 4—9 mm. wide, truncate, midrib well
developed below, somewhat reduced just below the receptacles, alae
thin and membranaceous, cryptostomata varying from few to 50-60
per sq. cm., mostly on the segments just below the receptacles, absent
in the older parts, very small but conspicuous on drying; receptacles
numerous, regularly and deeply furcate, or, rarely, single, mostly
complanate, narrow, acute, 2-3 cm. long; conceptacles relatively
numerous.

Growing on rock ledges exposed to heavy surf, in the upper
littoral belt. Sitka, Alaska.

Gardner, Genus Fucus, 1922, p. 33, pi. 28.

678 University of California PuMications in Botany \you 8

This form has been collected but once. It grew in abundance on
a small rocky island in front of Sitka harbor and well up in the littoral
belt, exposed to the heavy action of the surf. The plants growing in
the most exposed places are quite arborescent in character, the alae
wearing away up to the last two or three segments, leaving the much
thickened midrib. In depressions and less exposed situations, the
alae are usually wider and remain longer on the fronds. The plants
in this group become exceedingly mucilaginous on being soaked out
after a thorough drying.

3. Fucus edentatus De la Pyl.

Fronds usually narrow, more or less flaccid, regularly dichotomous,
olive green to olive brown or yellowish, usually very black on drying ;
segments mostly decidedly linear, at times cuneate, midrib distinct,
percurrent, cryptostomata and caecostomata few or none, receptacles
mostly narrow, linear-lanceolate, acuminate, inflated or complanate.

Growing in the middle and upper littoral belts. From Sitka,
Alaska, to Coos Bay, Oregon.

De la Pylaie, Flor. Terre-Neuve, 1829, p. 84.

Key to the Forms

1. Midrib prominent, percurrent, segments reduced at each forking

3. f. costatus (p. 680)

1 . Midrib less prominent, segments not reduced at each forking 2

2. Receptacles and segments widely divergent 2. f. divergens (p. 679)

2. Receptacles and segments not widely divergent 3

3. Fronds over 25 cm. high, receptacles often inflated 1. f. hesperius (p. 678j

3. Fronds less than 25 cm. high, not inflated 4

4. Fronds 9-15 cm. long, receptacles acute, 2-2.5 cm. long. 4. f. acutus (p. 680)

4. Fronds 12-22 cm. long, receptacles subulate, 2.5-4.5 cm. long

5. f. divaricatus (p. 681)

1. Fucus edentatus f. hesperius Gardner

Plate 103

Fronds arborescent, 30-40 cm. high, narrow, coriaceous, dicho-
tomous, dark olive green, black on drying, stipe cylindrical; segments
strict, linear, 5-10 mm. wide, with truncate apices and well developed,
prominent, percurrent midribs, cryptostomata absent or extremely
rare ; receptacles mostly complanate, or in part much inflated, bi- tri-
furcate, linear, 3-4.5 cm. long, apices acute ; conceptacles numerous.

1925] Setchell-Ganbier: Melanophjircae 679

Growing- on rock ledges exposed to the action of the surf, in the
upper third of the littoral belt. Cape Arago, at the entrance to Coos
Bay, Oregon.

Gardner, Genus Fucus, 1922, p. 28, pi. 21.

This form grew in abundance in company with F. furcatus f.
angustus. The two groups of plants were, however, in slightly dififer-
ent altitudes in the belt, and were not indiscriminately intermixed.
When the two sets of plants are dried, they are easily distinguishable
by the smoothness of the segments, forma hesperius being smooth
and usually shining, while forma angustus is rough, due to caecosto-
mata, and is of a duller color.

2. Fucus edentatus f. divergens Gardner

Fronds coriaceous, smooth and glossy, 28-38 cm. high, regularly
dichotomous, dark olive green, black on drying ; segments divergent,
7-11 mm. (up to 15 mm.) wide, cuneate below, linear above, reduced
somewhat above each forking, terminal truncate, growing point incon-
spicuous, midrib very distinct, percurrent, cryptostomata and caeco-
stomata absent or very sparse ; receptacles 3-6 cm. (up to 12 cm.) long,
definitely delimited, much wider than the segments, single or bifurcate,
and mostly widely divergent, apices acuminate or acute ; conceptacles
numerous and prominent.

Growing on rock ledges in the lower littoral belt. Kanaka Bay,
San Juan Island, Washington.

Gardner, Genus Fucus, 1922, p. 29, pi. 22.

This form of edentatus is closely related to forma hesperius. The
fronds average somewhat wider, more robust, the angles between the
segments are very much wider and more rounded, and the receptacles
mostly deeply bifurcate or single, widely diverging. The average
width of the segments is about 10 mm., but a few specimens were
found with extremely narrow segments, about 3 mm. These were
profusely branched, with wide angles, and small diverging receptacles.
Also a few specimens were found with the characters of the form, but
were 15 mm. wide. These few specimens are to be considered as the
extremes in individual variation in this particular character.

680 University of California Piihlications in Botany [Vol. 8

3. Fucus edentatus f . costatus Gardner

Fronds slender, subcoriaceous, 15-25 cm, high, regularly dicho-
tomous, yellowish brown, segments distinctly linear, relatively long,
reduced in width above each forking, widest 5-8 mm., terminal 2-4
mm., midrib highly developed, percurrent, alae narrow and mem-
branaceous, cryptostomata sparse, prominent ; receptacles definitely
delimited, simply or mostly bifid, 20-35 cm. long, apices acute ; con-
ceptacles moderately abundant and conspicuous.

Growing on rocks in the lower third of the littoral belt. Lower
Puget Sound region, Washington.

Gardner, Genus Fucus, 1922, p. 30, pis. 23, 24. Fucus evanescens
f. angustus, Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc),
no. 926 ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 284.

The plants distributed in the Phycotheca Boreali-Americana as
no. 926 are from the type locality, but were taken from the upper
limit of the belt of distribution when the tide was well in ; they are
considerably battered and worn away, and thus cannot be said to be
in typical condition.

This form resembles closely F. evanescens f. angustus Kjellm. in
width of fronds, and was so referred as is cited above. Subsequent
to the foregoing publications, material of the species, collected and
determined by Kjellman on the Vega expedition, has been received at
the Herbarium of the University of California (no. 132699). The lack
of a prominent percurrent midrib, almost complete absence of crypto-
stomata, and especially the very small receptacles of Kjellman 's
material, indicate unmistakably that our plant is of a different lineage.

4. Fucus edentatus f. acutus Gardner

Fronds 9-15 cm. high, arising from a relatively broad, flat hold-
fast, dichotomous, angles acute, olive green, black on drying ; segments
linear, 3-4 mm. Avide, apices truncate, midrib very prominent, percur-
rent, alae relatively narrow, persistent, cryptostomata sparse; recep-
tacles single or deeply bifurcate, tapering at both ends, 2-2.5 cm. long.

Growing on sandstone in the middle littoral belt. Bellingham
(Fairhaven), Washington.

Gardner, Genus Fucv^, 1922 p. 31, pi. 25. Fucus infiatus f.
linearis, Setchell and Gardner, Alg. N.W. Amer., 1903, p. 280.

The very dark color on drying, the absence of caecostomata, and
the sparseness of cryptostomata in this form seem amply sufficient to

1925] Setchcll-Gardner: Melanophyceae 681

ally it with the edentatus group rather than with the seemingly more
imperfectly defined group inflatus. The linear arrangement of the
cryptostomata in two rows along the prominent midrib and the
relatively small, pointed receptacles distinguish it from all other forms
of edentatus.

5. Fucus edentatus f. divaricatus Gardner
Plate 104

Fronds fragile, usually 12-22 cm., but occasionally up to 32 cm.
high, regularl}' dichotomous, with wide angles, stipe and lower
branches nearly terete, dark olive brown, receptacles frequently yellow-
ish ; segments narrowly linear, reduced in width slightly at each
forking, terminal, acute, 2-5 mm. wide, midrib highly developed to
the apices, alae almost absent, cryptostomata few and unevenly dis-
tributed ; receptacles definitely delimited, awl-shaped or slightly
flattened. 2.5-4.5 cm. long, mostly simple or deeply furcate ; con-
ceptacles conspicuous.

Growing on ledges of sandstone in the lower littoral belt. Belling-
ham, Washington, Comox and Ucluelet, Vancouver Island, British
Columbia.

Gardner, Genus Fucus, 1922, p. 31, pi. 26. Fucus inflatus f. fili-
formis, Setchell and Gardner, Alg. N.W. Amer., 1903, p. 281 ; Collins,
Mar. Alg. Vancouver Isl., 1913, p. 111.

4. Fucus evanescens Ag.

Fronds moderately robust, sometimes arborescent, usually decid-
edly coriaceous, dichotomous or in part subsecund, olive brown, to
yellowish above ; segments often quite foliaceous and crisped, in some,
narrow, cuneate to linear, midrib more or less distinct, in some cases
vanishing more or less in the terminal segments, cryptostomata few
to many, scattered ; receptacles very variable in shape and size, long
and narrow to short and blunt, bi- tri-furcate.

Growing in the littoral and upper sublittoral belts. From Bering
Sea to Coos Bay, Oregon.

Agardh, Sp. Alg., vol. 1, part 1, 1820, pp. 92, 93.

Fucus evanescens was established by C. A. Agardh in 1820 (loc.
cit.) based upon material collected by Chamisso. The type specimen
is in the herbarium of J. G. Agardh under no. 00299, labeled Kam-
tschatka with a query. The name was chosen to signify the vanishing
of the midrib toward the outer ends of the segments. Much of the

682 University of California Publications in Botany ['Vol. 8

material, however, which has since been assigned to the species, even
by J. G. Agardh, does not in the least conform to this character. The
name has been used quite extensively to designate plants growing on
both the north Atlantic and the north Pacific shores. The original
illustration by Agardh in Icones algarum ineditae, 1821, pi. 13,
fig. 1, is of a fragment of a frond with complanate receptacles, and
was probably from an immature plant. This plate was reproduced by
Gardner in The genus Fucus on the Pacific Coast of North America,
1922, plate 1, figure 2.

We have selected evanescens as the name best suited to include a
large aggregation of forms which appear in the northern portion of
our region, necessarily having to extend and modify Agardh 's original
description.

Key to the Forms

1. Typical mature fronds short, up to 15 cm 2

1. Typical mature fronds over 15 cm. high 4

2. Plants growing in salt marshes and tide flats 21. f. nanus (p. 698)

2. Plants attached to rocks by a disk-shaped holdfast 3

.3. Plants dendroid, segments usually much eroded 11. f. dendroides (p. 691)

3. Plants flabellate, segments cuneate 20. f. cuneatus (p. 698)

4. Average width of fronds less than 15 mm 5

4. Average width of fronds more than 15 mm 12

4. Average width of fronds unknown 18

5. Midrib absent or indistinct 19. f. ecostatus (p. 697)

5. Midrib distinct, usually percurrent 6

6. Receptacles simple to deeply bifid, usually blunt 7

6. Receptacles more or less decompound 10

7. Receptacles with distinct margin free from conceptacles

10. f. marginatus (p. 690)

7. Receptacles without a free margin 8

8. Receptacles widened suddenly, almost pedicellate.. 17. f. contractus (p. 695)

8. Receptacles less conspicuously pedicellate 9

9. Cryptostomata none or few 2. f. macrocephalus (p. 684)

9. Cryptostomata 20-25 per square centimeter 18. f. oregonensis (p. 696)

10. Fronds 4-5 mm. wide 16. f. angustus (p. 694)

10. Fronds more than 5 mm. wide 11

11. Receptacles more or less cornute 12. f. cornutus (p. 692)

11. Receptacles not cornute 8. f. flabellatus (p. 689)

12. Receptacles bi- tri-furcate, wide, spreading, acute. .6. f. stellatus (p. 687)

12. Receptacles variously divided, not stellate 13

13. Receptacles very large, tumid, mucilaginous 2. f. magnificus (p. 686)

13. Receptacles narrower, mostly complanate, at times inflated or mucilaginous. ...14

14. Receptacles complanate, up to 22 cm. long 9. f. longifructus (p. 689)

14. Receptacles much shorter 15

15. Receptacles with a distinct margin free from conceptacles... 5. f. robustus (p. 687)

15. Receptacles without free margin 16

16. Receptacles simple to deeply bifurcate, blunt ...3. f. pergrandis (p. 685)

16. Receptacles variously branched acute or acuminate 17

17. Cryptostomata absent or rare 13. f. rudis (p. 692)

192-5] SetcheU-Gardner: MeJanophyceae 683

17. Cryptostomata 12-25 per square centimeter 7. f. intermedius (p. 688)

18. Fronds fruiting in dilTerent zones 15. f. limitatus (p. 694)

18. Fronds fruiting in the same terminal zone 19

19. Fronds irregularly branched and distorted 14. f. irregularis fp. 693)

19. Fronds regularly dichotomous 1. f. typicus (p. 683)

1. Fucus evanescens f. typicus Kjellm.

Fronds 15-25 cm. high, coriaceous, dichotomous, dark brown ; seg-
ments cuneate to sublinear, midrib moderately distinct, percurrent,
cryptostomata few, small ; receptacles mostly complanate, deeply
furcate, segments obovate to linear-acuminate.

Growing on rocks in the upper littoral region. Harvester and
Kadiak Islands, Alaska.

Kjellman, Om Spets. Thall., II, 1877ff;, p. 3 ; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 282 ; Gardner, Genus Fucus, 1922, p. 51,
pi. 56.

Setchell and Lawson, no. 5122 (Herb. Univ. Calif., no. 99136),
Harvester Island, Uyak Bay, Alaska ; Rigg, no. 100, Kadiak Island,
Alaska.

We have a photograph of Kjellman 's type specimen of this form.
It represents a plant considerably smaller than the plants which we
have allied with it from the Alaskan waters. No. 5122, in particular,
is much more robust than Kjellman 's description calls for. The
receptacles are much longer and wider, but this may possibly be
accounted for by difference in age. The plants collected by Rigg are
smaller and more nearly coincide with the description. This photo-
graph shows Kjellman 's plant to be about 10 cm. high, possessing
distinct midribs in manj^ of the segments, and to have relatively small
receptacles. There is a plant in the Herbarium of the University of
California, sheet no. 132618, contributed by Kjellman and collected
on Spitzbergen in "1872-73," labeled in Kjellman 's handwriting,
"Fucus evanescens Ag.," which is almost a duplicate of the type speci-
men mentioned above. There is also a plant in the same herbarium,
collected in 1868 from the same locality and determined by J. G.
Agardh as F. evanescens Ag., whose fronds are about twice as wide
as those of the Kjellman plants. Otherwise all of these three collections
of plants are very much alike.

The type specimen of F. evanescens Ag. is in the herbarium of
J. G. Agardh at Lund under no. 00299. It has been examined by one
of us (Setchell) who finds that the plant is slightly smaller than the

684 University of California Publications in Botany [Vol. 8

plant referred to in our Algae of Northwestern America, under f.
typica, from Harvester Island, Alaska. The latter plant is in the Her-
barium of' the University of California under no. 99139. It differs only
slightly from the wide form of F. evanescens from Spitzbergen, men-
tioned above, having more numerous, larger, and better developed
receptacles. There is much greater disparity of size between this speci-
men and Kjellman's f. "typica" than there is between Kjellman's f.
"typica" and his f. limitaia. If we admit the specimen determined
by J. G. Agardh from Spitzbergen as belonging with f. typica Kjell-
man, then the Harvester Island specimen which is so close to it had
probably better be allied with it at present, until more is known of
the forms from that island. However, it also seems very close to
Kjellman's f. " corfiuta," but has wider fronds and receptacles than
the type specimen of that form.

We are referring here a series of plants collected on Kadiak Island,
Alaska, by G. B. Rigg, no. 100. These plants are only slightly wider
than the type specimen of Kjellman, and have numerous well developed
receptacles. They appear to be the closest in all characters to the type,
and we are taking them to be the best representatives of f. typicus
thus far discovered in Pacific coast waters.

2. Fucus evanescens f. macrocephalus Kjellm.

Fronds subcaulescent, 12-18 cm. high, subcoriaceous, dark brown
below varying to light brown or yellowish above ; segments linear to
slightly cuneate, 5-12 mm. wide, midrib well developed, percurrent,
alae rather thin, cryptostomata absent or sparse ; receptacles distinctly
delimited, complanate in part, but mostly very tumid and mucila-
ginous, light yellow, 2-3.5 cm. long, 1-2 cm. wide, simple, retuse, or
bifid ; conceptacles conspicuous.

Growing in the middle of the littoral region. Bering Sea to
Juneau, Alaska.

Kjellman, Om Beringh. Algflora, 1889, p. 34; De-Toni, Syll. Alg.,
1895, p. 202; Saunders, Alg. Harriman Exp., 1901, p. 432, pi. 62,
fig. 1 ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 282.

Saunders (1901, p. 432) reports this form as growing at Puget
Sound, Annette Island, Wrangell, Juneau, Sitka, Glacier Bay, Prince
William Sound, Cook Inlet, Kukak Bay, and Shumagin Islands, and
states: "This is the most abundant seaweed on the northwest coast."

1^25] Setchell-Gardner : Melanophyceae 685

We have not been able to examine any of the specimens of Saunders'
collections, if such exist. There are no numbers quoted, and hence
we are not attempting to place them. Most of the plants, particularly
of the southern portion of the range, undoubtedly belong to f,
magnificus Gardner,

3. Fucus evanescens f. pergrandis Kjellm.

Fronds caulescent, robust, subcoriaceous, up to 40 cm. high, regu-
larly dichotomous, dark brown below to yellowish brown above, black
on drying ; segments elongated, linear to slightly cuneate, 1-2 cm, wide,
midrib distinct, percurrent, cryptostomata varying from none to 20
per sq, cm. ; receptacles numerous, mostly complanate, occasionally
inflated, mucilaginous, 1-2.5 cm. wide, 2.5-4 cm. long, entire to bifid ;
conceptacles numerous, large.

Growing on boulders in the middle and upper littoral belts.
Unalaska, Alaska, to Puget Sound, Washington.

Kjellman, Om Spetsb. Thall., II, 1877«, p. 3 ; De-Toni, Syll. Alg.,
1895, p. 203 ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 284 ;
Collins, Mar. Alg. A^ancouver Isl., 1913, p. Ill ; Gardner, Genus Fucus,
1922, p. 46, pi. 47.

Setchell and Lawson, nos. 3284, 4049 (Herb. Univ. Calif., nos.
99125, 99123), Amaknak Island, Alaska, and in Collins, Holden and
Setchell, Phyc. Bor.-Amer., Exsicc, no. XL VI (sub F. platycarpus) ;
Rev. Albin Johnson, no. 5701 (Herb. Univ. Calif., no. 99126), Yakutat
Bay, Alaska; Gardner, no. 2230 (Herb. Univ. Calif., no. 201136),
Juneau, Alaska; Townsend, no. 5773 (Herb. Univ. Calif., no. 99129),
Kyska Island, Alaska ; Butler and Policy, no. 20 ( Herb. Univ. Calif.,
99127), Port Renfrew, British Columbia.

The type locality of this form is Spitzbergen, where Kjellman
says it grows at a depth of several fathoms. There are three fragments
of the form in the Herbarium of the University of California under
no. 132622, from the Spitzbergen Islands, determined by Kjellman.
Although authentic, they do not entirely coincide with Kjellman 's
descriptions and type specimens. Apparently they are not representa-
tive specimens of this form.

686 University of California Publications in Botany [Vol. 8

4. Fucns evanescens f. magnificus Gardner

Fronds foliaceous, 20-30 cm. high, subcoriaceous, dichotomous or
in part secund, dark brown below, yellowish above, olive brown on
drying; segments comparatively short, linear to cuneate, more or less
twisted, 2.5-3.5 cm. wide, midrib slightly evanescent above, crypto-
stomata variable, up to 50 per sq. cm., usually conspicuous with tufts
of exserted paraphyses when young; receptacles definitely delimited,
very tumid, mucilaginous, simple to decompositely furcate, blunt or
acute, very variable in width and length, mostly yellow ; conceptacles
moderately numerous and very conspicuous.

Growing in great profusion in the middle and upper littoral
regions. Juneau, Alaska, to Puget Sound, Washington.

Gardner, Genus Fucus, 1922, p. 48, pis. 51, 52.

So far as our observations extend, this form seems to be the most
abundant of all the forms included in the above mentioned region. It
varies widely in the shape of the receptacles. Some receptacles are
simple, others decompositely furcate, many quite obtuse, others
decidedly acute, but all are definitely delimited and swollen almost to
the bursting point with mucilaginous substance. The segments are
wide and foliaceous, not narrowing at the forkings, and the tendency
to the strict habit of growth often makes them quite contorted.

One of the distinguishing characters is the sparsity of receptacles
maturing at any one time. This habit, marked in only a few of our
forms, should receive more study in the field. Our opinion is that
the fruiting season extends over a much longer period than in most
of the other forms. In the summer season, when we have observed
them, one may find on the same individual receptacles varying from
completely mature, to others just beginning to form, and many more
sterile segments. The fronds fork. One branch of the dichotomj-
metamorphoses directly into a receptacle and the other continues to
grow, and it in turn may fork one or more times without fruiting.
This makes the mature receptacle appear to be lateral in origin,
whereas in reality it originates just as all others do. This seems to be
the condition prevailing in Thuret's F. platycarpus. Plate 16 in the
Etudes Phycologiques well illustrates this condition, but illustrates
also another condition which does not seem to occur in f. magnificiis,
viz., that the suppressed fruiting segments are alternate, whereas in f.
magnificus they are secund. We do not know how regularly the con-
dition prevails in F. platycarpus, but this condition is too constantly
prevalent in f. magnificus to be overlooked as a diagnostic character.

1925] SetcheJI-Gardner: Melanophyceae 687

5. Fucus evanescens f. robusms S. and G.

Plate 107

Fronds distinctly caulescent, 15-25 cm. high, much contorted, with
distinct stout stipe and holdfast, dichotomous, terminal portions
foliaceous, olive green to dark olive brown ; segments broadly cuneate,
terminal lobes rounded, 15-28 mm. wide, midrib distinct, slightly
reduced near the apices, alae membranaceous, cryptostomata sparse ;
receptacles broad, oblong, ovate or obcordate, very variable in size,
1.5-3.5 cm. long, complanate, or much inflated, not mucilaginous, with
distinct margin free from conceptacles w^hich are large and projecting.

Growing on rocks in sheltered, shaded localities along extreme
high-tide limit. St. Paul Island, Alaska, to Friday Harbor, San Juan
Island, Washington.

Setchell and Gardner, Alg. N.W. Amer., 1903, p. 283. Fucus
platycarpus? Setchell, Alg. Prib. IsL, 1899, p. 593; Gardner, Genus
Fucus, 1922, p. 47, pis. 49, 50.

The type specimen was collected by Miss Ida M. Rogers, no. 5724
(Herb. Univ. Calif., no. 99133), at Sitka, Alaska. One of us (Gardner)
collected it at Sitka in June, 1910, when it was in full fruit, and at
Friday Harbor in July of the same year. The type and the two co-type
specimens have all of the receptacles complanate, and are apparently
immature, but in the other collections mentioned many of the recep-
tacles are much inflated. The species is one quite free from mucilage
as compared with other species of Fucus growing in the same localities
but much lower down in the littoral belt. It seems to have become
fixed as a form, and has the distinct habit, or character, of being able
to persist in the upper two feet of the littoral belt, in which habitat
it is necessarily uncovered the greater part of the twenty-four hours
each day. It inhabits rock ledges, either steep or sloping, yet it is
strictly confined to the upper, very narrow, belt. Its fronds are
lighter and somewhat more fibrous than those of any other form.

6. Fucus evanescens f. stellatus Gardner

Fronds decidedly flaccid, usually contorted, 25-35 cm. high, dicho-
tomous or subdichotomous, light brown below, yellowish above, dark
olive brown on drying, stipe and holdfast relatively small ; segments
relatively short, cuneate, wider toward the apices, 1.5-2.5 cm. wide,
midrib decidedly reduced above, cryptostomata few and inconspicuous ;

688 University of California PuhlicaUons in Botany [^^ol. 8

receptacles definitely delimited, complanate to moderately tumid,
deeompositely furcate, mostly spreading and acuminate, 3-4.5 cm.
long ; conceptacles numerous, conspicuous.

Growing on boulders, logs, etc., in the lower littoral belt. Sack-
man's Point, near Tracyton, Kitsap County, Washington.

Gardner, Genus Fucus, 1922, p. 49, pi. 53. Fuciis evanescens,
Tilden, Amer. Alg. (Exsicc), no. 235.

Tilden's no. 235 seems to belong here, although the specimen in
Setchell's copy of her American Algae is only a fragment and just
beginning to produce receptacles.

The distinguishing character of this form is the deeompositely
furcate receptacles, often widely divergent. As many as eight diver-
gent apices have been observed with a common base. The fronds are
decidedly flaccid and dissolve rather readily in fresh water after
being dried.

7. Fucus evanescens f. intermedius Gardner

Fronds foliaceous, flaccid, 12-18 cm. high, light brown to yellowish,
dark brown on drying, holdfast and stipe small, angles broad, rounded ;
segments linear, reduced above each forking, 1-2 cm. wide, crypto-
stomata 15-25 per sq. cm., midrib narrow, prominent, percurrent, alae
relatively wide, membranaceous ; receptacles broad at the base, 1-2-
furcate, acuminate or acute, complanate or tumid with mucilage,
definitely delimited ; conceptacles small, numerous.

Growing on rocks in the middle of the littoral belt. East Sound,
Orcas Island, Washington.

Gardner, Genus Fucus, 1922, p. 44, pi. 44.

This form seems unmistakably connected, through its narrow
specimens, with F. evanescens f. costatus on one side, and, on the other
side, through its widest specimens, it seems not unlike certain narrow
specimens of F. evanescens f. pergrandis. It differs from the former
in having wider segments not perceptibly reduced in width, as in
f. costatus, above the forkings, and in having wider, much more robust
and blunt receptacles. From the latter it differs in being much less
robust throughout, in having fewer cryptostomata, and in having much
more delicate and membranaceous alae.

1^^-^J SetcheU-Gardner: 3IeJanopJnjceae 689

8. Fucus evanescens f. flabellatus Gardner

Fronds 15-25 cm. high, subcoriaceoiis, siibdichotomous or, in part,
secund, light brown to yellowish brown, holdfast small, stipe small,
2-3 cm. long ; segments linear to very slightly cuneate, 10-15 mm. wide,
strict, apices rounded, slightly truncate, alae relatively thin, midrib
slightly reduced below the receptacles, cryptostomata few, small,
inconspicuous ; receptacles fusiform, 2-3.5 cm. long, simple to bifurcate,
not definitely delimited.

Growing on sandstone ledges in the middle of the littoral belt.
Bellingham, Washington.

Gardner, Genus Fucus, 1922, p. 44, pi. 43.

The majority of the plants seemed scarcely mature in July, when
they were collected, but enough of them were selected that appeared
mature to be fairly certain that the measurements given for the recep-
tacles are sufficiently accurate. The distinguishing character of the
form is the pronounced fan shape of the whole frond when spread out,
and particularly of groups of segments, due to the secund method of
branching.

The cryptostomata are relatively few and inconspicuous, and there
is approximately the same number of caeeostomata ; thus in this char-
acter this form is on the border line between F. evanescens and
F. furcatus. Probably the tendency is in the direction of the degenera-
tion of these organs. The other characters, however, seem sufficiently
well represented to ally it with F. evanescens.

9. Fucus evanescens f. longifructus S. and G.

Frond with short stipe and firm holdfast, somewhat caulescent
below, foliaceous above, up to 45 cm. high, dark olive brown ; segments
long and narrow, strict, 1-2 cm. wide, midrib moderately conspicuous,
slightly evanescent, cryptostomata absent or very sparse; receptacles
usually complanate, not definitely delimited, 2-2.5 cm. wide, 1-3 times
forked, with the divisions long, linear or sometimes recurved and
pointed, up to 22 cm. long ; conceptacles very large and scattered.

Growing on stones in the upper littoral belt. Orca to Juneau,
Alaska.

Setchell and Gardner, Alg. N.W. Amer., 1903, p. 283; Gardner,
Genus Fu<;us, 1922, p. 50, pi. 54.

690 University of California Publications in Botany [Vol. 8

Setchell and Lawson, no. 5151 (Herb. Univ. Calif., no. 99110),
Orca, Alaska. Collins, Holden and Setchell, Phyc, Bor.-Amer.
(Exsiec), no. 928; Setchell and Lawson, no. 5186 (Herb. Univ. Calif.,
no. 99109), Juneau, Alaska.

This form is unlike all other forms in the extremely long, linear-
lanceolate, regular, complanate receptacles, frequently spreading or
recurved at the apices. In external appearance it approaches F.
fwrcatus f. rigidus, but that form has an abundance of caecostomata,
the receptacles more decompositely furcate, and it is more cartilaginous.

10. Fucus evanescens f. marginatus Gardner

Fronds distinctly caulescent, 20-50 cm. high, regularly and
repeatedly dichotomous, decidedly rigid and coriaceous, much con-
torted, yellowish brown, very dark olive brown on drying, stipe terete,
tapering upwards among the branches; segments 9-15 cm. wide, strict,
not narrowing above forking, truncate, mostly long, linear, midrib
moderately prominent, slightly evanescent, alae wearing away un-
evenly, cryptostomata almost absent; receptacles 1.5-3 cm. long,
definitely delimited, single or deeply bifurcate, narrowly ellipsoidal,
marginate ; conceptacles very prominent.

Growing on rocks in the middle littoral belt. Sitka, Alaska.

Gardner, Genus Fucus, 1922, p. 42, pi. 42.

The distinguishing characters of this form are absence of con-
ceptacles from a complete margin of the receptacle, particularly of
the rounded terminal portion, coupled with extreme sparsity of both
cryptostomata and caecostomata.

It is difficult to decide to which species this form is most closely
related. It has evident affinities with Fucus evanescens Ag. and with
F. spiralis L. Its color and consistency, coupled with the slightly
vanishing midrib in the terminal segments, seem to ally it with F.
evanescent, but, if allied with this species, it would be through f.
dendroides of Stroemfelt as its nearest relative. We have not seen
the type nor any authentic specimens of f. dendroides, but since no
one has questioned the validity of the form as belonging to F.
evanescens, we are retaining it as such and grouping certain Alaska
specimens with it. The narrow, rather long, considerably contorted
and profusely branched fronds, the blunt and somewhat fusiform
receptacles, and the dendroid habit certainly suggest its close simi-
larity with f. dendroides. However, we are inclined to keep it distinct

1925] Setchell-Gardner : Melanophyceae 691

on account of its decidedly more robust habit, the practical absence
of cryptostomata, and the much lighter color than that of the other
forms within our region which we have placed with f . dendroides.

11. Fucus evanescens f. dendroides Stroem.

Fronds distinctly caulescent, dense, coriaceous, 7-15 cm. high,
rigid, terete at the base and for some distance among the main
branches, tapering upward, the narrow alae having worn away leaving
only the much enlarged midrib, flabellate-dichotomous, in part sub-
secund, profusely forked, dark olive green to olive brown, brown on
drying; segments strict, somewhat contorted, narrow, 5-8 mm. wide
in depauperate specimens, cuneate to slightly linear, widest at the
truncate apices, growing point slightly depressed, midrib prominent,
percurrent ; cryptostomata sparse or entirely absent ; receptacles very
abundant, single, retuse to bifurcate, subfusiform to ovoid, consid-
erably swollen, 1-3 cm. long; conceptacles numerous, emitting rela-
tively long fascicles of paraphyses.

Growing on rocks exposed to fairly heavy surf, in the upper and
middle littoral belts. Agattu Island to Sitka, Alaska.

Stroemfelt, Om Algenveg. vid Islands Kuster, 1886, pp. 35, 36,
pi. 3; De-Toni, Syll. Alg., 1895, p. 203; Setchell and Gardner, Alg.
N.W. Amer., 1903, p. 284; Gardner, Genus Fucus, 1922, p. 41, pi. 41.

Townsend, nos. 5755, 5756 (Herb. Univ. Calif., nos. 99105, 99106),
Agattu Island, Alaska; Gardner, no. 2227 (Herb. Univ. Calif., no.
201148), Sitka, Alaska; Setchell and Lawson, no. 4052 (Herb. Univ.
Calif., no. 99095, sub. Fucus evanescens f. bursiger), Summer Bay,
Alaska (cf. Setchell and Gardner, 1903, p. 285).

Stroemfelt does not mention the height of the type in his diagnosis,
but his illustration, natural size, is 14 cm. high. The size of our plants,
their narrow contorted segments, the character of the receptacles, the
large firm holdfast and dendroid habit, agree so well with the descrip-
tion and figures of Stroemfelt as to leave little room for doubt as to the
affinity of the two. Comparison with the type material may, however,
show that the two sets of plants are distinct, as Stroemfelt does not
mention the cryptostomata nor the caecostomata, nor does he mention
the color, which is very striking in our plants. As the plants hang on
the rocks after the receding tide, the shape, appearance, color, and
abundance of receptacles reminds one of settled swarms of Italian bees.

692 University of California Publications in Botany [Vol.8

12. Fucus evanescens f. cornutus Kjellm.

Fronds subcoriaceous, 20-30 cm. high, regularly dichotomous, dark
olive green to dark olive brown ; segments strict, linear or very slightly
cuneate in the narrower specimens, 5-8 mm. wide, apices truncate,
midrib well developed, prominent, very slightly evanescent above,
alae narrow, cryptostomata very sparse, small ; receptacles distinctly
delimited, mostly complanate, simple or bifurcate, blunt to acuminate,
apices slightly laterally reflexed, 2-4 cm. long ; conceptacles numerous,
conspicuous.

Growing on rock ledges in the lower littoral belt. Yakutat Bay,
Alaska.

Kjellman, Om Beringh. Algflora, 1889, p. 34; De-Toni, Syll. Alg.,
1895, p. 202; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 283
(in part) ; Gardner, Genus Fiicus, 1922, p. 50, pi. 55.

Rev. Albin Johnson, no. 5719 (Herb. Univ. Calif., nos. 99103,
99104), Yakutat Bay, Alaska.

This form was first discovered on Bering Island and described by
Kjellman {loc. cit.). Comparison with a photograph of the type and
with a small portion of the type specimen seems to make it fairly
certain that the two specimens collected by Albin Johnson, and cited
above, are correctly placed, although they are much larger in all parts
than Kjellman 's description calls for. More study of material in the
northern waters is highly desirable to make certain this determination.
Neither in the type nor the other material referred to is the cornute
habit very pronounced.

13. Fucus evanescens f. rudis Kjellm.

*

Fronds somewhat caulescent, coriaceous, dark brown, dichotomous ;
segments strict, cuneate, 1-2 cm. wide, reduced at the forking, terminal
lobes truncate, midrib narrow but distinct to the apices, cryptostomata
absent or rare ; receptacles relatively large, 3.5-5 cm. long, rarely
complanate, mostly very tumid and mucilaginous, not definitely
delimited.

Growing in the lower littoral and upper sublittoral regions. Bering
Sea to Skagway, Alaska.

Kjellman, Om Beringh. Algflora, 1889, p. 34; De-Toni, Syll. Alg.,
1895, p. 202 ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 282 ;

1^25] SetcheU-Gardner: Melanophyceae 693

Gardner, Genus Fucus, 1922, p. 53, pi. 57. Fuciis vesiculosics Postals
and Rupreeht, Illus. Alg., 1840, p. 12, pi. 25 (cf. Kjellman, lac. cit.).

Setchell and Lawson, no. 5121 (Herb. Univ. Calif., no. 99131),
Harvester Island, Uyak Bay, Alaska. Fucus evanescens f. maero-
ccpJialus, Setchell and Lawson, no. 5178 (Herb. Univ. Calif., nos.
99117, 99118), Orca, Alaska; A. L. Bolton (Herb. Univ. Calif., 99115),
Skagway, Alaska.

Kjellman cites plate 30 of Postels and Rupreeht, lilustrationes
algarum, as representing this form. Plate 30 is labeled "Co^nstanUnea
rosa marina," but has in addition a small plant of Fucus. This plant
has long, acuminate receptacles. Plate 25 of the same work, to which
Kjellman probably refers, is labeled Fiicus vesicidasus and has decom-
positely furcate receptacles with long, acuminate apices. The plants
referred here by Setchell and Gardner (1903, p. 282) have the char-
acteristic receptacles referred to above. Comparison with a photo-
graph of Kjellman 's type specimen hardly bears out his conclusion
regarding the illustration in Postels and Rupreeht. The fronds are
reduced in width above each forking. Some of the receptacles are
relatively wide, others are very narrow, and those of Kjellman 's type,
which we have not seen, are very blunt. Some doubts must be enter-
tained as to the limits of this form. We are referring here plants from
Orca and from Skagway, cited above, as more nearly coinciding with
Kjellman 's type.

14. Fucus evanescens f. irregularis Kjellm.

Fronds about 15 cm. high, branching for the most part irregularly
dichotomous; segments narrow, at times scarcely extending beyond
the inconspicuous midrib, apices truncate; receptacles definitely
delimited, ovate, obovate, oblong, or obcordate ; conceptacles small.

Kjellman, Om Beringh. Algflora, 1889, p. 35.

A photograph of the type specimen indicates that this form,
segregated by Kjellman on Bering Island, is hardly to be considered
a distinct entity but rather a battered and distorted plant of some
other form. To our knowledge it has not been recognized since its
original discovery ; but if a valid form, it is quite likely to occur on
the North American side of the Pacific Ocean, hence we are including
it here as a possibility.

694 University of California Publications in Botany [Vol. 8

15. Fucus evanescens f. limitatus Kjellm.

Fronds caulescent or subcaulescent, submembranaeeous, dicho-
tomous, spreading, with rounded angles, yellowish brown, darker brown
on drying; segments short, cuneate-linear, apices rounded, truncate,
midrib distinctly reduced upward, cryptostomata few ; receptacles dis-
tinctly delimited, subpedicilate, seemingly lateral, entire or bifurcate,
ovoid-ellipsoidal, 12-18 mm. long, 8-12 mm. wide.

Growing in the upper sublittoral belt. Port Clarence to Norton
Sound, Alaska.

Kjellman, Om Beringh. Algflora, 1889, p. 34 ; De-Toni, Syll. Alg.,
1895, p. 202 ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 282 ;
Gardner, Genus Fucus, 1922, p. 54, pis. 58, 59.

Kjellman, Port Clarence, Alaska; R. C. McGregor, no. 5683 (Herb.
Univ. Calif., no. 99107), Besboro Island, Norton Sound, Alaska.

The only material of which we are aware that has previously been
referred to this form since its establishment by Kjellman is that
referred by Setchell and Gardner (loc. cit.) from Norton Sound, and
the same material, two small fragments of plants, is again referred
here to the same form, though with some doubt. It is evidently a
small form, and according to Kjellman, grows in the sublittoral belt,
an unusual habitat for small forms, or even other forms outside of the
Arctic regions.

16. Fucus evanescens f. angustus Kjellm.

Fronds subdichotomously branched, 20-30 cm. high, 4-5 mm. wide,
coriaceous or submembranaeeous; segments linear or cuneate-linear,
elongated, midrib distinct below, becoming inconspicuous or almost
vanishing at the apices, cryptostomata prominent, variable in number ;
receptacles small, oblong, ovoid to broadly ellipsoid, obtuse or acute
inflated or more or less complanate ; conceptacles hermaphroditic,
antheridia at times sparse ; color black on drying.

Kjellman. Algenveg. Murm. Meer., 1877, p. 27 ; Gardner, Genus
Fucus, 1922, p. 56.

This form of evanescens, first described by Kjellman, has not been
clearly identified with any specimens from our region, but is here
included because of its close resemblance to certain of our forms, and
hence the probability of its occurrence within our waters would seem
to warrant its inclusion.

1925] SetcJieU-Gardner: Melanophyceae 695

Yendo (1907, p. 16, })1. 1, fig. 1) identifies a plant of tlie Japanese
waters with this form. Setchell and Gardner (1903, p. 284) also
identify plants from East Sound and Fairhaven, Washington, with
this form, and specimens from East Sound were distributed in Collins,
Holden and Setchell 's Phycotheca Boreali-Americana, no. 926. On
comparison of our plants with a sheet of plants in the Herbarium of
the University of California, no. 132699, collected on the Vega expedi-
dition near Tjapka and contributed and labeled by Kjellman, Fucus
evanescens f. angustus, it has seemed best to change the determination
and to place our plants under Fucus edentatus f. costatus, under which
a detailed account is given. The plants illustrated by Yendo differ
decidedly from the Kjellman specimens referred to above, particularly
in the size of the receptacles and the prominence of the midrib. It
may be doubted whether this form really extends so far south on
either coast of the Pacific Ocean as was previously supposed, but its
occurrence farther north may well be expected.

17. Fucus evanescens f. contractus Kjellm.

Fronds 5-25 cm. high, slightly caulescent, subcoriaceous, irregularly
dichotomous or subsecund, dark brown to yellowish ; segments mostly
strict, cuneate-linear below, linear above, 3-10 mm. wide, apices
truncate, midrib distinct below, vanishing above, cryptostomata mod-
erately abundant, 15-20 per sq. cm., inconspicuous; receptacles com-
planate, distinctly delimited, 1.5-3.5 cm. long, ellipsoidal or obcordate,
single or bifid ; conceptacles few, but prominent.

Growing in the littoral region. Bering Sea, Alaska.

Kjellman, Om Beringh. Algflora, 1889, p. 34 ; De-Toni, Syll. Alg.,
1895, p. 202 ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 284 ;
Gardner, Genus Fncus, 1922, p. 55.

Setchell, nos. 5239, 5252 (Herb. Univ. Calif., nos. 99097, 99101),
St. Michael, Alaska; McGregor, nos. 5673, 5679 (Herb. Univ. Calif.,
nos. 99099, 99100), Golofin Bay, Alaska. Not Gardner, no. 90 (Herb.
Univ. Calif., no. 99096), Whidbey Island, Washington, sub F.
evanescens f. hursigcr (cf. Setchell and Gardner, 1903, p. 285),

Kjellman does not mention in his description of this form the
decided and sudden difference between the width of the segments and
the receptacles which they bear. This difference makes the receptacles
appear stipitate, since they are over twice as wide as the base of the
segments. Presumably this is the character upon which the form is
based.

696 Universitij of California Publications in Botany [Vol. S

No. 5252 of Setcliell (Herb. Univ. Calif., no. 99101) from St.
Michael, Alaska, quoted above, is as near the type as could be desired
in every particular. The plants are about 10 cm. high, thus repre-
senting the upper limits in size as stated by Kjellman. The plants
under the other numbers referred to are all larger in all parts than
the measurements given by Kjellman, but they seem too closely linked
to the type. to warrant separation without further studies of quantities
of material in the field. There is not the sudden widening of the
receptacles in any of the latter forms as in the type. The receptacle
character seems m.ore closely allied to that of i.macrocephalus.

, 18. Fucus evanescens f. oregonensis Gardner

Plate 106

Fronds distinctly caulescent, 15-20 cm. high, subcoriaceous, dicho-
tomous below, subsecund above, yellowish brown ; segments narrow,
sublinear or slightly cuneate, 4—7 mm. wide, flaccid, smooth, midrib
narrow and distinct below, slightly evanescent above, cryptostomata
small, inconspicuous, 20-25 per sq. cm. ; receptacles tumid, distinctly
delimited, much wider than the segments bearing them, single or bifid,
broadly fusiform, apices very obtuse ; conceptacles moderately
conspicuous.

Growing on floats, piles, stones, etc., in the middle and upper
littoral belts. East side of Coos Bay, opposite North Bend, Oregon.

Gardner, Genus Fucus, 1922, p. 40, pi. 40.

This form seems decidedly distinct from all other forms previously
described. Judging from its size, color, character of the midrib, and
its general consistency, its nearest relatives are to be found in the
evanescens group, and close to forma flabellatus. The plants are
smaller, segments narrower and regularly dichotomous instead of
subsecund, and the receptacles are shorter and blunter than in that
form. The only locality from which it has been reported is the one
given above, where it grew in profusion. Doubtless it may be found
in other localities on the same bay.

ii'2o] Setchell-Gardner : Melanophyceae 697

19. Fucus evanescens f. ecostatus Gardner

Fronds submembranaceous, 12-18 cm, high, dichotomous or sub-
sec und, yellowish brown, darker brown on drying, stipe and holdfast
fragile ; segments strict, cuneate, 5-12 mm. wide, growing point incon-
spieiioiis, midrib very inconspicuous throughout, nearly vanishing in
the terminal lobes, alae thin and membranaceous, cryptostomata very
few and inconspicuous; receptacles variable, inflated or complanate,
mostly narrow, bifid, acuminate ; coneeptacles inconspicuous.

Growing on sandstone, wooden floats, etc., in the upper littoral
belt. Coos Bay, Oregon.

Gardner, Genus Fucus, 1922, p. 39, pi. 39.

Forma ecostatus is quite abundant at various localities from the
mouth of Coos Bay up to its headwaters where several streams of fresh
water flow^ in. This form seems to be definitely fixed, and does not
appear to grade into any of the forms that are found growing outside
of the bay. More investigation, however, may show that it is only a
depauperate form of some definite species; on the other hand, it may
prove to be a distinct species. It varies considerably in size and width
of the fronds. The receptacles, though mostly pointed, are at times
wide and blunt ; at least there are specimens growing in the same
habitat M'hich have the general characters but differ only in the char-
acter of the receptacles, and in such cases the segments are generally
wider. They are connected, however, by almost imperceptible grada-
tions. The size of the plants may be somewhat influenced by the sub-
stratum, mostly wood and soft sandstone, neither of which is suited
to holding plants for a very great length of time. They usually do not
persist in such localities beyond a single fruiting period. If the
vanishing of the midrib is to be considered as having any diagnostic
value, this form eminently belongs with the evanescens group. The
midrib in some specimens is scarcely discernible in any part of the
frond, and in all it vanishes in the terminal segments.

698 University of California Putlications in Botany [Vol. 8

20. Fucus evanescens f. cuneatus Gardner

Fronds small, 5-10 cm. high, membranaceous, regularly dicho-
tomous, olive brown ; basal segments decidedly cuneate, terminal seg-
ments almost linear, 4—8 mm. wide, apices truncate, midrib incon-
spicuous, slightly evanescent above, cryptostomata few, inconspicuous ;
receptacles complanate or occasionally swollen, deeply bifid, usually
acuminate, not definitely delimited, large relative to the size of the
plant ; conceptacles relatively large and numerous.

Growing on sandstone in the upper littoral region. Near Empire,
Coos Bay, Oregon.

Gardner, Genus Fucus, 1922, p. 39, pi. 38.

One becomes somewhat exasperated in attempting to deal with such
forms as the one mentioned above. Ordinarily it might be considered
a depauperate form of some other well developed and characteristic
form. There were several square rods of fruiting plants growing on
very gently sloping sandstone near high-tide level. There were no
other plants growing near, and there was seemingly no reason why,
from the standpoint of anchorage, they should not grow much farther
down in the littoral belt. The reduced size could not be due to super-
abundance of fresh water, for several much larger forms were found
growing from three to six miles farther up the bay where the water
is much less saline. Many plants have well-developed mature
receptacles.

Its general characters seem to favor its being grouped with F.
evanescent, although it will fit practically as well into the edentatus
group. AVe are placing it with the former until more can be learned
of its life-history.

21. Fucus evanescens f. nanus Kjellm.

Fronds minute, 4-5 cm. high, 2-5 cm. wide, without distinct stipe
and usually without holdfast, branching very irregularly, considerably
twisted, color variable, yellowish brown to olive green, fruiting
sparsely ; segments extremely variable in shape and size, midrib slightly
developed below, usually vanishing above, cryptostomata few or
absent ; receptacles entire or bifid, ovate-ellipsoidal or with divergent,
blunt or acuminate apices.

1925] Setchell-Gardner: Melanophyceae 699

Growing mostly on mud flats at extreme high-tide limit, or even
considerably above, among various salt marsh plants, covered more
or less by fresh water. Yakutat Bay, Alaska, to Puget Sound,
Washington.

Kjellman, Om Spetsb. Thall., II, 1877a, p. 4; De-Toni, Syll. Alg.,
1895, p. 203 ; Setchell and Gardner, Alg. N.W. Amer., 1903, p. 285 ;
Gardner, Genus Fiicus, 1922, p. 37, pi. 37.

Rev. Albin Johnson, no. 5712 (Herb. Univ. Calif., no. 99121),
Yakutat Bay, Alaska; Gardner, no. 2256 (Herb. Univ. Calif., no.
201154), Sitka, Alaska; T. C. Frye (Herb. Univ. Calif., no. 132931),
Tokeland, Washington.

After considerable observation in several different localities, we
have come to the conclusion that f. nanus cannot be considered as a
distinct form of any one species, but is in reality composed of a mix-
ture of dwarfed plants of whatever species or forms of species that
happen to inhabit the region. The dwarfs are always found on mud
flats, salt marshes, and quite commonly at the mouths of rivers and
smaller streams where such mud flats and marshes may occur. At
extreme high tides, frequently, a great abundance of plants, of what-
ever species of Fucus happens to be growing in the vicinity, are thrown
up and lodge among other plants that grow there. The spores are shed,
and, being in a moist place, they germinate and persist for a longer
or shorter time. The plants are always much gnarled and distorted,
and rarely come to fruit. At the mouth of the Indian River at Sitka,
Alaska, is a favorable place for the study of such dwarfed plants.
There are many different forms of Funis growing in the vicinity, and
a great many plants are constantl}^ being cast up on the gradually
sloping flat at the mouth of the stream. There are enormous numbers
of dwarfed plants in all possible stages of development and distortion.
One can come to no other conclusion than that these are dwarfed
plants, that they are the offspring of whatever plants may by chance
be thrown up there, and that their specific identity cannot be traced,
growing as they do in such abnormal environmental conditions.

5. Fucus nitens Gardner
Plate 102

Fronds moderately flaccid, coriaceous, 30-40 cm. high, regularly
dichotomous, dark brown to yellowish brown, very dark on drying ;
segments strict, 9-18 mm. wide, smooth and glossy, cuneate, terminal

700 University of California Publications in Botany [Vol. 8

wider, slightly ovate, truncate, growing point very slightly depressed,
midrib moderately prominent, cryptostomata absent or very sparse ;
receptacles yellowish, complanate when young, frequently becoming
tumid at maturity, bi- tri-furcate, 4-7 cm., rarely 10 cm. long;
conceptacles large.

Growing on boulders and rock ledges. San Francisco Bay, Cali-
fornia.

Gardner, Genus Fiicus, 1922, p. 26, pis. 18, 19.

This relatively small group of Fucus plants, although very much
circumscribed in its distribution, seems so distinctly marked off in its
combination of characters from other species, particularly from those
in the southern portion of our range, that it is worthy of specific rank.
The combination of characters that distinguish this species consists
of the following : relatively long and narrow, smooth and glossy fronds,
strict, even overlapping habit of the terminal and subterminal seg-
ments, the dark brown color with yellowish receptacles at maturity,
absence of caecostomata, and the cryptostomata, when present, incon-
spicuous. Its affinities with F. edentattis may possibly be traced, but
they seem too remote to merit much serious consideration.

66. Pelvetia Dec'ne and Thur.

Fronds of tough, firm consistency, flexuose, the whole plant when
young, and the young terminal growing parts considerably flattened,
without midrib, becoming more or less terete with age, especially
toward the base, arising from a solid, disk-shaped holdfast; growing
region apical ; branching dichotomous, usually abundant ; reproductive
organs, antheridia and oogonia, developed in conceptacles limited to
the terminal, metamorphosed parts of the branches, the receptacles ;
oogonia developing two viable gametes, or eggs ; monoecious.
Decaisne and Thuret, Rech. sur Antherid., 1845, p. 12,
This genus was founded on the Fucus canaliculatus of Linnaeus
(Syst. Nat. II, 1759, p. 716) based largely upon the fact that the
oogonium produces but two viable eggs instead of eight, the character-
istic number for the genus Fucus. Eight nuclei are formed in the
oogonium, but six of them are extruded between the two eggs at the
time of their formation and become functionless. The egga are
fertilized outside of the oogonia as in the genus Fucus.

1^25] Seichell-Gardner : MeJanophyceae 701

Pelvetia fastigiata (J. Ag.) De-Toni

Mature fronds terete below, compressed above, narrowly linear,
profusely branched, branching often beginning just above the base,
diehotomous, fastigiate, 2-4 dm. (up to 10 dm.) high, 5-10 mm. diam.
below, tapering above ; color yellowish brown to dark olive green ; the
central portion of the fronds consisting of long, branched, colorless
filaments with thick gelatinous walls, surrounded by a layer of large
parenchymatous cells, becoming smaller toward the surface consisting
of a single layer of small, compact cells ; oogonia on one-celled stalks,
broadly elliptical, dividing longitudinally or occasionally obliquely
into two (very rarely into three) oospheres; antheridia very numerous,
on short branches in the basal portion of the conceptacles, paraphyses
not at all, or only slightly, extruded ; receptacles moderately swollen,
occasionally forked, variable in shape, some conical, others fusiform.

Always growing on rocks, in the middle of the littoral belt.
Extending from Coos Bay, Oregon, to the w^est coast of Lower Cali-
fornia (Ensenada).

De-Toni, Syll. Alg., vol. 3, 1895, p. 215; Gardner, Nuclear Extru-
sion, 1910, p. 130, pi. 17, figs. 11-16; Collins, Holden and Setchell,
Phyc. Bor.-Amer. (Exsicc), nos, 176 and CXIII. Fucus (Pelvetia.)
fastigiata Farlow, Anderson and Eaton, Alg. Exsicc. Amer.-Bor.,
no. 110. Fiwus fastigiatus J. Ag., Symb. I, 1841, p. 3 ; Harvey, Ner.
Bor.-Amer., 1851, p. 68, pi. 3 A.

The plants of Pelvetia fastigiata vary considerably in size. So
far as has been observed, they reach their greatest length in southern
California, where at San Pedro plants 10 dm. long have been observed.
Usually when the plants are growing w^here they are subjected" to a
heavy surf, the fronds become much thicker and much more rigid and
less branched than those growing in more sheltered situations. This
is the typical condition of plants on the west end of the Monterey
peninsula, the type locality. Here the branches are w^despreading
and the angles much rounded, and some of the fronds measure 2 cm.
wide near the base.

702 University of California Publications in Botany [Vol.8

Pelvetia fastigiata f, gracilis S. and Gr.

Plate 78a

Plants profusely and dichotomously branched, central branches
becoming much longer than those on the outside of the mass, giving
to the whole plant a more or less spherical outline ; frequently branches
arise from near the base and develop a mass of short branchlets ;
fronds slender, the terminal branchlets 1-3 mm. diam. ; fruiting in
the summer and autumn.

Growing in the middle of the littoral belt. Carmel Bay, Pacific
Grove, and Santa Catalina Island, California.

Setchell and Gardner, in Gardner, New Pac. Coast mar. alg., I,
1917, p. 386.

In the above mentioned localities, plants grow in groups, quite
separate from the typical P. fastigiata. The plants as a whole are to
be distinguished chiefly by their very slender fronds throughout as
compared with the typical form.

67. Pelvetiopsis Gardner

Fronds arising from a disk-shaped holdfast, frequently the hold-
fasts confluent, cylindrical at the base soon becoming flattened and
more or less concave-convex and contorted, ecostate, receptacles con-
spicuous, terminal, almost cylindrical and acuminate or flattened and
blunt ; oogonium producing but one viable gamete, or egg, the other
seven nuclei extruded in a single, small, non-viable sphere cut off from
the lower part of the oosphere ; cryptostomata sparse on the young
plants ; hermaphroditic.

Gardner, Nuclear Extrusion, 1910, p. 127, New Fucaceae, 1913,
p. 320.

This genus is well marked both macroscopically and microscopically.
It is more closely related to the genus Pelvetia than to the genus Fucus
in its gross morphological characters, but differs from both in the
production of but a single viable gamete in the oogonium. In this
latter respect it is like the genus Hesperophycus, but differs decidedly
from that genus in its gross morphological characters.

19--5J Setchell-Gard^ier: Melanojjhyceae 703

Pelvetiopsis limitata (Setchell) Gardner

Two fairly well defined forms of this species have been found along
the California coast, but not jjrowing together.

Pelvetiopsis limitata f. typica Gardner

Plate 46, fig. 6

Fronds narrow, 8-10 cm. long, when young beginning to fruit
before branching and when only 2-3 em. long; receptacles nearly
cylindrical, often long-attenuate ; conceptacles conspicuous.

Growing along high-tide level or above, where the surf dashes
against rocks. Ranging from Vancouver Island to central California.

Gardner, Nuclear Extrusion, 1910, p. 127, pi. 16, figs. 1-7, New
Fucaceae, 1913, p. 321, pis. 38, 39, figs. 8-30. Pelvetia fastigiata f.
limitata Setchell, in Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. 1238. Pelvetia fastigiata Tilden, Amer. Alg. (Exsicc),
nos. 233 and 517,

Pelvetiopsis limitata f. lata Gardner

Fronds comparatively robust, 15-18 cm. long, frequently much
contorted, receptacles flat, wide and mostly blunt, in age often inflated.

Growing usually near high-tide limit. Observed along the coast of
central California, from Tomales Point to Plalf Moon Bay.

Gardner, Nuclear Extrusion, 1910, p. 127, New Fucaceae, 1913,
p. 321, pis. 40, 41, figs. 31-43; Collins, Holden and Setchell, Phyc.
Bor.-Amer. (Exsicc), no. CXIV.

68. Hesperophycus S. and G.

Fronds complanate, arising from a solid, disk-shaped holdfast,
dichotomous, with distinct percurrent midrib, a short compressed stipe
and linear segments, oogonia developing but one viable gamete, or
egg, seven nuclei falling to the bottom of the oosphere being cut off and
forming a non-viable part.

Setchell and Gardner, in Gardner, Nuclear Extrusion, 1910, p. 127,
New Fucaceae, 1913, p. 317.

704 University of California Publications in Botany i'^oi.. 8

Hesperophycus Harveyanus (Dec'ne) S. and G.

Frond 2—1 dm. high, dichotomous, usually one of the dichotomies
outgrowing the other, forming several main axes, some of the laterals
with several branches close together becoming flabellate; color dark
olive green, or yellowish brown when in exposed situations ; segments
5-10 mm. wide, with a row of cryptostomata extending longitudinally
on either side of the midrib, having an abundance of extruding para-
physes ; receptacles simple but in part bifurcate, 1.5-3 cm. long,
cylindrical to ellipsoidal.

Growing on rocks in a restricted belt a few feet below high-tide
limit. The present known range is from Monterey Bay to Lower
California (Ensenada).

Setchell and Gardner, in Gardner, Nuclear Extrusion, 1910, p. 127,
pi. 16, figs. 8-10, NW Fucaceae, 1913, pp. 317-320, pis. 36-37 ; Collins,
Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no. CXII. Fucus
Harveyanus Dec'ne, Voyage Venus, 1864, p. 9, Altas, 1846, pi. 4.
Fucus ceranoides f. Harveyanus Tilden, Amer. Alg. (Exsicc), no. 339.

FAMILY 25. SAEGASSACEAE de-toni (lim. hut.)

Fronds differentiated into holdfast, stipe and more or less indis-
tinguishable from it the main rhachis of the frond which bears the
variously modified or differentiated branches arising on all sides of
the rhachis, and usually far surpassing it in length ; stipe and main
rhachis perennial, the growing region at the apex of the rhachis some-
what obscure, primary branches fruiting, then disintegrating back to
the rhachis, leaving scars on the constantly elongating axis; oogonia
usually few and large ; receptacles small, cylindrical, or slightly
complanate, terminal or subterminal.

De-Toni, Syst. Uebers. Fucoid., 1891, p. 174.

Key to the Genera

1. Vesicles absent in our species 69. Blossevillea (p. 705)

1. Vesicles present 2

2. Vesicles single 73. Sargassum (p. 711)

2. Vesicles seriate 3

3. Stipe short in our species, merging into the hapteres , . . . 70. Cystophyllum (p. 706)

3. Stipe long, persistent 4

4. Stipe decidedly angular 72. Cystoseira (p. 70S)

4. Stipe terete, more or less geniculate 71. Halidrys (p. 707)

1925] SetcheU-Gardner : MelanopJiyceae 705

69. Blossevillea Dec 'ne (orthog. mut.)

Erect fronds either arising directly from a mass of branched
hapteres or from other prostrate "runners" attached by branched
hapteres, usually divided into several main portions which are terete
or slightly flattened and two-edged ; these main divisions usually
developing numerous side branches, either on all sides or pinnate
along the two edges of flattened forms, in such species the frond
bending at each node, giving the whole frond a marked geniculate
appearance ; the terminal ramuli give rise to the receptacles ; air
vesicles may or may not be present, when present non-septate.

Decaisne, Sur Thalass., 1840, p. 409 ; Gardner, New Fucaceae, 1913,
p. 325, et seq.

In the last mentioned publication, a discussion is given as to the
reasons for the use of the term, Blossevillea of Decaisne, instead of
CystopJiora of J. G. Agardh, for the plant we are here including in
our account.

Blossevillea Brandegeei S. and G.

Erect fronds developing from short, irregularly branched, pros-
trate filaments attached to rocks by small, disk-shaped holdfasts ;
fronds cylindrical or subcylindrical at the base, gradually becoming
flattened, with a midrib, occasionally forked, 23-35 cm. high, 4-6 mm.
wide, alternately pinnatifid, producing a geniculate appearance
throughout the whole plant ; pinnae arising from the margins only,
slightly midribbed, twice or thrice pinnatifid, 5-8 cm. long; vesicles
absent ; dried plants black and very brittle ; receptacles terminating
the ultimate ramuli, 10-15 mm. long; conceptacles mammiform,
usually in two fairly well defined rows on each side of the slightly
flattened receptacles ; one oosphere in each oogonium ; monoecious.

Known only from Guadalupe Island, off the coast of Lower
California.

Setchell and Gardner, in Gardner, New Fucaceae, 1913, p. 325,
pi. 46.

We have no data as to the abundance of this species, nor as to its
habitat. It has not been reported since its discovery by Dr. Edward
Palmer in 1897, on the above mentioned island.

706 University of California Publications in Botany [Vol. 8

70. Cystophyllum J. Ag.

Fronds arising from a broad, flattened, irregular, parenchymatous
holdfast, several fronds apparently arising from the same holdfast,
the main stipe soon disappearing as such ; main fronds comparatively
long, giving rise to very small, flattened branches, and to numerous
long, filiform, nearly terete branches; these long branches bearing
numerous, short, lateral branches, variously dissected into ramuli
bearing small, usually solitary, vesicles beyond which the receptacles
develop oosphores single in an oogonium ; plants perennial.

J. G. Agardh, Sp. I, 1848, p. 228.

When J. G. Agardh founded the genus Cystophyllu-m {loc. cit.),
he enumerated and described nine species, all of which had been
previously published under other genera. The first species mentioned
in his account, the type of the genus, was C. onustum founded on
Fucus o-nu^tus of Mertens (Sur plusieurs espec. du Fu-cus, 1819, p.
183) and the Sargassum onustum of Agardh (Sp. Alg., 1820, p. 32).
He placed it in the family Fucaceae, in which it was retained by Kjell-
man in Engler and Prantl (1893, pp. 279 and 283). De-Toni (1895)
shifted the genus to the much modified family, Sargassaceae, in which
we are retaining it.

Cystophyllum geminatum (Ag.) J. Ag.

Stipe arising from a solid, conical, fibrous disk, at first producing
numerous, long, cjdindrical, filiform with alternate branches, 6-24 dm.
long, later producing several alternate side branches similar to the
main stipe, these in turn producing the filiform branches ; the filiform
branches produce numerous, short, alternate, or fasciculate branches
producing in turn, near their bases a few, alternate, linear or spatulate,
ribless branches (leaves), and toward their apices ramuli terminated
mostly by single, broadly fusiform air vesicles ; these at first apiculate,
later the receptacles developing from the apiculate part ; plants
dioecious.

Growing in the lower littoral and upper sublittoral belts. Extend-
ing from the Bering Sea, Alaska, to Puget Sound, Washington.

J. G. Agardh, Sp. Alg., vol. 1, 1848, p. 232 ; Setchell and Gardner,
Alg. N.W. Amer., 1903, p. 285. Cystophyllum Lepidium Harvey,
Coll. Alg. Vancouver Island, 1862, p. 163 ; Saunders, Alg. Harriman
Exp., 1901, p. 432; Tilden, Amer. Alg. (Exsicc), no. 232. Cystoseira

19-^] SetcheU-Gardner: Melanophyceae 707

Lepidium Rupreclit, Tange Oeliot. Meeres, 1851, p. 347 ; Collins,
Holden and Setchell, 'Phyc. Bor.-Amer. (Exsicc), no. XLVII. Cysto-
seira gemimita Agardh, Syst. Alg., 1824, p. 286.

Since publishing our account in the Algae of Northwestern
America, we have not been able to secure any additional data by which
to separate the four species of the genus Cystaphyllum, accredited to
our region and which we united into one species under the foregoing
combination.

71. Halidrys Lyngb.

Plants consisting of a solid holdfast, a stipe, flattened and terete
fronds ; the flattened fronds alternate on the stipe, these metamorphos-
ing into pinnately branched filiform branchlets; cysts in a series,
developing in the flattened fronds, leaving a margin on either side of
the series; these fronds terminating in branched receptacles; one egg
cell in each oogonium ; monoecious or dioecious ; perennial, the fruiting
fronds dying back to the stipe each year.

Lyngbye, Hydrophyt. Dan., 1819, p. 37.

But two species of this genus, as restricted by Greville (Alg, Brit.,
1830, p. xxxiv), are at present known, H. siliquosa (L.) Lyngbye, on
the Atlantic coast of Europe, and the following species on the coast
of southern California.

Halidrys dioica Gardner

Stipe arising from a flattened, solid, warted ba.se, terete, bent at the
"nodes," 40-50 cm. long, 4—6 mm. diam., solid, flexible, not forked;
primary pinnae annual, arranged alternately upon the stipe, always
arising just below the growing point, 10-18 dm. long, flat and linear
at the base, pinnately branched above, the linear portion 10-25 cm.
long, 8-12 mm. wide ; secondary branches or pinnules sessile, flat, acute
or obtuse at the apex, the oldest ones entire, the younger ones variously
notched and incised, the incisions increasing in depth, and the seg-
ments increasing in length toward the apex of the branch, the upper-
most becoming pinnately branched ; the upper secondary branches
develop flattened, margined, lanceolate, acuminate, short pediceled air
vesicles, divided into 5-9 chambers, slightly constricted at the divi-
sions, the oldest rounded at the base and mucronate at the apex ; the
apex of the younger ones developing farther into branched filiform
receptacles and these in turn bearing many conceptacles ; plants peren-

708 University of California Publications iri Botany [Vol. 8

nial, dioecious, the two sexes being indistinguishable bj- external
morphological characters; color varying from dark brown to olive
green.

Growing in the littoral and sublittoral belts on exposed rocks, or in
the sublittoral belt in quiet water. San Pedro, Redondo and at Avalon,
Santa Catalina Island, southern California and at least as far south
as Ensenada, Lower California.

Gardner, New Fucaceae, 1913, p. 323, pis. 42-45 ; Collins, Holden
and Setchell, Phyc. Bor.-Amer. (Exsicc), nos. CXVI, CXVII.

72. Cystoseira Ag.

Frond differentiated into holdfast, stipe and branches of various
orders ; holdfast somewhat fibrous and woody, solid or more or less
cavernous, conical in general outline ; usually each holdfast giving rise
to one main stipe, decidedly angular, with terminal meristem, and
giving rise on all sides to a few main divisions having terminal grow-
ing points ; the primary branches arising from the main stipe and its
divisions and far surpassing them in length, flattened and pinnately
lobed at the base, resembling certain fern leaves, but divided above
into more or less filiform branches and branchlets bearing the recep-
tacles at their termini, and vesicles, usually seriate, in branchlets of
different orders ; oosphere single in the oogonium ; plants monoecious
or dioecious, perennial.

Agardh, Sp. I, 1820, p. 50.

Agardh enumerated fifty-seven species under his newly founded
genus. Cystoseira ericoides came first in the list and may be con-
sidered the type of the genus. It was founded on Fucus ericaides of
Linnaeus (Sp. pi. 1763, p. 1631). Agardh 's combination is still
retained and the plant is well illustrated by Harvey in Phycologia
Britannica (vol. 3, pi. 265).

Key to the Species

1. Fronds robust throughout, 2-4.5 m. long 1. C. osmundacea (p. 709)

1. Fronds shorter, up to 1.5 m. long, more slender 2

2. Fructiferous ramuli aggregated into short, dense clusters, vesicles 6-8

mm. diam 2. C. Setchellii (p. 710)

2. Fructiferous ramuli not densely aggregated, vesicles 2-3 mm. diam

3. C. neglecta (p. 710)

1925] Setchell-Gardner: Melanophyceae 709

1. Cystoseira osmundacea (Menz.) Ag.

Fronds robust, perennial, up to 6 m. long; stipe nearly triangular
in cross-section, somewhat geniculate, having numerous prominent
scars caused by the disintegration of the primary branches; flattened
portion in the lower part of the primary branches with deep rounded
sinuses and jjrominent pinnae, rounded at the outer ends, very closely
resembling the fronds of certain ferns, gradually reduced in size
toward the outer end of the rhachis and finally merging into the long,
branched, filiform branches ; vesicles 5-12 seriate, 4-6 mm. diam.,
mostly in the branches having only the receptacular ramuli beyond;
receptacles conspicuous, abundant, often much branched, terminating
the ramuli; plants dioecious.

Growing in tide pools in the lower littoral belt and in the upper
sublittoral belt, usually in moderately sheltered localities.

Central Oregon (Coos Bay) to Lower California (Ensenada).

Agardh, Sp. I, 1820, p. 69 ; Gardner, New Fucaceae, 1913, p. 333.
Fucus osmundaoeus Menzies, in Turner, Hist. Fuc, vol. 2, 1809, p. 92,
pi. 105.

The type locality of the species is Port Trinidad, Humboldt County,
California. Two forms, fairly constant in structure, have been noted
as follows :

Cystoseira osmundacea f. tjrpica Gardner

This form comprises the shorter specimens, more robust throughout,
usually with fewer (5-8) vesicles in a series. It grows, as a rule, in
more exposed localities in tide pools along low-tide level, and seems to
be limited largely to the central and northern parts of the range.

Gardner, New Fucaceae, 1913, p. 335, pis. 51, 52.

Cystoseira osmundacea f. expansa (Ag.) Setchell

Forma expansa includes the more delicate and slender, much
elongated (up to 6 m. long) specimens with smaller vesicles and a
larger number (7-12) in a series. It is most luxuriant in the southern
part of the range.

Setchell. in Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. XLVIII ; Gardner, New Fucaceae, 1913, p. 336, pi. 53.
Cystoseira expansa Agardh, Syst. Alg., 1824, p. 290. Cystoseira
osmundacea Collins, Holden and Setchell, loc. cit., nos. XCVI and
CXV a, not h. Both distributions are small parts of plants.

710 University of California Publications in Botany [Vol. 8

2. Cystoseira Setchellii Gardner

Stipe developing from an irregularly conical-shaped solid base,
flexible, cartilaginous, irregularly and sparingly forked, 2-3 dm. long,
8-10 mm. diam., triangular in cross-section, giving rise to numerous,
alternate, flattened branches at short intervals, always from just below
the growing point; flattened branches coriaceous, annual, deciduous,
thus producing prominent scars on the stipe, serrated at the base,
deeply and pinnately lobed or divided in the middle, 3-4 times pin-
nately divided into filiform ramuli at the apex; the pinnate segments
often in turn becoming serrated, lobed, and divided; color of the
living plant dark brown, turning black when dry ; receptacles terminal
on the ramuli, 15-25 mm. long; vesicles 1-3 seriate, terminal one fre-
quently mucronate, spherical, containing internal, longitudinal, white
fibers, without a margin, 6-8 mm. diam., situated at the base of the
serrated segments, or at the base of the filiform branches ; whole plant
6-8 dm. long; perennial; dioecious, the two sexes being very similar
in external appearance.

Growing in the sublittoral belt. San Pedro and Redondo, southern
California.

Gardner, New Fucaceae, 1913, p. 329, pis. 47-50. Cystoseira osmiin-
dacea, Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no.
CXV b.

3. Cystoseira neglecta S. and G.

Stipe arising from an irregular, conical-shaped, somewhat fibrous
holdfast, flexible, rather sparingly forked, 5-15 cm. long, 4-6 mm.
diam., triangular in cross-section, giving rise to flattened branches
alternately arranged just below the terminal growing point ; flattened
branches coriaceous, flexible, becoming rigid and brittle when dry,
dying back after fruiting, leaving a permanent angular scar on the
stipe, when young divided into rounded alternate lobes by a deep
sinus rounded at the base ; the lobes of the -lower and middle portion
of the branches becoming 2-3 times pinnatified, and the upper lobes
becoming repeatedly divided into cylindrical branches terminating in
numerous receptacles; air vesicles developing on separate filiform
branches, solitary or few in a series but separated 1-3 mm. from
each other, spherical, smooth, 2-3 mm. diam., whole plant 4—7 dm.
long ; color of living plant light brown, turning black when dry ;
perennial ; dioecious.

19-5] SetcheU-Gardner: Melanophyceoe 711

Growing in the upper sublittoral belt. Avalon, Santa Catalina
Island, California.

Setchell and Gardner, in Gardner, New Pac. Coast Mar. Alg. I,
1917, pp. 388-390, pis. 34, 35.

73. Sargassum Ag.

Plants attached by a more or less irregular, warty, solid, paren-
chymatous base, or by numerous' stolon-like growths from the main
axis, at most but a few decimeters high ; frond variously branched, con-
sisting of a main basal part, or stipe-like portion terete or slightly
angled remaining short with transition region near the end, bearing
few to many branches on all sides, alternately arranged ; these main
branches more or less elongated, filiform, terete or slightly angled,
more or less branched, developing short, flattened, generally more or
less horizontal, sterile branches (leaves) with midrib and cryptosto-
mata ; the receptacles and vesicles, variously arranged and modified,
developing in the axils of these leaves ; plants monoecious or dioecious ;
oogonia large, containing a single egg.

Agardh, Sp. I, 1820, p. 1.

In treating of the species of Sargassum found within our territory,
we are accepting the genus as established by C. A. Agardh in Species
Algarum in 1820, following the arrangement and restriction of the
species as proposed by J. G. Agardh in his Species Sargassorum. 1889,
and further modified and amplified by A. Grunow in Additamenta,
1915, 1916. There is still much need of further investigation of the
species growing in the Gulf of California and around the islands in
the Pacific Ocean off the coast of Mexico.

Key to the Species

1. Leaves on all orders of branches decompound 1. S. Palmeri (p. 712)

1. Leaves simple 2

2. Vesicles, leaves, and receptacles intermixed, forming a heteroclyte cyme.. 3
2. Vesicles, leaves, and receptacles not intermixed, not forming a hetero-
clyte cyme 6

3. Receptacles two edged, apex and margin serrate-dentate.. 5. S. Bryantii (p. 714)

3. Receptacles terete 4

4. Leaves slightly flattened to filiform 3. S. guardiense (p. 713)

4. Leaves flattened, asymmetrical, dentate 5

5. Vesicles and receptacles not spinose 2. S. acinacifolium fp. 713)

5. Vesicles and receptacles more or less spinose 4. S. lapazeanum (p. 714)

6. Receptacles ancipitally angulate, margins serrate-dentate, conceptacles

on lateral surfaces 7

6. Receptacles cylindrical, more or less papillate, conceptacles on all sides .11

712 Vniversity of California PuhJications in Botany [Vol.8

7. Cryptostomata conspicuous 8

7. Cryptostomata minute or none 9

8. Branches and branchlets muricate 6. S. horridum (p. 715)

8. Branches and branchlets smooth 7. S. Marchantae (p. 715)

9. Leaves lanceolate, acuminate, spinulose-dentate 8. S. Liebmanni (p. 716)

9. Leaves more or less asymmetrical 10

10. Primary branches long and very slender, branchlets sparse

10. S. insulare (p. 717)

10. Primary branches more robust, branchlets abundant

9. S. Agardhianum (p. 716)

11. Cryptostomata absent 12

11. Cryptostomata sparse or abundant 13

12. Leaves blunt 11. S. Brandegeei (p. 718)

12. Leaves acute 16. S. herporhizum (p. 720)

13. Fronds muricate 13. S. polyacanthum f. americanum (p. 718)

13. Fronds smooth 14

14. Leaves narrowly lanceolate to jBliform, ecostate, vesicles ellipsoidal....

14. S. Johnstonii (p. 719)

14. Leaves lanceolate, with percurrent midrib, vesicles subspherical to

ovate 17. S. paniculatum (p. 721)

14. Leaves lanceolate, with percurrent midrib, vesicles spherical 15

15. Crj^ptostomata sparse, inconspicuous, receptacles acuminate

.12. S. sinicola (p. 718)

15. Cryptostomata abundant, conspicuous, receptacles blunt :

15. S. cylindrocarpum (p. 720)

1. Sargassum Palraeri Grunow

Plate 41, fig. 56, and plate 94

Frond arising from a solid, rugose, more or less disk-shaped hold-
fast, 4.5-7 dm. high ; stipe terete, verrucose, up to 18 cm. long, bearing
2-5 terete or slightly angled, alternate branches at its summit disin-
tegrating after fruiting ; the primary branches giving rise to sterile
segments (leaves) alternately arranged and often densely crowded,
dendroidally and dichotomously dissected into 15-25 slightly flattened
segments 2-3 cm. long, subcostate ; older branches and at times the
ramuli, more or less muriculate ; numerous lateral terete secondary
branches arising in the axils of the leaves, longer below, shorter above ;
these bearing reduced leaves similar to those on the main fronds, pro-
ducing vesicles and the receptacles in their axils ; vesicles usually
solitary, smooth, spherical, 3.5-6 mm. diam., on the ends of pedicels
as long or longer than their diameter ; receptacles densely racemose,
substipitate, often with a vesicle or reduced leaf as a part of the group ;
the fruiting fronds often appearing nude as the leaves wear away ; con-
ceptacles conspicuous ; dioecious ; plants perennial.

Grunow, Add. cog. Sargass., 1915, p. 338 ; Gardner, New Pac.
Coast Mar. Alg. II, 1918, p. 448. Sargassiim pilulifericm Farlow,

19-5] Setchell-Gardner: MeUmophycem 713

Anderson and Eaton, Alg'. Exsicc. Amer. Bor., no. 102 ; Collins. Holden
and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 537a (not 537&). Not
C. Agardh, Sp. I, 1820, p. 27. Not Yendo. Fucaceae Japan, 1907,
p. 54. Not Funis pilidifer Turner, Fuci, vol. 1, 1808, p. 145, pi. 65.
Sargassum dissect if olium Setchell and Gardner, in Gardner, New Pac.
Coast Mar. Alg. I, 1917, p. 386.

2. Sargassum acinacifolium S. and G.

Basal parts unknown ; branches terete, smooth ; leaves 12-16 mm.
long, asymmetrical, the upper margin concave and smooth, the lower
margin and apex coarsely dentate, ecostate, cryptostomata absent ;
vesicles situated at the base of the receptacles or more rarely among
the receptacles, subspherical, smooth, apiculate, 1.5-2.5 mm. diam., on
pedicels shorter than the diameter ; receptacles 2-3 times forked,
nearly cylindrical, not spiny, acuminate, more or less denticulate
toward the apices.

Cast ashore, Guaymas ( ?), Mexico. »

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 732, pi. 21,
fig. 82.

This species of Sargassum is a near relative, apparently, of S.
lapazeainim and of S. Brymitii, but it differs sufficiently in leaf,
bladder, and receptacle characters to warrant giving it separate
characterization.

3. Sargassum guardiense S. and G.

Basal parts unknown ; primary branches up to 5 dm. high ; second-
ary branches long and slender ; branches and ramuli all smooth, terete ;
leaves slightly flattened, to filiform, ecostate, margins smooth, crypto-
stomata absent or rare ; vesicles situated at the base of the fructiferous
ramuli or among the branches of the receptacles, subspherical to
slightly cylindrical, 2-4 mm. long, smooth, tapering at both ends,
apiculate, on pedicels shorter than the diameter ; receptacles 1-3 times
forked, terete, not spinose, 5-8 mm. long, forming with the vesicles
short heteroclyte cymes.

Cast ashore at Angel de la Guardia Island, Gulf of California.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 732, pi. 19,
fig. 64.

Unfortunately we have no complete specimens of this seemingly
very well defined species of Sargassum. The character of the holdfast

71-1 University of California Piiblications in Botany [Vol. 8

and of the main stipe, which are in many instances of much value in
classification, cannot be stated. We have several clean primary
branches which are in good fruiting condition and otherwise seem-
ingly characteristic. The loose, open character of the branching and
the shape and size of the vesicles remind one very much of some mem-
bers of the genus Cystophyllum J. Ag. It has perhaps its nearest
relative in 8. carpophyllum but differs in leaf characters and in the
vesicles being regularly lateral or terminal to the receptacles.

4. Sargassum lapazeanum S. and G.

Fronds 4.5-6 dm. high, arising from a solid parenchymatous disk ;
stipe 1.5 cm. long; primary branches 5-7, terete, smooth, giving rise
to numerous long, slender, secondary branches ; leaves 0.5-1.5 cm.
long, asjmimetrical, widest toward the apices, with very short petioles,
the basal half of the upper margin smooth and concave, the remainder
of the blade sharply dentate, midrib inconspicuous, cryptostomata
abundant and conspicuous ; vesicles scattered among the receptacles,
ellipsoidal, 1-2 mm. long, transformed from the base of a leaf, mostly
crowned by the remnants of the blade ; receptacles 4-7 mm. long, 2-3
times forked, branches strict, spinulose, intermixed with leaves and
receptacles forming a heteroclyte cyme.

Cast ashore. La Paz, Lower California.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 733, pi. 20,
fig. 74.

^t5-

5. Sargassum Bryantii S. and G.

Basal parts unknown ; branches terete, more or less contorted ;
leaves 6-12 mm. long, 0.5 as broad as long, ecostate, asymmetrical, the
upper margin concave and smooth, the lower margin and the end
unevenly serrate ; cryptostomata few and irregularly placed ; vesicles
numerous along the ramuli or, more rarely, intermixed with the
receptacles, subspherical, marginate when young, spinose, short- petio-
late; receptacles short, 4-8 mm. long, 1.5-3 mm. broad, irregular
cylindrical below, blunt or pointed, at times slightly spinose and
crowned with a rudiment of a leaf.

Cast ashore near La Paz, Lower California.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 733, pi. 21,
fig. 83.

The characters of the receptacles do not agree in every particular
with those given by J. Agardh (Sp. Sargas. Austral.) in his key to

1925] Setchell-Gardner : Melanophyceae 715

Eusargassum. They are not regularly two-edged and serrate-dentate
along the margins. Many of them are slightly flattened above, some-
what spinose, and crowned with a leaf rudiment. The species fits into
this section better than into any other.

6. Sargassum horridum S. and G.

Basal parts unknown ; branches and branchlets muricate, leaves
linear-lanceolate, acute, midrib percurrent, margins deeply and doubly
serrate ; cryptostomata numerous and conspicuous on the leaves, stems,
and vesicles ; vesicles sparse, occupying the position of leaves near the
base of the ramuli or scattered among the receptacles, spherical,
4-8 mm. diam., short-petiolate ; receptacles decompoundly ramose,
decidedly spinose.

Cast ashore. La Paz, Lower California,

Setchell and Gardner, Mar. Alg., Gulf Calif., 1924, p. 734, pi. 20,
figs. 65, 66.

Like the majority of our specimens from the Gulf of California,
the specimens of this species have neither holdfast nor stipe. Pre-
sumably manj^ of them grow only in the sublittoral belt and collectors
have observed onlj^ such specimens as have been cast ashore, and these
are usually fragmentary. Otherwise the specimens of S. horridum are
in excellent condition.

7. Sargassum Marchantae S. and G.

Basal parts unknown ; primary branches 4.5-6 dm. high, terete,
smooth ; secondary branches numerous, densely fructiferous ; leaves
4-6 cm. long, 3-5 mm. wide, linear-lanceolate, acute, midrib percurrent,
margins irregularly serrate-dentate ; cryptostomata numerous and con-
spicuous; vesicles sparse, spherical, on short pedicels near the base of
the ramuli or near the base of the branching receptacles, 4-6.5 mm.
diam., smooth ; receptacle several times forked, occasionally one fork
developing into a leaf or a vesicle forming a "heteroclyte cyme ( ?),"
but all others fructiferous and from a single pedicel, supported near
the base of a leaf, cylindrical, blunt, spinulose, the whole cyme 10-15
mm. long.

Cast ashore, Eureka, near La Paz, Lower California.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 735, pi. 19,
fig. 63.

Sargassum Marchantae is probably closely related to 8. Liehmanni
J.Ag.

716 Vniversity of California Puhlications in Botany [Vol. 8

8. Sargassum Liebmanni J. Ag,

Plate 45, fig. 81

Fronds about 30 cm. high, arising from a parenchymatous disk,
stipe terete, very short ; primary branches filiform, slender, smooth ;
leaves lanceolate, acuminate, costate, cryptostomata absent, margins
undulate, spinulose-dentate; vesicles spherical, submarginate, apicu-
late, sparse, on pedicels shorter than their diameter, smooth ; recep-
tacles densely aggregated into a racemose cyme, more or less twisted,
triangular, with each angle bearing a row of spinescent teeth.

West coast of Mexico.

J. Agardh, Nya Alg., 1847, p. 8, Sp. Sargass. Aust., 1889, p. 91,
pi. 5; Grunow, Ad. cog. Sargass., 1915, p. 398.

The exact locality whence the type material of this species was
obtained by Liebmann is not known. Agardh states "Pa mexicanska
stranden af Stillen Oceanen. ' ' Possibly the locality is St. Augustin. It
apparently has not been seen since the original discovery was made.

9. Sargassum Agardhianum Farlow

Plate 39, fig. 46; plate 41, fig. 55, and plate 92

Plants anchored by a firm, expanded, warty, parenchymatous hold-
fast ; perennial ; fronds 25-35 cm. high, rarely up to 9 dm. high, terete,
slender, simple at first but later branching a few millimeters above the
base; branches few to many, filiform, terete or slightly angled, much
surpassing the main frond or stipe which remains comparatively short ;
as the lower branches fruit and disintegrate, new ones arising from the
transition region above; the fronds and branches giving rise either to
leaf -like segments alternately arranged, linear-lanceolate, with sharply
toothed margins and apex, and with a midrib and scattered crypto-
stomata, or to filiform segments beset with sharp teeth ; in the axils of
the segments either short branches bearing clusters of small vesicles,
2-6 in number, or longer secondary branches bearing leaf -like seg-
ments and vesicles in turn ; vesicles on pedicels 1-1.5 mm. long,
spherical or slightly ellipsoidal, smooth or in part slightly margined
and toothed, with few cryptostomata, apiculate, 1.5-2.5 mm. diam, ;
receptacles in small clusters in the axils of certain segments, simple or
bearing few sharp tooth-like projections; conceptacles conspicuous.

Growing in the lower littoral and upper sublittoral belts. Southern
California and Lower California (Ensenada).

1925] • S etch ell-Gardner: Melanophyceae 717

Farlow, List of Mar, Alg. U. S., 1876, p. 706 (nomen nudum), in
FarloAV, Anderson and Eaton, Alg. Exsicc. Amer.-Bor., no. 103, MS ;
Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), nos. 179 and
CXVIII; J. G. Agardh, Sp. Sargass. Aust., 1889, p. 93; Grunow, Add.
cog. Sargass., 1915, p. 401.

The type locality of S. Agardhianum is San Diego, California.
The form of the species is considerably modified by habitat. It is
common in rather shallow rockpools in the lower half of the littoral
belt or in deeper pools higher up. In these situations it is always
covered with water and grows short and stocky with the secondary
lateral branches more highly developed. When growing in quiet
water in the sublittoral belt, it becomes much more elongated and
attenuated, reaching a length of nearly a meter. It grows in pro-
fusion on the harbor side of the government breakwater at San Pedro,
California, but is wholly absent on the exposed side. The antheridia
and oogonia in this species are often in separate conceptacles on the
same individual, or even in the same receptacle.

10. Sargassum insulare S. and G.

Fronds 7-9 cm. high, arising from a parenchymatous disk ; stipe
small, 5-10 mm. long; primary branches cylindrical throughout, 1-2
mm. diam., smooth, moderately and alternately branched ; leaves 1-2
cm. long, about half as wide as long, asymmetrical, the upper margin
concave and mostly smooth, the lower margin and apex convex and
crenate or dentate, ecostate ; cryptostomata sparse ; vesicles inter-
mingled with the receptacles, spherical or subspherical, 1.5-2.5 mm.
diam., short petiolate, often crowned by the remnant of a leaf; recep-
tacles moderately branched, standing on a single pedicel on the base of
a leaf, irregular in shape, clothed with scattered blunt spines, some-
times crowned by a rudiment of a leaf.

Growing on rocks in the upper sublittoral belt. San Marcos
Island, Gulf of California.

Setchell and -Gardner, Mar. Alg. Gulf Calif., 1924, p. 735, pi. 20.
figs. 67, 68, and pi. 21, fig. 78.

718 University of California PuMications in Botany [Vol. 8

11. Sargassum Brandegeei S. and G.

Basal parts unknown ; branches and branchlets terete, smooth,
^Yithol^t cryptostomata ; branching rather dense in the upper parts ;
leaves 15-25 mm. long, 4-8 mm. wide, apices blunt, base cuneate,
margins deeply serrate, cryptostomata absent ; vesicles spherical, small,
2-3 mm. diam., smooth, apiculate or crowned by a rudiment of a leaf,
supported by pedicels mostly shorter than their diameter, occupying
positions of leaves toward the base of the ramuli, or scattered among
the receptacles ; receptacles in short dense racemes, with short distinct
pedicels below but with sessile branches above, mostly blunt.

Cast ashore, Guaymas (?), Mexico.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 736, pi. 21,
fig. 79.

12. Sargassum sinicola S. and G.

Basal parts unknown ; branches and branchlets smooth, terete ;
leaves linear-lanceolate, acute, margins serrate-dentate, midrib per-
current, cryptostomata sparse, inconspicuous ; vesicles numerous,
mostly borne near the base of pedicel supporting the receptacles,
spherical, smooth, 3-5 mm. diam., pedicels equaling the diameter;
receptacles 1-3 times forked, the lower pedicellate, the upper sessile,
cylindrical, with acuminate apices, not spinose.

Cast ashore, Eureka, near La Paz, Lower California.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 736, pi. 20,
fig. 73.

This species seems nearly related to S. podocanthum Sond. and to
8. spinuligerum Sond. but the leaves are much more ' ' glandular ' ' and
the receptacles much more branched.

13. Sargassum polyacanthum f. americanum S. and G.

Basal parts unknown ; branches and branchlets up to 1.5 mm. diam.,
moderately muricate, leaves 3-4 cm. long, narrowly lanceolate, acute,
margins serrate-dentate, midrib percurrent, cryptostomata sparse ;
vesicles spherical, smooth, up to 8 mm. diam., occupying positions of
leaves along the ramuli or at times supported by a leaf; receptacles
1-2 times forked, 2.5-4 mm. long, obtuse-conical.

Cast ashore. La Paz ( ?), Lower California.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 736.

1925] SetcheU-Oardner: Melanophyceae 719

This form stands very close to S. spmuligerum Sond. The leaves
are longer and narrower than in that species. There are also other
specific differences.

14. Sargassum Johnstonii S. and G.

Basal parts unknown ; primary branches relatively robust, terete,
smooth, up to 8 dm. long, secondary branches numerous, densely
crow^ded with fructiferous ramuli ; leaves narrowly lanceolate, ecostate,
margins sparsely denticulate, cryptostomata almost absent, 1.5-2.5 cm.
long, 2-4 mm. wide ; vesicles, smooth, narrowly elliptical, merging
gradually below into a short petiole, crowned by a mucron or a
remnant of a blade, scattered along the fruiting rhachis among the
receptacles, 3-5 mm. long, on pedicels shorter than their length ;
receptacles single or 2-3 times forked, nearly cylindrical, mostly blunt,
with slightly denticulate apices.

Cast ashore, Georges Island, Gulf of California.

Setehell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 737, pi. 20,
fig. 72 and pi. 21, fig. 80.

Sargassum Johnstonii is related to *S^. galapagense Grun. but differs
in having elongated, long-apiculate vesicles, and slightly denticulate
receptacles and in details of leaf characters.

Sargassum Johnstonii f. laxius S. and G.

Basal parts unknown; primary branches up to 13.5 dm. long,
secondary branches very much less frequent and much longer than in
the species ; leaves filiform, 1-2 cm. long, cryptostomata sparse, incon-
spicuous ; vesicles subspherical, mostly long-mucronate ; receptacles
1-2 times forked, rarely simple, not denticulate.

Cast ashore at Guaymas, Mexico,

Setehell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 737, pi. 21,
figs. 75 and 81.

This form differs from the species in the following particulars :
the secondary branches and fructiferous ramuli are very much more
widely scattered and several times longer, the leaves are narrower, in
fact they are filiform, the vesicles are very much shorter, about one
and a half times as long as broad, and the receptacles are less branched
and rarely, if ever, denticulate at the apices.

720 University of California Puhlications in Botany [Vol. 8

Sargassum Johnstonii f . gracile S. and G.

Basal parts unknown; branches of all orders \'ery slender, long,
and wide apart, up to 1 mm. diam. ; leaves filiform ; vesicles narrowly
ellipsoidal.

Cast ashore, Guaymas, Mexico.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 738, pi. 21,
fig. 76.

This differs from the species in being decidedly more slender
throughout, much less frequently branched, and in having the branches
much longer and more delicate, and in having filiform leaves. In these
respects they more nearly coincide with f. laxius but are much more
delicate throughout than in that form. The receptacles are young in
the specimens at hand. The vesicles are practically the same in form
as those of the species.

15. Sargassum cylindrocarpum S. and G.

Basal parts unknown ; primary branches and branchlets smooth,
terete ; leaves 5-8 cm. long, 3-5 mm. wide, linear-lanceolate, acute,
serrate-dentate, midrib percurrent, cryptostomata abundant and con-
spicuous ; vesicles 5-8 mm. diam., spherical, smooth, on stipes shorter
than the diameter, occupying positions of leaves toward the base of
the fructiferous ramuli ; receptacles several times forked, cylindrical,
blunt, 1-2 cm. long.

Cast ashore, La Paz, Lower California.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 738, pi. 21,
fig. 77.

16. Sargassum herporhizum S. and G.

Fronds 6-8 dm. high, attached at first by a small parenchymatous
disk, later the short stipe giving rise to erect primary branches in
part and to horizontal branches which in turn develop attaching
branches, hapteres, below and to erect branches above ; primary
branches slender, terete, smooth, densely clothed with leaves and with
scattered secondary branches below and with fructiferous branches
above ; lower leaves sublinear, upper linear-lanceolate and acute, lower
3-5 mm. broad, upper 1-2 mm. broad, midrib percurrent, margins
sparsely denticulate, cryptostomata absent ; vesicles numerous, scat-
tered among the receptacles, spherical to subspherical, smooth, rarely

1925] Setchell-Gardner : Melanophyceae 721

apiculate, 1-2 mm. diam., on pedicels as l()n<i- as or longer tiian the
diara. ; receptacles short, 5-10 mm. long, siibcylindrical, acuminate,
only sparsely branched, tuberciilate with conspicuous conceptacles ;
plants very dark on drying.

Growing in the upper sublittoral belt. Georges Island, Gulf of
California.

Setchell and Gardner, Mar. Alg. Gulf Calif., 1924, p. 739, pi. 20,
figs. 69-71.

17. Sargassum paniculatum J. Ag.

Plate 46, fig. 5

Basal parts unknown ; main branches and branchlets relatively
slender, terete, smooth ; secondary branches arising near the middle
of the primary branches, longest below, gradually diminishing upward,
the fructiferous ramuli arising on the base of the leaves of the
secondary branches ; leaves linear-lanceolate, acute, midrib percurrent,
margins acutely denticulate-serrate, in part doubly so, cryptostomata
scattered, or in a fairly regular row on each side of the midrib, incon-
spicuous; vesicles subspherical to ovate 2-4 mm. long, smooth, on
pedicels as long as or longer than the diameter, occupying positions of
leaves at the base of the fructiferous ramuli or scattered among the
receptacles ; receptacles cylindrical, acuminate, more or less branched,
forming loose racemes.

Guadalupe Island, off the coast of Mexico.

J.Agardh, Sp. Alg., vol. 1, 1848, p. 315, Sp. Sargass. Aust., 1889,
p. 122, pi. 12, figs. 1-3 ; Grunow, Add. cog. Sargass., 1916, p. 177.

We are including this species in our account, based upon a portion
of a frond collected by Dr. E. Palmer at Guadalupe Island, Lower
California, Mexico, communicated to the Herbarium of the University
of California (no. 170615) by Dr. W. G. Farlow, and are placing it in
8. paniculatum on the authority of Th. Reinbold. who examined the
specimen. The plant agrees fairly well with the descriptions and
figures of J. Agardh {loc. cit.).

We have no knowledge of the "Sargassum fuJigmosum var. (?)
Calif ornica" described by Grunow, 1916, p. 173. "Hab. ad littora
Californiae (leg. Askenasy)," and hence are not including it in our
flora until we obtain further information concerning its structure
and distribution.

722 University of California Publications in Botany [Vol. 8

BIBLIOGRAPHY

Adanson, M.

1763. Families des plantes. Paris.

Agardh, C. a.

1810-1812. Dispositio algarum Sueciae, D. 1-4. Lund.

1817. Synopsis algarum Scandinaviae, adjecta dispositione universali

algarum. Lund.
1820-1828. Species algarum rite cognitae cum synonymis, differentiis speci-
ficis et descriptionibus succinctis.
1820. Vol. 1, part 1, pp. 1-168.
1822. Vol. 1, part 2, pp. 169-531.
1828. Vol. 2, part 1, pp. i-lxxvi and 1-189.
[The Lund edition is dated 1820, but the " Gryphiswald " edi-
tion is dated 182L]
1821. Icones algarum ineditae, fase. 2. Stockholm.
1824. Systema algarum. Lund.

Agardh, J. G.

1841. In historian! algarum symbolae. Continuatio prima. Linnaea, vol.

15, pp. 443-457.
1847. Nya Alger fran Mexico. Oefvers. af Kongl. Vet.-Akad. Forhandl.,
Arg. 4, no. 1. Stockholm.

[Often quoted as "Alg. Lieb. " See note, Setchell and Gard-
ner, 1920, p. 307.]
1848-1876. Species genera et ordines algarum. Lund.
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1867. De Laminarieis. Lunds Univers. Arsskrift, vol. 4. Lund.

1868. Bidrag till Kannedomen af Spetsbergens Alger. Tillagg till fore-

gaende afhandling. Kongl. Sv. Vet.-Akad. Handl., vol. 7, no. 8.

Stockholm.
1872. Bidrag till Kannedomen of Gronlands Laminarieer och Fucaceer.

Kongl. Sv. Vet.-Akad. Handl., vol. 10, no. 8. Stockholm.
1878-1890'. Till Algernes systematik, Nya didrag. Lunds Univ. Arssk.
1873. Forsta afdelningen, vol. 9.

1882. Andra afdelningen, vol. 17.

1883. Tredje afdelningen, vol. 19.
1885. Fjerde afdelningen, vol. 21.
1887. Femte afdelningen, vol. 23.
1890. Sjette afdelningen, vol. 26.

1889. Species Sargassorum Australiae, descriptae et dispositate, etc. Kongl.

Sv. Vet.-Akad. Handl., vol. 23, no. 3, pis. 1-31. Stockholm.
1892. Analecta algologica.

1894. Contr. 1, Contr. IL

1896. Contr. III.

1897. Contr. IV.
1899. Contr. V.

Anderson, C. L.

1891. List of California marine algae with notes. Zoe, vol. 2, pp. 217-225.
San Francisco.

1894. Some new and some old Algae but recently recognized on the Cali-
fornia coast. Zoe, vol. 4, pp. 358-362.

1^25] Setchell-Gardner : Melanophyceae 723

Areschoug, J. E.

1840. Algae scandinavicae exsiccatae. Ed. 1 (not seen).

1861-1879. Algae scandinavicae exsiccatae. Series nova, fasc. i-ix. XJpsala.

1842. Algarum minus rite cognitarum pugillus primus. Linnaea, vol. 16,

pp. 225-236, pi. 8.

1843. Algarum (Phycearum) minus rite cognitarum pugillus secundus.

Linnaea, vol. 17, pp. 257-269, pi. 9.
1846. Enumeratio phycearum in maribus Scandinaviae crescuntium. Sectio

prior, Fucaceas continens. Acta Eeg. Soc. Sci. Upsala, vol. 13.
1850. Phycearum quae in maribus Scandinaviae creseunt, enumeratio, sectio

posterior Ulvaceas continens. Nov. Acta Reg. Soc. Sci. Upsala,

vol. 14, pp. 385-454.
1853. Virginia, ett nytt algslagte. Oefvers. Kongl. Vet.-Akad. Forhandl.,

nos. 7, 8, pp. 145-147.
1866-1884. Observationes phycologicae. Nova Acta Reg. Soc. Sci. Upsala,

ser. 3.

1866. Part 1. De Confervaceis nonnullis, vol. 6.

1875. Part 3. De Algis nonnullis scandinaviae et de conjunc-

tione phaeozoosporarum Dictyosiphonidis hippuroidis, vol. 10.

1883. Part 4. De Lamiuariaceis nonnullis, vol. 12.

1884. Part 5. De Laminariaceis nonnullis (continuatis), vol.
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1873. Om de Skandinaviska Algformer, som aro narmast beslagtade med

Dictyosiphon foeniculaceus, eller Kunna med denna lattast for-

blandas. Botaniske Notiser, no. 6, pp. 161-171.
1876. De tribus Laminarieis et de Stephanocystide osmundaceae (Turn.)

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65-73.
1881. Beskrifning pa ett nytt algslagte, Pelagophycus, horande till Lami-

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1898. On the structure and development of Soranthera Post, and Rupr.

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Batters, E. A. L.

1888. A description of three new marine algae. Jour. Linn. Soc. Bio., vol.
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BORGESEN, F.

1902. The marine algae of the Faeroes, in Warming, Botany of the Faeroes,
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1914. The marine algae of the Danish West Indies. Part 2, Phaeophyceae.
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1920. The marine algae of the Danish West Indies. Vol. 2, Rhodophyceae,
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BORNET, E.

1891. Note sur quelques Ectocarpus. Bull. Soc. Bot. France, vol. 38, pp.

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1892. Les algues de P.-K.-A. Schousboe, recoltees au Moroc & dans la

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724 University of California Puhlications in Botany [Vol. 8

BoBY DE Saint Vincent, J. B. (M. A. G.)

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1822. Vol. 1.

1823. Vol. 4.

1824. Vol. 6.

1826. Vol. 9, vol. 10.

1827. Vol. 13.

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Brandt, E. P.

1923. Potash from kelp; early developments and growth of the giant kelp,
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Briquet, J.

1906. Regies internationales de la nomenclature botanique adoptees par
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1891. The plurilocular sporangia of Asperococcus hullosus and Myriotrichia
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1891. Notes on New England Marine Algae V. Bull. Torr. Bot. Club, vol.

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1896. Notes on New England Marine Algae VII. Bull. Torr. Bot, Club,

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1913. The marine algae of Vancouver Island. Canada Geological Survey.

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Collins, F. S., Holden, I., and Setchell, W. A.

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1853. Algues marines du Finistere. Fasc. 1-3 (not seen).

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1925] Setchell-Gardner : MeJanophyceae 725

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De la Pylaie, B.

1829. Flore de Terre-Neuve et des iles Saint Pierre et Miclon avec figures

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Derbes. A., et Solier, A. J. J.

1850. Sur les organes reproducteurs des algues. Ann. Sci. Nat., ser. 3, Bot.,

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De-Toni, J. B. (G. B.)

1891. Systematische Uebersicht der bisher bekannten Gattungen der echten

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1741. Historia muscorum, in qua circiter sex centae species veteres et
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DlLLV^YN, L. W.

1802-1809. British Confervae, or colored figures and descriptions of the
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Draparnaud, J. P. R.

1801. Discours relatifs a I'histoire naturelle. Montpellier.

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1830. Botanicon Gallicum seu Synopsis Plantarum in Flora Gallica Descrip-

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1822. Commentationes botanicae (Observationes botaniques) (1823). Tournay.

English Botany, or colored figures of British plants with their essential char-

1790-1814. acters, synonyms, and places of growth, to which will be added

occasional remarks by James Edward Smith. The figures by

James Sowerby. Vols. 1-36. London.

1843. Supplement, vol. 3.

726 University of California Piiblications in Botany [Vol. 8

Farlow, W. G.

1875. List of the marine algae of the United States with notes on new

and imperfectly known species. Proc. Am. Acad. Arts and Sci.,
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1876. List of the marine algae of the United States. Eeport of the U. S.

Fish Comm. for 1875, pp. 691-717. Washington.
1881. The marine algae of New England and adjacent coast. Report of
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1889. On some new or imperfectly known algae of the United States, I.

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Farlow, W. G., Anderson, C. L., and Eaton, D. C.

1877-1889. Algae Exsiccatae Americae-Borealis. Fascicles 1-5. Boston.

Flora Danica

Icones plantarum sponte nascentium in regnis Daniae et Norvegiae et in
ducatibus Slesvici, Holsatiae et Oldenburg, ad illustrandum opus
de iisdem plantis, regie jussu exarandum, Flora Danicae nomine
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1768. Vol. 3, fasc. 7.

1770. Vol. 3, fasc. 9.

1771. Vol. 4, fasc. 10.
1778. Vol. 5, fasc. 13.

[For a detailed statement of the dates, etc., of this work, see
Pritzel, 1851, p. 214, sub. (Eder.j
FOSLIE, M.

1890. Contributions to the knowledge of the marine algae of Norway, I,

East-Finnmarken. Tromso Museums Aarshefter, vol. 13.

1891. Ihid. II, vol. 14.

1892. List of the marine algae of the Isle of Wight. Det. Kgl. norske

Vidensk. Selsk. Skrifter, pp. 267-282. (Pp. 1-16, reprint.)

Trondhjem.
1894. New or critical Norwegian Algae. Det. Kgl. norske Vidensk. Selsk.

Skrifter, pp. 1-31, pis. 1-3 (reprint). Trondhjem.
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1825. Systema orbis vegetabilis. Primas lineas novae constructionis peri-

clitatur Elias Fries. Part 1, Plante homonemeae. Lund.
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Elias Fries.
Frye, T. C.

1918. The age of Pterygophora californica. Publ. Puget Sound Biol. Sta.,

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Gardner, N. L.

1910. Variation in nuclear extrusion among the Fucaceae. Univ. Calif.

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1917. New Pacific Coast marine algae, I. Ibid., vol. 6, no. 14, pp. 377-416,

pis. 31-35.

1918. New Pacific Coast marine algae, II. Ibid., vol. 6, no. 16, pp. 429-454,

pis. 36, 37.
1922. The genus Fucus on the Pacific Coast of North America. Ibid., vol.
10, no. 1, pp. 1-180', pis. 1-60.

1925] SetcJiell-Gardner: 3IeIanophyceae 727

Gleerup, C. W. K.

1880. Enumerantur plante Scandinavicae. Point-Verzeichniss Gewachse
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Gmelin, S. G.

1768. Historia Fucorum.
Gobi, C.

1874. Die Brauntange (Phaeosporeae und Fucaceae) des finnischen Meer-
busens. Mem. I'Acad. Imp. des Sci. de St. Petersbourg, ser. 7, vol.
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Gray, J. E. (see also S. G. Gray),
Gray, S. F.

1821. A natural arrangement of British plants, etc. Vols. 1, 2. London.

[J. E. Gray, son of S. F. Gray, did the systematic work in the

two volumes which came out under the foregoing title with S. F.

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Greville, E. K.

1823-1829. Scottish Cryptogamic Flora, or colored figures and descriptions
of ciyptogamic plants belonging chiefly to the order Fungi, and
intended to serve as a continuation of English Botany. Vols. 1-6.
l]dinburgh.

[For dates of volumes, see Setchell and Gardner, 1920, p. 313.]
1830. Algae Britannicae, or descriptions of the marine and other inarti-
culated plants of the British Islands, belonging to the order, Algae,
with plates illustrative of the genera. Edinburgh,
1830. Synopsis generam algarum cum enumeratione specierum cognitarum,
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Griggs, K. F.

1906. Benfrewia parvula, a new kelp from Vancouver Island. Postelsia,

pp. 247-274, pis. 16-19. St. Paul, Minn.

1907. Cymatliere, a kelp from the west coast. Ohio Naturalist, vol. 7, no. 5,

pp. 89-96, pi. 7.
1909. Juvenile kelps and the recapitulation theorj', I. American Naturalist,

vol. 43, pp. 7-30.
1909a. The sporophylls of Lessotiiopsis. Ohio Naturalist, vol. 9, no. 4,

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Grunow% a.

1915-1916. Additamenta ad cognitionem Sargassorum. K. K. Zool.-Bot.

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1915. Vol. 65, pp. 329-448.

1916. Vol. 66, pp. 1-48, 136-185.
Guernsey, J.

1912. Notes on the marine algae of Laguna Beach. First Ann. Eept. of
Laguna Beach Marine Lab., Pomona College, Calif., pp. 195-218.
Guignard, L.

1892. Observations sur I'appareil mucifere de Laminaricees. Ann. Sci. Nat,
Bot., ser. 7, vol. 15, pp. 1-46.

Gunnerus, J. E.

1766-1772. Flora norvegica, observationibus praesertim oeconomicis panos-
que norvegici locupletata,
1766. Part 1.
1772. Part 2.

728 Vniversity of California Publications in Botany [Vol.8

Harvey, W. H.

1830-1841. Algae, in Hooker and Arnott's Botany of Captain Beechey's

voyage. (Cf. Journ. Bot., vol. 31, p. 298 for dates of issue.)
1838. The genera of South African plants, arranged according to the

natural system. Cape Town.
1841. A manual of the British marine algae, containing generic and specific
descriptions of all the known British sea-weeds, Avith plates to
illustrate all the genera. London.
1849. Idem., ed. 2.
1846-1851. Phycologia Britannica. London.

[For bibliographical note concerning dates of publication of
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1852. Nereis Boreali-Americana. Part I, Melanospermeae.
1858-1863. Phycologia Australica. London.

1858 vol. 1, 1859 vol. 2, I860' vol. 3, 1862 vol. 4, 1863 vol. 5.
1862. Notice of a collection of algae made on the northwest coast of North
America, chiefly at Vancouver Island by Dr. David Lyall in the
years 1859-1861. Jour. Proc. Linn. Soc. Bot., vol. 6, pp. 157-177.
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1883-1885. Die Meeresalgen Deutschlands und Oesterreichs, in Eabenhorst's
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[Issued in 10 Lieferungen between 1883 and 1885.]

Hauck, F., and Eichteb, P.

1885-1896. Phykotheka Universalis (Exsicc). Fasc. 1-15. Leipzig.

Hoffman, Edna

1911. Fructification of Macrocystis. Univ. Calif. Publ., Bot., vol. 4, pp.
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Hooker, W. J.

1833. The English flora of James Edward Smith. Class XXIV, Crypto-
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Hooker, J. D.

1844-1847. The Botany of the Antarctic Voyage of H. M. Discovery Ships,
Erebus and Terror, etc. Vol. 1, Flora Antarctica.

[For dates of issuance of the various parts see B. Daydon
Jackson, in the Journal of Botany, 1912, pp. 284, 285.]
Howe, M. A. '

1911. Phycological Studies V, Some marine algae of Lower California,

Mexico. Bull. Torr. Bot. Club, vol. 38, pp. 489-514, pis. 27-34.
1914. The marine algae of Peru. Mem. Torr. Bot. Club, vol. 15, pp. 1-185,

pis. 1-66.
1918. Algae, in Britton, Flora of Bermuda, pp. 489-540.
1920. Algae, in Britton and Millspaugh, Bahama Flora, pp. 553-618. New
York.
Hudson, G.

1762. Flora Anglica; exhibens plantas per regnum Britanniae sponte

crescentes, distributas secundum systema sexuale, cum differentiis

specierum, synonymis auctorum, nominibus incolarum, solo locorum,

tempore florendi, ofiicinalibus pharmacopaeorum. Ed. 1. London.

1778. Flora Anglica. Ed. 2. London.

1925] Setchell-Ganhter: Melanophijceae 729

Johnston, E. C.

1912. Brief notes on the kelps of Alaska, in Fertilizer resources of tlie
United States, 62 Congr., 2 Sess., Sen. Doc. 190, pp. 214-216.
KiBBE, Alice

1915. Some points in the structure of Alaria fisUdosa. Puget Sound Mar.
Sta. Publ., vol. 1, no. 8, pp. 43-57, pis. 8, 9.

Kjellman, F. E.

1872. Bidrag till Kannedomen om Skandinaviens Ectocarpeer och Tilop-

terider. Stockholm.
1877. Ueber die Algenvegetation des Murmanschen Meeres on der West-

kiiste von Nowaja Semlja und Wajgatsch. Nova Acta Reg. Soc.

Sci., ser. 3, vol. extra-ord. (no. 12). Upsala.
1877a. Om Spetsbergens marina Klorofyllforande Thallophyter, II. Bihang

till Kgl. Sv. Vet.-Akad. Handl., vol. 4, no. 6. Stockholm.
1880. Ehodospermeae und Fucoideae, in Gleerup, C. W. K.,- Enumerantur

plantae Scandinavicae. Points-Verzeichniss Gewachse Scandi-

naviens. Heft 4. Lund.
1883. The algae of the Arctic Sea. Kongl. Sv. Vet.-Akad. Handl., vol. 20,

no. 5. Stockholm.

1889. Om Beringhafvets Algflora. Bihang till Kongl. Sv. Vet.-Akad.

Handl., vol. 23, no. 8.
1889a. Undersokning af nagra till slagtet Adenocystis Hook. fil. et Harv.
Hanforda Alger. Bih. till Kgl. Sv. Vet.-Akad. Handl., vol. 15, afd.
3, no. 1.

1890. Handbok i Skandinaviens hafsalgflora. I, Fucoideae. Stockholm.
1891-1893. In Engler und Prantl, Die natiirl. Pflanzenfam., 1 Teil, 2 Abt.

60 Lieferung —

Phaeophyceae, 1891, pp. 176-181;

Ectocarpaceae, 1891, pp. 182-189;

Choristoeai-paceae, 1891, pp. 190-191;

Sphacelariaceae, 1891, p. 192;
86 Lieferung —

Sphacelariaceae, 1893, pp. 193-197;

Encoeliaceae, 1893, pp. 197-204;

Striariaceae, 1893, pp. 204-208;

Desmarestiaceae, 1893, pp. 209-211;

Dictyosiphonaceae, 1893, pp. 212-214;

Myriotrichiaceae, 1893, pp. 214-215;

Elachistaceae, 1893, pp. 216-221;

Chordariaceae, 1893, pp. 221-230;

Stilophoraceae, 1893, pp. 230-233;

Spermatochnaceae, 1893, pp. 233-235;

Sporachnaceae, 1893, pp. 236-239;

Ralfsiaceae, 1893, p. 240;
97 Lieferung —

Ralfsiaceae, 1893, pp. 241-242;

Laminariaceae, 1893, pp. 242-260;

Lithodermataceae, 1893, pp. 260^262;

Cutleriaceae, 1893, pp. 262-265;

Tilopteridaceae, 1893, pp. 265-268;

Fucaceae, 1893, pp. 268-288;

730 University of California Publications in Botany ["Vol.8

141 Lieferung —

Fucaceae, 1896, pp. 289-290;

Dictyotales, Dictyotaceae, 1896, pp. 291-297.
1893a. Om Fucoideslagtet Myelophycus Kjellm. Bih. till Kgl. Sv. Vet.-Akad.,
Handl., vol. 18, ad. 3, no. 9.

Kjellman, F. E., and Svedelius, N.

1910. Encoeliaceae, iw Engler und Prantl. Die natiirl. Pflanzenfam., Nachtr.
z. 1 Theil, 2 Abt., pp. 154-158.

KUCKUCK, p.

1891. Beitrage zur Kenntniss einiger Ectocarpus-Arten der Kieler Fohrde.

Bot. Centralblatt, vol. 48.

1892. Ectocarpus siliculosus Dillw. sp. forma varians n. f. eiu Beispiel fiir

ausserordentliche Schwankungen der plurilocularen Sporangieu-
form. Ber. d. d. Bot. Gesell., vol. 10, pp. 256-258.

1894. Bermerkungen zur marinen Algenvegetation von Helgoland. Wissen-
schaft. Meeresuntersueh. Biolog. Anstalt auf Helgoland, n. F.,
vol. 1, part 1, pp. 225-263.

1899. Beitrage zur Kenntniss der Meeresalgen, 5-9, in Wissenschaftliche
Meeresuntersuchungen herausgegeben von der Kommission zur
deutschen Meere in Kiel und der biologischen Anstalt auf Helgo-
land. N. F., dritter Band. Abteilung Helgoland. Heft 1. Kiel
und Leipzig.

KUETZIKG, F. T.

1843. Phycologia generalis, oder Anatomie, Physiologic und Systemkunde

der Tange. Leipzig.
1843a. Ueber die Systematische Eintheilung der Algen. Linnaea, vol. 17,

pp. 75-107.
184.5. Phycologia germanica d. i. Deutschlands Algen in biindigen Be-

schreibungen, nebst einer Anleitung zum Untersuchen und Bestim-

men dieser Gewachse fiir Anfanger. Nordhausen.
1845-1871. Tabulae Phycologicae.

1845-

1849.

Vol; 1.

1861.

Vol. IL

1850-

-1852.

Vol. 2.

1862.

Vol. 12.

1853.

Vol.

3.

1863.

Vol. 13.

1854.

Vol.

4.

1864.

Vol. 14.

1855.

Vol.

5.

1865.

Vol. 15.

1856.

Vol.

6.

1866.

Vol. 16.

1857.

Vol.

7.

1867.

Vol. 17.

1858.

Vol.

8.

1868.

Vol. 18.

1859.

Vol.

9.

1869.

Vol. 19.

1860.

Vol.

10.

1871.

Index.

[For bibliographical notes, see Setchell and Gardner, 1919,
pp. 118, 119, and 1920, pp. 316.]
1849. Spe,cies Algarum. Leipzig.

KUXTZE, O.

1891-1898. Revisio genenim plantarum vascularium omnium atque cellu-
larium multarum secundum leges nomenclaturae Internationales
cum enumeratione plantarum exoticarum in itinere mundi collec-
tarum. Wiirtzburg.

1891. Parts 1 and 2. Algae, part 2, pp. 877-930.

1898. Part 3: 2.

1925] Setchell-Gardner : Melmwphyceae 731

Kylin, H.

1907. Studien ueber Algenflora der schwedischen Westkiiste. Akad.

Abhandl. Upsala.
1916. Ueber den Generationswechsel bei Laminaria digitata. Svensk. Bot.

Tidsskrift, vol. 10, pp. 551-561.
1918. Studieu ueber die Entwicklungsgeschichte der Phaeophyceen. Svensk

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Lamouroux, J. V. F.

1809. Observations sur la physiologie des algues marines, et description de

cinq nouveaux genres de cette famille. Nouveau Bull, des Sci.

par la Soc. Phil, de Paris, vol. 1, pp. 330-333, pi. 6, fig. 2 A, B, C, D.

180'9a. Exposition des caracteres du genre Dictyota et tableau des especes

qu'il renfei-me. Jour, de Bot., vol. 2, pp. 38^4.
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Jour, de Bot., vol. 2, pp. 129-135.
1813. Essai sur les genres de la famille des thalassiophytes non articulees.
Ann. du Mus. d'Hist. Naturelle par les professeurs de cet etab-
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[Usually quoted from the reprint which is paged from 1-84.
The plates are numbered the same in both.]
Le Jolis, a.

1855. Examen des especes confondues sous le nom de Laminaria digitata,
Auct., suivi de quelques observations sur le genre Laminaria.
Trans. Soc. Nat. Cherbourg, vol. 3.
1863. Liste des algues marines de Cherbourg. Mem. Soc. Imp. Sci. Nat. de
Cherbourg, vol. 10, p. 1. Paris.

[Tlie paper was published as a separate in 1863 before it
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1880. Eeprint of the foregoing.

Algues marines de Cherbourg (Exsicc), fasc. 5. Paris.
Leman, D. S.

1822. Laminaria, in Levrault, Dictionnaire des sciences naturelles, vol. 25,

p. 189.
1826. Pyaiella, ibid., vol. 44, p. 127.

LiGHTrOOT, J.

1777. Flora Scotia, or a systematic arrangement, in the Linnaean method,

of the native plants of Scotland and the Hebrides. London.
Link, H. F.

1809. Nova plantarum genera e classe Lichenum, Algarum, Fungorum, in

Schrader, Neues Jour. f. d. Bot., vol. 3B, p. 1.
Linnaeus, C.

1737. Genera plantarum, eorumque characteres naturales secundum numerum,

figuram, situm et proportionem omnium fructificationis partium.
1745. Flora suetica, exhibens plantas per regnum Sueciae crescentes, sys-

tematice cum differenteiis specierum, synonymis autorum, nomini-

bus incolarum, solo locorum, usu pharmacopoeorum. Ed. 1.

Stockholm.
1755. Ibid. Ed. 2.
1753. Species plantarum, exhibentes plantas rite cognitas, ad genera relatas,

cum differentiis specificis, nominibus trivialibus, synonymis

selectis, locis natalibus secundum systema sexuale digestas. Ed. 1.

Stockholm.

732 , University of California Publications in Botany [Vol. 8

1763. lUd. Ed. 2.

1758-1759. Systema naturae per regna tria naturae, secundum classes,

ordines, genera species, cum characteribus, differentiis, synonymis,

locis. 2 vols., pp. 1-1384.

Lyngbye, H. C.

1819. Tentamen hydrophytologiae Danicae, continens omnia hydrophyta
cryptogamia Daniae, Holsatiae, Faeroae, Islandiae, Groenlandiae
hucusque cognita, systematice disposita, descripta et iconibus
ilhistrata, adjectis simul speciebus Norvegicis. Copenhagen.

McClatchie, a. J.

1897. Seedless plants of southern California. Proc. So. Calif. Acad. Sci.,
vol. 1, pp. 337-395.

Mackay, J. T.

1836. Flora hibernica, comprising the Flowering Plants,, Ferns, Characeae,
Musci, Hepaticae, Lichens and Algae of Ireland, arranged accord-
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MacMillan, C.

1900. Observations on Lessonia. Bot, Gaz., vol. 30, pp. 318-334, pis. 19-21.

Chicago.
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22-26. St. Paul, Minn.
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Magnus, P.

1874. Die botanischen Ergebnisse der Nordseefahrt vom 21 Juli bis 9
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Meneghini, G.

1843. Alghe Italiane e Delmatiche. Fasc. 4. Padova.

Mertens, H.

1829. Zwei botanisch-wissenschaftliche Berichte vom Dr. Heinrich Mertens,
.... Erster Bericht iiber verschiedene Fucus-Arten, an den Vater,
Prof. Mertens in Bremen. Linnaea, vol. 4, pp. 43-58.

Mertens, J. C.

1819. Sur plusieurs especes de Fucus, neuvelles, ou pen connues, observees

dans la collection du Museum. Mem. du Museum d'Hist. Nat.,
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Mitchell, Margaret O.

1893. On the structure of HydroclatJirus Bory, in Murray, Phycologieal
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Mueller, O. F.

See Flora Danica.

Muenscher, .W. C.

1917. A key to the Phaeophyceae of Puget Sound. Puget Sound Marine
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Nees ab Essenbeck, C. G.

1820. Horae physicae berolinenses.

1^25] Setchell-Gardner: Melanophycede 733

NiEUWLAND, J. A.

1917. Critical notes on old and new genera of plants, no. 9. Am. Midland

Nat., vol. 5, no. 2, p. 30.
Hid., no. 10, pp. 50-52.

NORDSTEDT, O.

1911. Algological Notes 5-7. Botaniske Notiser, p. 263. Lund.

(Eder, G. C.

See Flora Danica.

OKAMtTBA, K.

1899-1903. Algae Japonicae Exsiccatae. Fasc. 1, 2.

1901. Illustrations of the marine algae of Japan. Vol. 1, no. 2. Tokyo.

1907. Icones of Japanese algae, vol. 1, no. 4. Tokyo.

1908. Ihid., vol. 1, no. 6. Tokyo.
1910. Ihid., vol. 2, no. 7. Tokyo.
1913. lUd., vol. 3, no. 2. Tokyo.

1918. Ihid., vol. 4, no. 3. Tokyo.

Olafsen, E., and Povelsen, B.

1772. Eeise igiennem Island, beskreven af Eggert Olafsen. 2 vols., pp.
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Oltmanns, F.

1904-190'5. Morphologie und Biologie der Algen.

1904, vol. 1; 1905, vol. 2. Jena.
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1917. North Pacific coast species of Desmarestia. Puget Sound Mar. Sta.

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Pfeiffer, L.

1873-1875. Nomenclator botanicus. Nominum ad finem anni 1858 publici
juris factorum, classes, ordines, tribus, familias, divisiones, genera,
subgenera vel sectiones designantium enumeratio alphabetica.
Adjectis auctoribus, temporibus, locis systematicus apud varios,
notis literariis alque etymologicis et synonymis.

1873. Vol. 1, part 1.
1875. Vol. 1, part 2.

1874. Vol. 1, part 1.
1874. Vol. 2, part 2.

POSTELS, A., and Euprecht, F.

1840. Illustrationes algarum in itinere circa orbem jussu imperatoris
Nicolai I. atque auspiciis Navarchi Friderici Liitke annis 1826,
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collectarum. St. Petersburg.

Pringsheim, N.

1862. Beitrage zur Morphologie der Meeresalgen. Physikal. Abhandl. der
Kon. Akad. d. Wissensch. Berlin.

734 University of California Pnhlications in Botany [Vol.8

Pritzel, G. a.

1851-1872. Thesaurus literaturae botanicae omnium gentium inde a rerum
botanicarum initiis ad nostra usque tempora, quindecim millia
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1864-1868. Flora Europaea algarum aquae dulcis et submarinae. Leipzig.

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1865. Vol. 2.
1868. Vol. 3.

Kafinesque-Schmalz, C. S.

1814. Precis des decouvertes et travaux somiologiques entre 1800-1814, ou
choix raissone de ses principales decouvertes en zoologie et en
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Bathbone, Mart

1904. Notes on Myriactis Areschougii and Coilodesme californica. Jour.
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1828. Conspectus regni vegetabilis per gradus naturales evoluti. Leipzig.

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1888. Die braunen Algen (Fucaceen und Phaeosporeen) der Kieler Bucht.

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Part 2, 1892.
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der Kommission zur wissenschaftlichen Untersuchung der deutschen
Meere in Kiel. Heft 1, Berlin.

[Both of these publications bear the same date on the title
page, and we have no knowledge as to which actually antedates
the other. Under Ealfsia verrucosa in the Algenflora, p. 48, E-einke
cites "taf. 5, 6," of the Atlas. It may seem from this citation
that the Atlas appeared first.]

1890. L'^^ebersicht der bisher bekannten Sphacelaciaceen. Bericht Deutsch.

Bot. Gesell., vol. 8, Heft 7, pp. 201-215,

1891. Beitrage zur vergleichenden Anatomie und Morphologic der Sphace-

lariaceen. Bibl. Botanica, vol. 5, Heft 23.
1903. Studien zur vergleichenden Entwicklungsgeschichte der Laminaria-
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Eosenthal, 0.

1890. Zur KenntnisH von Macrocystis und Thallassiophyllum. Flora, vol. 73,
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Eosexvikge, L. K.

1893. Gronlands Havalger. Meddelelser om Gronland, vol. 3, pp. 765-981,

pis. 1, 2. Copenhagen.
1910. On the marine vegetation from Northeast Greenland. Medd. om

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192.3] SetcheJl-Gardner : Melan^phyceae 735

EOTH, A. G.

1797-1806. Catalecta botanica quibus plantae novae et minus cognitae
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1797. Fase. 1.
180O. Fasc. 2.
1806. Fasc. 3.

EOUSSEL, H. F. A. DE

1806. Flora du Calvados ct terrains adjacents, oomposee suivant la methode
du Jussieu. Ed. 2.

EUPRECHT, F. J.

1848. Bemerkungen ueber den Ban und das Wachsthum einiger grossen
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1851. Tange des Oehotskischen Meeres. Middendorff 's sibirische Eeise,

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[For explanation of dates of this publication, see Setchell and
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1852. Xeue oder unvollstandig bekannte Pflanzen aus dem nordlichen

Theile des Stillen Oceans. Mem. de I'Acad. Sci. Nat., vol. 7, pp.
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Sauxders, De A.

1895. A preliminary paper on Costaria with description of a new species.
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1898. Phycological Memoirs. Proc. Calif. Acad. Sci., ser. 3, Bot., vol. 1,

pp. 147-168, pis. 12-32. San Francisco.

1899. New and little known brown algae of the Pacific coast. Erythea,

vol. 7, pp. 37-40, pi. 1.

1901. Papers from the Harriman Alaska Expedition, 25, The Algae. Proc.
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1901o. A new species of Alaria. Minn. Bot. Stud., vol. 2, part 5, pp. 561-562,
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Sauvageau, C.

1892. Sur quelques Algues pheosporees parasites. Jour, de Bot., vol. 6,
pp. 36-43, 5.5-59, 76-80', 90-96, 97-106.

1895. Note sur I'Ectocarpus pusillus Griffiths. Jour, de Bot., vol. 9, pp.

274-291.
1895t/. Sur les sporanges pluriloeulaires de \'" Asperococcus comprcsstis"
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1896. Eemarques sur la reproduction des Pheosporees. Ann. Sci. Nat.,

8 ser., Bot., vol. 2, pp. 223-274.
1896fl. Sur la nature des sporanges en chapelet de " VEctocnrpiis confer-

voides." Jour, de Bot., vol. 10, pp. 140-144.
1896ft. Sur I'Ectocarpus virescens Thuret, et ses deux sortes de sporanges

pluriloeulaires. Jour, de Bot., vol. 10', pp. 98-107 and 113-126.
1896c-1897a. Observations relatives a la se.xualite des Pheosporees. Jour.

de Bot., vol. 10, pp. 357-367, 388-398, and vol. 11, pp. .5-14.

736 University of California PuhJications in Botany [Vol.8

1897. Sur quelques Myrionemacees. Ann. Sci. Nat., 8 ser., Bot., vol. 5,

pp. 161-288.

1898. Idem. Eeprint, pp. 1-130.

1900-1904. Kemarques sur les Sphacelariacees. Jour, de Bot.

1900. Vol. 14, pp. 1-51.

1901. Vol. 15, pp. 51-167.

1902. Vol. 16, pp. 167-228.

1903. Vol. 17, pp. 228-332.

1904. Vol. 18, pp. 332-348.

1914. Idem. Eeprint, pp. xii and 634. Bordeaux.

1915. Sur la sexualite heterogamique d'une Laminaire (Saccorhua iulbosa).

C. E. de I'Acad. des Sci., vol. 161.
1917. Sur un nouveau type d'alternance des generations chez les Algues
brunes (Dictyosiphon foeniciilaceus). C. E. de I'Acad. Sci., vol. 164,
pp. 829-831. Paris.

SCHRADER, H. F.

1903. Observations on Alaria nana sp. nov. Minnesota Bot. Stud., vol. 3,
part 2, pp. 157-165, pis. 2.3-26.

SCHUH, E. E.

1900. Bhadinocladia, a new genus of brown algae. Ehodora, vol. 2, pp. Ill,

112, pi. 18.

Setchell, W. a.

1891. Concerning the life history of Saccorhiza dermatodea (De la Pyl.)

J. Ag. Proc. Am. Acad. Arts and Sci., vol. 26, pp. 177-217, pis. 1, 2.
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iaceae. Trans. Conn. Acad., vol. 9.
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i896a. The Elk Kelp. Erythea, vol. 4, no. 12, pp. 179-184, pi. 7.

1899. Algae of the Pribilof Islands, in Jordan, Fur seals and fur-seal

islands of the North Pacific Ocean, vol. 3, pp. 589-596, pi. 95.
Washington.

1901. Notes on algae I. Zoe, vol. 5, nos. 6-8, pp. 121-129.

1905. Eegeneration among kelps. Univ. Calif. Publ. Bot., vol. 2, no. 5, pp.

139-168, pis. 15-17.
1905«. Post-embryonal stages of the Laminariaceae. Univ. Calif. Publ. Bot.,

vol. 2, no. 4, pp. 115-138, pis. 13, 14.
1908. Nereocystis and PelagopJiycus. Bot. Gaz., vol. 45, pp. 12.5-134. Chicago.
1908a. Critical notes on Laminariaceae. La Nuova Notarisia, ser. 19, pp.

90-101.
1912. Algae novae et minus cognitae I. Univ. Calif. Publ. Bot., vol. 4,

no. 14, pp. 229-263, pis. 25-31.
1912a. The kelps of the United States and Alaska, in Fertilizer resources

of the United States. 62 Congr., 2 Sess., Sen. Doe. 190, pp. 130-178.
1924. The vegetation of Tutuila Island. Carnegie Inst. Wash., Publ. no.

341. Washington.

Setchell, W. A., and Gardner, N. L.

1903. Algae of Northwestern America. Univ. Calif. Publ. Bot., vol. 1,

no. 3, pp. 165-418, pis. 17-27.
1919. The marine algae of the Pacific Coast of North America. Part I,

Myxophyceae. Univ. Calif. Publ. Bot., vol. 8, no. 1, pp. 1-138,

pis. 1-8.

1925] Setchell-Gardner: Melanophyceae 737

1920. The marine algae of the Pacific Coast of North America. Part II,
Chlorophyceae. Univ. Calif. Publ. Bot., vol. 8, no. 2, pp. 139-374,
pis. 9-33.

1922. Phycological Contributions.

II. New species of Myrionema.

III. New species of Compsonema.

IV. New species of Hecatonema.

V. New species of Pylaiella and Strehlonema.
VI. New species of Ectocarpus.
Univ. Calif. Publ. Bot., vol. 7, no. 11, pp. 333-426, pis. 32-49.
1924. The Marine Algae, in Expedition of the California Academy of
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Skottsberg, C.

1907. Zur Kenntniss der Subantarcktischen und Antarktisclien Meeres-
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Smith, A. L., and Whitting, F. G.

1895. Notes on the sori of Macrocystis and Postelsia, in Murray, Phyc. Mem.,
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Smith, J. E.

1790. See English Botany.

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1795-1801. Nereis Britannica; continens species omnes Fucorum in insulis
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1797. Fasc. 2, pp. ix-xviii.

1801. Fasc. 3, pp. xix-xl and 1-112 with appendix, pis. 1-17
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1816. Idon. Ed. 2. Oxonii.

1809. Tentamen marino-cryptogamicum. Mem. Soc. Nat., vol. 2, pp. 90-98.
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Steudel, E. T.

1840-1841. Nomenclator botanicus sen synonymia plantarum universalis,
numerans ordine alphabetic* nomina atque synonyma, turn generica
turn speeifica, et a Linnaeo et a recentioribus de re botanica
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nova elaborata, et aucta.
Ed. 1, 1821.
Ed. 2, part 1, A-K, 1840, pp. 1-852; part 2, L-Z, 1841, pp. 1-810.

Stizenberger, E.

1860. Dr. Ludwig Rabenhorst's Algen Sachsens, resp. Mittel-Europas.
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Stroempelt, H. F. G.

1886. ' Cm Algenvegetationen vid Islands Kuster. Akademisk Afhandling.
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738 University of California Publications in Botany [Vol.8

1886(7. Einige fiir die Wissenschaft iieue Meeresalgen aus Island. Bot. Sekt.

af Naturvet. Studentsallsk i Upsala, in Bot. Centralblatt, vol. 26,

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no. 9, pp. 381-384, pi. 3.

Taylor, W. E.

1922. Recent studies of Phaeopliyceae and their bearing on classification.
Botanical Gazette, vol. 74, no. 4, pp. 431-441. Chicago.
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1850. Eecherches sur les zoospores des algues et les antheridies des crypto-

games. Ann. Sci. Nat., 3 ser., Bot., vol. 14, pp. 214-260, pis. 16-31.

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Thuret, G., and Borxet, E.

1878. Etudes phycologiques. Paris.

TiLDEN, Josephine E.

1894-1909. American Algae (Exsicc).

1894. Century I.

1896. Century II.

1898. Centuiy III.

1900. Century IV.

1901. Century V.

1902. Century VI.

1909. Century VII, fascicle 1.
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1804. Eemarks upon the Dillenian Herbarium. Trans. Linn. Soc. (London),

vol. 7, pp. 101-115 (read, April 19, 1803).
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1808. Vol. 1.

1809. Vol. 2.
1811. Vol. 3.
1819. Vol. 4.

Velley, T.

1795. Coloured figures of marine plants, found on the southern coast of
England; illustrated with descriptions and observations accom-
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To which is prefixed an inquiry into the mode of propagation
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Vickers, Anna

1908. Phycologia Barbadensis. Iconographie des algues marines recoltees
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"Weber van Bosse, Anna

1910. Notice sur quelques genres nouveaux d 'algues de I'archipel malaisien.
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1805-1810. Beitrage zur Naturkunde. In Verbindung mit Ihren Freunden
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13--5] SetcheU-Gardner: Melanophyceae 739

Woodward. T. J.

1797. Observations upon the generic character of Ulva, with the descrip-
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Wyatt, Mary

Algae Danmonienses, or dried specimens of marine plants principally
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1903. HedophjiUuni spirale, sj). nov., and its relation to Thalassiophyllum and

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1913. Some new algae from Japan. Nyt Magazin for Naturvidensk., vol.

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1914. Notes on algae new to Japan II. Botanical Magazine, vol. 28, no.

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1919. A monograph of the genus Alaria. Jour. Coll. Sci. Imp. Univ. Tokyo,

vol. 43, Art. 1, pp. 1-145, pis. 1-19.

1920. Novae algae Japonicae. Bot. Mag., vol. 34, pp. 1-12. Tokyo.

EXPLANATION OF PLATES

PLATE 34

Dictyota Binghamiae J. Ag.

Fig. 1. Habit sketch of a robust plant. X 0.5.

Fig. 2. Cross-section of a frond showing the character of the cells at the
margin. X 125.

Dictyota flabellata (Collins) S. and G.

Fig. 3. Habit sketch of a part of a frond. X 0.5.

Neurocarpus zonarioides (Farlow) Howe

Fig. 4. Cross-section of a frond showing the position of the fruit, the alae
(shortened) and the thickened central portion. X 45.

Zonaria Farlowii S. and G.

' Fig. 5. Habit sketch of a terminal fragment of a frond showing the method
of development. X 1.

[740]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER] PLATE 34

PLATE 35

Punctaria occidentalis S. and G.

Fig. 6. Cross-section of a frond showing the character of the cells, the
character and position of the zoosporangia, and two groups of hairs. X 250.

Dictyota flabellata (Collins) S. and G.
Fig. 7. Cross-section through a group of oogonia. X 325.

JSalorhipis Winstonii (Anderson) Saunders

Fig. 8. Cross-section showing groups of hairs and superficial zoosporangia.
X 125.

Taonia Lennehackerae Farlow

Fig. 9. Cross-section of a frond showing its structure and the position of
the oogonia. X 125.

Fig. 10. Surface view. X 65.

Neurocarpiis zonarioides (Farlow) Howe
Fig. 11. Cross-section showing the characters of the cells. X 200;

Myrionema strangulans Grev.

Fig. 12. A portion of the margin of the basal disk of cells showing the
method of cell division and branching of the filaments. X 125.

[742]

UNIV. CALIF. PUBL. BOT. VOL. 8

fSETCHELL-GARDNER I PLATE 35

0?&#?.<So|°„o

CD

%

r-

0

PLATE 36

Dietyota flaiellata (Collins) S. and G,

Fig. 13. A group of mature hairs. X 125.

Fig. 14. Young plants growing in position from tetrasporangia before
division. X 125.

Fig. 15. Same as fig. 14, further advanced. X 125.

Fig. 16. Cross-section of a frond showing the early stage in the develop-
ment of hairs and young tetrasporangia. X 125.

Fig. 17. A mature tetrasporangium. X 125.

HeterocJiordaria abietina (Eupr.) S. and G.

Fig. 18. Portion of a cross-section of a frond showing zoosporangia, para-
physes, and hairs. X 250.

Fig. 19. A group of gametangia as seen in cross section. X 250.

Zonaria Farlowii S. and G.

Fig. 20. A longitudinal section at the margins (a) and through a young
sorus (&). X 125.

Neurocarpus zonarioides (Farlow) Howe

Fig. 21. Habit sketch of a portion of a frond showing the distribution of
the sori along the mid-rib. X 2.

Balfsia californica S. and G.

Fig. 22. Section perpendicular to the surface of a frond showing the char-
acter of the basal cells, and zoosporangia and paraphyses in various stages of
development. X 125.

[744]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER] PLATE 36

PLATE 37

Spliacelaria californica Sauv.

Fig. 23. Habit sketch showing the method of branching. X 10.

Fig. 24. Cross-section in the mature part. X 125.

Fig. 25. End view of a frond. X 125.
X 125.

Fig. 26. Portion of the main frond and a branch with one i^ropagulum.

Fig. 27. A branch with a terminal zoosporangium. X 125.

Sphacelaria siibfusca S. and G.
Fig. 28. A terminal branch with propagula and a free propagulum. X 125.

Sphacelaria furcigera Kuetz.

Fig. 29. Tenninal portion of a ramulus with a short hair and two propagula.
X 125.

Punctaria liesperia S. and G.
Fig. 30. Habit sketch. X 1.

Pylaiella Uttoralis f.

Fig. 32. Fruiting ends of ramuli showing opposite branching and inter-
calary zoosporaugia. X 125.

[746]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER I PLATE 37

PLATE 38

Elachistea fucicola (Velley) Aresch.

Fig. 33, Habit sketch showing *he relative size of the solid basal portion
to the free filaments. X 8.

Fig. 34. A portion of the fruiting region showing zoosporangia, paraphyses,
and the base of one of the assimilating filaments. X 125.

Fig. 35. A few cells of the basal, widest, and apical portions of an assimi-
lating filament. X 125.

PJmeostrophion irregulare S. and G.
Fig. 36. Cross-section of a mature frond. X 250'.

Endarachne Binghamiae J. Ag.
Fig. 37. Cross-section. X 125.
Fig. 38. Fascicle of hairs and a group of cells in surface view. X 125.

Neurocarpus zonarioides (Farlow) Howe
Fig. 39. Group of hairs. X 200.

[748]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER] PLATE 38

PLATE 39

Soranthera ulvoidea f. typica S. and G.

Fig. 40. Section through a sorus showing zoosporangia, paraphyses, and
hairs. X 250.

Fig. 41. Section through a sorus. X 75.

Fetrospongium rugosum (Okamura) S. and G.

Fig. 42. A few filaments, pei-pendieular to the surface, showing method of
branching, character of the cells, rhizoidal filaments, and zoosporangia. X 125.

Fig. 43. Habit sketch. X 0.5.

Chordaria flagelliformis f. typica Kjellm.

Fig. 44. Portion of a. section showing hairs, paraphyses, and characters of
the cells. X 250.

Scytosiphon Lomentaria f. typicus S. and G.

Fig. 45. Port'on of section with gametangia and one paraphysis (?). X 250.

Sargassum Agardhianum Farlow

Fig. 46. A segment of the frond sho\\'ing the character of a leaf, vesicles,
and receptacles. X 2.5.

[750]

UNIV. CALIF. FUBL. BOT. VOL. 8

[SETCHELL-GARDNERl PLATE 39

PLATE 40

DictyosipJion foeniculaceus (Huds.) Grev.

Fig. 47. Habit sketch of a terminal portion of a frond. X 2.

rig. 48. Tip of a branch showing the character of the surface and hairs.
X 125.

Fig. 49. Portion of a cross-section of a frond showing the character of the
cells, the base of a hair, and a zoosporangium. X 250.

Myelophycus intestinalis f . tenue S. and G.

. Fig. 50. Cross section of a frond showing zoosporangia and paraphyses.
X 250.

Myrionema strangulans Grev.
Fig. 51. Basal filaments bearing erect filaments and zoosporangia. X 250.

Leathesia difformis (L.) Aresch.

Fig. 52. A few surface filaments showing a hair, gametangia and "para-
physes," with large terminal cells filled with chromatophores. X 250.

[752]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNERl PLATE 40

52

PLATE 41

Streblonema pacificum Saunders

Fig. 53. Plants in various stage of development creeping among the
sporangia of the host, with gametangia projecting beyond the host. X 250.

Hapterophycus canalictilatus S. and G.
Fig. 54. Section of the thallus through a cryptostoma highly magnified.

Sargassum Agardhianum Farlow
Fig. 55. Habit sketch of the end of a branch. X 1.

Sargassum Palmeri Grunow.

Fig. 56. Piece of a frond bearing a dissected leaf, a vesicle, and a group
of receptacles. X 1.5.

Cliordaria dissessa S. and G.

Fig. 57. Section perpendicular to the surface showing the character of the
cells, hairs, paraphyses, and zoosporangia. X 125.

[754]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNERl PLATE 41

PLATE 42

Gobia simplex (Saunders) S. and G.

Fig. 58. View of internal filaments extending longitudinally and a group of
surface filaments showing hairs, paraphyses, and zoosporangia, X 250.

Mgira virescens (Carm.) S. and G.

Fig. 59. Surface filaments bearing zoosporangia and hairs. X 250.
Fig. 60. Surface filaments bearing a hair and zoosporangia. X 125.

Scytosiphon Lomentaria f. cylindricus maculatus S. and G.

Fig. 61. Longitudinal section of a part of a frond showing the characters
of the interior and the surface cells. Not fruiting. X 250.

Fig. 62. CToss-section of a portion of a frond showing the characters of the
cells, paraphyses, and gametangia in various stages of development. X 250.

[756] :

1

i

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER] PLATE 42

PLATE 43

Zonaria Farlowii S. and G.

Fig. 63. Habit sketch indicating parallel and radiating lines of cells.
Diagrammatic.

Leathesia amplissima S. and G.

Fig. 64. Section perpendicular to the surface of the frond showing the
characters of the cells, hairs, paraphyses, and zoosporangia. X 250.

Leathesia difformis (L.) Aresch.

Fig. 65. Section perpendicular to the surface of the frond showing the
characters of the cells, hairs, paraphyses, and gametangia. X 250.

Fig. 66. Section perpendicular to the surface of the frond showing the
character of the cells, zoosporangia, gametangia, hairs, and paraphyses X 250.

Leathesia nana S. and G.

Fig. 67. Section through the frond perpendicular to the surface showing
the character of a few filaments bearing hairs, gametangia, and paraphyses.
X 125.

[758]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER I PLATE 43

PLATE 44

Hea Fascia f.

Fig. 68. Cross-section of a frond at the edge showing gametangia in
continuous sori. X 190.

Ilea Fascia f.

Fig. 69. Cross-section of a portion of a frond through several sori on both
sides. X 100.

Ilea Fascia f.

Fig. 70. Cross-section through a portion of a frond showing gametangia
and paraphyses (?). X 500.

Ilea Fascia f.

Fig. 71. Cross-section of a portion of a frond showing mature gametangia
and paraphyses (?). X 125.

Scytosiphon Lomentaria (Lyngb.) J. Ag.

Fig. 72. Cross-section through different parts of the frond: near the base;
farther up; mature gametangia. X 125.

Ilea Fascia f.
Fig. 73. Cross-section through gametangia in sori on both sides. X 125.

Scytosiphon Lomentaria f. complanatus minor S. and G.

Fig. 74. Cross-section through the margin showing short filaments extend-
ing into the cavity and a sorus of gametangia extending around the margin.
X 125.

Scytosiphon Lomentaria f. typicus S. and G.

Fig. 75. Cross-section showing the character of the cells, gametangia, hairs,
and paraphyses (?). X 250.

[760]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNERl PLATE 44

68

PLATE 45

Coilodesme rigida S. and G.
Fig. 76. Section perpendicular to the surface of the frond. X 125.

Coilodesme hulligera Stroemf.

Fig. 77, Surface view. X 250.
Fig. 78. Cross-section. X 250.

Punctaria chartacea S. and G.
Fig. 79. Cross-section of a sterile frond. X 250.

Sargassum Liebmaimi J. Ag.

Fig. 81. A segment showing the character of the stem, leaves, fruit, and
vesicles. X 2.

Colpomenia sinuosa f.

Figs. 82, 83. Cross-section showing unusually long gametangia. X 250.

Fig. 84. Cross-sections. X 250'.

Fig. 85. Cross-section showing paraphyses and enlarged at outer end.
X 250'.

Fig. 86. Cross-section showing some gametangia divided. X 250.

[762]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER] PLATE 45

82

i^cm

®^

78

ggsaooas^^^

«Q^^^

a

^^^Q^asaoDooee

77

79

81

PLATE 46

Coilodesme californica (Eupr.) Kjellm,

Fig. 1. Surface view. X 250.
Fig. 2. Cross-section. X 250.

Coilodesme corrugata S. and G.
Fig. 3. Surface view. X 250.
Fig. 4. Cross-section. X 250.

Sargassum paniculatum J. Ag.

Fig. 5. Segment showing the characters of the stem, leaves, receptacles,
and vesicles. X 1.5.

Pelvetiopsis limitata f. typica Gardner
Fig. 6. Diagrammatic section through a conceptacle. Stained.

Ectocarpus confervoides f. variabilis Saunders
Fig. 7. Hamulus showing gametangia and zoosporangia (?). X 250.

[764]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER] PLATE 46

^

^i

PLATE 47

Myriogloia Andersonii (Farlow) Kuckuck.

Fig. 8. Habit sketch of the tip of a frond, highly magnified.

Fig. 9. Young, colorless filaments giving rise to young cortical filaments
and an assimilating hair. Highly magnified.

Fig. 10. Segment of the surface showing zoosporangia and clavate para-
physes. X 375.

Meneghiniella Brandegeei S. and G.

Fig. 11. Fascicle of gametangia, highly magnified.

[766]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER] PLATE 47

I

PLATE 48 ;

)
Myriogloia Andersonii (Farlow) Kuckuck.

Figs. 12, 13, 14. Segments of the surface of three different collections i

showing variations in shapes and sizes of the different parts. X 375. '

[768]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER] PLATE 48

PLATE 49

Ectocarpus Farlcsii S. and G.
Fig. 15. Portions of a frond taken at different levels. X 250.

MenegMmella Brandegeei S. and G.

Fig. 16. Portion of the surface of a frond showing cortical assimilating
filaments and gametangia. Highly magnified.

Myriogloia Andersonii (Farlow) Kuckuck.

Fig. 17. Segment of the surface of a filament showing stipitate zoo-
sporangia and irregularly clavate, uncinate assimilating filaments. X 375.

Punctaria Jiesperia S. and G.

Fig. 18. Sections of a sterile, and of a fruiting frond, respectively show-
ing protruding conical gametangia. X 250.

Punctaria expansa S. and G.
Fig. 19. Section showing gametangia and hairs. X 250.

[770]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER] PLATE 49

¥

18

16

PLATE 50

Chlanidophora abyssicola S. and G.

Fig. 1. Two specimens of mature plants. Natural size.

i

Fig. 2. Young aplanosporangium in the distromatic portion of the frond. '

The two small cells below represent cross-sections of hairs. |

Fig. 3. Gross-section showing three mature aplanosporangia with numerous ■

aplanospores and two paraphyses. ;

Fig. 4. Surface view of a portion of the growing margins of a frond show-
ing the method of cell division. i

Fig. 5. Portions of four cross-sections showing the character of the cells in

different parts of the frond. The upper is the younger, near the growing margin. i

The others are successively older toward the base. j

Fig. 6. Longitudinal section through the margins. ]

Fig. 7. Longitudinal section in the older part of the frond showing the i

origin and prostrate position of hairs. Figures 2-7. X 250. |

. \

Phaeostrophion irregulare S. and G. ]

Fig. 8. Cross-section of the frond with zoosporangia. X 250. '

[772]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNERl PLATE 5C

8

PLATE 51

Ectocarpus socialis S. and G.

Fig. 12. Small group of plants showing rhizoids, a few cells with chro-
matophores, and the positions and shapes of the zoosporangia. X 125.

Fig. 13. Selection of parts of filaments showing the positions and relative
shapes and sizes of the gametangia. X 125.

Ectocarpus cylindricus f. codiopMlus S. and G.

Fig. 14. Group of young plants showing both zoosporangia and game-
tangia, the two parts of ramuli on the left representing these organs on the
same plant. X 60.

Ectocarpus Taoniae S. and G.

Fig. 15. Portion of a creeping filament showing erect filaments and the
positions and relative shapes and sizes of the gametangia on both the creeping
and the erect filaments. X 250.

Ectocarpus afflnis S. and G.
Fig. 16. Diagrammatic representation of a small group of plants on their host.
Fig. 17. Portions of filaments illustrating positions, relative shapes and
sizes of gametangia. X 225.

This plate is a reprint of Setchell and Gardner, Univ. Calif. Publ. Bot., vol. 7,
p. 420, pi. 46, 1922.

[774]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNERl PLATE 51

^5>^0^'

PLATE 52

Strehlonema anomalum S. and G.
Fig. 1. Part of a plant freed from its host bearing gametangia. X 250.

Figs. 2, 3. Fragments of plants bearing zoosporangia, some of which contain
mature zoospores. The cells of the main filaments ai"e divided longitudinally.
X 250.

Strehlonema Johnstonae S. and G.

Fig. 4. Parts of plants showing characteristic creeping and erect filaments
and gametangia. X 125.

Strehlonema rugosum S. and G.

Fig. 5. Section through the host perpendicular to its surface, showing the
papillate character caused by the presence of Strehlonema. Diagrammatic.

Fig. 6. Surface view of figure 5. Diagrammatic.

Fig. 7. Section showing the structure of the Strehlonema plants and their
relation to the host. X 250.

Strehlonema corymhiferum S. and G.
Fig. 8. Fragments of typical plants freed from their host. X 250.

This plate is a reprint of Setchell and Gardner, Univ. Calif. Publ. Bot., vol. 7,
p. 400, pi. 43, 1922.

[776]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNERl PLATE 52

PLATE 53

Hecatonema clavatum S. and G.

Fig. 1. Section through a typical mature plant showing penetrating rhizoids
from the basal layer, the leavate, sterile, ereet filaments, and typical game-
tangia. X 250'.

Fig. 2. Same as figure 1, but showing "ascocysts. " X 250.

Fig. 3. Small fragment without rhizoids. X 125.

Fig. 4. Segment of the base at its margin. X 250.

Hecatonema Lawsonii S. and G.

Fig. 5. Two segments of the base showing frequent radial divisions of the
cells. X 250.

Fig. 6. Young plant before any erect filaments have begun to develop.
X 250.

Fig. 7. Typical gametangia and true hairs. X 250.

This plate is a reprint of Setchell and Gardner, Univ. Calif. Publ. Bot., vol. 7,
p. 382, pi. 40, 1922.

[778]

UNIV. CALIF. PUBL. EOT. VOL. 8

[SETCHELL-GARDNER] PLATE 53

PLATE 54

Compsonema secundum S. and G.

Fig. 1. Three fragments of plants showing small gametangia on short
pedicels from a basal filament and complex terminal gametangia with mostly
secund lateral developments. X 125.

Fig. 2. Fragment of a plant showing a modification in which the game-
tangia are mostly temiinal and erect, and only rarely possessing lateral pro-
tuberances. X 125.

Compsonema pusillum S. and G.

Fig. 3. Three fragments of plants showing shapes and positions of zoospo-
rangia and gametangia. Character of the creeping and of the erect filaments,
and a feAv branches pushing between the surface cells of the host. X 125.

Compsonema secundum f . terminale S. and G.

Fig. 4. Fragment of a plant showing the characteristic terminal and large
lateral gametangia. X 125.

Fig. 5. Filament which seems to be producing zoosporangia below an empty
gametangium. X 125.

Compsonema tenue S. and G.

Fig. 6. Fragments of plants showing erect filaments, true hairs, and zoospo-
i-angia (?) and gametangia sessile on the creeping filaments. X 250.

This plate is a reprint of Setchell and Gardner, Univ. Calif. Publ. Bot., vol. 7,
p. 372, pi. 37, 1922.

[780]

UNIV. CALIF. PUBL. BOT. VOL. 8

[SETCHELL-GARDNER] PLATE 54

PLATE 55

Myrioncma minutissimum S. and G.

Fig. 1. Two groups of gametangia, the left, youuger, and the right, older.
X 125.

Fig. 2. Segment of the base showing the typical splitting of the terminal
cells. X 250.

Fig. 3. Small groups of gametangia showing different lengths of pedicels.

X 250.

Myrionema attenuatum f. doUiforme S. and G.

Fig. 4. Fragment showing relatively large erect sterile filaments and rela-
tively small and few gametangia. X 250.

Fig. 5. Fragment showing the opposite condition to that of figure 4. X 250.

Myrionema attenuatum S. and G.

Fig. 6. Fragment taken near the margin of the frond. X 2'50'.
Fig. 7. Fragment taken near the center of the frond. X 250.

Myrionema foecundum f. divergens S. and G.
Fig. 8. Sections taken from four different places in the frond., X 125.
Fig. 9. Segment of the base at the margin of the frond. X 250.

Myrionema compsonematoides S. and G.

Fig. 10. Section of a characteristic frond showing the erect sterile filaments,
gametangia, and "ascocysts. " X 125.

Myrionema hecatonematoides S. and G.

Fig. 11. Five fragments showing the nature of the base, the shapes, sizes,
and positions of the gametangia, and the erect filaments. X 125.

Myrionema primarium S. and G.

Fig. 12. Section through a typical frond and its host perpendicular to the
latter. X 250:

This plate is a reprint of Setchell and Gardner, Univ. Calif. Publ. Bot., vol. 7,
p. 352, pi. 34, 1922.

[782]

UNIV. CALIF. PUBL. BOX. VOL. 8

fSETCHELL-GARDNERl PLATE 55

PLATE 56

A. Laminaria Farlowii Setehell

From a photograph of a plant showing typical buUations, but abnormal as to
the deep splitting of the blade.

B. Costaria costata (Turn.) Saunders
From a photograph of a typical plant of moderate dimensions.

[784]

INDEX

Univ. Calif. Publ. Bot., vol. 8, part III

Titles of papers, and names of new taxononiic units, also pages where
diagnoses appear are iu boldface.

Aeniaea, 433.

Aciileatae, 563.

Adenocystis (Lessonii var. ?) califor-
nica, 580.

^gira, 508, 544, 546, 547.

virescens, 547; figures showing,
opp. 756.
-lEgiraceae, 507, 508, 509, 543, 558,

570.

Agarum, 593, 609, 613, 614.

lirassieaeforme, 614.

cribrosum, 614, 615, 616; plate
showing, opp. 798.

Delisei, 614.

esculentum, 614.

fimbriatum, 604, 616; plate show-
ing, opp. 814.

Gmelini, 614, 615.

pertusum, 614, 615.

platvneurum, 614.

Pylaii, 614, 638, 639, 641.

quinquecostatum, 614.

Turneri, 614, 615.

Agareae, 609.

Alaria, 448, 614, 631, 634, 635.
cordata, 640.
crispa, 642.
curtipcs, 640.
dolicliorhachis, 642, 643.
fistulosa, 421, 644.

f. platyphylla, 645.

f. stenophylla, 645.
grandifolia, 644.
lanceolata, 637, 642, 643.
laticosta, 643.
macroptera, 643.

marginata, 459, 637, 640, 641, 644;
plate showing, opp. 804.

f. nana, 637.
nana, 636, 637, 640.
praelonga, 637, 638, 641.

f. nana, 637.
Pylaii, 638, 639, 641.

f. longipes, 639.

f. typiea, 639.
taeniata, 642.
tenuifolia, 449, 638, 639.

f. amplior, 639.

f. tvpica, 638.
valid;!, 643, 644.

f. longipes, 644.

Alariaceae, 613, 633.

Alarieae, 633.

Analipus, 550, 575.
fusifonnis, 575, 570.

Aplanosporeae, 649.

Arthrocladia, 554, 558, 559.

Arthrothamnus, 619, 620.

bifidus, 619, 620.

Bongardianus, 619.

kurilensis, 620.

radicans, 620.

spirale, 619.
Ascocyelus, 455, 460, 461, 465, 469.

baltieus, 465, 466.

gloliosus, 470.
Asperoeoccaceae, 522, 530', 587.
Asperococcus, 524, 525, 650.

eompressus, 523, 524.

Cystoseirae, 579, 583.

Bachelotia, 401, 402.
Bacillariales, 388.
Blossevillea, 705.

Brandogeci, 705.
Callithanmion, 432.
Castagnea, 544, 546, 548.

divaricata, 574.

fistulosa, 546.

niediterranea, 546.

polycarpa, 545, 546.
Ceramium confervoides, 412.
Chaetomorpha aerea, 422.
Chactopteris, 393, 397.

plumosa, 398.
Chaetophoroideae, 454.
Chilionema ocellatum, 460.
Chlanidote, 658.
Chlanidophora, 658.

abyssicola, 658; figures showing,
opp. 772.

microphylla, 658.
Chlorophyceae, 388, 445.
fhlorospermeae, 454.
Chnoosi3ura, 552, 553.

atlantica, 553.

fastigiata, 553.
var. pacifica, 553.

implexa, 553.
pacifica, 430, 553.
pannosa, 553.

[889]

Index

Chnoosporaceae, 552.
Chorda, 592.

Filum, 531, 592.
Lomentaria, 532.
Chordaceae, 591.

Chordaria, 507, 543, 570, 571, 572.
574, 576, 577, 588.
abietina, 550, 551, 577.
Andersonii, 557.
Cabrera, 572.

dissessa, 574, 575; figure showing,
opp. 754; plate showing, opp. 822.
divaricata, 572, 574, 575.
rilum, 572.

flagelliformis, 507, 543, 550, 557.
570, 572.
f. chorda eformis, 573.
f. ramuseulifera, 573.
f. typica, 573; figure showing,
opp. 750.
gracilis, 573; plate showing, opp.
840.
• Gunjii, 550.
Lomentaria, 531.
rotunda, 572.
Chordariaceae, 453, 455, 507, 543,

549, 550, 570, 571.
Chordariales, 388, 550, 570, 578.
Chordarieae, 454, 571, 577.
Choristocarpaceae, 431, 649.
Cladophora, 461.

albida, 418.
Cladosiphon, .544, 545, 546, 548.
balticus, 576.
erythraeum, 549.
fistulosum, 545.
mediterraneus, 544, 545, 546.
zostericola, 548, 549.
Cladostephus verticillatus, 392.

Codiuni Brandegeei, 440.
cuneatum, 439.

fragile, 425, 433, 437, 438, 439.
Setchellii, 424, 433.

Coilodesmaceae, 570, 571, 577.

Coilodesme, 570, 577, 578, 581.

amplissima, 580.

bulligera, 552, 578, 581, 582 ; figures
showing, opp. 762.

californica, 579, 580, 581, .583,
584; figures shoAving, opp. 764;
plate showing, opp. 844.

corrugata, 582; figures showing,
opp. 764.

Cystoseirae, 578, 583.

linearis, 583, 584.

polygnampta, 581, 582; plate show-
ing, opp. 836.

rigida, 584; figures showing, opp.
762, 830.

sitchcnsis, 580, 583, 584.

Coilodesmeae, 577.

Coilonema, 586, 587.
Colophermum, 407, 408.
Colpomenia, 535, 539, 552.

sinuosa, 539; figures showing, opp.
762.
f. deformans, 487, 542.
f. expansa, 540.
f. expansissima, 541.
f. tuberculata, 541, 542.
f. typica, 540.
tuberculata, 541.
Compsonema, 455, 467, 473, 474, 475,
477, 478, 479, 480, 481, 482, 483.
485, 488, 491.
coniferum, 479.
dubium, 479.
fasciculatum, 478.
fructuosum, 476.
gracile, 473, 476, 486.
immixtum, 487.
intrieatum, 482, 483.
myrionematoides, 474.
numniuloides, 475.
pusillum, 477; figure showing, opp.

780.
ramulosum, 484.

secundum, 483, 484; figures show-
ing, opp. 780.
f. terminale, 484; figures show-
ing, opp. 780.
serpens, 479, 480.
sessile, 474.
speciosum, 485.

f. piliferum, 485.
sporangiiferum, 481, 482.
streblonematoides, 481.
tenue, 478; figure showing, opp.
780.
Conferva albida, 418.
foeniculacea, 589.
fueicola, 504.
granulosa, 426.
littoralis, 401, 403.
scutulata, 502.
siliculosa, 411.
tomentosa, 417, 418.
Constantinea rosa marina, 693.
Corycus, 514.
Corynephora, 510.

marina, 511.
Corynophlaea, 507, 508, 510.

baltica, 510.

Corynophlaeaceae, 507.

Costaria, 593, 609, 614, 622, 623.

costata, 459, 472, 604, 608, 610,

611; figures showing, opp. 784,

830.

Mertensii, 610, 611.

quadrinervia, 610.

reticulata?, 610, 622.

Turneri, 609, 610, 611.

var. pertusa, 610, 612.

[890]

Index

duoria verrucosa, 498.

Cumagloia Andersonii, 441, 442, 444.

Cutleriales, 388, 389, 399.

Cyclosporeae, 662.

Cylindrocarpus, 507, 508.

mieroscopicus, 502, 507, 508, 509.
rugosa, 510.

Cymnthnere, 593, (i07, 608.
triplicata, 604, 608.

Cymathaereae, 606.

Cystophora, 705.

Cystophyllum, 578, 579, 706, 714.
gemiiiatum, 478, 580, 583, 584, 706.
hakodateiise, 578.
Lepidium, 584, 706.

Cystoseira, 578, 579, 708.
amplissima, 582.
ericoides, 708.
expansa, 709.
geminata, 707.
Lepidium, 579, 706.
negleeta, 582, 710.
osmuiidaeea, 432, 435, 580, 709, 710.

f. expansa, 709.

f. typica, 709.
Setchellii, 710.
thyrsigera, 579.

Delesseria sanguinea, 466.

Desmarestia, 414, 554, 558, 559, 560,
569.
aculeata, 562, 564, 565.
subgen. Dichloria, 561.
subgen. Eudesniarestia, 563, 565.
farcta, 562.
foliaeea, 569.
herbacea, 416, 446, 566; plate

showing, opp. 848.
intermedia, 564, 565.

var. fuscescens, 565.
latifron.s, 563, 564,565; plate show-
ing, opp. 852.
latissima, 568, 569.
ligulata, 566, 567; plate showing.

opp. 846.
media, 561, 562, 56,5.

var. tenuis, 561.
munda, 567, 568, 569; plate show-
ing, opp. 850.
paeifiea, 562.
tabacoides, 569.
viridis, 562.

Desmarestiaceae, 558.

Desmarestiales, 388, 389, 399, 554,
570.

Desmia herbacea, 567.

Desmotrichum, 514, 515, 516, 517.
balticum, 516.
undulatum, 516.

[891]

Dichloria, 559.

Dictyoneurum, 610, 621, 622, 623.
californicum, 622; plate showing,
opp. 812.

Dictyopteris, 656.
zonarioides, 656.

Dictyosiplion, 576, 586, 587, 650.
Chordaria

f. gelatinosa, 590.
foeniculaceus, 531, 574, 588, 589,
590; figures showing, opp. 752.
f. americaiius, 588.
hippuroides, 589, 590.
hispidi-s, 588, 589.
tenuis, 588, 589.

Dictyosiphonaceae, 587.

Dictyosiphonales, 388, 389, 399, 570,
586, 587.

Dictyota, 651.

Bartayresiana, 654.

Binghamiae, 652,653; figures show-
ing, opp. 740.

crenulata, 655.

dichotoma, 649, 653.

flabellata, 652; figures showing,
opp. 740, 742, 744.

hesperia, 655.

Johnstonii, 653, 654.

Kunthii, 652.

liturata, 654.

marginata, 654.

Pappcana, 654.

pinnatifida, 654.

sandvicensis, 655.

Vivesii, 654.

Dictyotaceae, 649, 650, 651.
Dietyotales, 387, 388, 649, 650.
Dilophus, 651, 653.
Diplostromium, 515, 517.
Ecklonia, 654.
radiata, 646.

Ecklonieae, 645.

Ectocarpaceae, 398, 400, 403, 453,
455, 458, 487, 502, 503, 508, 509.

Ectocarpales, 388, 389, 391, 398, 399.

461, 570, 649.
Ectocarpeae, 400.
Eetocarpineae, 398.
Ectocarpus, 401, 407, 408, 409, 413.
418, 443, 452, 477, 478, 479, 480,
481, 483, 502, 587.
acuminatus, 435.
acutus, 415, 416, 419, 425.
aecidioides, 451.
affinis. 432; figures showing, opp.

774.
amphibius, 412.
breviartieulatus, 429, 430.
Bryantii, 440.
chantransioides, 430, 431, 508, 509.

Index

chitonicola, 436.
commensalis, 424.
confervoides, 411,
421, 425.

412, 413, 415.

f.
f.
f.
f.
f.
f.
f.

acuminatus, 416.

nanus, 415.
parvus, 413.
pygmaeus, 415.
subulatus, 411, 412.
typicus, 414, 416.
variabilis, 415,418,423; figure
showing, opp. 764.
corticulatus, 418, 419, 425.
cylindricus, 432, 437, 438.
f. acmaeophilus, 433.
f. codiophilus, 433, 438; figure
showing, opp. 774.
draparnaldiaeformis, 412.
Duchassaingianus, 428.
ellipticus, 436.
eramosus, 424, 425.
flagelliferus, 422, 423.
flocculiformis, 438.
fruetuosus, 419.
globifer, 437, 438.
gonodioides, 439, 440.
granuLitus

f. corticulatus, 419.
granulosoides, 427, 431.
granulosus, 420, 426, 429, 431.

f. corticulatus, 426.
hamatus, 430.
hemisphericus, 430, 434, 502, 508.

f. minor, 434.
littoralis, 408.

5 protensus, 401, 403.
luteolus, 425.
macroceras, 412.
Mesogloiae, 423.
Mitchellae, 428.
mueronatus, 419, 420.
oviger, 428, 429.
Padinac, 420.
paradoxus

var. pacificus, 435.
Parksii, 427; figure showing, opp.

770.
penicillatus, 416.
Saundersii, 434.

siliculosus, 401, 410, 411, 412, 413.
parvus, 413.
f. subulatus, 411, 412.
f. typicus, 414.
f. varians, 408.
simulaiis, 422.
socialis, 437; figures showing, opp.

774.
subulatus, 411.
Taoniae, 427; figure showing, opp.

774.
terminalis, 415, 421.
tomentosoides, 417.
tomentosus, 417, 418.
viresceus, 416, 428, 437.

[892]

Egregia, 613, 647.
laevigata, 648.

f. borealis, 649.
Menziesii, 432, 459, 464, 524, 647,
648, 649.

Egregieae, 647.
Eisenia, 645, 646.

arborea, 646.

bicyclis, 646.

Elachisteaceae, 454, 493, 502, 503,

507.

Elachistea, 502, 503, 505.

fucicola, 503, 504, 505; figures

showing, opp. 748.
lubrica, 504, 505.
pulvinata, 507.
scutulata, 503.

Elachisteae, 502.
Encoeliaceae, 552.
Encoelium clathratum, 543.

Cystoseirae, 583.
Endarachne, 530, 538, 585.

Binghamiae, 536, 538; figures show-
ing, opp. 748, 838.
Eudesmarestia, 559.
JEudesme, 544, 545, 546, 548.
virescens, 545, 546, 547.
Zosterae, 546.
Eupylaiella, 401, 402.
Flagellates, 388.
Eucaceae, 663.

Fucales, 387, 388, 390, 650, 663.
Fucus, 435, 504, 663, 664, 702.
Agarum, 615.
bifidus, 620.
bursigerus, 676.
ceranoides

f. Harveyanus, 704.
clathratus, 543.
Clathrus, 614.
costatus, 609, 610.
cribrosus, 615.
edentatus, 678, 700.
f. acutus, 680.
f. costatus, 680, 695.
f. divaricatus, 681; plate show-
ing, opp. 880.
f. divergens, 679.
f. hesperius, 678; plate showing,
opp. 878.
ericoides, 708.

evaneseens, 670, 681, 683, 684, 688,
689, 690, 698.
f. angustus, 680, 694, 695.
f. bursiger, 691, 695.
f. bursigera, 677.
f. contractus, 695.
f. cornuta, 684.
f. cornutus, 692.
f. costatus, 688.

Index

f. cuneatus, C98.

f. dcndroides, 690, 691.

f. ecostatus, 697.

f. flabollatus, 689.

f. intermedins, 688.

f. irregularis, 693.

f. limitatus, 694.

f. loiigifructus, 689.

f. maerocephalus, 684, 693.

f. magiiificus, 669, 686.

f. margiiiatus, 690.

f. nanus, 698.

f. oregonensis, 696; plate show-
ing, opp. 884.

f. pergrandis. 670, 685, 688.

f. robustus, 687; jilate showing,
opp. 886.

f. rudis, 692.

f. stellatus, 687.

f. typica, 684.

f. tvpicus, 683.
Fasc'ift, 536.
fastigiatus, 701.
Fihim, .592.
flagelliformis, 572.
flavus, 660.
fungiformis, 494, 499.
fureatus, 483, 664, 665, 689.

f. alibreviatus, 672; plate show-
ing, opp. 868.

f. angustus, 673, 679. '

f. eontortus, 668.

f. cornutus, 671.

f. elongatus, 667.

f. latifrons, 670; plate showing,
opp. 874.

f. linearis. 672.

f. luxurians, 666, 667; plate
showing, opp. 870.

f. nigricans, 671.

f. reflexus, 669, 676.

f. rigidus, 668, 670, 690; plate
showing, opp. 872.

f. typieus, 665, 667.

f. variabilis, 669.
Harveyanus, 704.
herbaceus, 567.
inflatus

f. edentatus, 665.

f. filiformis, 681.
Luetkeanus, 624.
membranaf-eus, 673.

f. abbreviatus, 676.

f. acuminatus, 677.

f. latissimus, 675; plate show-
ing, opp. 882.

f. limitatus, 675.

f. obtusus, 677.

f. typieus, 674.
Menziesii, 647, 648.
nitens, 699; plate showing, opp.

876.
onustus, 706.

osmundaceus, 709.
pilulifer, 713.
platycarpus, 685, 686, 687.

f. magnificus, 686.
pseudoaculeatus, 565.
pyriferus, 627.
radiatus, 046.
saccharinus, 595.
spiralis, 690.
Tournefortii, 660.
vesiculosus, 693.

Gigartina radula, 468, 478, 479, 480.
Giraudia, 502.

Glossophora Kunthii, 652, 653.
Gobia, 576, 587.

baltica, 576, 577.

simplex, 576, 577; figures showing,
opp. 756, 828.

Gonodia, 505.

JoluKstonii, 506, 507.
Mart'hantae, 506, 507.
moniliformis, 507.
pulvinata, 507.

Gracilaria. confervoides, 427.
Gymnosorus, 659.

Hafgygia

Andersoni, 605.
Bongardiana, 605.

f. subsessilis, 619.
Sinclairii, 598.

Halicoceus, 663.

Halidrys, 578, 663, 707.

dioica, 435, 584, 707.

siliquosa, 707.

Haligenia, 591.

Haliseris polypodioides, 656.

Haloglossum, 523, 524, 525.
Griffithsianum, 524.

Halorhipis, 523, 524.

Winston ii, 523, 524, 525; figure
showing, opp. 742.
Halosaccion, 505.
Halothrix, 502.
Halyseris, 656.

Hapalospongidion gelatinosum, 493.
Haptorophyeus, 499, 500.

canaliculatus, 500; figure showing.

opp. 754; plate showing, opp.

826.

Heeatonema, 455, 467, 473, 476, 488,

490, 491.

clavatuni, 491; figures showing, opp.

778.
diffusum, 486.
Lawsonii, 489; figures showing.

opp. 778.
maculans, 488.
speeiosum, 485.
variabile, 490.

[893]

Index

Hedophylleae, 616,
Hedophvllum, 617, 620.

sessile, 451, 475, 617, 618, 619;

plate showing, opp. 818.
subsessile, 593, 618, 619.
Heterochordaria abietina, figures
showing, opp. 744.

Helminthocladia calvadosii, 453.
Herbaccae, 565.
Hesperophycus, 702, 703.
Ilarveyaiius, 434, 446, 704.

Heterochordaria, 550, 552.

abietina, 550, 551, 577; figures
showing, opp. 744; plate show-
ing, opp. 854.
Heterochordariaceae, 549, 570, 571,
576.

Himanthalia, 663.
lorea, 486.

Homoeostrichus, 659, 660.

Homoeostroma, 515, 516.

latifolia, 520.

latifolium, 519.

lobatum, 519.

undulatum, 522.

Hydroclathrus, 541, 542, 650.
cancellatus, 543.
clathratus, 542, 543.

Ilea, 535, 585, 586.

Fascia, 535; figures showing, opp.
760.
f. caespitosa, 537.
f. debilis, 537.
f. filiformis, 537.
f. typiea, 537.
f. zosterifolia, 537.

Laminaria, 593, 594, 607, 636.
Agardhii, 596.

Andersonii, 449, 452, 604, 605, 646.
Anthrothamnus? longipes, 597.
apoda, 618.
atrofulva, 606.
bifida, 620.
Bongardiana, 601, 604, 605, 606.

619.
bullata, 600, 601.

f. simplex, 601, 602.
complanata, 467, 596, 597; plate

showing, opp. 786.
costata, 609, 611.

cuneifolia, 594, 600, 601; figures
showing, opp. 790; plate show-
ing, opp. 792.
f. amplissima, 603.
f. angusta, 602.
f. subsimplex, 602.
debilis, 517.
dentigera, 604.

f. brevipes, 604.
dermatodea, 609.

Despreauxii, 639.

ephemera, 463, 599, 603; plate

showing, opp. 788.
Farlowii, 468, 599; figure showing,

opp. 784.
fissilis, 606.
groenlandica, 451, 601.
latifolia, 600.
longicruris, 451.
longipes, 597, 598.
musaefolia, 639.
nigripes, 606.
pallida, 458.
personata, 599, 600; plate showing,

opp. 794.
platyloba, 605.
platymeris, 601, 604, 605, 606, 619.

f. complanata, 606.

f. compressa, 606.

f. elliptica, 606.

f. oblonga, 606.

f. reniformis, 606.

f. subteres, 606.

f. taeniata, 606.
porra, 630, 631.
Pylaii, 641.
remotifolia, 639.
Euprechtiana, 597.
saccharina, 451, 595, 598, 600.

f. complanata, 597.

f. linearis, 596.

f. membranacea, 596.
sessilis, 618.

Sinelairii, 459, 597, 598.
solidungula, 599, 600.
taeniata, 605.
triplicata, 608.

Laminariaceae, 414, 587, 593, 607.
616, 621, 636.

Laminariales, 387, 389, 390, 399, 590,
593, 621.

Laminarieae, 594.

Laminarius, 594.

Leathesia, 508, 510, 511.

amplissima, 513; figure showing,

opp. 758.
difformis, 511, 513; figures show-
ing, opp. 752, 758.
nana, 511; figure showing, opp. 758.

Leathesiaceae, 507, 508, 509.

Leptonema, 502.

Lessonia, 622, 623, 631, 632.
fuscescens (?), 632.
littoralis, 631, 632.
nigrescens, 632.
repens, 597.
Sinelairii, 598.

Lessoniaceae, 613, 621, 631.

Lessonieae, 621, 623.

Lessoniopseae, 631.

[894]

Index

Lessoiiiopsis, 621, 631, 633.

littornlis, 632, 633, 636; plates
showing, opp. 806, 808.
Liebmnnnia, 447, 544, 545.

Levillei, 548.

Posidoiiiae, 545, 546, 548, 549.

Linckia Zosterae, 546, 547.
Linkia clavata, 495, 496.
Literature cited, Melanophyceae, 722.
Litosiphon, 514.

Lithodernia, 494, 500.
fatiscens, 500, 501,
Hgnieola, 501.

Lithodermataceae, 493.

Macrocysteae, 623, 626, 629.

Macrocystis, 626, 627, 628, 630.

integrifolia, 628; plate showing,
opp. 796.

Menziesii, 648.

olitusa, 648.

pyrifera, 457, 468, 627, 628, 629;
plates showing, opp. 800, 802.
Melanophyceae, 387, 388, 389, 408.
414, 445, 454, 549, 570, 571, 650.
Melanospermeae, 387, 454.
Mencghiniella, 545, 546, 548.

Brandegeei, 548, 549; figures show-
ing, opp. 766, 770.

erythraea, 549.

zostericola, 549.
Mesogloia, 544.

Andersonii, 554, 555, 556, 557.

fistulosa, 545, 546.

natalensis, 556.

simplex, 576, 577.

vermic'ulata, 544.

vireseens, 545, 547, 548.
Mesogloiaceae, 453, 454, 544.
Mesospora, 494.
Microcoryne, 507, 508.
Mierospongium, 455, 492.

gelatinosum, 493.

globosum, 470.

Saundersii, 492, 493.

Myelophycus, 527, 550, 576.

cacspitosum, 527, 528.

intestinale, 527, 528.

f. tenue, 528; figure showing,
opp. 752.
Myriactis, 505, 507. v

Sargassi, 507.
Myrioeladia, 544, 556.

callitricha, 554, 555, 556.

capensis, 554, 556.

Chorda, 554, 555.

grandis, 554.

Kuromo, 554.

Lovenii, 555.

natalensis, 554.

Sciurus, 554, 555.

Myriocladieae, 555.
Myriogloea, 554.
Myriogloeeae, 555.
Myriogloia, 555, 556.

Andersonii, 416, 423, 555, 556, 557;
figures showing, opp. 766, 768,
770; plate showing, opp. 824.
Myriogloiaceae, 554, 565, 558, 570.
Myrionenia, 448, 452, 454, 455, 460,
467, 468, 473, 475, 476, 477, 481,
483, 487, 488, 490, 496.
attenuatum, 468; figures showing,
opp. 782.
f. doliiforme, 468; figures show-
ing, opp. 782.
bnltieum, 465.

f. californic'um, 465.
f. pedicellatum, 466.
clavatum, 495.
compsonematoides, 467; figure

showing, opp. 782.
corunnae, 458, 464.
f. angulatum, 459.
f. sterile, 460.
f. uniforme, 458, 459.
foecundiim, 460, 464, 465.
f. californicuni. 470.
f. divergens, 463; figures show-
ing, opp. 782.
f. niajus, 463.
f. ramulosum, 462.
f. simplicissimum, 461, 463, 467,

489.
f. subulntum, 462.
globosum, 470.
f. affine, 471.
hecatonematoides, 467; figure show-
ing, opp. 782.
minutissimum, 464; figures show-
ing, opp. 782.
oliscurum, 472.
orbiculare, 460.
(Phycocelis)
balticum, 465.
foecunda, 460.
globosum, 470.
phyllophilum, 469, 470, 490.
primarium, 456, 457; figure show-
ing, opp. 782.
f. acuminatum, 457.
setiferum, 470.
speciosum, 487.

strangulans, 454, 461, 471, 472,
489; figures showing, opp. 742.
752.
vulgare, 461, 489.

Myrionemaceae, 453, 454, 455.

Myrionemntaceae, 453, 454, 458, 480.

487, 493, 494, 503.
Myrionemeae, 455.
Mytilus, 406.
Myxophyceae, 388.

r895]

Index

Nematophloea, 515, 517.
Nereia, 545.

Nereocvstis, 623, 624, 629, 630, 631.
645.
gigantea, 630, 631.
Luetkeana, 419, 429, 447, 450, 458.
460, 461, 462, 464, 467, 474,
475, 476, 477, 481, 482, 484,
485, 486, 489, 490, 491, 624.
priapus, 624.

Neurocarpus, 655.
Cokeri, 656.

zonarioides, 656; figures showing,
opp. 740, 742, 744, 748, 862.

Nostochinae, 454.
Odonthalia, 526.
Omphalophyllum, 388, 513, 514.
Opospermum, 503.

Padina, 657, 661.

deusta, 499.

Durvillaei, 661; plate showing, opp.
858.
Panthocarpus, 401, 402.
Pelagophycus, 623, 624, 629, 630, 631.

giganteus, 630.

porra, 630.

Pelvetia, 700, 702.

fastigiata, 434, 701, 702, 703.
f. gracilis, 702; figure showing,

opp. 828.
f. limitata, 703.

Pelvetiopsis, 702.
limitata, 703.
f. lata, 703.

f. typiea, 425, 703; figure show-
ing, opp. 764.

Peridiniales, 388.

Petalonia, 535.

Fascia, 536.
Petrospongium, 508.

Berkeleyi, 509, 510.

rugosum, 509; figures showing, opp.
750.
Phaeophyeeae, 388.
Phaeosaccion, 388, 513, 514.
Phaeosporales, 389.
Phaeosporeae, 389, 390.
Phaeostrophion, 585.

irregulare, 586; figures showing.
■ opp. 748, 772, 842.

Phaeozoosporeae, 389.
Phaeurus, 558, 559.
Phloeospora, 529.
tortilis, 529.

Phycocelis, 455, 460.
fecunda, 451, 461.
maculans, 488.
reptans, 461.

Phycodendron, 594.
Phycolapathum, 515, 517.
Phycophyta, 388.
Phycopteris cuneata, 661.
Phyllitis, 535.

Fascia, 536, 538.
PhvUospadix, 428, 465, 466, 511, 518.

" 536.
Phyllospora Menziesii, 648.
Pleiadophora, 655.
Pleurophycus, 593, 606, 607, 608.
Gardneri, 406, 607; figure showing,
opp. 832.
Pogotrichum, 514.
Porphyra naiadum, 445.
Postelsia, 623, 624.
californica, 625, 626.
palmaeformis, 406, 625, 626, 633,
636; plate showing, opp. 810.
Pterygophora, 631, 632, 634.

californica, 634, 635; plate show-
ing, opp. 820.
Punctaria, 512, 514, 515, 517, 519.
524, 650.
chartacea, 521; figures showing,

opp. 762, 834.
expansa, 521, 522; figure showing,

opp. 770.
fissilis, 518.
glacialis, 518, 520.
hospej'ia, 517; figures showing, opp.

746, 770.
latifolia, 514, 515, 517, 519, 520.

521.
lobata, 519.
occidentalis, 520; figures showing,

opp. 742, 832.
plantaginea, 515, 516, 518.
tenuissima, 515, 516.
undulata, 516.
Winstonii, 524.

Punctariaceae, 513, 514, 530.
Punctarieae, 513.
Pylaiella, 401, 408, 413, 418, 483.
curta, 402.
fulvescens, 407.
Gardneri, 405.

littoralis, 402, 404, 407, 408, 429;
figure showing, opp. 746.
var. divaricata, 404.
var. firma, 404.
var. opposita, 404.
var. varia, 404.
nana, 406.
Postelsiae, 406.
tenella, 406, 407.
unilateralis, 402, 404.

Postelsia, 406.

palmaeformis, 406.

[896]

Index

Ealfsia, 45-1, 455, 493, 494, 496.

Boriietii, 496.

calif ornica, 497; figure showing,
opp. 744.

clavata, 495, 496, 497.

deusta, 499, 500.

extensa, 500.

fuiigiformis, 494, 499.

hesperia, 495, 498.

verrucosa, 496, 497, 498.
Ealfsiaceae, 454, 493.
Eenfrewia parvula, 462, 603.
Ehadinocladia, 515, 517.
Ehodoniela, 526.
Eliodomelaccae, 403.
Ehodopliyceae, 445, 650.
Ehodynienia, 505.

palmata, 488.

Eivularia Zosterae, 546.
Eupreclitiella, 550.
filiformis, 576.

Saccharina, 594.
Saccorhiza, 591.
Sargassaceae, 663, 704.
Sargassum, 662, 711.
aciiiacifolium, 713.
Agardhiauum, 716, 717; figures
showing, opp. 750, 754; plate
showing, opp. 856.
Brandcgeei, 718.

Bryantii, 713, 714.
carpopliyllum, 714.
cylindrocarpum, 720.
dissectifolium, 713.
fuliginosum

var.(?) californica, 721.
galapagcnse, 719.
guardiense, 713.
lierporhizum, 720.
horridum, 506, 715.
insulare, 506, 717.

Johnstonii, 719.

f. gracile, 720.

f. laxius, 719.
lapazeanum, 713, 714.

Liebmanni, 715, 716; figure show-
ing, opp. 762.

Marchantae, 715,

onustum, 706.

Palmeri, 712; figure showing, opp.

754; plate showing, opp. 860.
paniculatum, 721; figure sliowing,

opp. 764.
piluliferum, 553, 712.
podocanthuni, 718.
liolyacanthum

f. amerif-anum, 394, 718.
sinieola, 718.
spinuligerum, 718, 719.

Scytosiphon, 530, 531, 535.
bullosas, 542.
liipjiuroides, 590.

Lomentaria, 531, 532; figure show-
ing, opp. 760.
var. complanata, 532.
f. coniplanatus, 536.
major, 534.

minor, 534; figure showing,
opp. 760.
f. cylindricus

maculatus, 534; figures show-
ing, opp. 756.
major, 533.
minor, 533.
f. typieus, 532, 533; figures show-
ing, opp. 750, 760.
tortilis, 529.
Scytosiphonaceae, 514, 528, 530', 552.
Scytosiphoneae, 530, 577.
Scytothamnaceae, 570, 571.
Scytothamnus, 570, 587.
australis, 552.

Solenia (Enteromorpha), 454.
Soranthera, 525, 650.
ulvoidea, 525.
f. dififormis, 526.
f. typica, 526; figures showing,
opp. 750, 838.

Spermatochnaceae, 587.
Spermatoehnus, 555.
Sphacelaria, 392, 393.
arctica, 394.
brevicornis, 394.
californica, 395; figure showing,

opp. 746.
cirrhosa, 396.
cornuta, 394.
didichotoma, 397.
divaricata, 397.
fusca, 396.

furcigera, 396; 397; figure show-
ing, opp. 746.
plumula, 395.

var. californica, 394.
racemosa, 393.

var. arctica, 394.
subfusca, 395, 396; figure showing,

opp. 746.
vaiialiilis, 397.
Sphacelariaceae, 391, 392.

Rphacelariales, 388, 389, 390, 391.
399, 570.

Spinularia latifrons, 564.

Sporochnus medius, 565.

Stictyosiphon, 514, 528, 529.

adriaticus, 529.

tortilis, 529, 530.

[897]

Index

Stilophora, 545, 552, 650.
Stragularia, 454, 494.
Stragulariaceae, 493.
Streblonema, 408, 440, 443, 445, 452,
453, 476, 477, 481.
aeeidioides, 451.

f. pacifieum, 450, 451, 475.
aiiomalum, 441, 442, 443, 444;

figures showing, opp. 776.
corymbifcrum, 441, 442, 443, 444;

figure showing, opp. 776.
evagatum, 449.
faseieulatum, 442, 443.
investiens, 452.
irregulare, 447.
Johnstonae, 441, 442, 444; figure

showing, opp. 776.
luteolum, 425.
minutissimum, 442, 447.
myrionematoides, 446, 450, 452.
pacifica, 448.

pacifieum, 448, 449; figure show-
ing, opp. 754.
penetrale, 446, 450.
Porphyrae, 445.
rugosum, 449; figures showing, opp.

776.
scabiosujn, 450.
transfixum, 446.
volubilis, 443.
vorax, 444, 445.

Strepsithalia, 453, 455, 507, 508.
investiens, 453.

Striaria, 528, 529, 530.
attenuata, 530.

Striariaceae, 514, 528.

Strigoearpus, 653.

Stypopodium, 659.

Atomaria, 657.

lobatum, 660.

Symphoricoccus, 502.

Taenia, 656, 657.
Atomaria, 658.
australasica, 658.

Lennebackerae, 420, 657; figures
showing, opp. 742, 864.

Thalassiophyllum, 593, 613, 620.
Clathrus, 613; plate showing, opp.
816.

The Marine Algae of the Pacific
Coast of North America, III.
Melanophyceae, 383.

Tilopteridaceae, 649.

Tilopteridales, 649.

Tremella difformis, 512.

Trinitaria confervoides, 565.

Tubicutis, 531.

Ulva, 472.

Atomaria, 657.

plantaginifolia, 515.

Priapus, 624.

serrata, 657.
Villania, 659.
Virginia, 624.

Palma-Maris, 626.
Virides, 561.
Zonaria, 656, 657, 659.

Atomaria, 657.

deusta, 499.

Diesingiana, 661.

Farlowii, 660; figures showing,
opp. 740', 744, 758, 866.

flava, 660.

interrupta, 661.

Sinclairii, 660.

Tournefortii, 660.

Turneriana, 660.

zonalis, 660.

Zostera, 418, 420, 461, 518, 520.
marina, 519.

Page 515, line 31.

Page 518, line 33.

Page 565, line 7.

Page 576, line 1.

Page 610, line 33.

Page 638, line 22.

Page 646, line 36.

EEKATA

For tenuissimum read tenuissima.

For glaciale read glaeialis.

For aculeae read aculeata.

For Eupreclatiolla read Euprechtiella.

For Dictyoneuron read Dictyoneurum.

For Aguarum read Agarum.

For Andersonia read Andersonii.

[898]

UNIV, CALIF. PUBL BOT VOL. 8

SETCHELL-GARDNER I PLATE 56

PLATE 57

Laminaria complanata (S. and G.) Setehell
From a i^liotograph of a typical plant showing flattened crooked stipe.

[786]

UNIV. CALIF. PUBL, BOT. VOL 8

[SETCHELL-GARDNER I PLATE 57

PLATE 58

Laminaria ephemera Setchell

From a photograph of a series of plants to show variation in the splitting
of the blade, difference in its width, and character of its base.

[788]

UNIV CALIF PUBL. BOT VOL 8

I SETCHELL- GARDNER I PLATE 58

PLATE 59

A. Laminaria cuneifolia J. Ag.

Prom a photograph of a plant showing a short fiat stipe, cordate base, and a
split blade free from bullae.

B. Laminaria cuneifolia J. Ag.

From a photograph of a plant with two rows of bullae witliin the margin,
leaving a smooth central strip and showing entire blade.

[790]

UNIV. CALIF. PUBL. BOT. VOL. 8

;SETCHELL-GARDNER I PLATE 59

B

PLATE 60

Lm'iiinaria cuneifoUa J. Ag.
From a photograph of a plant showing a short blade bullate all over.

[792]

UNIV. CALIF. PUBL. BOX. VOL. 8

[SETCHELL-GARDNER I PLATE 60

PLATE 61

Laminaria personata S. and G.
From a photograph of a tyxjical phuit with three small specimens at the base.

[794]

UNIV. CALIF. PUBL. BOT. VOL. 8

SETCHELL-GARDNER 1 PLATE 61

PLATE 62
Macrocystis integrifolia Bory.

From a photograph of a plant showing the character of the hapteres and
flattened ' ' rhizome. ' '

[796]

UNIV. CALIF. PUBL. BOT. VOL. 8

SETCHELL -GARDNER I PLATE 62

PLATE 63
Aganim cribrosum Bory.

From a photograph of a short, wide specimen showing the distribution and
irregular form of the perforations and the very ample scrolls at the base.

[798]

UNIV. CALIF. FUBL. BOX. VOL. 8

[SETCHELL-GARDNER I PLATE 63

'^^^^

PLATE 6i \

Macrocystis pyrifera (Turner) Ag.

rrom a photograph of a series of young plants showing early stages in
development.

[800]

UNIV. CALIF. PUBL. BOX. VOL. 8

ISEI CHELL-GARDNER I PLATE 64

PLATE 65
Macrocystis pyrifera (Turner) Ag.

Eepresentiug an early 'stage in the development in advance of those repre-
sented on plate 64.

[802]

UNIV. CALIF. FUBL. BOT. VOL. 3

1 SETCHELL -GARDNER I PLATE 65

PLATE 66

Alaria marginata Post and Eupr.

From a photograph of a plant niedium in all of its characters, with most of
the sporophylls removed.

[804]

UNIV CALIF FUBL. BOT. VOL 8

[SETCHELL-GARDNER 1 PLATE 66

PLATE 67

Lessoniopsis littoralis (Farlow and Seteliell) Eeiuke

From a photograph of a young phuit showing the method of branching by the
splitting of tlie blade from the base upwards.

[806]

UNIV. CALIF. FUBL. BOT. VOL. 8

I SETCHELL-GARDNER I PLATE 67

PLATE 68

Lessoniopsis liiioralis (Farlow and Setchell) Eeinke

From a photograx^li of a fragment of a plant showing the method of branch-
ing and the origin and character of tlie sporophylls.

[808]

UNIV. CALIF. PUBL. BOX. VOL. 8

I SETCHELL-GARDNER I PLATE 68

PLATE 69

Postelsia palmaeformis Eux^r.

From a pliotogiapli of a group of typical plants with many of the sporophylls
removed.

[810]

UNIV. CALIF. PUBL. BOT. VOL. 8

I SETCHELL-GARDNER I PLATE 69

PLATE 7C

Biciijoneurum californicum Rupr.

From a photograiih of a group of young plants showing the method of origiiL
of new blades by splitting in the transition, meristematic region.

SlL'

UNIV, CALIF. FUBL. BOT, VOL, 8

ISETCHELL GARDNERl PLATE 70

PLATE 71

Ac/arum jimlriaium Harvey

From a photograph of half of a plant and of the basal part of another, the
latter showing well the character of the stipe.

[814]

UNIV. CALIF. FUBL. BOT. VOL. 8

tSETCHELL-GARDNER I PLATE 71

A <• ' 1

PLATE 72

Thallassiophylhim CJatlirus (Gmel.) Post, and Rupr.
From a photograph of a typical plant.

[816]

UNIV CALIF PUBL. BOT VOL 8

SETCHELL-GARDNER I PLATE 72

PLATE 73

Hedopliylluin sessile (Ag.) Setehell
From a photograph of a typical plant.

[818]

UNIV. CALIF, FUBL BOT. VOL. 8

I SETCHELL-GARDNER I PLATE 73

PLATE 74

Pierygophora californica Enpr.
From a xJliotograph of a typical young jjlant.

[820]

I

UNIV. CALIF. FUBL. BOT. VOL. 8

[SETCHELL-GARDNER I PLATE 74

'^w^^!^^.

PLATE 75

Chordaria dissessa S. and G.
From a photograph of a fragment of a x^lant.

[822]

UNIV CALIF FUBL BOT VOL. 8

[SETCHELL-GARDNER I PLATE 75

PLATE 76

Myriogloia Andersonii (Farlow) Kuckuck

From a photograph of a typical young plant, showing the myriads of
assimilating hairs.

[824]

UNIV. CALIF PUBL. BOX. VOL. 8

f SETCHELL-GARDNER I PLATE 76

I

p

*»-^-.

PLATE 77

Hapteropliycus canaliculatus Setehell
From a photograph of a group of typical plants, taken from aljove and from below.

[826]

UNIV. CALIF. PUBL. BOT. VOL. 8

ISETCHELL-GARDNER I PLATE 77

PLATE 78

A. Pelvetia fastigiata f . gracilis S. and G.
From a photograph of a typical plant Avith most of the branches removed.

B. Gobia simplex (Saunders) S. and G.
Prom a photograph of a group of typical plants removed from their host.

[828]

UNIV. CALIF. FUBL. BOX. VOL. 8

[SETCHELL-GARDNER I PLATE 78

PLATE 79

A. Costaria costaia (Turn.) Saunders

From a photograph of a form with very wide, short blade .and long, wide
stipe.

B. Coilodesme rigida S. and G.
From a photograph of a group of young plants in position on the host.

[830]

I

UNIV CALIF. FUBL BOX. VOL. 8

[SETCHELL-GARDNER I PLATE 79

PLATE 80

A. Pleurophycus Gardueri Setehell and Saunders
From a photograph of a typical plant.

B. Functaria occidentalis S. and G.
From a photograph of a typical plant.

[832]

UNIV CALIF. PUBL. BOX, VOL. 8

fSETCHELL-GARDNER I PLATE 80

"— M>aflf

PLATE 81

Punctaria chartacea S. and G.
From a photograph of three plants representing variations in size.

834 J

UNIV. CALIF, PUBL. BOX. VOL. 8

[SETCHELL-GARDNER I PLATE 81

PLATE 82

Coilodesme polygnampia S. and G.

From a photograph of a group of typical plants.

[836;

UNIV. CALIF. FUBL. BOT. VOL. 8

SETCHELL-GARDNER I FLATE 82

PLATE 83

A. Endaracline Binghamiae J. Ag.
From a protograph of a series of plants representing shapes and. sizes.

B. Soranthera ulvoidea f. typica S. and G.
From a photograph of a group of dried plants.

[838]

UNIV. CALIF, PUBL, BOT. VOL. 8

[SETCHELL GARDNER 1 PLATE 83

A

PLATE 84

Cliordaria gracilis S. and G-.

From a x^hotograiili of a series of typical i^laiits.

[840]

UNIV. CALIF. PUBL. BOT. VOL.

ISETCHELL GARDNER! PLATE 84

PLATE 85

Phaeostrophion irregulare S. and G.
From a photograph of a series of old plants.

[842]

UNIV, CALIF. PUBL. BOT. VOL, 8

I SETCHELL- GARDNER I PLATE 85

PLATE 86

Coilodesme californica (Eupr.) Kjellm.

From a photograph of a group of young plants in position on the host.

[844]

UNIV. CALIF. FUBL. BOT. VOL. 8

I SETCHELL-GARDNER I PLATE 86

PLATE 87

Besmarestia ligulata (Ligiitf.) Lamour.
From a photograph of a typical phint of moderate size.

[846]

UNIV CALIF PUBL BOX VOL. 8

[SETCHELL-GARDNER I PLATE 87

PLATE 88

Desmnrestia heriacea (Turner) Lamour.

From a photograph of a typical young plant with hairs still in position.

[848]

UNIV CALIF, FUBL, BOX. VOL. 8

I SETCHELL-GARDNER I PLATE 88

PLATE 89

Desmarestia munda S. and G.

From a lahotograph of a part of a typical plant showing tlie characteristic
toothed margins.

[850]

UNIV. CALIF. FUBL. BOT. VOL. 8

SETCHELL -GARDNER I PLATE 89

PLATE 90

Desmarestia latifrons (Rupr.) Kuetz.

From a photograph of the basal part of an old frond and the upper part of
a younger frond.

I8.-.2

UNIV CALIF FUBL, BOT VOL. 8

fSETCHELL-GARDNER I PLATE 90

PLATE 91

Heterochordaria abietina (Eupr.) S. and G.
From a photograph of a group of typical mature phmts.

[854]

UNIV. CALIF. FUBL. BOT. VOL. 8

[SETCHELL-GARDNER I PLATE 91

PLATE 92

Sargassum Agardhianum Farlcw
From a pliotograph of a typical plant.

[S.16]

UNIV. CALIF, PUBL. BOX, VOL. 8

[SETCHELL-GARDNER I PLATE 92

PLATE 93

Padina Durvillaei Bory
From a photograph of a typical plant.

[858]

UNIV CALIF, PUBL BOT. VOL 8

SETCHELL-GARDNER I PLATE 93

PLATE 94

Sargassum Palmeri Gruiiow

From a photograph of a form with very finely dissected "leaves."

[830]

UNIV. CALIF FUBL. EOT. VOL. 8

[SETCHELL-GARDNER I PLATE 94

PLATE 95

Neurocarpus sonarioides (Farlow) Howe
From a photograph of a typical phmt.

[862]

UNIV CALIF. PUBL. BOT. VOL, 8

ISETCHELL- GARDNER I PLATE 95

PLATE 96

Taonia Lennebaclerae Farlow
From n photograpli of a typical plant.

[834J

UNIV. CALIF, FUBL, BOT, VOL. 8

SETCHELL G.ARDNER I PLATE 96

PLATE 97

Zonaria Farlowii S. and G.
From a pliotogrr.ph of a typical ]ilant.

[866J

UNIV. CALIF. PUBL. BOX. VOL. 8

! SETCHELL-GARDNER I PLATE 97

yiP^

PLATE 98

Fucus furcatus f. allreviatus Gardner

From a photograph of fresh material of the type specimen, showing rela-
tively large, Avide, and scarcely matured receptacles. Three-fourths natural size.

[868]

UNIV. CALIF, FUEL. EOT VOL S

I SETCHELL-GARDNER 1 PLATE 98

PLATE 99

Fucus f'urcatus f. luxurians Gardner

From a photogi'aph of the tyj)e specimen, a young plant fruiting for the
first time, showing many, sterile tei-minal segments. One-third natural size.

[870]

UNIV. CALIF. FUBL. BOX. VOL. 8

SETCHELL-GARDNER I PLATE 99

PLATE 100

F'ucus fnrcatiis f. rigidus Gaidner
From a photograph of the type specimen. One-third natural size.

[872]

UNIV, CALIF. PUBL. BOT. VO!_. 8

[SETCHELL-GARDNER 1 PLATE 1 OO

PLATE 101

Fucus furcatus t. latifrons Gardner

From the photograph of the type specimen, a young plant just beginning to
fruit. Two-fifths natural size.

[874J

UNIV. CALIF. PUBL. BOX. VOL. 8

SETCHELL-GARDNER I PLATE 101

PLATE 102
Fitcits nitens Gardner

From a photograph of the type specimen showing mature receptacles, mostly
swollen. Two-fifths natural size.

[87(5 1

UNIV. CALIF. FUBL. BOT. VOL. 8

I SETCHELL-GARDNER I PLATE 102

PLATE 103

Fucns edentaius f.. liesperius Gardner
From a photograph of a portion of a plant, the type. One-third natural size.

[878]

UNIV CALIF. FUBL. BOT. VOL. 8

I SETCHELL-GARDNER 1 PLATE 103

PLATE 104

Fucus edentatus f. divaricates Gardner

From a photograph of a fruiting plant. Three-fifths natural size.

[880]

UNIV. CALIF. PUBL. BOX. VOL. 3

SETCHELL-GARDNER I PLATE 1 04

PLATE 105

Fucus memhranaceus f. latissimus Gardner

From a jjhotograpli of the type specimen. Oue-tliird natural size.

[882]

UNIV. CALIF. PUBL. EOT. VOL. 8

SETCHELL GARDNER] PLATE I05

PLATE 106

Fucus evanescens f. oregonensis Gardner

From a i)liotograpli of the tyj^e specimen. One-half natural size.

[884]

UNIV. CALIF. PUBL, BOT. VOL. 8

.SETCHELL-GARDNER I PLATE I06

PLATE 107

Fucus evanescens f. rolmstus S. and G.
From a photograph of a portion of a fresh plant. One-third natural size.

[886]

UNIV. CALiF. PUBL. BOX. VOL. 8

fSETCHELL GARDNERI PLATE 107

UNIVEBSITY OF OALIFOBNIA PUBLICATIONS— (OonUnued)

Vot 6. 1914^1919.

1. Parasitic Florideaa^ by William Albert SetcheU. Pp. 1-34, plates 1-6.

April, 1914 ^^^.....„„__ w~~, — .- .ae

2. Phytomorula regularis, a Symmetrical Protopbyte Belated to Coelastrum, by

Cbarles Atwood Eofoid. Pp. 85-40, plate 7. April, 1914 M

3. Variation in Oenothera ovata, by KatheriBe Layne Brandegee. Pp. 41-50,

plates 8-9. June, 1914 . „ , — .10

4. Plantae Mexicanae Purpusianae. VI, by Townsbend Stitb Brandegee^ Pp.

51-77. Jnly, 1914 ^^^^^.^..^ J6

5. Tbe Seinaia Assemblage, by William Albert Setchell. Pp. 79-152, plates

10-16. October, 1914 . _. .76

6. Notes on Pacific Coast Algae. I, Tylaiella Postelsiae, n. sp., a New Type in

the Genus Pylaiella, by Carl Skottsberg. Pp. 153-164, plates 17-19. May,

7. New and Noteworthy Callfomian Plants. II, by Harvey Monroe Hall. Pp.

165-176, plate 20. October, 1915 _. .15

8. Plantae Mexlcanae Purpusianae. VII, by Townshend Stlth Brandegee. Pp.

177-197. October, 1915 . :.... :..... , .25

9. Floral Belations among the Galapagos Islands, by A. L. Kroeber. Pp.

199^220. March, 1916 „ _ .20

10. The Comparative Histology of Certain Califomian Boletaceae, by Harry S.

Yates. Pp. 221-274, plates 21-25. February, 1916 „ : J50

11. A Bevision of the Tuberales of California, by Helen Margaret GKlkey. Pp.

275-356, plates 26-30. March, 1916 __.___._ .80

12. Species Novae vel Minus Cognltae, by T. S. Brandegee. Pp. 357-361. May,

15. Plantae Mexlcanae Purpusianae. vm, by Townshend Stith Brandegee.

Pp. 263-375. March, 1917 ... .15

14. New Pacific Coast Marine Algae. I, by Nathaniel Lyon Gardner. Pp. 377*

416, plates 31-35. June, 1917 .. . ,.. ,..-j^ ..„..._„- ^ .40

16. An Account of the Mode of Foliar Abscission in Citrus, by Bobert W.

Hodgson. Pp. 417-428, 3 text figures. February, 1918 JO

16. New Pacific Coast Marine Algae. II, by Nathaniel Lyon Gardner. Pp. 429^

454, plates 36-37. July, 1918 .... .26

17. New Pacific Coast Marine Algae, m, by Nathaniel Lyon Gardner. Pp.

455-486, plates 38-41. December, 1918 „........^ .„...^.. .86

18. New Pacific Coast Marine Algae. IV, by Nathaniel Lyon Gardner. Pp.

487-496, plate 42. January, 1919 .15

19. Plantae Mexlcanae Purpusianae. IX, by Townshend Stlth Brandegee. Pp.

497-504. November, 1919 . .06

Vol 7. 1918-1922.

1. Notes on the Califomian Species of Trillium L. I, A Beport of the General

Besults of Field and Garden Studies, 1911-1916, by Thomas Harper Good>
speed and Bobert Percy Brandt. Pp. 1-24, plates 1-4. October, 1916 .86

2. Notes on the Califomian Species of Trillium L. II, The Nature and Occur-

rence of Undeveloped Flowers, by Thomas Harper Goodspeed and Bobert
Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 „. ,16

3. Notes on the Califomian Species of Trillium L. m. Seasonal Changes in

Trillium Species with Special Eeferenoe to the Reproductive Tissues, by
Bobert Percy Brandt. Pp. 39-68, plates 7-10. December, 1916 ™ .30

4. Notes on the Califomian Species of Trillium L. IV, Teratological Varia-

tions of Trillium sessile var. giganteum H. & A., by Thomas Harper Good-
speed. Pp. 69-100, plates 11-17. January, 1917 _...._ JO

6. A Preliminary List of the Uredlnales of California, by Walter 0. Blasdale.

Pp. 101-157. August, 1919 .. _. „ £0

6, 7, 8. A Bubber Plant Survey of Western North America. X Chrysothamnus
Tiauseosus and Its Varieties, by Harvey Monroe HalL n. Chrysil, a New
Bubber from Chrysothamnus nauseosus, by Harvey Monroe Hall and
Thomas Harper Goodspeed. HL The Occurrence of Rubber In Certain
West American Shrubs, by Harvey Monroe Hall and Thomas Harper
Goodspeed. Pp. 159-278, plates 18-20, 8 figures in text. November, 1919. 1.2B

UKIVSBSITY OF OAUFOBNIA PUBLIOATIONS— (ConUaned)

9. Phycologlcal Contributioas, I, by William iUbert Setchell and Natbaaiel

Lyon Gardner. I>p. 279-324, plates 21-!51. April, 1920 ...^... -.........^«-~~ JSO

10. Plantae Mexicanae Puxpusiaaae. Z^ by Townsbend Stitb Brandegee. Pp.

825-331. December, 1920 . ;„^..,...._~.^ — ™^......,....^.™,.__.— „ ^ JO

11. Fbycologlcal OontributiMis n to VL New Species of: 11. Myrionema;

m. Compsonema; IV. Hecatonema; V. Pylaiella and Streblonema; VI,
Ectocarpus. By WiUiam Albert Setchell and Natbaolel Lyon Gardner.
Pp. 333-426, plates 32-49. May> 1922 ...... —_._-..... — ^^._,„^- . ..^ IJBO

12. Notes on Pacific Coast Algae. IL On the Oalifomlan "Delessetla Querel»

f oUa^ ' ' by Carl Skottsberg. Pp. 427-436* plate 50. June, 1922 . — .™-.._^ ^5
18. TTndescribed plants mostly from Baja California, by Ivan Murray Johnston.

Pp. 437-446. August, 1922 — ...,..-v.....„............... -...—. .25

14. Morphology, Development, and Economic Aspects ot Schizophyllum com-
mune Fries, by Frederick Monroe Bsidg. Pp. 447-498, plates 51-61.
August, 1922 ...J .80

V©L 8. 1919-.

The Marine Algae of the Pacifiic Coast of North America, by WiUiam
Albert Setchell and Nathaniel Lyon Gardner.

Part I. Myxophyceae. Pp. 1-138, plates 1-8. November, 1909 $L5d

Part n. Chlorophyceae. Pp. 139-374, plates 9-33. July, 1920... 2.75

Index to Parts I and II. Pp. 375-382.
Part III. Melanophyceae. Pp. 383-898, plates 34-107. June, 1925... 5.00

ToL 0. A Beport upon the Boreal Flora of the Sierra Nevada of California, by Frank

Jason Smiley. Pp. 1-423, platee 1-7. October, 1921 ..... ..... ^.. 5.00

Vol 10. 1922-1924.

1. The Genus Fucvs on the Pacific Coast of North America, by Nathanid Lyon

Gardner. Pp. 1-180, plates 1-60. April, 1922.............. ..„.. 2.25

2. Plantae Mexicanae Purpusianae, 33, by Townshend Stith Brandegee. Pp.

181-188, 1 figure in text. November, 1922............ ....... :............ ......:............. .15

3. A Revision of the Calif omian Species of Lotius, by Alice H. Ottley. Pp.

189-305, platea 61-82, maps 1-10. September, 1923................... ;.........„„:. 2.00

4. Notes on a CoUectioa of New Zealand Hepaticae, by William Henry Pear-

son. Pp. 307-370, plates 83-103.
6. More New Zealand Hepaticae, by William Henry Pearson. Pp. 3737392,
plates 104-109.

Nos. 4 and 5 in one cover. June, 1923 1.25

6. Parasitic Florideae, H, by Williak Albert SetchelL Pp. 393-396.

7. A Revision of the West North American Species of Callophyllis, by

William Albert SetcheU. Pp. 397-4Q1.

Nos. 6 and 7 in one cover. May, 1923 „.:........... .26

8. Plantae Mexicanae Purpusianae, XH, by Townshend Stith Brandegee. Pp.

403-421. October, 1924 ..i :....... 1 ..:....:.....:_.. ..........L...... .25

9. New Species of Plants from Indo-China, by Elmer D. Merrill. Pp. 423- V

430. October, 1924 ...:.....„.. ;..._....... ...........L... _... .28

VoL 11. 1922-.

1. Interspecific Hybridization in NiootiaTw,. I. On the Eesuits of Backcrossing

the F, Sylvestris-Tdbacum Hybrids to Sylvestris, by Thomas Harper Good-
speed and Boy Elwood Clausen. Pp. 1-30. August, 1922 _. _ .tf

Vol 12. 1924-^

L Lichenes a W. A. Setchell et H. £. Parks in Insula Tahiti a 1922 Collectl,

scripsit Edv. A. Vainio. Pp. 1-16. January, 1924 ..„ .36

2. Beport upon a CoUeotion of Ferns from Tahiti, by WiUiam B. Mazwo.

Pp. 17-44, plates 1-6. April, 1924 _...._ ..._. ........ . .45

3. Tahitian Mosses, CoUected by W. A. Setchell and H. B. Parks; Determined

by V. F. Brotherus. Pp. 45-48. September, 1924 '........., .26

VoL 13. 1924-.

L Phycological Contributions, VII, by William Albert SetcheU and Nathaniel

Lyon Gardner. Pp. 1-13. October, 1924 ........_. .„.._ .25

2. Eemizonia congesta, a Genetic, Ecologic, and Taxonomic Study of the Hay-
field Tarweeds, by Ernest Brown Babcock and Harvey Monroe HaU-
Pp. 15-100, plates 1-7, 4 figures in text. December, 1924 ....1 1.36

New York Botanical Garden Librar

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