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Committee on Publication
Barton W. Evermann Chairman and Editor
C. Hart Merriam David White
A. D. Hopkins Lyman J. Briggs
PROCEEDINGS
OF THE
Washington Academy of Sciences
Vol. XII
1910
WASHINGTON January-December, 19 io
■y -, ,^o.
4-li'ri-
AFFILIATED SOCIETIES
Anthropological Society of Washington.
Biological Society of Washington.
Botanical Society of Washington.
Chemical Society of Washington.
Columbia Historical Society.
Entomological Society of Washington.
Geological Society of Washington.
Medical Society of the District of Columbia.
National Geographic Society.
Philosophical Soclety of Washington.
Society of American Foresters.
Washington Society of the Arch^ological Institute of America.
Washington Society of Engineers.
WASHINGTON ACADEMY OF SCIENCES
OFFICERS FOR 1910 President
Charles D. Walcott
Vice-Presidents
From the Anthropological Society Walter Hough
Archcpological Society Mitchell Carroll
Biological Society Theodore S. Palmer
Botanical Society David White
Chemical Society H. W. Wiley
Engineers Society Bernard R. Green
Entomological Society A. D. Hopkins
Foresters Society Gifford Pinchot
Geographic Society Henry Gannett
Geological Society F. L. Ransome
Historical Society Jas. Dudley Morgan
Medical Society Louis Mackall
Philosophical Society Robert S. Woodward
Corresponding Secretary Treasurer
Frank Baker Arthur L. Day
Recording Secretary
Bailey Willis
Managers Class of 1911
Barton W. Evermann L. O. Howard O. H. Tittmann
Class of 191 S
Frederick V. Colville Geo. M. Kober E. W. Parker
Class of 1912
L. A. Bauer C. F. Marvin C. Hart Merriam
vu
STANDING COMMITTEES FOR 1910
Meetings David T, Day, Chairman
J, S. DiLLER
L. 0. Howard E. B. Rosa E. A. Ballock W. J. Humphreys
Finance
E. W. Parker, Chairman Arthur L. Day
A. H. Brooks Thomas H. Kearney George R. Putnam
Rules
James Dudley Morgan,
Chairman Walter Hough O. H. Tittmann
Policy A. D. HovKiNS, Chairman David T. Day David White A. H. Brooks Walter Hough J. N. Rose
F. W. Clarke Bailey Willis
Publication
Barton W. Evermann, Chairman Lyman J.Briggs A. D. Hopkins C. Hart Merriam David White
Building
Geo. M. Kober, Chairman J. Howard Gore J. N. Rose Willis L. Moore
J. S. DiLLER
Affiliation
Bailey Willis, Chairman
E. W. Parker G. K. Burgess
Membership Henry Gannett, Chairman
F. W. Clarke L. O. Howard Geo. M. Kober C. W. Hayes
Vlll
CONTENTS
PAGB
On the manner of locomotion of the Dinosaurs, especially Diplodocus, with remarks on the origin of the birds . i
The Lichen flora of the Santa Cruz Peninsula, California 27
The Polytrichaceae of Western North America . . .271
Index 329
IX
ILLUSTRATIONS
Plates
I. The form and attitudes of Diplodocus 26
Text-Figures
1. Section through foot of Tcstudo 2
2. Left acetabuhim 8
3. Acetabulum of Hzard Metapoceros g
4. Femur of Diplodocus 10
5. Proximal end of femur of Diplodocus 10-12
1. Catharinea crispa 277
2. Catharinea angustata 278
3. Catharinea undulata 279
4. Catharinea selwyni 280
5. Oligotrichum parallelum 283
6. Oligotrichum aligerum 284
7. Oligotrichum incurvum 285
8. 0. incurvum and O. i. latifolium 287
9. Psilopilum glabratum 289
10. Bartramiopsis lescurii 291
11. Polytrichadelphus lyallii 293
12. Pogonatum contortum 296
13. Pogonatum capillare 298
14. Pogonatum urnigerum 300
15. Pogonatum alpinum 302
16. Pognatum alpinum, variety 305
17. Pogonatum alpinum, varieties 306
18. Polytrichum gracile 310
19. Polytrichum attenuatum 311
20. Polytrichum ohioense 313
21. Polytrichum inconstans 314
22. Polytrichum commune 316
23. Polytrichum commune and varieties 318
24. Polytrichum jinsenii 319
25. Polytrichum yukonense 320
26. Polytrichum sexangulare 321
27. Polytrichum juniperinum 323
28. Polytrichum strictum 324
29. Polytrichum hypcrboreum 326
30. Polytrichum piliferum 327
xi
PROCEEDINGS
OF THE
WASHINGTON ACADEMY OF SCIENCES
Vol. XII, No. i, pp. 1-25. Pl. i, Figs. 1-7 February 15, 1910.
ON THE MANNER OF LOCOMOTION OF THE DINO- SAURS ESPECIALLY DIPLODOCUS, WITH REMARKS ON THE ORIGIN OF THE BIRDS.
By Oliver P. Hay.
In a paper published some months ago (Amer. Naturalist, vol. xliii, 1908, pp. 672-681) the writer advanced the proposition that thesauro- podous dinosaurs, especially Diplodocus, did not walk, as the elephants do, with the body high up from the ground and with the legs straight or nearly so, and moving in approximately perpendicular planes, but rather as do the crocodiles, with the body low down, and with the thighs standing well out from the animal's sides. While I was further considering the subject I received from my friend Dr. O. Abel, of Vienna, a paper' in which, while endorsing my views regarding the nature of the food of Diplodocus and the manner of taking it, he en- deavors to show that I am in error as to the bodily pose and the manner of locomotion of the sauropods. Dr. Abel maintains that the accepted views of the way in which these animals walked is the correct one and he finds support for this view in the structure of the feet. He accepts Hatcher's opinion that Diplodocus and Brontosaurus were digitigrade and argues that therefore they walked as represented in Hatcher's restoration of the reptile. The evidences that they were digitigrade are found in the belief, probably correct, that the upper ends of the metatarsals and metacarpals were not arranged in a straight line, but
• Verhandl-zool.-botan. Gessellsch. Wien, 1909, pp. 1 17-123. Proc. Wash. Acad. Sci. , February , 19 10.
2 OLIVER P. HAY
in an arc of a circle; further, that the feet were entaxonic, that is, had the inner digits more strongly developed than the outer ones.
Now, it is the writer's opinion that these evidences of digitigrady will hardly stand a test. The hinder feet of the bear are certainly plantigrade and yet the metatarsals are arranged very distinctly in an arc of a circle. On the other hand, the tiger and the hyaena are digitigrade, but their metatarsals are almost in a plane. Various animals will, I think, be found to transgress Dr. Abel's rule, as one may see by looking through a collection of skeletons. Furthermore, if it is desired to see an entaxonic foot in which the metatarsals are arranged in an arc of a circle and which is nevertheless plantigrade one has only to examine the foot of the human skeleton.
FIG. I SECTION THROUGH HIND FOOT OF TESTUDO. XL 05^, ASTRAGALUS; W^f. 2, METATARSUS OP SECOND DIGIT; ph. T, ph. 2, FIRST AND SECOND PHALANGES; t., TARSAL OF SECOND ROW; tib., TIBIA.
The writer is not disposed to deny that Diplodocus and its relatives were more or less digitigrade; but this digitigrady, through perhaps equal to that of the hinder foot of the elephant, does not prove that these reptiles walked like the elephant. The land tortoises of the genus Testudo have the feet constructed much like those of the elephant, being provided with a thick pad of skin, muscles, tendons, and con- nective tissue under the astragalus and the metatarsals and applying only the ungual phalanges to the ground. Nevertheless the legs of these reptiles stand out from the sides of the body as I have supposed that those of Diplodocus did. A figure (Fig. i) is here presented showing a section made through the hind foot of T. tahulata. Unfor- tunately I have not been able to find or make a similar section through
THE POSE AND LOCOMOTION OF DIPLODOCUS 3
the hind foot of the elephant; but, to judge from various mounted skeletons and from good figures of others, one can hardly suppose that the heel of the elephant is lifted farther from the ground relatively than that of the tortoise.
I grant that Dr. Abel's efforts are along a line where they are needed. Those who believe in the mammal-like gait of Diplodocus ought to give their reasons therefor. I do not assert that reasonable argu- ments for their view cannot be produced, but hitherto the correctness of this view has been assumed. The subject is a difficult one and needs to be studied from various points of view and by all who have the opportunity. And in studying the movements of animals one soon learns that they can assume so many positions that one may be at loss, in the case of an extinct creature, to determine which positions were the usual ones.
In the primitive condition the limbs of the Tetrapoda stand out at right angles with the body,- and in approximately this position they are found in most Amphibia and Reptilia. When these animals are walking, the humerus and the femur move backward and forward mostly in horizontal planes. In most mammals, on the contrary, the humerus is turned backward against the thorax and the femur forward against the flank. The hand, which otherwise would be directed backward, is turned forward by the crossing of the bones of the lower arm. The movements of arm and leg are then mostly in sagittal planes. In the duckbill and the echidnas the limbs have retained the position found in most reptiles.
Now, among all the reptiles that live today there are none, except perhaps the chameleons, that have attained even an approach to the condition found among the mammals.
It is evident that before the close of the Jurassic there existed both carnivorous and herbivorous dinosaurs that went about habitually on only their hinder legs; but it is by no means necessary to believe that the immediate ancestors of these bipeds walked first like mammals and afterwards like birds. It is well known that certain lizards can run swiftly on their hind legs, the fore legs and the tail being held free from the ground. Furthermore, as may be seen from W. Saville-
' Huxley, Anat. Vert. Animals, 1872, p. T,y, Flower's Osteology of the Mammalia, 1885, p. 362.
4 OLIVER P. HAY
Kent's figures' the hinder limbs are not carried backward and forward in sagittal planes like those of mammals.
It seems not diflEicult to understand the history of the attainment of the bipedal habit among lizards and dinosaurs. When the fore- legs of a quadrupedal reptile are of nearly the same length and have the same structure as the hind legs there seems to be no good reason why the animal cannot run as fast on four legs as on two. However, the hinder limbs, being nearer the center of gravity of the animal, receiving more of the weight, and being more devoted to propulsion of the body, are likely to become larger and more powerful, while the fore legs may become more or less reduced, with or without special modification for other purposes. If now a reptile whose fore legs have become relatively much shorter than the hinder ones has occas- sion to run with the greatest possible speed, it is likely to find that the fore legs cannot take as long steps as the hinder ones; and naturally it endeavors to get them out of the way by lifting them up in the air.
This practice would be of great advantage and would tend to become fixed. The reduced limbs might then become modified for other purposes or undergo further reduction. In the beginning, the femora would stand out from the body, giving the animal a wide tread. In time, however, the knees might be drawn closer to the flanks, the tread would become narrower and the pace more rapid. At no stage, however, would the reptile walk like a quadrupedal mammal; and no argument in favor of such a gait or Diplodocus can b : deduced from bipedalism in lizards.
If the mammal-like gait of Diplodocus be insisted upon on the ground of straightness of the femur it may be pointed out, as I did in the article in the American Naturalist, that the femora of sphenodon and of lizards, animals that creep, are straight. If it be contended that it is in the heavy-bodied animals that a straight femur is corre- lated with a lifting of the body from he ground during locomotion, it may be permitted to recall that the femora of Allosaiirus and Tyran- nosaurus, great carnivorous dinosaurs, are distinctly bent. The femora of Trachodon are straight, while those of Campiosauriis and Laosaurus are curved. Curvature of the femur seems, therefore, to have no relation to size of body or erectness of pose. The femora of
» Nature, vol. 53, 1895, pp. 396-397.
THE POSE AND LOCOMOTION OF DIPLODOCUS 5
crocodiles, little and great, are curved; as were too those of their prede- cessors, Aetosaurus, of the Triassic, and of Alligalorellus, of the Juras- sic, the former with femora hardly four inches long, the latter with these bones about an inch in length.
Diplodocus has been erected on column-like legs partly because it has been supposed that the great weight of its body required this. However, the legs of animals are not straight in proportion to the the weight of their bodies. The legs of the largest camels seem not to be straighter than the legs of the llamas. Some rhinoceroses and some oxen have very heavy bodies; nevertheless, their femora lack much of being in line with their tibia and these much of being in line with the metapodials. Certainly it is not because of the immense weight of the body that the legs of a man are straight.
There must, of course, be a limit to the size of an animal that can move itself about on land, in whatever position; but it may be sug- gested that a reptile that could not walk about as crocodiles do, rest- ing at least now and then, its body on the ground, could not well have erected itself when once it had lain down. That the largest crocodiles are far from the limit of active movement on the land may be judged from the following extract taken from W. Saville-Kent.*
The celerity with which a huge 25-footer, as witnessed by the writer in the Norman River, North Queensland, will make tracks for and hurl itself into the water, if disturbed during its midday siesta by the near impact of a rifle bullet, is a revelation.
It must be further taken into consideration that the weight of a crocodile 25 feet long, with short, thick neck, large head, long body, and heavy tail, would be much greater than that of a sauropod of the same length, in which most of the length is composed of slender neck and comparatively slender tail.
It is generally conceded that such carnivorous dinosaurs as Allo- saurus, Dryptosaurus, and Tyrannosaurus, and such herbivorous forms as Trachodon and Campiosaurus walked bipedally erect. If now comparison be made of the femora of any of these with those of the sauropods great differences will be noted. The shaft of the former appears to be more elaborately modeled and to consist of finer and harder bone; all the articular surfaces are smooth and they carry the
* Living Animals of the World, p. 547.
6 OLIVER P. HAY
conviction that the original surfaces, barring a thin layer of cartilage, are preserved ; there is a definite head, separated from the shaft by a distinct neck and nearly filling the acetabulum; and there is a definitely formed trochanter major. In the Sauropoda, on the contrary, the shaft seems to be composed of coarser bone; the articular surfaces are rough and show that they were covered by a thick layer of cartilage; the head merges imperceptibly into the supposed great trochanter and into the shaft; and the head lacks much of filling the acetabulum. In its low stage of differentiation the femora of the sauropods resemble greatly those of the crocodiles and are hardly above those of the lizards. They furnish no warrant for the belief that their possessors walked in mammalian fashion.
The structure of the foot of Diplodocus indicates that this reptile walked in a way very different from that in which the bipedal dinosaurs walked. In the latter the foot had the third toe most strongly de- veloped (mesaxonic); in the sauropods the two inner toes were the strongest, the third somewhat weaker, while the other two were greatly reduced. This difference of structure must have had its history and its meaning. That the feet of Diplodocus were shortened and more or less digitigrade indicates that they were employed for walking, not at all for swimming. The feet of the crocodiles are to be regarded as entaxonic, the inner digits being of stouter build, although slightly shorter than the third; but here the digits are elongated and webbed to assist in swimming. When the animal is walking, the pressure comes against principally the inner side of the foot. The trionychid turtles have the three inner digits most strongly developed and clawed; the others are slender and unarmed. The clawed digits are, of course, the ones employed for excavating hiding places in the sand and mud and getting foothold in walking and running; and these turtles are, for moderate distances, rapid and powerful runners on the land and on the bottoms of streams.
It is true that the foot of man is entaxonic and is directed nearly forward, but its history is wholly different from that of the sauropod foot. It is certain that the ancestors of man were climbing animals, with hallux strongly developed and opposable to the other digits. Being later employed for locomotion on the ground, the foot under- went a transformation to its present form. The form assumed at any time by an organ must depend greatly on the form previously pos-
THE POSE AND LOCOMOTION OF DIPLODOCUS 7
sessed. Doubtless the Sauropodaand the Theropoda started out with the same pedal outfit, and there seems to be no reason for supposing that the former passed through an arboreal stage and back into an ambulatory stage.
The position of the trochanter major of the sauropods is open to question and there are differences of opinion. Marsh* regards as this trochanter the outer upper angle of the femur, including a part of the rough surface forming the proximal end of the bone. Hatcher's view (Mem. Carnegie Mus., I. p. 46) appears to be the same. Osborn* has identified as the trochanter the rough surface which descends for some distance below the upper end of the femur on the fibular border. Neither of these views seems to the writer satisfactory. If the femora of the Triassic dinosaurs described by v. Huene in his monograph, Die Dinosaurier der europdischen Triasformation, be examined it will be found that the trochanter in question is placed at a considerable distance below the head of the bone, on the dorsal surface, and near the fibular border. In the more highly specialized dinosaurs of the Jurassic the trochanter is a distinct process arising from the position described and ascending nearly to the level of the head. In such dinosaurs as Trachodon and Triceratops the trochanter has reached the outer upper angle of the femur, and is well separated from the head by a distinct neck. The writer believes that in the sauropods the trochanter occupied the same primitive position that it has in the Triassic Theropoda. It is not essential that it should be represented by a process or even by any unusual roughness, as is shown by the femur of the crocodile.
This being the case, what explanation is to be made of the outer portion of the rough surface on the proximal end of the femur ? The writer believes that it forms a part of the head of the bone and entered into the acetabulum. The matter will be discussed. In order to illustrate a possible position of the femur in the acetabulum a figure is here presented (Fig. 2). This has been obtained by placing a section of the proximal end of the femur, taken from Hatcher's figure in Memoirs of the Carnegie Museum, vol. I, p. 46, in the acetabulum as shown in the same writer's figure in the second volume of the same
" Dinosaurs N. A., PI. XVI, fig. 3, t.
•Mem. Amer. Mus. Nat. Hist., i, p. 211, fig. 14.
8
OLIVER P. HAY
Memoirs, plate IV, fig. 2. The so-called head of the femur is toward the left, against the pubic process. According to this figure, there was room in the acetabulum for the femur, standing at right angles with the pelvis, so that it could rotate on its longer axis and could swing backward and forward. Such movements would be required in case the reptile walked as does the crocodile. In the execution of these movements it would probably happen, as it does in the lizards, that some part of the head would at times be outside of the acetabulum, in order to show the resemblance of this joint in the lizards to the one
FIG. 2 LEFT ACETABULUM, CONTAINING SECTION OF PROXIMAL END OF femur; THIS SECTION SHOWN BY HEAVY LINE. X tV) *^-' ILIUM; tsch.,
ischium; pub., pubis.
depicted, a drawing (Fig. 3) is shown of the acetabulum and head of the femur of Metapoceros.
However, the articulation at the hip was probably not effected in just this way. It appears that in some cases the proximal end of the femur is wider than the acetabulum. Dr. E. S. Riggs informs me that in Apalosaurus {Brontosaurus) and Brachiosaiirus the upper end of the femur is about 23 inches wide, exceeding the fore-and-aft diameter of the acetabulum by 3 or 4 inches. I do not regard this fact as wholly irreconcilable with the view illustrated by figure 2, the head of the femur having sometimes a greater diameter than the acetabulum, as in the land tortoises. Nevertheless, I will not argue the matter. A somewhat different arrangement at the articulation is more probable.
THE POSE AND LOCOMOTION OF DIPLODOCUS 9
Certain principles must be regarded as indisputable. One of these is that primitively, in the common ancestor of the dinosaurs, the crocodiles, and the lizards, probably in the early dinosaurs themselves, the whole proximal end of the femur constituted the anatomical head. Another is that before there could be any such structures and confor- mations of these as w^e find at the hip joint of Allosaurus, for instance, or of Trachodon, every possible stage from the one just described must have been passed through. Through countless generations the thigh must gradually have assumed a more and more forward position in habitual locomotion. While muscles and nerves were being trained
FIG. 3 ACETABULUM OF LIZARD METAPOCEROS, CONTAINING SECTION OF HEAD OF FEMUR. X 2. SECTION OF FEMUR SHOWN BY HEAVY LINE. ALSO
SIDE VIEW OF FEMUR X 2. Fcm., FEMUR; U., ILIUM; isch., ischium; ptib.,
PUBIS.
to this end the femur must have been developing a projecting head, that part of the proximal end on the fibular side was being excluded from the acetabulum, and the rotation of the proximal end of the femur around a perpendicular axis was being changed to rotation around a horizontal axis, which in mammals would pass through both femoral heads. Now, as regards the hinder leg and the hip joint, at what stage in the long journey indicated above, do we find Diplodocus? Obviously those who believe that this animal ought to be set up on its legs in the way seen in drawings, plaster restorations, mounts of the actual bones, and the plaster facsimiles of the skeleton that are being distributed over the world, must hold that Diplodocus had reached
lO
OLIVER P. HAY
practically the ultimate, or mammalian stage. The writer believes that it had attained only the first station in the journey.
A study of the femora of the sauropods shows that the proximal end varies somewhat in shape. Usually it is more or less truncated or it is slightly concave toward the fibular side and convex toward the tibial side. Figure 4 represents in outline a side view of the proximal half of the bone, as represented by Hatcher. As already stated, the
FIG. 4 OUTLINE OF SIDE VIEW OF PROXIMAL END OF FEMUR OF DIPLODOCUS.
proximal border is very rough, as shown by figure (Fig. 5) also taken from Hatcher. Undoubtedly this was covered by a thick layer of cartilage. Cope (Amer. Naturahst, xii, 1878, p. 84) says that if the layer of cartilage were ossified it would be an epiphysis, like that of the mammals. Figure 6 presents the same outline as does figure 4,
FIG. 5 PROXIMAL END OF FEMUR OF DIPLODOCUS. h, THE SO-CALLED HEAD
but to it there has been added a dotted line which is intended to indi- cate the writer's view of the form of the upper end of the femur when the cap of cartilage was present. The stage of development reached by the animal was that at which a femoral head was being developed on the tibial side of the bone and the fibular border was being freed from the articular cup. Although the whole proximal end may, in some genera, have been too broad to enter the cavity
THE POSE AND LOCOMOTION OF DIPLODOCUS
II
the greater part did so enter. Doubtless, when the leg was extended forward, a considerable part of the cartilage covered surface on the fibular border was out of the cup, and when the leg was directed back- ward the rounded anterior part of the head was out. This is exactly what happens in the lizard and, for that matter, in most animals. The head of the femur of Diplodocus, compared with that of the crocodile, differed in having its long axis coincident with the plane through both condyles; while in the crocodile the head is twisted from the plane mentioned about 75°. Figure 7 represents the same humerus as figure 6, but lines are drawn across the head to show the varying relations of the bone to the acetabulum. The line aa may
FIG. 6 PROXIMAL END OF FEMUR OF DIPLODOCUS. X A- THE DOTTED LINE SHOW THE LIMITS OF THE CARTILAGE.
be regarded as a section through the acetabulum when the leg is thrown far forward; bb, when the leg is at right angles with the body; cc, when the leg is thrown well backward. Of course, as the leg is swung from front to rear, the femur will turn also on its long axis. As is well known, the acetabulum of the Sauropoda is widely open in the skeleton. I am not aware that any one has discussed the way in which in life this opening was filled. It seems improbable that it was shut simply by membrane, for this would have been too yielding to the pressure of the head of the femur, if inserted as generally sup- posed. It seems most probable that the opening was occupied by a mass of cartilage, an imossified portion of that common cartilage from which were developed the ilium, the pubis, and the ischium. This would have formed a firm concave bed on which the convex head
12
OLIVER P. HAY
of the femur could rotate. If the femur was inserted as the writer supposes it was, its pressure would have been exerted mostly against the bony side-walls of the acetabulum and but little against the tissue filling the inner opening.
In his splendid monograph on Die Dinosaurier der europdischen Triasformation Dr. v. Huene has presented numerous restorations of the Triassic carnivorous dinosaurs (Pis. IC-CX). In order to show the author's conception of their modes of progression, three species, Plateosaurus reinigeri, Thecodontosaurus antiquus and Anchi-
FIG. 7 PROXIMAL END OF RIGHT FEMUR, WITH ITS CAP OF CARTILAGE, AND HORIZONTAL SECTION THROUGH ACETABULUM. dd, SECTION OF ACETABU- LUM; aa, LINE CORRESPONDING TO dd WHEN LEG IS THROWN FORWARD; bb, LINE CORRESPONDING TO dd WHEN LEG IS AT RIGHT ANGLES WITH BODY; CC. LINE CORRESPONDING TO dd WHEN LEG IS THROWN BACKWARD.
saurus colurus are restored each in two positions, walking on all fours and on their hinder extremities only. Dr. v. Huene has the following to say (p. 291) regarding the position of the hinder hmbs:
Das Femur passt in der Weise in den Acetabularschnitt, dass das ver- breiterte medial abstehende Proximalende nicht transversal unter dem Ileum liegt, sondern schrag nach vorn und medial gerichtet ist (daher wendet sich auch das Knie etwas auswarts).
Notwithstanding this explanation, one is struck by the very mam- mal-like position of the body and the limbs of these reptiles in the quadrupedal pose. Elbows and knees are drawn well towards the sides and the digits are directed straight forward. At least, the pose of these restorations is quite different from that of any living reptiles.
THE POSE AND LOCOMOTION OF DIPLODOCUS I3
One of these species, Anchisaurus colurus was described by Marsh from the Triassic of the Connecticut Valley, and he published a restoration of the skeleton in his work The Dinosaurs of North America, PI. IV. Dr. R. S. LulF has identified this dinosaur as the maker of the tracks known as Anchisauripus dananus (Hitch.) This identification is extremely interesting, in case it can be substantiated. The bones of the hind foot of Anchisaurus colurus fit accurately in the tracks named. These tracks are placed close to or on the line along which the animal was moving, the "line of direction" (Beckles), and there are, in the several specimens known, no indication of impres- sions of either the fore feet or the tail.
A study of the various printed restorations of this species reveals an animal of elongated body, with limbs not greatly unlike those of a crocodile, the hinder legs being a little longer in proportion to the length of the body than in the crocodile, while the fore legs are about three-fourths the length of the hinder ones. In the crocodile the fore limb is little more than two-thirds as long as the hinder.^^ As com- pared with the hind foot of the crocodile that of Anchisaurus is a little longer. Now, with this view of the creature, what is there in it to lead one to suppose that it erected itself on its hinder limbs, unless it were on rare occasions; and on such occasions would it not have borne itself as did the running lizard figured by Saville-Kent ? What one is asked to believe is that it bore itself so loftily that it is never found to have put its hands on the ground or to have dragged its tail in the mud. Furthermore, this reptile walked with all the skill and the circumspection of the heron and the barn-yard fowl; for each foot was brought forward and placed very near or on the line of direction and thus immediately under the center of gravity. This is very different from the way in which Saville-Kent's lizard ran; for when a foot was advanced it was placed far from the line of direction and at the same time the tail was jerked violently toward the same side, in order to bring the center of gravity over the advanced foot. The dinosaur in question seems to have had no other use for its tail than to serv'^e as a counterpoise to the weight of the head and trunk.
Omitting the feet, the legs of most birds consist of three long seg- ments, viz: the femur, the tibia, and the tarsometatarsus. The
' Mem. Bost. Soc, Nat. Hist., v. p. 487.
' DoUo, Bull. Mus. roy. d'Hist. nat. Belgique, ii, 1883, p. 107.
14 OLIVER P. HAY
relatively short femora diverge downward so that the knees are almost always farther apart than are the great trochanters, some- times much farther. Nevertheless the feet in walking are generally placed on the line of direction, a result brought about through the con- vergence of the elongated middle and lower segments of the two legs. If they are not brought close to this line, as in the short-legged ducks and geese, the walk becomes a waddle.
The femur of Allosaurus, of the Upper Jurassic, possesses a head that projects strongly inward; and this was provided with a well- defined smooth articular surface, which is elongated transversely to the animal and convex from front backward. The surface of the ilium against which this head fitted is also smooth. Now the confor- mation of the head of the femur and the ilium is such that the femur must have diverged considerably from its fellow, thus widely separat- ing the knees. The tibia is shorter than the femur, and the inner condyle appears to stand lower than the outer. The metatarsus is relatively short. I see no way, therefore, for the feet to be brought, except with unusual effort, near the line of direction in walking or near each other in standing. The limbs of Allosaurus may be com- pared to those of the penguins, although in Allosaurus the femora may not have been directed so strongly forward and the feet may have been more digitigrade. It would probably be very difficult for the penguin to plant its feet one in front of the other in walking. I believe therefore that Allosaurus had a wide trackway and that when it walked and ran it preserved its equilibrium by whisking its tail from side to side.
Examination of a femur, accompanied by the tibia and the fibula, in the U. S. National Museum, apparently that of Tyrannosaurus, shows the same form of the head of the femur that is found in Allo- saurus, thus making it probable that this dinosaur also had a straddling gait. Professor Osborn (Bull. Amer. Mus. Nat. Hist., xxii, p. 293) presents a figure of the femur of Tyrannosaurus. He says that the plane of the head makes an angle of 45° with the axis of the vertebral column, and that therefore the distal ends of the femora are approxi- mated. Whether the angle is in front of or behind the head of the femur is not stated. In Allosaurus the head is directed inward and forward. The effect of this would certainly be to throw the knees outward and to plant the foot farther away from the line of direction.
THE POSE AND LOCOMOTION OF DIPLODOCUS 1 5
The convergence of the femora is rare even among the mammals. If Professor Osborn is right the hind legs of Tyrannosaurus had attained the human stage in the respect mentioned.
Another potent reason for believing that the dinosaurs just named, together with Iguanodon and Trachodon, walked with a wide tread is found in the form of the body. In mammals the abdomen is usually contracted posteriorly, so that between the thighs it is shallow, per- mitting the femora to remain parallel with each other or even to con- verge. Therefore, in walking, the feet are placed near or on the line of direction. In the birds the baggy abdomen descends between the thighs and spreads these, thus requiring the convergence of the long lower segments to bring the feet together. The kangaroos have the abdomen much like that of the birds; and in them the thighs are found to diverge toward the knees, but the long tibiae permit the feet to be placed close to each other in standing and leaping. In Allosaurus and Iguanodon the belly came down nearly to the knees and passed backward between the thighs into the tremendous tail. It must be that the knees were much farther apart than the upper ends of the femora were and that the tread was wide. The writer is further of the opinion that in the bipedal dinosaurs the femora were directed more strongly forward than they are usually placed in restorations, although not so much so as in birds. This position would tend to reduce the height of the reptiles and would make the thighs more divergent.
In a paper published by Mr. William H. Ballou (Century Mag., Iv, 1897, pp. 15-23), but the facts and suggestions of which were furnished by Professor E. D, Cope, there is a figure representing two individuals of Hadrosaiirus {Trachodon) mirabilis. One of these is on the shore, resting on its hind legs and haunches, the other is standing and feeding in the water. By examining these restorations, made by Mr. Charles R. Knight, one may judge regarding the proba- bility that these reptiles could leave a straight row of tracks behind them.
Mr. S. H. Beckles* has described and figured some series of large footprints found in the Wealden near Hastings, England. These have been identified by DoUo (Bull. Mus. roy. d'Hist. nat. Belgique,
"Quart. Jour. Geol. Soc, x, 1854, 456, pi. xix.
1 6 OLIVER P. HAY
ii, 1883, p. 117, pi. iii, fig. 8) as the tracks of Igiianodon mnntelli. A study of these footprints shows that in the case of the series designated by cc the length of the step was close to 5 feet while the width of the trackway was about 2 feet 2 inches. The tips of the inner toes came, however, pretty close to the line of direction. It must be observed that in all of these tracks the toes are turned inward so much that the axis of the middle toe prolonged passes through the next imprint in front, made by the opposite foot. Now, I find no reason for supposing that in life the toes were so directed inward. None of the figures of Iguanodon so represent them; nor are the toes thus placed in any of the restorations of Tracliodon. The explanation of the matter seems to be that the reptile, if reptile it was, was lounging leisurely along, with short steps, and, to keep its equilibrium, was swinging its body around a perpendicular axis passing through the pelvis, the tail being thrown in one direction, the trunk in the opposite. In this way the feet would be planted not far from the line of direction and pointing toward it. Had the animal been running, the feet would have been planted farther from the line of direction and with toes directed forward.
Now, if these conclusions regarding the gait of the Upper Jurassic and Upper Cretaceous carnivorous dinosaurs are justified, is it prob- able that the Triassic Anchisaurus colurus, with an equally hea\y abdomen and with less elongated and more primitive limbs, had the ability to walk, just as a bird does, accurately placing one foot directly in front of the other and under the center of gravity? It seems to the writer that we need more proof of it. If it could so walk, one might inquire what was the useof all the modifications undergone by the dinosaurs up to the end of the Cretaceous. It seems most probable that Anchisaurus walked usually in crocodilian or lacertilian style, with, however, the femora drawn somewhat more closely to the sides. Now and then, when in great haste and for short distances, it was probably able to progress bipedally in an awkward fashion. In the same category may be placed some of the European dinosaurs figured by Dr. v. Huene, such as Thecodontosaurus anliquus and the species of Plateosaurus. Others, as Pachysaurus ajax and Massosaurus carinaius, probably walked more or less habitually on their hinder limbs, but with a wide trackway and with much swinging of the tail from side to side.
Dr. v. Huene's statement of his view of the manner of insertion of
THE POSE AND LOCOMOTION OF DIPLODOCUS I7
the femur has been quoted above. To the writer it seems probable that the whole proximal end of the bone constituted the head and was inserted into the acetabulum, as in lizards and crocodiles, and that the thigh was directed outward still more than Dr. v. Huene has supposed.
What then made those bird-like tracks that are so abundant in the sandstones of the Connecticut River valley ? Why not birds, indeed ? Although remains of birds have not yet been found in Triassic rocks there can be little doubt that these animals had already freed them- selves from the dinosaurs. Already long before the close of the Juras- sic the hinder limbs of birds had, as we learn from Archceopteryx taken on its present form, with doubtless ability to plant its footsteps on the line of direction. This limb was at that early time far in advance of the hind leg of the dinosaurs of even the Upper Cretaceous; and it was doubtless even in the Triassic far in advance of the limb of the dinosaurs of that time. No bird remains have been found where those famous tracks occur, it is true. It is also true that nearly loo kinds of tracks have been distinguished, while only 8 or lo species of dinosaurs have been discovered in the North American Triassic; and of these only one has had its tracks identified. Therefore, it seems to the writer entirely reasonable to suppose that those bird-like tracks, even some of them that show the presence of fore feet and tail, were really made by birds. For if the birds diverged from the dinosaurs early in the Triassic their wings were as yet probably unfitted for con- tinuous flight in the air. Many of them were probably running ani- mals and some of them may still have retained a tendency to grow to a large size. Success in flying necessitated in later times a reduc- tion in size of body. In the Trias the hands had not yet become reduced and transformed through the development of great pinion feathers, and they may have been at times applied to the ground in walking and resting. The tail was yet long, little befeathered, and might drag on the ground and leave a trail. And it must not be regarded as wholly certain that the tracks of large bipedal animals of later times were made by dinosaurs. There may have been in the Jurassic and the Cretaceous, as well as in the Tertiary, running birds of even greater size than the largest moa, whose foot was hardly inferior in size to that of many dinosaurs. On the other hand, such dinosaurs as Compsognathus and Hallopus may have walked like Proc. Wash. Acad. Sci. February, 19 lo.
1 8 OLIVER p. HAY
birds, but the remains of such are found in the Triassic no more than those of birds.
If now such Theropoda as Anchisaurus colurus, more advanced probably in every respect than the Sauropoda ever were, did not walk habitually erect, like mammals, on either two or four legs, but pro- gressed either in more or less crocodilian manner on all fours or in a straddling way on the hind legs, is it probable that the sauropods ever walked high up on four legs in the jaunty manner in which they have been represented ? It is to be considered that these great herbivorous reptiles possessed a huge abdomen, deep and probably broad, which extended backward and merged into the tail, necessitating the diver- gence of the relatively long femora. The outer surfaces of the pubic and ischiadic bones were clothed with great masses of muscles, as were too the insides of the femora. Assuming that the legs were as straight as they have been represented, the feet could have been hardly closer together than the knees, probably considerably farther apart. A bulky animal walking thus could preserve its equilibrium only by either swaying the body from side to side, to throw it over the ad- vanced foot, or throwing the tail toward that side. In the case of the fore foot the long neck might be used to preserve the balance. One might amuse and instruct himself by working out the movements of the animal according as it was walking, trotting, pacing, or per- chance galloping.
The writer is not willing to assert that Diplodocus and its relatives never straightened out their legs, thus lifting themselves well above the ground, and never walked thus. Even the crocodiles have been known to do this, as a rare occurrence.*'" In the U. S. National Museum there is a specimen of the Florida crocodile mounted in this position. The femora are directed forward and outward, the tibiae downward. The feet are widely separated as a mechanical neces- sity. What is disputed by the present writer, is that this was the cus- tomary attitude of the sauropods; and their great bulk makes it doubtful if it was ever assumed.
The writer is of the opinion that the feet of the primitive dinosaurs had the inner digits somewhat more strongly developed than the median and outer ones; that is, they were entaxonic, not mesaxonic,
10
Hornaday, Two Years in the Jungle, pp. 55, 266.
THE POSE AND LOCOMOTION OF DIPLODOCUS I9
resembling in this respect the feet of the crocodiles. A reason for this conclusion is found in the fact that all the feet of the sauropods are entaxonic and also the fore feet of the earliest known theropods. It is therefore more probable that the hinder feet of the latter reptiles became mesaxonic from an entaxonic condition than that their fore feet and both fore and hind feet of the sauropods should be transformed. That the manus of the theropods was entaxonic may be seen from Marsh's figure of the fore foot of Anchisaurus colurus and A. polyzelus (Dinosaurs N. A., pis. ii, iii) and from Dr. v. Huene's figures. Furthermore, the hinder feet of the early theropods present plain indications of a former entaxonic arrangement. The foot of Ammosaurus^^ shows a very stout first digit, not greatly shorter than the others, while the second does not fall behind the third and fourth in diameter of the bones, little in length. The superiority of the second to the third seems to have been retained in Allosaurus. When the hind leg began to be drawn forward against the side and the weight of the body was throwTi to a greater extent on the median digits a stimulus appears to have been given to the development of the third digit, while the first, relieved to some extent of its former duty, became reduced and turned backward.
In the later theropods the manus also became mesaxonic. This is seen in Marsh's restoration of the skeleton of Ceratosaurus (op. cit., pi. xiv). Mr. C. W. Gilmore, who has recently mounted this skeleton has shown me the remains of the one hand preserved. Most of the phalanges are missing. There are present four metacarpals, and there are no traces of the fifth in the rock. The first is considerably reduced, the second is the largest. Thus, there is evidence that all the feet of the carnivorous dinosaurs became transformed from the entaxonic to the mesaxonic condition. It further appears that the sauropods retained the primitive condition of the feet, fore and hinder, more persistently than did the other groups of the order.
For reptiles that progress by creeping, having the humerus and the femur at right angles with the body in the middle of the step, the entaxonic condition seems most effective. It is found in the croco- diles and the turtles, being especially well displayed in the triony- chids and the land tortoises. In reptiles the first digit is usually
" Marsh, op. cit. pi. iii, fig. 6.
20 OLIVER P. HAY
retained long after the disappearance of the fifth. In the lizards, however, the fifth is often larger than the first, a condition depend- ent perhaps on their habit of climbing about on rocks and trees. In the mammals, on the other hand, it is the first digit that earliest suffers reduction.
An attempt has already been made on a previous page to account for the origin of the bipedal habit in reptiles. Evidences are present, it is beheved, which show that bipedalism in the dinosaurs was not due to specialization of the anterior limbs. If an examination be made as to the relative lengths of the fore and the hinder limbs in the carnivorous dinosaurs, it will be found that in Anchisaurus colurus the fore limb is about three-fourths as long as the hinder; in Plateosau- rus quenstedti about two- thirds; in Pachysaurus ajax, about one-half. These are Triassic dinosaurs. In Ceratosaurus, of the Upper Jurassic, the fore hmb is only about two-fifths as long as the hinder. In Tyrannosaurus, of the Upper Cretaceous, the fore limb is diminutive, in case the humerus found with the specimen really belonged to it.^^ As we have seen, the great pollex of the late Triassic forms had become much reduced in the Upper Jurassic species. Therefore, in place of specialization, the whole limb suffered degeneration. If now it be asserted that bipedalism in the theropods was occasioned by speciali- zation of the fore limb for other purposes than locomotion, we shall have the case presented of an organ which, as soon as it was free to continue its specialization, began to degenerate. Without doubt however, the fore limb continued to be used for various purposes, just as the ostrich continues to use its diminutive wings.
Various opinions have been expressed regarding the origin of the Sauropoda. Marsh^^ expressed the opinion that the group included the most primitive forms of dinosaurs. Baur" held that the Sauro- poda had no close relationships to the other reptiles usually classed with them as dinosaurs. Osborn'^ believes that it is possible to derive the sauropod type from a primitive quadrupedal theropod type. In his work already so often quoted, Dr. v. Huene expresses
'^ Osborn. Bull. Amer. Mus. Nat. Hist., xxii, pi. xxxix.
'' Dinosaurs N. A., p. 164.
'* Amer. Naturalist, xxv, p. 450.
" Nature, vol. 73, 1906, p. 284.
THE POSE AND LOCOMOTION OF DIPLODOCUS 21
his view that the sauropods were derived from the carnivorous dino- saurs. He sums up his conclusion as follows (p. 351):
Die Sauropodcn ein friihes Theropoden-Stadium festhalten and fixiren und so eine gleichartige und relativ wenig weiterbildungsfahige Masse bilden, die sich wohl nur infolge des Riesenwuches bis zum Schluss der Kreidezeit behaupten konnte.
Regarding the time of origin of the Sauropoda Dr. v. Huene has the following to say (p. 351):
in der Zeit zwischen dem Schluss der Trias und dem Auftreten von Dystroph?cus im alteren Jura ist die erste Umpragung zum Sauropoden- Typus erfolgt.
Dr. V. Huene calls attention to the numerous characters common to the Theropoda and the Sauropoda, and he believes that the latter inherited these common characters from the former suborder. Such a derivation would, the present writer holds, require extremely impor- tant modifications in the structure of the early Theropoda. The hind foot had, at the end of the Trias, become decidedly mesaxonic, with the hallux greatly reduced and probably somewhat turned back- ward. To create the foot of Diplodocus, for example, the hallux and the second digit must have been stimulated to increased growth ; that is, the foot must have been made entaxonic; whereas, the upright gait that is usually attributed to Diplodocus ought to have increased the size of the middle digits and further reduced the hallux. The meta- tarsals that had become lengthened had to be shortened. The fore limb, that in the late Triassic theropods had become reduced in length, sometimes greatly so, must have taken on renewed vigor and increased size. All the modifications that had been attained and all the ten- dencies established that looked toward making bipeds out of these theropods had to be reversed.
Probably little or no importance can be attached to the fact that no remains of sauropods have yet been encountered in the Triassic deposits. It is certain that but a small proportion of the animals that made those Connecticut Valley tracks have left us other traces of their existence. Then, it is extremely probable that comparatively few of the residents of that region were accustomed to parade on those desolate and dangerous tidal flats. The sauropods especially, being slow-footed plant-eaters, would naturally have sought localities where
22 OLWER P. HAY
there were fewer long-legged enemies and where the grazing was more satisfying.
To the writer, therefore, it appears most reasonable to suppose that the Sauropoda were a more primitive stock than the Theropoda and that the latter were derived from the early Triassic representa- tives of the former. Those primitive sauropods were no doubt far smaller than any of the group that are known to us. They probably had shorter necks, although with no fewer vertebrae; the vertebrae were less complexly constructed than those of their Jurassic descendants, and fewer of these had coosifised to form the sacrum. The digits, too, were probably longer and the outer ones were less reduced. We can hardly doubt that they crawled on their bellies.
The conviction has been expressed that bipedalism in the dinosaurs was caused by the relative reduction of the fore limbs. On the other hand, the writer believes that bipedalism among the birds was the result of specialization of the fore limbs. These different tendencies gave the signal for the parting of the dinosaurs and the birds. The birds were the gainers by the separation. They secured all that the dinosaurs got and far more besides. The two groups separated at an early period, early in the Triassic, possibly even in the Permian. It was undoubtedly at a time when the members of neither the one group nor the other had begun to walk on the hinder legs only. The feet, fore and hinder, were yet entaxonic. The hinder fifth digit was probably somewhat reduced, while the hallux was large and directed forward. Not until after the divergence of the two groups did the legs of the birds begin to be turned against the flanks and the body to be lifted from the ground. As greater and greater pressure began to be thrown on the middle digits the hallux began to be dwarfed and to be relegated to the hinder part of the foot. Archccopteryx shows that the hand had been entaxonic, for in it the two outer digits had wholly disappeared; while the pollex, though somewhat reduced, was yet large and functional.
It seems quite certain that the differentiation of the fore limb was initiated by the appearance of incipient feathers in the form, perhaps, of enlarged scales, which stood out from the ulnar side of the arms. The presence of these feathers, or scales, led to the flapping of the wings in the air, not conversely. Perhaps the individuals on which these rudimentary feathers first appeared were accustomed to clamber
THE POSE AND LOCOMOTION OF DIPLODOCUS 23
about over rocks and shrubs and the h'mbs of trees. Possibly the primitive birds, although not more than many lizards, strictly arboreal, often found safety and repose amid the branches and leaves of the Triassic ferns, calamites, and conifers. Possessing a fringe of feath- ers on their arms, they soon found these of advantage when they were running or making leaps to catch their prey or to escape capture by their enemies. When once they had made this discovery, the race entered on the conquest of the realms of the air.
It will be observed that the writer, in opposition to Dr. Francis Nopcsa'" holds that the primitive birds became bipedal while they were learning to fly and because of it, instead of becoming so long before the flying habit was initiated. It will be observed that the fore limbs of Dr. Nopcsa's ''Pro-avis" are already greatly reduced, and it might be questioned whether such limbs could be rejuvenated. It is certain that the ostriches have for untold generations been flap- ping their wings, to aid in running, but these limbs have steadily degenerated.
As believed by Dr. v. Huene, the Orthopoda probably took their origin from the Theropoda. If the views expressed by the present writer are true or approach truth, birds came on the arena before either of the suborders of dinosaurs just named; and hence most of the characters which have suggested relationship between the birds and the dinosaurs, which characters have been so clearly presented by Dollo and Nopcsa in the papers already quoted, have all arisen independently in the two groups as a result of their starting from the same goal and speeding in nearly the same direction. On the other hand, the sauropods are nearest the stock from which sprang the birds, and it is in their skeletons that we must seek for the primitive common characters.
To the writer it seems probable that the avidinosaurs were not amphibious animals, but dwellers on the land. It is not likely that wings were developed on animals that lived much in the water. The Theropoda and the Orthopoda continued to inhabit the land, although this did not prevent them from seeking their food in swamps or from refreshing themselves in the water. After the sauropods had attained such bulk that locomotion on the land became troublesome they
" Proc. Zo5l. Soc. London, 1907, p. 234.
24 OLIVER P. HAY
betook themselves to the streams, in order to enjoy the advantages of easier transportation; and then they became still more massive. Had they originally been aquatic and had they continued so, their feet would have remained more like those of crocodiles, less digitigrade and less shortened than they were in Diplodocus.
In his paper on the relationships of the birds and the dinosaurs^^ Professor Osbom says:
Thus tridactylism is correlated with rapid bipedal progression, the inner and outer digits sufifering reduction.
In formulating this apparently important generalization Professor Osborn did not qualify it with the statement that most of the so-called tridactyl animals are really tetradactyl, the hallux being present and usually functional. Nor could he have had before him the skeleton of any of the sloths, animals that are strictly tridactyl behind, but which are neither bipedal nor endowed with great speed. Tridactyl- ism prevailed among the extinct horse-like perissodactyls and is a characteristic of modern tapirs. On the other hand, there may exist swift bipedal progression independently of tridactylism. The ostrich makes rapid headway with only two toes, one might almost say, with a toe and a half. The kangaroos are wonderful bipedal leapers, whose functional digits are reduced to two, the fourth and the fifth. Man may be justly counted among the swift runners, trained individ- uas making their mile in four and a quarter minutes, and he possesses a pentadactyl entaxonic foot. No bipedal artiodactyl is recalled, but, as illustrating a possibility, one must not forget to mention Pan, the shepherd god of old Arcady. From which considerations it may be concluded that the bipedal rapid runners have adopted no standard form of foot.
Accompanying the present paper is a drawing (PI. I) which is intended to represent the habits of Diplodocus, especially as regards its habitual pose of body and its manner of locomotion, as conceived by the writer. This drawing was executed by Miss Mary Mason Mitchell, after consultation with the author of the paper. Two indi- viduals are in the foreground. One is collecting food from the sur- face of the water; the other has the head high in air and is jealously
" Amer. Naturalist, xxxiv, 1900, p. 796.
THE POSE AND LOCOMOTION OF DIPLODOCUS. 25
regarding the approach of another, which is swimming. In the far distance is a fourth specimen lying stretched out at full length on the bank.
In the paper published by Mr. Ballou, referred to on page 15, there is a figure which represents a group of four individuals of Amphicaiias latus, a dinosaur closely related to Brontosaurus and attaining a length of from 60 to 80 feet. These animals are shown as walking about on the bottom of a river, feeding on the vegetation there and rising on their hind legs to reach the air. The idea here suggested is adopted by Professor Osbom^® as correct. Mr. Knight, under Professor Osborn's direction, has made a restoration of Bronto- saurus*^ in which the same idea regarding the habits of the sauropods is inculcated. In this restoration a number of individuals, otherwise invisible, are sticking their heads out of the water. The ability of any large animal to walk thus submerged must depend on its having a massive skeleton, as have the hippopotamus and the manatee. In Diplodocus, on the contrary, almost every conceivable device has been employed to reduce the weight of the skeleton. The great vertebrae contain large and small internal cavities, while externally the processes are canned into thin plates and buttresses and the centra are deeply excavated on each side. Moreover, as has been shown by Hatcherj^"* the limb bones are hollow. It would seem to have been hardly more possible for Diplodocus to walk about immersed in water than it would be for a man to do the same. Even if the reptile could have remained sunken, any pressure by the feet in the effort to walk would have sent it to the surface.
After the text and the drawings of this paper had been completed the writer received the Scientific American of November 6, 1909, in which is printed a popular article on the attitude of Diplodocus. In this article mention is made of a paper on this subject recently pub- lished by Dr. Gustav Tornier of Berlin, a paper not previously seen by the present writer. Unfortunately too, he has not seen the original papers of Messrs. Drevermann and Boule. No numbers of the Umschau, of Frankfort, for the present year are accessible.
"Bull. Amer. Mus. Nat. Hist., x, p. 220.
•» Amer. Mus. Jour., V. p. 68.
*' Mem. Carnegie Mus., i, p. 53, fig. 23.
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Vol. XII, No. 2, pp. 27^269. May 15, 1910.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA,
CALIFORNIA.*
By Albert W. C. T. Herre.
Wer hat je die Flechten, wer hat die Moose gezahlet, Deren Friihling beginnt, wen Froste den Herbst entblattern, Deren iippiger Wuchs die Scheitel atherischer Alpen Da, wo sie Flora verlast, mit Tausend Farben bekleidet?
J. G. Herder.
The present paper is a synopsis of the author's studies of the syste- matic limitations and relationships of the lichens of the Santa Cruz Peninsula of California.
As already explained in earlier papers (Proc. Wash. Acad. Sci., vol. vii, p. 325, et seq., and Botanical Gazette, vol. xliii, pp. 267- 273), the Santa Cruz Peninsula forms a natural biological region, and as such naturally commends itself to the study of the naturalist interested in geographical distribution.
The region here treated is a roughly triangular area lying west of San Francisco Bay and the broad, originally treeless Santa Clara- San Benito valleys, and north of Monterey Bay and the Pajaro river. Rising from sea level along most of its border, its surface is greatly broken by a spur of the Coast Range, the Santa Cruz mountains, which rise at their highest point to an elevation of 3793 feet. Within this region, measuring no more than 90 miles in length and tapering from about 35 miles in breadth at the south to perhaps 6 at the Golden
* A Thesis presented as a part of the requirements for the Degree of Doctor of Philosophy at Leland Stanford Junior University, California.
Proc. Wash. Acad. Sci., Maj% 1910.
28 HEERE
Gate, are to be found the dense redwood forest and the naked ocean rock, the cold, foggy mountain crag and the bare, bhstering expanse of sand-dune, the monotonous salt marsh and the impenetrable chaparral.
The earliest collector of lichens in California was Archibald Menzies, a Scotch botanist and collector, who visited the northwest coast of America during the years from 1779 to 1796; in November and Decem- ber, 1792, he visited San Francisco Bay, Santa Clara, and Monterey and obtained specimens which were described by Acharius and also supplied Tuckerman with material more than 50 years later. Charles Wright, botanist of the North Pacific Expedition, collected a number of lichens at various points in the peninsula in 1855 and 1856.
The most important collections were those made by H. N. Bolander, who collected over a great part of the peninsula, and in fact over most of California, discovering a large number of most remarkable lichens during the years from 1863 to 1875.
Since Bolander's time no special work has been done on the lichens of the Santa Cruz region, though more or less important collections were made by the following: Dr. C. L. Anderson, of Santa Cruz; Dr. W. G. Farlow, who published a valuable set of Californian lichens, part of which were collected in the Santa Cruz mountains; Dr. L. M Underwood; Dr. Marshall Howe; and C. F. Baker.
The present paper describes 307 species and subspecies; but it is believed that further investigation will raise this number very materi- ally. In fact there are in the author's herbarium many specimens which he has as yet been unable satisfactorily to determine in the absence of authentic material for comparison, and literature which is not at present accessible.
In the matter of generic nomenclature, the treatment of the best students of lichens of the present day has been followed. In the case of species names the earliest recognizable name found in accessible authorities has been adopted.
Synonymy too often is the bugaboo of science, and the habit of many of our most eminent lichenologists of the past, as Nylander and Tuckerman, of changing names to suit their opinions, has not improved matters. No scientist, however eminent, has any right to change a name because it is inapplicable or denotes a character which may be common to a number of species within a genus. A name should have
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 29
an appropriate and distinctive meaning, but it need have no special significance and may be totally devoid of meaning or may even be misleading, as when minima is applied to the largest species of a genus. In the present unsettled condition of botanical nomenclature one is often in doubt as to what to do, but it seems clear that the law of priority should be observed.
Perhaps no more important work could be done than the careful overhauling of the synonymy of American lichens by someone who has access to the published exsiccata of Europe and America as well as the literature of the subject.
The author has endeavored to avoid the old conception where the word species was almost a generic term, including a large number of subspecies, varieties, and forms. Nature has no clean-cut, sharply drawn definitions, and perhaps in no organisms are the actual varia- tions and gradations so numerous or more puzzling than in lichens. But, nevertheless, a species should be a pretty distinct and well defined group in which the degree of variability is relatively small. Those forms which present constant differences in the field, or in structure, may be regarded as distinct species, while instead of giving every minor variation a varietal name we should rather work out the eco- logical factors producing them, and not overload an already too bur- densome synonymy.
While every part of the Santa Cruz Peninsula has been visited many times, certain localities naturally, have been found the richest in numbers or rarity of species. Perhaps first of these stands the region at the head of Devils Canon, a wild region where is found the largest mass of bare rock in the peninsula, and where there is a nearly vertical descent of perhaps 800 feet. Other localities offering peculiar attractions to the collector are the cliffs of the Golden Gate, and of the ocean shore from Point San Pedro to Pigeon Point, and the sand- stone ridges of Castle Rock and vicinity. But as a matter of fact there is not a caiion winding down to the ocean, not a group of old forest trees, not an insignificant reef of igneous rock outcropping in the foothill pastures, but will amply repay the intelligent efforts of any collector.
The Santa Cruz Peninsula is peculiarly rich in endemic species, and although collections in other parts of the state may considerably extend the range of some of them, it is probable that a goodly number
30
HEERE
may never be found elsewhere. The climatic conditions which cause this have already been discussed by the author elsewhere. Yet these same climatic conditions also give us an exceedingly diversified lichen flora, and it is believed that the present work can therefore be used as a manual for the identification of lichens over a great part of the western half of the United States.
In taking up the study of lichens, while the habit and general macroscopic structure is of importance and nothing can quite take the place of careful field work, the student must also be careful and accurate in the microscopic examination of all material. For this examination careful sections should be made both of thallus and fruit. A comparison of sections of the thallus of two plants may show con- stant differences when the fruit is similar in structure. The algae, too, need special attention, since some of the features relied upon by algologists may be absent when living under the conditions found within the lichen thallus. This is especially true of some of the fila- mentous blue-green algae.
As a corroborative test in the determination of species, one often finds certain chemical tests of considerable value. For this purpose a bit of the cortex, medulla, or apothecium is subjected to the action of potassium hydrate, or KOH, using a 25 per cent or 50 per cent solution. In the same way a saturated solution of calcium chloride, CaCl202 is used, either by itself or applied immediately after KOH. The ordinary solution of iodine used in the botanical laboratory, designated as I, is used frequently, most often in the examination of sections of apothecia.
Results of the above tests are not to be considered as sufficient to separate species when there are no other differences, but are secondary characters, to be considered with other characters based on structure or primary differences. Personally, the author has found the tests to be quite uniform and reliable, even when examining specimens obtained from regions thousands of miles apart, or growing on dif- ferent substrata.
In examining the spores of lichens, the beginner or general student must be cautious about observing and measuring any spores which may happen to be within the field of the microscope. In addition to the spores of various fungi which occur on the surfaces of all plants, the thallus of some lichens is frequently infested with parasitic fungi.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 3 1
while the apothecia of others are covered with minute parasitic apothecia. For example, the thallus of Heppia guepini is commonly the host of a parasitic Endococcus. As Heppia is very often sterile and the apothecia are not visible to the naked eye, one is exceedingly apt to be confused by the Endococcus spores unless very careful sec- tions are made.
Nylandcr, Tuckerman, and others, described the minute apothecia covering the disk of the fruit of many lichens as parasitic Buellias, Lecideas, and the like. But as they never contain algae, and have no thallus of their own, they are undoubtedly parasitic fungi, and accordingly are not considered in the present work.
In the preparation of this paper the author has been assisted at all times, and especially in the study of the Lecideaceae, by his fellow worker, Dr. H. E. Hasse, of Sawtelle, California, who has given his time and energy wi hout stint.
To my friend and teacher. Dr. Alexander Zahlbruckner, curator of the botanical section of the Imperial Natural History Museum of Vienna, Austria, I wish to express my gratitude for help while study- ing in the Museum and collecting with him in the Styrian Alps. His masterly treatment of lichens in Engler and Prantl's Die Naturlichen Pfianzenfamilien has been followed in this paper.
To Dr. W. G. Farlow of Harvard University I am indebted for many favors and the gift of valuable specimens, while to him and Mr. A. B. Seymour I owe the privilege of examining the Tuckerman Herbarium.
To the authorities of the British Museum and Kew Garden I am indebted for courtesies while examining the herbaria there.
The veteran Californian botanist, Volney Rattan, long time pro- fessor in the San Jose State Normal School, generously gave me a considerable collection of Californian lichens, presented him by H. N. Bolander; this collection has been of great service in deciding many difficult points.
Prof. Bruce Fink, of Miami University, Oxford, Ohio, has kindly determined a set of my collections of Cladonias, a labor of love which I greatly appreciate.
To Dr. William Trelease, the U. S. National Museum and the Bureau of Plant Industry, at Washington, D. C, and to the Botanical
32
HEERE
department of the University of California, I am indebted for the loan of literature otherwise inaccessible.
My sincere thanks are due Prof. William Russell Dudley, head of the department of systematic botany at Leland Stanford Junior Univer- sity, for timely assistance and helpful criticism in many ways, espec- ially in the final preparation of this paper. To him I dedicate this work, as long since he called my attention to the distinctiveness of the Santa Cruz Peninsula as a biological region.
Los Gatos, California, October, 1908.
ARTIFICIAL KEY TO GENERA OF LICHENS OF THE SANTA CRUZ PENIN- SULA, CALIFORNIA.
Crustaceous Lichens.
A. Thallus absent.
B. On thallus of Pertusaria; apothecia top-shaped.
IX. Sphinctrina BB. On rock, wood, or bark.
C. Asci multisporous, spores minute XXVI. Biatorella
CC. Spores not more than 8.
D. Apothecia lecideine XIX. Lecidea
DD. Apothecia lecanorine XLIII. Lecanora
AA. Thallus more or less developed.
E. Apothecia pyrenocarpous, more or less globose, immersed, hemi- spherical or sessile, with a terminal pore. F. Algae Trentepohlia.
G. Paraphyses branched and twining IV. Arthopyrenia
GG. Paraphyses simple and free V. Porina
FF. Algae Pleurococcus.
H. Thallusuniformcrustaceous; spores 8, simple... T. Verrucaria HH. Thallus of sub-foliaceous to minute squamules.
/. Spores 8, simple, colorless II. Dermatocarpon
II. Spores 2, muriform, yellowish to brown. .III. Endocarpon EE. Apothecia not pyrenocarpous.
J. Apothecia more or less stalked, or sessile, spores extruded and forming a sporal mass or mazaedium covering the disk.
K. Apothecia containing algae IX. Sphinctrina
KK. Apothecia not containing algae. L. Apothecia on long stalks. M. Thallus and stipe greenish-yellow, powdery.
VII. Coniocybe MM. Thallus and stipe not yellow.
N. Spores bilocular VI. Caliciiim
NN. Spores 4-8 celled VIII. Stenocybe
LL. Apothecia short-stipitate or sessile.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 33
O. Apothecia pear or top-shaped, growing on Pertusaria
thallus, containing alga;; spores simple . . .IX. Sphinctrina
00. Apothecia crateriform, not containing alga;; spores
bilocular, or simple in i form X. Cyphelium
JJ. Apothecia without a mazasdium.
P. Apothecia usually linear or elongate, w'ith a fissure-like disk; seldom circular; algae Trenlepohlia. Q. Thallus very thin, uniform crustaceous. R. Apothecia without margin, more or less stellate, branch- ing, or irregular XII. Arthonia
RR. Apothecia with a margin.
5". Apothecia innate, fissure-Hke, with evident disk; spores
caterpillar-like, brown XIV. PhcBographis
SS. Apothecia sessile, not innate or fissure-like; apothecia
irregular, linear or ellipsoid XIII. Opegrapha
QQ. Thallus thick, irregular, of ten warty or sub-plicate; apothe- cia circular or nearly so, with both proper and thalline
margins XV. Dirina
PP. Apothecia more or less circular and dish or shield-like; never linear, though often angular or variously shaped from pressure or crowding. T. Algae blue-green {Cyanophycea). U. Algae Nostoc.
V. Apothecia biatorine or lecideine. .XXXV. Parmeliella
VV. Apothecia lecanorine XXXVII. Pannaria
UU. Algae not Nostoc. W. Algae Scytonema.
X. Thallus squamulose to small-foliaceous, mostly of parenchyma: spores simple, colorless.
XXXIV. Heppia XX. Thallus crustaceous, coralloid, to small squamu- lose, spores colorless, 2-8 locular.
XXXVI. Placynthium WW. Algae not Scytonema.
Z. Algae Stigonema; thallus microscopically fruticose,
of terete filaments XXVIII. Zahlbrucknera
ZZ. Algffi Gloeocapsa; thallus diffract-crustaceous to
minutely squamulose XXXI. Pyrenopsis
TT. Algae not blue-green.
a. Algae Trenlepohlia '. XVII. Lecanactis
aa. Algae Pleurococcus, Protococcus, or Palmella.
b. Spores simple, colorless, minute, exceedingly numerous; thallus of scales or squamules, with usually innate apothecia.
XXVII. Acarospora bb. Spores not excessively numerous.
c. Spores bi-locular, topically polari-locular, or becoming muri- form by interpolation of cross-walls. d. Spores colorless, polar-bilocular.
e. Apothecia lecanorine LVIII. Caloplaca
34 HERRE
ee. Apothecia biatorine or lecideine LVII. Blastema
dd. Spores brown, bilocular to muriform.
/. Apothecia urceolate; spores muriform from beginning.
XVIII. Diploschistes ff. Apothecia not urceolate.
g. Spores with a distinct halo. . . .XXIII. Rhizocarpon gg. Spores without a halo.
h. Apothecia lecideine LXI. Buellia
hh. Apothecia lecanorine LXII. Rinodina
cc. Spores simple to multilocular, but not polar-bilocular; color- less. J ,. |^
i. Apothecia lecideine.
j. Spores simple XIX. Lecidea
jj. Spores bi- to multilocular,
k. Spores bilocular XX. Catillaria
kk. Spores 4-16 locular
/. Thallus without cortex, areolate or uniform.
XXI. Bacidia II. Thallus with cortex; of swollen or plicate warts or squamules . .XXII. Toninia it. Apothecia lecanorine.
m. Apothecia single or grouped, immersed in thai- line warts; disk very narrow; spores large to
very large XLII. Pertiisaria
mm. Apothecia solitary, not immersed in thalline
warts; disk relatively broad. n. Spores simple.
0. Paraphyses free, simple; spores small to medium. p. Spermatia thread-like; thallus variously colored but not orange-yellow.
XLIII. Lecanora pp. Spermatia ellipsoid; thallus orange-yel- low XL VII. Candelariella
00. Paraphyses branched and entwined; spores
quite large XLIV. Ochrolechia
nn. Spores bilocular.
q. Sterigmata exobasidial. r. Thallus orange-yellow; spermatia ellipsoid.
XLVII. Candelariella rr. Thallus not orange-yellow; spermatia
thread-like XLV. Lecania
qq. Sterigmata endobasidial ; thallus marginally lobed or of more or less lobate squamules.
XLVI. Placolecania
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 35
Foliaceoiis Lichens.
A. Thallus gelatinous when wet; color always dark; algae blue-green iCyanophyce^.)
B. Algae Scytonema XXX. Polychidium
BB. Algae Nostoc.
C. Thallus dark green without cortical layer XXXII. CoUema
CC. Thallus usually lead-colored, with distinct cortical layer.
XXXIII. Lcptogmm A A. Thallus not gelatinous when wet.
D. Apothecia never present. E. Thallus dark.
F. Plant apparently black, the surface coveied with black isidia.
XXXIX. Sticta FF. Plant dark brown, sub-f ruticose ; the ascendant, irregularly
cut lobes with narrow white edge L. Cetraria
EE. Thallus green or pale.
G. Yellowish green with gray soredia; beneath villous or with
naked pale spots XXXVIII. Lobaria
GG. Plant more or less orbicular, often very large, grey, yellowish or bright green; beneath black, usually brown-maigined,
more or less black-fibrillose XLIX. Parmclia
DD. Apothecia usually present and abundant.
//. Thallus attached at a single point near the center by an umbili- cus. /. Apothecia visible to naked eye; thallus large or of medium size. /. Apothecia adnate, gyrose; thallus brown. . .XXV. Gyrophora JJ. Apothecia immersed, appearing as minute dark specks on the
ashy gray thallus II. Dermatocarpon
II. Apothecia not visible to naked eye; thallus very small, dark olive or blackish brown, expanded or ascendant.
XXXIV. Ileppia HH. Thallus attached by numerous rhizoids, not umbilicate. K. Apothecia adnate on undei side ot marginal lobes.
L. Algaae Nostoc; spores 2-4 locular XL. Nephroma
LL. Algae Protococcus; spores simple LI. Nephromopsis
KK. Apothecia always on upper surface of thallus. M. Thallus bright yellow or oiange.
N. Apothecia chestnut; spores simple, colorless.
L. Cetraria NN. Apothecia yellow or orange. 0. Spores polai-bilocular, colorless, 8.
LIX. X author ia
00. Spores simple, 16 to 60 XLVIII. Candelaria
MM. Thallus not bright yellow 01 orange.
P. Thallus horizontal, orbicular or variously lobed; undei surface with veins or cyphels. Q. Under surface with small white cyphels, villous or fleecy XXXIX. Sticta
36 HERRE
QQ. Under surface without cyphels.
R. Thallus pale villous beneath with large, pale,
naked spots XXXVIII. Lobaria
RR. Thallus pale or whitish beneath, with brown veins and fibrils; apothecia adnate on tips of more or less elongate lobes.
XLI. Pelligera. PP. Without veins or cyphels on under surface.
6'. Spores bi-locular, brown LXIV. Physcia
SS. Spores simple, colorless.
T. Thallus flat, appressed; under surface brown or black, more or less black fibrillose; apothecia
scattered over surface XLIX. ParmeUa
TT. Thallus sub-fruticose, compressed; apothecia marginal or on tips of ascendant lobes.
L. Cetraria
Fruticose Lichens.
Plants more or less erect and shrub-like, or drooping and pendulous.
A. Thallus of two kinds: (i) a horizontal, more or less leafy or granulose one; (2) a more prominent, erect, and caulescent one, really stalks for the apothecia but apparently the plant; simple, and club, cup, or funnel-shaped, or slender and much branched; apothecia scarlet or
brown XXIV. Cladonia
A A, Thallus uniform, not two-fold. B. Apothecia globose, terminal; plant tufted, shrub-hke, gray.
XL SphcBrophorus BB. Apothecia dish or shield-Hke; terminal, marginal, or more rarely scattered. C. Thallus hair-like.
D. Black or brown, like tangled mats of fine hair.
LIV. Alecioria DD. Color not black or brown.
E. Thallus erect or decumbent, densely tufted, intricately branched, terete, gray, sterile; on maritime rocks.
XVI. Dendrographa EE. Thallus coarser, gray or pale straw-color, rarely red; tufted or pendulous, becoming enormously elongated; apothe- cia concolorous or pale tan, with fibrillose margin.
LVI. Usnea CC. Thallus not hair-like. F. Plants not gray or green. G. Thallus brown or black. H. Sooty black, very small, compact, sterile: on vertical
sandstone walls XXIX. Ephebe
HH. Greenish black or brown, spreading, compressed; apothe- cia abundant, terminal L. Cetraria
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 37
GG. Thallus yellow.
/. Spores simple, colorless; thallus bright lemon-color; apothe-
cia chestnut LIII. Letharia
II. Spores polar-bilocular, colorless; plants and apothecia reddish yellow or orange. /. Thallus erect, lax, pendulous or decumbent; on trees and
maritime rocks LX. Theloschistcs
J J. Thallus short, rigid, becoming decumbent; on mari- time rocks LVIII. Cahplaca
FF. Plants gray, green, or pale. K. Apothecia present.
L. Apothecia concolorous; thallus tufted, compressed or terete, or elongate, pendulous, and greatly compressed.
LV. Ramalina LL. Apothecia not colored like thallus.
M. Apothecia black, pruinose or naked; thallus mar- ginally fibrillose and fuzzy . .LXIY. Anaplychia MM. Apothecia not pruinose or black.
N. Apothecia chestnut; lobes long, ascendant, white
beneath, on trees L. Celraria
NN. Apothecia yellowish or dusky; plants very short, stout, erect, rigid, sub-crustaceous; on maritime
rocks XLIII. Lecanora
KK. Apothecia absent.
0. Lobes narrow, ascendant, margined with stout, usually
branching fibrils LXIV. Anaptychia
00. Lobes not marginally fibrillose.
P. Sub-crustaceous; short, stout, terete; powdery, simple
or branched; on maritime rocks . .XLIII. Lecanora
PP. Thallus pendulous or erect ; more or less white sorediate ;
on trees and shrubs LII. Evernia
LICHENES.
ASCOLICHENES.
Composed of Fungi belonging to the Ascomycetes, living in union (symbiosis ?) with Algae, these two distinct classes of plants apparently forming a morphological and physiological unit.
I. Pyrenocarpe^: Apothecia globular, usually opening at the summit by a minute pore.
II. Gymnocarpe^: Apothecia more or less open and the disk exposed, circular and shield-like or dish-like, elongated and variously shaped, or crater-like.
38 IIERRE
PYRENOCARPE^.
Thallus crustaceous, squamulose, rarely foliaceous, more rarely fruticose. Gonidia of Pleurococcus, Chroolepus or Trentepohlia, Phyllactidium, Nostoc, or Sirisiphon algae. Soredia are absent or very rare. Apothecia globular or hemispherical, opening only by a pore at the summit, immersed in the thallus with only the apex protruding, or sessile; naked or more or less covered by a thalline layer; solitary, or confluent and forming a stroma; a proper exciple or margin, known as the perithecium, pale to black, closed or open below (dimi- diate) ; within this a more or less distinct envelope, the amphithecium, enclosing a globose hymenium, the nucleus of many authors, which is soft, gelatinous, and often contains algae or oil drops. Paraphyses simple, or branched and then twining and net-like; commonly soon gelatinizing and apparently lacking. The Angiocarpous lichens of many authors.
KEY TO FAMILIES.
Thallus with Pleurococcus or Palmella alga.
Thallus crustaceous, without cortex V errucariacecs
Thallus foliaceous or squamulose, cortex present — Dermatocar pacecB
Thallus with Trentepohlia alga, crustaceous, the apothecia solitary or
sub-confluent but not forming stroma PyrenulacecR
VERRUCARIACE^.
Thallus uniform crustaceous, growing upon or within the upper layers of the substratum, without cortex, the gonidia of Pleurococcus or Palmella algae. Apothecia single, erect, with an apical pore.
I. Verrucaria (Web.) Th. Fr.
Verrucaria Weber, Prim. Flor. Hols. p. 85.
Verrucaria Th. Fries, Gen. Heterol. Europ. 109. 1861.
Verrucaria A. Zahlbr., Ascolichenes, 54. 1907.
Thallus crustaceous, rimose areolate, or powdery, commonly with an evident hypothallus, rarely sorediate, usually upon the substratum, rarely within. Apothecia entirely immersed, half sunken, or sessile; perithecium coal-black, horny, globular, flask-like, or hemispherical with the underside open; paraphyses soon gelatinizing; spores 8, ellipsoid, oval, or globose, one-celled, colorless or rarely brown.
About 100 species, often difficult to define.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 39
KEY TO SPECIES.
A. Thallus thin to very thin.
B. Black, resembling a smear of black paint; on maritime rocks.
5. melas BB. Not black. C. Very thin and powdery, or mostly obsolete; forming white spots
on limestone 6. calciseda fusca-spora
CC. Of minute ashen or gray granules; on sandstone.
4. miiralis AA. Thallus more or less areolate or scaly; from thin becoming thick. D. Color pale.
E. Areoles thick, bluish gray 7. stanfordi
DD. Color dark to the naked eye.
F. Thallus areolate or scaly, ashy gray under lens ; black appearance
due to the numerous apothecia i . rupestris
FF. Thallus some shade of brown. G. Pale to dark olive brown, with a more or less greenish cast when
wet 2. viridula
GG. Dark brown to black, not becoming greenish when wet.
3. nigrescens
I. VERRUCARI A RUPESTRIS Schrader.
Verrucaria rupestris Schrad. Spicil. Fl. Germ. 109. 1794. Verrucaria rupestris Leighton, Brit. Angiocarp. Lich. 60. PL. 25. f. 4. 1851.
Thallus effuse, thin to thickish, apparently continuous, but really minutely fissured and areolate or scaly; black to the naked eye, but examination with the lens shows it to be ashy gray.
The black color is due to the numerous, large, prominent, hemis- pherical, and semi-immersed apothecia; perithecium thick, black, dimidiate; amphithecium thin, pale brown; hymenial gelatine blue with I; spores colorless to pale yellow, ellipsoid or sometimes pointed
, 8i-ii
at one end, ~ pL.
20-25
On rocks at Point Lobos, San Francisco, and elsewhere along the ocean shore; probably occurring throughout our territory. Com- monly distributed over the North Temperate region.
I also doubtfully refer here a plant on sandstone, Castle Rock Ridge, altitude 2000 feet, differing as follows:
Thallus areolate, fissured, the areoles distinct or running into a
40 HERRE
continuous crust, chaffy or microscopically corallinoid, dingy brown
in color; resembling a crustaceous Pannaria or Collemaceous lichen.
Apothecia not numerous or conspicuous, small, black, semi-im-
mersed; spores larger than in the previously described form, . ? fx.
24-30
In other respects agreeing.
Unlike anything in the accessible literature or specimens, and per- haps new.
2. VERRUCARIA VIRIDULA Ach.
Verrucaria viridula Acharius, Lich. Univ. 675. 1810.
Thallus forming a thickish, effuse, rough or uneven, chinky or fissured crust of minute, polygonal areoles, closely compacted; their surface smooth or rough or warty; color varying from pale to dark olive blackish brown, with a more or less evident greenish cast when wet; KOH-; CaCl30,-.
Apothecia usually but one in an areole, rather large,black, immersed, with only the apical portion visible; usually without an ostiolum; perithecium dimidiate, thick, black; amphithecium thin, black; hymenial gelatine bluish with I; spores broadly elliptical, colorless,
I2i — I??
often with one or two very large oil drops, — ~ fi.
24 - 34
Abundant on sandstone in the foothills and probably occurring all through the mountains.
A plant of Europe, North America, and northern Africa.
3. VERRUCARIA NIGRESCENS Pers. Verrucaria nigrescens Persoon, Ust. Ann. Bot. 14: 36. 1795.
Thallus effuse or sub-determinate, thin or becoming moderately thick, of minute areoles, compacted into a nearly uniform crust; color dark brown to black.
Apothecia numerous, of moderate size, hemispherical, more or less immersed and with the apex often covered by the thallus; the apical pore usually not visible; perithecium entire; hymenial gelatine pale salmon or reddish with I, or part reddish and part blue with I; spores ellipsoid to short and broadly ovate, often falsely bi-nucleolate or
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 4I
6 - 81V II - 13I ,8-15
containing; a large oil drop, a. — ^^ //, and n',
^ ^ ^14-18 'i9i - 23 19-29
according to Nylander, ~ a.
23 - 27 "^
On sandstone in the foothills and mountains. Common in Europe,
Africa, and North America.
4. VERRUCARIA IMURALIS Ach.
Vernicaria muralis Acharius, Meth. Lich. 115. 1803. Verriicaria muralis Tuck. Gen. Lich. 269. 1872. Verrucaria muralis Jatta, Lich. Syll. Ital. 1325. 1900.
Thallus thin, effuse, and disappearing, ashen or gray in color, of minute granules.
Apothecia, very small, scattering, black, pruinose or naked, hemis- pherical, rather prominent; the ostiolum very minute, hardly visible under the lens; perithecium dimidiate, black; amphithecium pale brown or yellowish; hy menial gelatine blue with I; spores ellipsoid,
colorless and pale yellowish, a.
13 - 24^
Rarely on sandstone on the highest peaks. Generally distributed over Europe and North America.
5. VERRUCARIA MELAS Herre, new species.
Thallus thin or very thin, effuse, microscopically areolate and intricately fissured, appearing to the naked eye as a daub of dead black paint; KOH— ; CaCL02 — .
Apothecia not numerous, at first low and covered by the thallus, becoming more prominent and emergent, hemispherical, finally sessile and sub-globose, the apical portion then irregularly depressed and the ostiolum comparatively large; perithecium entire, black; asci oval, clavate, and pear-shaped, the hymenial gelatine blue with I; spores
„. .J 8^ - I2i
ellipsoid, , «.
T-9h - 27 Rare; on rocks a few feet above the sea at Point Lobos, San Fran- cisco, associated with Arthopyrenia halodytes.
Strongly resembling Verrucaria maura in general appearance, but
42 HERRE
differing in the thinner thallus, the much larger and somewhat dif- ferently shaped spores, and the chemical reaction. A specimen of Verrucaria maura from Dr. Farlow, collected at Campobello, New
Brunswick, yields me oblong spores, measuring -~ 7 a, while the
I2i — 16
reaction with I is vinous red. (melas, ixiXac, black.)
6. VERRUCARIA CALCISEDA FUSCA-SPORA Herre, n. subsp.
Verrucaria calciseda DC. Fl. Fr. 2: 317. 1805.
Verrucaria calciseda Leighton, Lich. Fl. Grt. Britain, ed. 3, 458. 1879.
Thallus effuse, endolithic, very thin and powdery or obsolete, form- ing white spots on the rock.
Apothecia numerous, minute, black, immersed in tiny pits in the rock, becoming slightly protuberant; ostiolum depressed, circular, pore-like, or often irregularly fissured; perithecium broad, dimidiate, black; gelatinous thecium bluish or blue with I; asci clavate; spores ovoid or elliptical, from colorless becoming dusky and finally dark
brown, ^~ ^ «.
i7i - 28'^
Abundant on limestone near the summit of Black Mountain, alti- tude 2700 feet, mixed with Rinodina hischoffi, immersa, from which it is with difficulty externally distinguishable. The internal structure not satisfactorily made out owing to the difficulty of extracting the apothecia from their holes without breaking them.
Differs from the type in having dark spores. Leighton says (Brit. Angiocarpous Lichens, 60, 1851, pi. 25, f. 4) that the spores of V. rupestris are pale yellow, but elsewhere he and other authors call the spores of all this group colorless.
(fusca, dark; spora, oTzdpoc, seed.)
7. VERRUCARIA STANFORDI Herre, new species.
Thallus of thick, somewhat convex areoles, reaching a thickness of 2 mm., either separate or arranged in small groups or finally compacted into a deeply fissured, continuous, chinky crust; the surface very min-
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 43
utely granulose, of a handsome bluish gray color, shading off to ashen; KOH-; CaCl203-.
Apothecia one to several in each areole, the perithecium sunken in the thallus, globular, entire, thick, black; ostiole from sunken becom- ing elevated, circular, thickish; hymcnial gelatine blue vi^ith I; spores 8, simple, colorless at first, at last brownish, elongate-ellipsoidal,
6 - 8.C 14^ - 22
On rocks in the foothills near Stanford University, at an latitude of 150 feet. On a rock beside the road to Lick Observatory, alt. about 800 feet, Alt. Hamilton Range.
(stanfordi, because found near Stanford University.)
DERMATOCARPACEiE .
Thallus foliaceous or squamaceous, with a cortex of pseudoparen- chyma upon one or both sides; fastened to the substratum by an umbilicus, by rhizoids, or adherent by the medulla; gonidia of Pleuro- coccus algae. Apothecia solitary, erect, with a minute pore at the summit, more or less immersed in the thallus.
KEY TO GENERA.
Apothecia without gonidia II. Dermatocarpon
Apothecia containing gonidia III. Endocarpon
II. Dermatocarpon (Esch.) Th. Fr.
Dermatocarpon Eschweiler, Syst. Lich. Gen. 21. 1824. Dermatocarpon Th. Fries, Gen. Het. Europ. 103. 1861.
Thallus leafy and greatly expanded or reduced to areolate squa- mules; umbilicate, or appressed and adnate; the algal layer beneath the upper cortex. Apothecia solitary, erect, minute, immersed, or with only the apical portion emergent upon the surface, upon which they appear as minute specks; perithecium clear or black, globular or ovoid, without algae; paraphyses usually gelatinizing, rarely poorly developed and then branched and net-like; asci 8—16 spored; spores ellipsoid, simple, colorless.
Proc. Wash. Acad. Sci., May, 19 10.
44 IIERRE
KEY TO SPECIES.
Thallus of flat or flattish adnate squamules.
Perithecium black; thallus more or less olive or blackish brown; on
bark i . sqiiameella
Perithecium clear; squamules more or less reddish or chestnut brown ;
on earth 2. hepaticum
Thallus foliaceous, umbiHcate.
One-leaved, large 3. minatum
More or less polyphyllous
The convolute, complicate lobes ascendant, the plant more or less
cespitose 4. miniatum complicatum
Thallus pseudo-crustaceous, small, closely appressed.
5. aquaticum
Section CATOPYRENIUM.
Thallus squamulose; or minutely foliaceous, but not umbilicate; perithecium black.
I. DERMATOCARPON SQUAMEELLA (Nyl.).
Verrucaria squameella Nyl. in litt. to Dr. Hasse. 1897.
Thallus foliaceous, small to minute, irregularly lobate, with uneven, undulate surface; the more or less upturned margin crenate, some- times minutely orbicular-lobulate fringed; color dark olive or reddish brown to blackish; beneath whitish or pale, fastened to the substratum by numerous rhizoids.
Apothecia rather numerous, immersed, minute, pear-shaped or subglobose, the perithecium entire, dark brown or blackish; asci clavate to short ventricose; hymenial gelatine vinous red with I, but when KOH is applied first and then I the reaction is a quickly fading
2* — c; "6 — 7
blue; spores narrowly ellipsoid, — = ^ — /i; fx," Nyl. in
i2i - 17^ 15 - 18
liti.
Very rare; collected but once, on the bark of Quercus lobata, in the foothills 5 miles south of Stanford University. Originally collected by Dr. Hasse near Los Angeles, where it is also very rare.
Section ENDOPYRENIUM.
Thallus of adnate squamules or sub-foliaceous, but not umbilicate; perithecium clear.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 45
2. DERMATOCARPON HEPATICUM (Ach.) Th. Fr.
Endocarpon hepaticum Acharius, Lich. Univ. 298. 1810. Dermatocarpon hepaticum Th. Fr. Lich. Arct. 255. i860.
Thallus of small, rounded, closely adnata squamules, these becom- ing wavy; margin more or less incised or crenate, upturned, blacken- ing; usually distinct, but sometimes slightly imbricate; color varying from bright reddish brown to chestnut, and dull dusky brown.
Apothecia numerous, immersed, the ostioles minute, becoming pro-
tuberant with black margin; spores oval, -^ — fi.
12 — 14
On bare earth or in crevices of rocks in the foothills and mountains. In Tuckerman Herbarium (Bolander's collection) from San Fran- cisco and Oakland. Common on calcareous soil throughout the North Temperate Zone.
Section ENTOSTHELIA. Thallus foliaceous, attached by a central umbilicus.
3. DERMATOCARPON MINIATUM (L.) Mann.
Lichen miniatus Linn. Sp. PI. 2: 1149. 1753.
Dermatocarpon miniatum Mann, Lich. in Bohemia observ. p. . 1826.
Dermatocarpon miniatum Herre, Proc. Wash. Acad. Sci. 7 : 394. 1906.
Thallus medium to large, smooth, coriaceous, one-leaved or lobate, the margin rounded, undulate, or crenate, and more or less recurved; attached by an umbilicus; color whitish to bluish gray or occasionally brownish; sometimes granulose pruinose; beneath varying from a bright to a dark brown or black, smooth or minutely pustulate.
Apothecia very numerous, minute, scattered, immersed in the
thallus; opening by small dark or brown pores, which appear as specks
thickly distributed over the entire surface. Spores ellipsoid,
4i - 6i ^^ n.
9f - 17
On rocks, in shaded or damp situations. A common and con- spicuous lichen throughout the foothills and to the summit of the Santa Cruz range, the thallus reaching a diameter of more than four inches in specimens from San Bruno Mountain. Common in the
46 HERRE
mountains of Europe, northern Africa, North America, and New
Zealand.
(miniatum, red, of no application here.)
Of several named varieties one is common here.
4. DERMATOCARPON MINIATUM COMPLICATUM (Sw.)
Th. Fr.
Lichen complicatus Swartz, Nov. Act. Upsal. 4: 38. 1776. Dermatocarpon miniatum complicatum Th. Fr. Nov. Act. Soc. Sci. Ups. Ill, 3: 353. 1861.
Thallus small to medium, polyphyllous, densely compacted, the imbricate and complicate lobes rotund, convolute, and more or less ascendant, with recurved margin; the surface more or less roughened and wrinkled. Otherwise like the species.
Common in the foothills with the type, but most abundant on sun- nier or more exposed rocks.
5. DERMATOCARPON FLUVIATILE (Weber).
Lichen aquaticus Weis, PI. Crypt. 77, 1770. Not L. 1753. Lichen fluviatile Weber, Spec. Fl. Gott. 265. pi. ^. 1778, Dermatocarpon aquaticum A. Zahlbr. Annalen des K. K. Naturh. Hofm.
Band XVI, Heft I, p. 81. 1901. Dermatocarpon aquaticum Herre, Proc. Wash. Acad. Sci. 7 • 394- 1906.
Thallus small, thick, smooth, lobes densely imbricate and com- pacted; margin rounded, entire or crenate-lobulate; closely appressed, umbilicate; resembling an intricately convolute, adnate, crustaceous lichen; color dull gray or olive-brown; sometimes white granulose pruinose; beneath smooth, dark brown to dingy black.
Apothecia as in Dermatocarpon miniatum, but proportionately
larger and less numerous; spores ellipsoid, — ^ ~ jx.
14^ - i9i
Abundant on granite cliffs 200 feet above the sea, near Point San Pedro; occurring also on wet sandstone in Devils Cafton, altitude 2300 feet, and at the head of Kings Creek at about 2800 feet. Widely dis- tributed in the North Temperate region.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 47
III. Endocarpon (Hedw.) A. Zahlbr.
Endocarpon Hedwig, Descr. et Adumbr. Muse. 2 : 56, pi. 20. f. A. 1788. Endocarpon A. Zahlbr., Ascolichenes, 61. 1907.
Thallus of distinct squamules or leafy scales, closely adnata. Apothecia solitary, much as in Dermatocarpon, but the hymenium containing minute ellipsoid or circular green algae lying between the asci; perithecium dark to black; paraphyses gelatinizing; asci saccate or ventricose-clavate, with 1—6 spores which are elongate or ellipsoid, muriform-multilocular, from colorless becoming yellowish and dark brown.
Species few, widely distributed on earth and rocks.
KEY TO SPECIES.
Squamules flat, closely adnate i. pusillum
Squamules overlapping, with upturned margins 2. pallidum
Thallus with rounded or incised lobules more or less complicate-imbricate.
3. pulvinatum
I. ENDOCARPON PUSILLUM Hedwig.
Endocarpon pusillum Hedwig, Descri. et Adumbr. Muse. 2 : 56, pi. 20,
f. 8, 1788; Germany.
Thallus of small to minute squamules, more or less scattered, or occasionally forming a crust; flat, closely adnate, circular or with crenate or lobate margin; surface smooth or fiexuous; color passing from reddish brown to blackish brown; KOH— ; CaCUOj — .
Apothecia small, immersed, rather numerous, the minute ostiolum at length enlarged and pit-like; perithecium dark brown, entire; spores 2, one usually larger than the other, at length dark brown,
T-Zh - 24 -^^ fi.
24i - 56
Common on crumbly sandstone in the foothills, either alone or upon a cushion of Scytonema filaments, which also often grow over the thallus. Probably occurring all through the mountains wherever there is sufficient rock exposure.
Recorded from Europe, North America, Algeria, and New Zealand.
48 HERRE
2. ENDOCARPON PALLIDUM Ach.
Endocarpon pallidum Acharius, Lich. Univ. 301. 1810, Switzerland.
Thallus of small pale reddish or darker scales, these overlapping, flexuous, and more or less marginally upturned; the rounded lobes with more or less crenate margin; KOH— ; CaClzOz — .
Apothecia minute, immersed, opening to the surface by a pore which sometimes becomes enlarged; perithecium dark brown, entire;
spores 2, from pale becoming light brown, — //; Dr. Hasse gives
31.8-37
them as—— — « 45 - 52 " Rare, on stones in the foothills; found in Europe, North America,
and New Zealand.
3. ENDOCARPON PULVINATUM Th. Fr.
Endocarpon puhinatum Th. Fr. Lich. Arctoi, 257. i860. Polyblastia pulvinata Jatta, Syll. Lich. Ital. 562. 1900.
Thallus dark chocolate-brown, the squamules more or less imbri- cate, lobulate, with irregularly crenate or incised margin, the whole plant resembling a minute Gyrophora; apothecia very small, immersed, the perithecium more or less protruding, black; asci clavate; spores 2,
muriform multilocular, //.
14 - 17
On boulders in the foothills; rare. A lichen of arctic and alpine
Europe.
PYRENULACE^.
Thallus uniform crustaceous, without cortex or rhizoids; growing upon or within the substratum; the algae Trentepohlia. Apothecia single or confluent, erect, with a terminal pore; spermatia exobasidial.
KEY TO GENERA.
Paraphyses branched and twining, or becoming gelatinous ; spores 2 -6
celled IV. A rthopyrenia
Paraphyses unbranched, free; spores 2 — many celled. ...... .V. Porina
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 49
IV. Arthopyrenia.
Thallus thin, with an indistinct hypothallus as a rule. Apothecia with a globose or hemispherical, horny, black perithecium; para- physes permanent or gelatinizing; spores wedge-shaped, oval, or elongate, with usually blunt ends.
KEY TO SPECIES.
A . On maritime rocks /. halodytes
A A. On bark.
B. Spores bilocular, arranged in one row in the asci.
C. Paraphyses slender, distinct 6. conjormis
CC. Paraphyses threadlike, branched, entangled 5. hijormis
BB. Spores bilocular, often with a halo, variously arranged in the asci.
D. Paraphyses gelatinizing 2. analepta
DD. Paraphyses distinct.
E. Apothecia naked 3. analeptella
EE. Apothecia more or less pruinose 4. cinerea-prtiinosa
Section EUARTHOPYRENIA.
Perithecia usually hemispherical; paraphyses normally becoming gelatinous; spores usually with a gelatinous halo, constricted in the middle, of 2 dissimilar cells, either cell or both sometimes further divided.
I. ARTHOPYRENIA HALODYTES (Nyl.) Wedd.
Verrucaria halodytes Nyl. Enum. generale des Lichens, 142. 1858. Arthopyrenia halodytes Weddell, Excurs. lich. d. I'ile d'Yeu, 307.
1874. Arthopyrenia halodytes Olivier, Lich. de I'Ouest et Nord-O. France. 2 : 261. 1900.
Thallus efifuse, thin and uniform, or irregularly thickened, when it becomes somewhat fissured; blackish reddish or reddish brown; KOH - ; CaCl,02 -.
Apothecia very small and numerous, black, more or less immersed, or emergent and sub-globose; scattered or becoming clustered and crowded; paraphyses mostly lacking, rather thick and jointed when
present; asci ventricose, — ^ //; no reaction with I; spores
49 - 62
bilocular, ellipsoid, irregularly arranged in the asci, -^ — pL.
II- 14.7
so
HERRE
Abundant on precipitous rocks just above high tide, Point Lobos, San Francisco. Described from the northern coast of France.
2. ARTHOPYRENIA ANALEPTA (Ach.) Korber-
Lichen analeptus Ach. Lich. Suec. Prodr. 15. 1798. Verrucaria analepta Ach. Meth. Lich. 119. 1803. Arthopyrenia anakpta Korber, Syst. Lich. Germ. 367. 1855.
Thallus e£fuse, thin to obsolete, forming olive or coppery patches on bark.
Apothecia small to minute, usually not numerous, scattered, black, sessile or hardly innate, hemispherical to conical; perithecium dimidi- ate; paraphyses disappearing in a gelatinous mass; asci long, tubular,
- l_/x; spores 4, 6, and 8 in the asci, bilocular, ellipsoidal, con-
90 — 122
stricted at the middle, the divisions w^edge-shaped, one often larger
than the other, -i ^; once observed 3-locular; asci and their con-
17-25 tents yellowish-brownish with I.
Rare; on the bark of Schiniis molle (pepper tree) at Mayfield.
Originally described from Northern Europe, but found throughout
the world.
3. ARTHOPYRENIA ANALEPTELLA (Nyl.).
Verrucaria analeptella Nyl. Flora, 363. 1872.
Thallus very thin, effuse, smooth, continuous, shining; whitish, to drab and oHve; brownish with KOH; CaCUOz — .
Apothecia not very numerous, minute, semi-immersed, flattened hemispherical, spreading basally; ostiolum depressed, dot-like; peri- thecium dimidiate, thick, black; paraphyses free, distinct, branched,
• 1 , . 14 — 18.5 . 16
twmmg; asci tubular to ventricose, jx, averagmg ^- jx;
52 - 73 58
spores in 2 rows in asci, bilocular, or with several false septa, sur-
- o
rounded by a halo, oblong-clli])soidal, -^ ix; no reaction with I.
10 — 22
Rare; on the bark of Platanus racemosa, in Stevens Creek Canon.
Common in Europe on smooth-barked trees; near Arthopyrenia
analepta (Ach.), but differing in the distinct paraphyses.
THE LICHEN FLOILV OF THE SANTA CRUZ PENINSULA 5 1
DIDYMELLA FALLAX Wainio.
Artho pyrenia fallax Nyl. Flora, 363. 1872.
This fungus, long considered to be a lichen, but which is apparently not a lichen as it seems to lack alga?, is common on the trunks and limbs of smooth-barked trees in the foothills. It may be recognized as follows: Thallus thin, rather effuse, forming smooth, whitish, cream-colored or pale olive patches; sprinkled with the minute, black, sub-globose, sessile or half-innate apothecia; paraphyses hair-like, free, simple and straight or branched and more or less twining; I — ; asci elongate, brownish with I; spores bilocular or becoming 4-locular,
cons'ricted at the middle, ^^^^^ ^^fi .
12^ — 20
4. ARTHOPYRENIA CINEREO-PRUINOSA (Schaer.) Jatta.
Verrucaria cinereo-pruinosa Schaerer, Spicilegia, 342. 1836. Artho pyrenia cviereo-pruinosa Jatta, S\ll. Lich. Ital. 529. 1900.
Thallus diffuse, thin, uniform, pale gray to olivaceous; KOH yellow; CaCl202 — .
Apothecia scattered, black, small to minute, more or less immersed and sessile, hemispherical or sub-globose; covered with the epidermal thalline layer, and hence ashy-pruinose, or naked; perithecium dimi- diate, black, thickish; paraphyses branched and twining, or simple;
I — ; spores bilocular, pointed and slender ellipsoid, ^? — ji-
12 - 15
On the bark of Umbellularia californica, in Stevens Creek Cafion.
The above seems to be a variety of this European lichen, characterized
by mailer spores than the type.
Section ACROCORDIA.
Apothecia solitary, hemispherical, globular, or conical; paraphyses permanent, hair-like, twining and net-like; asci cylindrical or ventri- cose, the spores of 2 similar cells arranged in a single row.
52 HERRE
5. ARTHOPYRENIA BIFORMIS (Borr.) Mull. Arg.
Verruciria hifonnis Borrer, Eng. Bot. Suppl, 2617,/. i. 1829. Arthopyrenia hiformis Miill. Arg.
Thallus effuse, thin, uniform or nearly so, whits to dark gray in color; KOH yellow brown.
Apothecia black, numerous, of medium size, hemispherical or
sub-globose, the base immersed, with a minute pore at the apex; peri-
thecium entire, thick, black; paraphyses abundant, distinct, long and
thread-like, branched and entangled; asci tubular, mostly sterile with
12 granular contents, yellowish with I; a fertile ascus measured — ix\
62
spores bilocular, pointed-ellipsoid, — — p..
18-24
On the twigs of an oak in the hills west of Los Gatos, altitude about 1500 feet. Much resembling Arthopyrenia gemmata, but differs in the entire perithecium. Referred here with doubt, the apothecia and spores being larger than in European specimens in my herbarium. The reaction with I is also different from that given by Leighton. A. iformis is found pret y generally over Europe and Noth America.
6. ARTHOPYRENIA CONFORMIS (Nyl.).
Verruca ia conformis Ny lander, Flora, 257. 1864; France. Verrucaria conformis Leighton, Lich. Fl. Grt. Brit. ed. 3. 463. 1879.
Tha lus effuse, thin and uniform, finally chinky abo t the apothecia; silvery gray in color; brown with KOH ; CaCljOj— .
Apothecia very numerous, small to minute, hemispherical, semi- immersed, black; perithecium dimidiate but extending so far basally as to be nearly entire; paraphyses long, very slender and hair-like; asci long, tubular; I-; spores usually in a single row, slender ellipsoid, bilocular, constricted at the middle; each cell very strongly constricted, so that the spores appear 4-locular, as if made up of four balls, the end ones being much smaller than the others; from falsely 2 or 3-septate,
sometimes distinctly 3-septate, — fi.
8^—16
On the bark of Pseudotsuga taxifolia, in the mountains. Recorded from France and the British Isles.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 53
V. Porina (Ach.) Mull. Arg.
Porina Acharius, Lich. Univ. 60. 18 10. Porina Mull. Arg.,
Thallus uniform crustaceous, upon or within the substratum, with Trentepohlia algae. Apothecia simple, scattered, with clear or dark, entire or dimidiate perithecium, with an apical pore. Paraphyses simple, free. Spores 6 to 8, ellipsoid, spindle-shaped and needle-like, bilocular or multilocular, colorless.
A genus of wide distribution, with about 150 species dwelling on bark and rocks, of which but one seems to occur here.
I. PORINA CARPINEA (Pers.) A. Zahlbr.
Verrucarla carpinea Persoon, in Ach. Meth. Lich. 120. 1803. Verrucaria carpinea Schaerer, Enumer. Crit. Lich. Europ. 221. 1850. Porina carpinea A. Zahlbr., Ascolichenes, 66. 1907.
Thallus thin, uniform, effuse or sometimes limitate, pale ashy or greenish brown to brown; KOH leaves a brown stain.
Apothecia numerous, scattered, minute, black, often shining or apically polished, rather prominent, hemispherical or sometimes sub- globose; the minute apical pore readily seen; perithecium dimidiate, the amphithecium pale; spores fusiform, bilocular to quadrilocular,
15-18
Common on various trees in the foothills and mountains. Com- mon over the temperate regions of Europe and America.
GYMNOCARPEtE.
A. Disk of the apothecia more or less open; paraphyses forming a net" work over the asci and holding a protruding sporal mass which con- ceals the disk, the mazaedium of authors ConiocarpinecB
A A. Paraphyses and spores not forming a mazaedium.
B. Apothecia linear, elongate, ellipsoid, or rarely circular.
Graphidinecs BB. Apothecia circular Cyclocarpinece
CONIOCARPINE^.
Thallus crustaceous, leafy, or fruticose, without rhizoids; algae Pleurococcus, Protococcus, Stichococcus, or Trentepohlia. Apothecia with a more or less open disk; asci usually cylindrical and soon dis-
54
HERRE
appearing, the ripe spores then forming with the paraphyses a protub- erant, powdery mass, the "mazaedium, " which remains a long time; spores 8, colorless or dark, globose and simple, or septate and ellipsoid or elongate.
KEY TO FAMILIES.
A. Thallus crustaceous, without cortex.
B. Apothecia on a stalk or stipe, with a proper margin. .Caliciacece BB. Apothecia sessile, with a proper or a thalline margin.
Cypheliaceae
A A. Thallus fruticose, with a cortex; apothecia sessile. .Sph(Brophorace(Z
CALICIACE^.
Thallus crustaceous, with Proto-, Pleuro-, and Stichococcus algae, Apothecia usually a circular or globose head lifted on a slender stipe, which is rarely branched or with more than one head.
KEY TO GENERA.
A . Apothecia urn-like, on a very short thick stipe, practically sessile.
IX. Sphinctrina A A. Apothecia on a long stipe.
B. Spores globose, simple VII. Coniocyhe
BB. Spores septate.
C. Spores bilocular; disk broad VI. Calicium
CC. Spores 4-8 locular; disk very narrow or dot-Uke.
VIII. Stenocybe
VI. Calicium (Pers.) DeNotrs.
Calicium Persoon.
Calicium DeNotaris, Giom. Bot. Ital. an. 2, pi. i. 309. 1847.
Thallus crustaceous, thin to obsolete, warty, or of dusdike or mealy granules. Apothecia stalked, with a circular or topshaped head, with a flat or convex disk; proper margin black or pruinose; spores bilocular or sometimes with an indistinct septum and apparently one-celled, constricted at the middle, smoky to blackish brown.
Occurring in all parts of the world on dead or rotting wood, on bark, and also on rocks.
I. CALICIUM POPULNEUM De Brond. Calicium populneum De Brond. in Duby, Bot. Gall. 2 : 638. 1830. Thallus white or grayish white, thin, uniform, effuse; KOH yellow;
1
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 55
Apothecia minute, black, the stipe very short; capitulum sub- cylindrical to top-like; paraphyses short, thread-like; asci not properly observed; spores ellipsoid, simple or apparently so, but really becom- ing bilocular, with a narrow, almost invisible septum, best seen as a darker portion of the spore when examined under a medium power
of the microscope, «.
12 — 16
Rare; forming rather shiny, whitish patches on the smooth bark of Quercus agrifolia at Devils Canon, altitude 2300 feet.
In the absence of authentic material for comparison referred here with some doubt, but in all probability correctly.
2. CALICIUM CURTUM (?) Turn. & Borr.
Calicium curtum Turner and Borrer, Lich. Br. 148. 1839. Calicium curtum Crombie, British Lichens, i : 93. 1894.
Thallus thin, granulose, or occasionally wanting, whitish to dusky or blackish gray.
Apothecia small to moderate, usually abundant, black, the stipe usually stout, quite short; capitulum at first sub-cylindrical, then flattened, until they are much like the lower half of a top or an urn; disk broad, plane, black, with a concolorous, narrow, erect, entire margin; disk and margin finally concealed by the extruded spore mass; margin very narrowly whitish pruinose at times; paraphyses branched,
5 - 6i thread-like, entangled; asci narrow, cylindrical, [i; spores
ellipsoid to oblong, bilocular, nearly always constricted at the middle,
II - I4i
On an old fence on the Stengel ranch, near the head of Alpine Creek, altitude 1400 feet.
While differing in several important respects from the published descriptions of C. curtum, our plant is nearer to it than to anything else.
A lichen widely distributed in both the northern and the southern hemispheres.
56 HERRE
VII. Conicybe Ach. Coniocyhe Acharius, Vet. Ak. Handl. 283. 1816.
Thallus crustaceous, powdery to obsolete; algae Protococcus or Stichococcus. Apothecia on a long stipe, the head becoming globose from the mazsedium; spores simple, globose, yellowish or nearly colorless.
Found on wood and bark, exposed roots, and rarely on stone.
I. CONIOCYBE FURFURACEA (L.) Ach.
Mucor furfuraceus Linne, Sp. PI. Ed. 3, 1655. 1764. Coniocyhe furfuracea Ach. Vet. Ak. Handl. 288. 1816. Coniocybe furfuracea Tuck. Gen. Lich. 243. 1872.
Thallus indeterminate, of loose, powdery, scurfy granules; greenish- yellow or sulfur-colored.
Apothecia small, on long, slender, weak stems, globose, concolorous; the stems dark, their color concealed by sulfur-colored powder; sporal mass globose, from yellow becoming pale brown; paraphyses short, thread-like, from simple becoming branched; asci short, slender, cylindrical; spores simple, colorless or very pale brown, spherical, 2.5 to 4.9 [x in diameter.
On roots and earth on high, overhanging, shady banks along the road above Congress Springs, at an altitude of 500 to 600 feet. Widely distributed over Europe and North America, usually on exposed roots.
VIII. Stenocybe Nyl.
Stenocyhe Ny lander, Bot. Not. 84. 1854..
Thallus nearly obsolete, or the apothecia upon a foreign thallus. Apothecia long-stipitate, scattered, black, with globose or pear-shaped head, the disk at first closed, then dot-like, with a black margin. Spores ellipsoid to spindle-shaped, 2—8 celled, large, dark. Four species, found in Europe, California, and Japan.
I. STENOCYBE MAJOR Nyl.
Stenocyhe major Nylander, Bot. Not. 84. 1854.
Stenocyhe euspora Nyl. with Crombie, Journ. Bot. 272. 1882.
Stenocyhe euspora Crombie, Brit. Lich. i : 97. 1894.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 57
Thallus very thin, of white or gray granules, or altogether absent; apothecia also growing upon the thallus of Cyphelium inquinans.
Apothecia not abundant, small, shiny, black, the stipe slender or sometimes robust; capitulum hke a smooth cylindrical club in shape; margin thin, entire, erect, concolorous; disk from dot-like becoming small, circular, plane; section of fruit light brown, becoming violaceous- reddish basally; paraphyses abundant, hair-hke, about ^ /i thick; asci elongate cylindrical, 12 — 15 X 190 — 230 n; thecium bluish with I; spores dark brown, pointed-ellipsoid to elongate spindle-shaped, mostly 4-locular, but also simple and then containing large oil drops,
9^ — 14 or bilocular, trilocular, 5- and 6— locular; {J-
36.7 - 5ii
On the bark of P seudotsiiga iaxifolia, mingled with Cyphelium
inquinans, at Devils Canon, alt. 2300 feet. Found on the bark of
various Coniferae in the mountains of Europe.
IX. Sphlnctrina E. Fr.
Sphinctrina E. Fries, Syst. Orb. Veg. 120. 1825. Sphinctrina Crombie, Brit. Lich. i : 83. 1894.
Thallus obsolete, the apothecia growing upon crustaceous bark lichens, usually some species of Pertiisaria. Apothecia sessile or very short-stipitate, pear-shaped to globose, shiny black, at first closed with a dot-like, deeply impressed disk; spores simple or rarely bilocular, from colorless becoming dark.
"O
I. SPHINCTRINA TUB^FORMIS Mass.
Sphinctrina ttibcpformis Massalongo, Mem. Lich, 155. 1853.
Sphinctrina tuhceformis Tuck., Gen. Lich. 241. 1872.
Calicium microcephalum Tulasne, Mem. Hist. Lich, 78, pi. 15, f. 20.
1852. Sphinctrina microcephala Nyl. Mem. Soc. Cherb. 5* 91. 1857. SphifKtriiia microcephala Crombie, Brit. Lich. i : 84. 1894.
Thallus wanting; apothecia minute, globose to top-shaped, sessile or almost stalked; color a very dark polished brown; at first closed; when open the disk is very small, concave or plane, bordered by the thick, entire proper margin; asci slender, long, cylindrical; paraphyses septate, thread-like, simple or slightly branched; thecium bluish with
58 HERRE
I; spores simple, globose to ellipsoid, arranged in a straight row in the
9—9!
asci, dusky when mature, — — fi-
9I - 14I
On the thallus of Pertusaria pustulata, growing on the bark of Quercus agrifolia, at Santa Cruz. As yet not seen elsewhere. Oc- curring on the thallus of various Pertusarias in Europe and North America.
The name given by Massalongo must supersede that of Tulasne, since the name microcephala had already been applied to a member of the same genus by Smith, Eng. Bot. 5' 138 (1795?) and Turner and Borrer, Lich. Brit. 130. 1839.
CYPHELIACE^.
But one genus occurs with us, with characters as follows:
X. Cyphelium Th. Fr.
Cyphelium Th. Fries, Gen. Heter. Europ. 100. 1861.'
Thallus warty-crustaceous or smooth, uniform or radiately lobed at the circumference. Apothecia innate to elevated-sessile, hemis- pherical or globose, at first closed, at last open and often crateriform. Proper margin varying from a mere trace beneath the hymenium to well developed, when it may be accompanied by a thalline margin. Paraphyses few, thread-like; spores in a single row, in our species simple or bilocular.
Usually occurring on dead wood, the bark of large old trees, rarely on stone.
KEY TO THE SPECIES.
Section Cypheliopsis A. Zahlbr.
Spores simple, more or less globose i. holanderi
Section Eucyphelium A. Zahlbr.
Spores bilocular, usually constricted at the middle. A. Thallus uniform.
B. Thallus yellow or greenish yellow 2. tigillare
BB. Thallus white or whitish 3. inquinans
AA. Thallus marginally lobed.
C. On rocks.
D. Thallus thick; spores large 4. calif ornicum
DD. Thallus rather thin; spores small or medium 5. Jarlowi
CC. On dead wood.
E. Thallus thick; whitish or ashy; KOH — 6. occidentalis
EE. Thallus brownish yellow; KOH decided yellow . .7. andersoni
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 59
I. CYPHELIUM BOLANDERI (Tuck.) A. Zahlbr.
Acolium bolanderi Tuck. Lich. Calif. 27. 1866.
Acolium bolanderi Tuck. Syn. N. Am. Lich. II: 136. 1888.
Acolium bolanderi Cummings, Williams, and Seymour, Lichenes
Boreali-Americana, no. 141, San Diego, Calif. Cyphelium bolanderi A. Zahlbr. Ascolichenes, 84. 1907.
Thallus orbicular, passing into indeterminate, often large, patches or sometimes with the thallus almost disappearing; of convex wart- like areoles, sometimes sub-lobate at the circumference, usually sparsely distributed, rarely contiguous; fertile warts becoming greatly enlarged, so that they seem to be foreign; color whitish, and light to dark ashy gray; the fertile areoles darker and somewhat tawny or even yellowish brown; sometimes nearly the whole thallus passes into the same color; a black hypothallus sometimes evident; KOH yellow,
Apothecia innate, crateriform, the disk plane, black, more or less gray pruinose, the proper margin obsolete; paraphyses long, slender comparatively few in number, simple; I—; asci small, cylindrical; hypothecium narrow, brown, underlaid by a broad black band, the remains of the proper margin; spores spherical, simple, 7.5 to 16 /i in diameter; specimens in Tuck. Herbarium from San Bruno Mountain have spores 5 — 12 and 8 — 13 n in diameter; spores becoming pro- truded in a black, columnar mass.
Abundant throughout on various rocks. Reported only from the coast region of central and southern California.
(Named for H. N. Bolander, Californian educator, collector, and botanist.)
2. CYPHELIUM TIGILLARE (Ach.) Th. Fr.
Lichen tigillaris Ach. Lich. Suec. Prodr. 67. 1798. Cyphelium tigillare Th. Fries, Gen. Heter. Europ. loi. 1861.' Acolium tigillare Tuck. Gen. Lich. 238. 1872. »
Thallus indeterminate, often spreading very extensively, of closely appressed, irregular, often coalescing areoles, more or less fissured and plicate, or reduced to small, scattered, sub-lobate warts; color bright to dusky greenish yellow; KOH—; CaCLOj — .
Proc. Wash. Acad. Sci., May, 1910.
6o HERRE
Apothecia numerous, small, innate, in swollen warts, the disk plane, black, the narrow, entire margin soon excluded; paraphyses short, slender, entangled; I—; spores bilocular, broad ellipsoid to sub- 9f — 14.8
Abundant on old fences and rails in the Bay and coast regions and in the foothills. Found on dry dead wood and bark of conifers throughout Europe and North America.
{tigillare, pertaining to beams, because found growing on old beams.)
3. CYPHELIUM INQUINANS (Sm.) Trevis.
Lichen inquinans Smith, Engl. Bot. 12: t. 810. 1801. Cyphelium inquinans Trevis, Flora, 4. 1862. Calicium tympanellum Ach, Meth. Lich. 89. 1803. Acolium tympanellum Tuck. Gen. Lich. 238. 1872. Acolium tympanellum Cummings, Seymour, and Williams, Decades of N. Amer. Lichens, no. 234, Yosemite.
Thallus effuse, of white or whitish granules or irregular warty nodules, forming a more or less continuous crust of moderate thickness, or thin and dispersed, or nearly disappearing; KOH— ; CaClzOj — .
Apothecia small or of moderate size, numerous, sessile but not appressed and often sub-stipitate, plane, black; the disk naked or often gray or whitish pruinose; the entire margin black, or white pruinose, at length concealed by the protruding sporal mass which stains the fingers; paraphyses slender, simple or branched and twin- ing, i^ to 2// broad; hypothecium a broad straight brown-black band;
6-7^ asci cylindrical, pL\ thecium more or less light blue with I;
35 -49 spores broadly ellipsoid or oblong, bilocular, often constricted at the
9f — 14
middle, «; spores of a specimen from H. Sandstede,
12^ — 22
16 Oldenberg, Germany, are — /"•
On the bark of Pseudotsuga taxifolia, Devils Canon, alt. 2300 feet. In North America found only in California, but occurring in Algiers and throughout Europe.
{inquinans, filthy.)
THE LICHEN FLOR.A. OF THE SANTA CRUZ PENINSULA 6 1
4. CYPHELIUM CALIFORNICUM (Tuck.) A. Zahlbr. Trachylia californicum Tuck.
Acolium californicum Tuck. Lich. Calif. 27. 1866. Acolium californicum Tuck. Gen. Lich. 237. 1872. Cyphelium californicum A. Zahlbr. Ascolichenes, 84. 1907.
Thallus orbiculate to effuse, thick, uniform, with plicate irregular surface and crenate, radiately lobulate margin; often sub-imbricate or else fissured and lobulate-areolate; color white, whitish, and brown- ish gray, the margin much lighter-colored than the central portion; the fertile warts not so enlarged proportionately as in Cyphelium bolanderi, but exhibiting the same tendency to become brown; KOH yellowish; CaCLOj — .
Apothecia from small becoming medium or large, innate, crateri- form, the disk broad, concave or plane, black; not pruinose in the specimens seen; thecium colorless, the paraphyses very long, slender, and intricately entwined; asci slender, cylindrical; underlaid by a thick brown-black band, the remains of the proper margin, of the following shape,
spores dark, bilocular, constricted in the middle, the sporoblasts
9f — 15 10 - 18
approximate, «; according to Tuck., — u.
i3i - 25 ^' ^ 18 - 25 '^
Common on rocks in the San Bruno Hills, at from 500 to 1000 feet
altitude. Collected but once elsewhere, among some specimens of
Lecanora pinguis, 50 feet above the sea at Point Lobos, San Francisco.
Specimens examined in the Tuckerman Herbarium, the herbarium
of Dr. C. L. Anderson, and the author's own material collected by
Bolander. Probably confined to the coast of California.
5. CYPHELIUM FARLOWI (Tuck.) Herre. Acolium farlowi Tuck, in Tuck. Herbarium (1885); Monterey. Acolium farlowi in Anderson Herbarium?
This lichen has not been positively identified as yet by me from our territory, but probably occurs along the north shore of Monterey Bay. It differs from Cyphelium californicum, to which it is closely related, in its smaller and thinner thallus and in the much smaller spores which,
7i - 9^
according to Tuckerman, measure u.
12-15
62 HERRE
Specimens in the Tuck. Herb, were collected by Dr. W. G. Farlow at Monterey, in August, 1885, growing on rocks.
In the herbarium of Dr. C. L. Anderson of Santa Cruz, are several specimens labelled Acolium farlowi which were collected by him on shale about Santa Cruz. Probably some of these were identified by either Tuckerman or Farlow and are therefore correct, but the one now in my possession is Cyphelium bolanderi.
Apparently very rare and local in its distribution. (Named in honor of the eminent professor of cryptogamic botany at Harvard Univer- sity, Dr. W. G. Farlow.)
6. CYPHELIUM OCCIDENTALIS Herre, new species.
Acolium sp. Hasse determ,, in Baker, Pacific Coast Lichens, No.
436, 1902; Stanford University.
Thallus determinate, forming rounded or oval patches, or effuse and spreading extensively; of rounded thick areoles, vdth their sur- faces made up of many small rounded nodules, the whole forming a chinky, deeply fissured crust of a whitish or ashy gray color; KOH - ;CaCl202-.
Apothecia innate in swollen warts, from small becoming medium or large; disk black; the thickish, entire, white margin concealed by the protruding spore mass, the surface of which with age often becomes tinged more or less with green; the long slender paraphyses broader than in the other members of the genus occurring with us; thecium bounded basally by a very broad, sub-crescentic brown- black band; asci rather slender, cylindrical; spores bilocular, strongly
9! "~ 15-9 constricted at the middle, oblong-ellipsoid, -^ jJ-
On old fences in the mountains and formerly in the foothills, but the fences on which it occurred there some years ago have since been replaced by wire fences.
The protruding spore mass marks the fingers with black when touched, as is also the case with some other members of the genus.
7. CYPHELIUM ANDERSONI Herre, new species.
Thallus orbicular, thickish, margin radiately crenate-lobed, fis- sured crustaceous, the smooth surface plicate and rugulose con-
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 63
torted, passing at the center into areoles; color a pale brownish yellow; KOH gives a decided yellow; CaCl202 — .
Apothecia innate in large thick warts of a deeper yellow than the thallus, strongly resembling those of C. bolanderi; disk broad, black, plane, bordered by the persistent, rather broad, white and conspic- uous entire margin ; paraphyses numerous, long, slender, i — 2V broad, serpentine; asci very slender, 4 — 5 /^ broad, curved or straight, cyHndrical, the tip slightly pointed; thecium underlaid by a rather narrow, convex, blackish-brown band; I — ; spores con- stricted at the middle, bilocular, broadly ellipsoid to oblong, 8|- II 137 - 20
Here described from two specimens collected on old redwood boards near Santa Cruz, by Dr. C. L. Anderson.
A very singular plant, with a thallus much like that of C. californi- cum but with different apothecia, spores, and chemical reaction.
I take pleasure in naming this for the veteran algologist of Santa Cruz, colleague of Asa Gray and Tuckerman.
SPH^ROPHORACEiE.
We have but one genus of the family.
XI. Sphaerophorus.
Sphcerophorus Persoon in Ust. Ann. Bot. 7: 23. 1794.
Thallus erect, bushy, brittle, with cylindrical or flattened branch- es; alga Protococcus. Apothecia terminal, in globose sweUings of the tips of branches, which are at first closed and later open by an irregular fissure at the tip.
I. SPH^ROPHORUS GLOBOSUS (Huds.) Herre.
Lichen glohosiis Hudson, Fl. Anglica, 1 : 460. 1762. SpharopJiorus globosus Herre, Proc. W. Acad. Sci. 7: 393- 1906. Lichen globiferus Linne, Mantissa, 133. 1767. Sphcerophoriis glohifertis De Candolle, Fl. Fr. 2: 327. 1805. Sphcerophorus globiferus Tuck. Gen. Lich. 231. 1872. Sphcernphoron coralloides Persoon, Usteri Annal. d. Bot. 1: 23. 1794.
64 HERRE
Thallus fruticose, tufted and shrub-like, erect, branched, terete, smooth, with short, fine, and very numerous terminal branchlets, these often in clumps which shatter off very readily; color silvery gray or whitish, but varying to brownish or a decided brown; rarely reddish. AUke on all sides; KOH — . Medullary layer densely cottony; violet with I.
Apothecia terminal, within the swollen and globular tips of the fertile branches, which split open, exposing the globose apothecia; spores violet-black, simple, spherical, 7 — lo /< in diameter.
On trees, dead wood, and sandstone. On the Pacific side of the peninsula occurring from near sea-level to the summit of the range, but not descending on the Bay side more than a few hundred feet, remaining within the limits of the redwood and spruce forests. Occasionally found in great abundance. A strikingly handsome plant. Described from Europe and recorded from Arctic and cool temperate America, Madeira, and Australasia.
GRAPHIDINEiE.
Thallus crustaceous, uniform, without cortex or with a cortex on the upper side; or fruticose, erect or decumbent, with an evident cortex and medulla; alga Palmella, Trentepohlia, Phycopeltis or Phyl- laclidium.
Apothecia compressed, more or less linear, with small, fissure-like disk, are characteristic of the group. Proper margin well developed or absent; sometimes with a thalline margin; innate or sessile, rarely on a short stipe. Spores colorless or dark, variously shaped, but more often spindle or needle-shaped, and multi- or murilocular.
KEY TO FAMILIES.
A. Apothecia without margin ArthoniacecB
A A. Apothecia with margin. B. Thallus crustaceous.
C. Without cortex GraphidiacecB
CC. Cortex present on upper side DirmacecB
BB. Thallus fruticose, erect or decumbent Roccellacem
ARTHONIACEiE.
Thallus uniform crustaceous; gonidia of Palmella, Trentepohlia, or Phyllactidium algae. Apothecia forming irregular dots, lines, or spots, simple or branched; single or forming a stroma, without proper margin. But one genus occurs with us.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 65
XII, Arthonia.
Arthonia Ach. Lich. Univ. 25. 18 10. Arthonia A. Zahlbr., Ascolichenes, 89. 1907.
The alga is Trentepohlia. Apothecia innate, surrounded by the thallus, circular, dot-like, or irregularly stellate, or more or less elon- gate; spores variously shaped, oval, ellipsoid, wedge-Hke, or needle shaped; bi- or multilocular, the cells often of unequal size; colorless or rarely brown.
About 500 species, mostly of the warmer parts of the earth, on bark and rocks.
KEY TO SPECIES.
Thallus pinkish or reddish when wet; spores 4-locular punctiformis Thallus not changing color when wet; spores 4— locular radiata
I. ARTHONIA PUNCTIFORMIS Ach.
Arthonia punctiformis Ach, Lich. Univ. 141. 1810.
Arthonia punctiformis Willey, Synopsis Genus Arthonia, 41. 1890.
Thallus white or whitish, forming more or less determinate spots or blotches, thin, uniform, sometimes evanescent; when moistened becoming pinkish or reddish; no reaction with chemicals.
Apothecia small or minute, slightly elevated, rounded, oblong or irregular, black; epithecium granulose, blackish; thecium pale, the asci pear-shaped; reaction with I first bluish, than wine-red; accord- ing to Hasse "indigo-blue"; spores oblong-ovoid, 4-to 6-locular,
4-6 12 - 19^
Found fertile on Vaccinium on White Horse Creek, and on Alniis on the Gazos Creek, near the sea coast; I also refer here a sterile lichen which is abundant on the stems of Baccharis pillularis and other shrubs. A lichen of Europe and North America.
2. ARTHONIA RADIATA (Pers.) Th, Fr.
Opegrapha radiata Persoon, Ust, Ann. 29. 1794.
Arthonia radiata Th. Fr. Lich. Arctoi, 240. i860.
Arthonia radiata Willey, Synopsis Genus Arthonia, 44. 1890.
Thallus thin, uniform or shghtly scaly, determinate, often limited by a black hj^Dothalline line; color, various shades of gray to olive and fawn-color; no reactions with KOH or CaCl202.
66 HERRE
Apothecia numerous, linear, very irregularly shaped, branching, stellate, curved, or straight; innate, with sunken disk, or becoming somewhat elevated; brown to black; epithecium granulose, blackish; thecium colorless, the asci short, pear-shaped or almost oval; blue with I, then slowly turning to vinous red, only the asci stained
yellowish ; spores ovoid-oblong, 4-locular, -^ ^ and ^ fJ-
Abundant on trees throughout; an exceedingly variable plant, found all over the world.
I have referred to this species a large series of Arthonias which vary in several particulars, but which agree pretty well in spore characters.
GRAPHIDACE^.
Thallus uniform crustaceous, cortex absent or poorly developed, with Palmella or Trentepohlia alga, in our species only the latter. Apothecia usually linear, rarely circular or spot-Uke, sohtary, or aggregate, but not forming stroma; simple or branched, with a well developed proper margin, frequently with a thalhne margin; disk usually narrow and fissure-hke; paraphyses simple and unbranched or branched and twining; rarely gelatinizing. Spermatia exobasi- dial.
KEY TO GENERA.
Paraphyses branched, twining XIII. Opegrapha
Paraphyses simple, not twining XIV. Phceographis
XIII. Opegrapha.
Opegrapha Humboldt, Fl. Frib. 57. 1793.
Alga Trentepohlia. Apothecia innate, appressed or sessile, usu- ally more or less elongate, with a black proper margin; hypothecium clear or dark; spores 8, oval, elhpsoid, or spindle-shaped, straight or bowed, colorless, 2-18-locular.
Numerous species, on rocks, wood, and bark, distributed all over the world.
KEY TO THE SPECIES.
Spores 6-locular 3. varia
Spores 4-locular
On maritime rocks i . saxicola
On bark of trees 2. prosiliens
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 67
I. OPEGRAPHA SAXICOLA Ach.
Opegrapha saxicola Ach. Syn. Meth. Lich. 71. 1814. Opegrapha saxicola Stizenberger, Ueber die Stcinbew. Opegr.- Arten, 23. 1865.
Thallus effuse and very thin, or almost entirely disappearing, of minute whitish, gray, or yellowish granules.
Apothecia thickly or sparsely scattered, mostly bluntly ellipsoid, or nearly circular, or compressed and sub-linear; straight or crooked; dead black; margin at first thick and rounded, later thin, sharp- edged, the disk then broadly visible; epithecium granulose, blackish brown (reddish brown according to Stiz.); hypothecium broad, dark brown; thecium colorless, slightly wine-red with I; paraphyses threadlike, with tips not at all or very slightly thickened; asci oblong or sub-clavate; spores 4-locular, ellipsoidal or ovoid, straight
6 — 10 or rarely slightly curved, with a more or less evident halo, ,u;
according to Stizenberger they are "wasserhell bis braun, 20 - 30 mik. lang und 5-8 mik. dick. "
Spermogonia abundant, appearing as small or minute black dots; spermatia small, short, 3.5 - 4 A< long.
Abundant on sandstone near the old CUff House, San Francisco; originally described from Europe, where it is widely distributed.
Our plant is a variety, differing in several respects from typical 0. saxicola.
2. OPEGRAPHA PROSILIENS Stirton.
Opegrapha prosiliens Stirton, Grevillea, 3 : 36. 1874.
Opegrapha prosiliens Leighton, Lich. Fl. Grt. Britain ed. 3, 403. 1879.
Opegrapha prosiliens Hasse, Bull. So. Calif. Acad. Sci. 5:42. 1906.
Thallus white, whitish, yellowish white, and yellowish-greenish, thin to very thin, becoming almost or entirely obsolete, forming small, variously shaped distinct patches, or effuse and spreading indefinitely; KOH — ; CaCl202 — .
Apothecia black, prominent, straight or curved, narrowly oblong or ovoid, the margin broad, rounded, or rarely thin and erect, when the disk is visible; disk ordinarily but a crevice; epithecium granu- lose, dark brown; hypothecium from very dark brown merging into black; paraphyses very slender, much branched and entwined, their
68 HERRE
tips clear or slightly darkened; bluish with I; asci subcylindrical or
12? — 22
clavate, usually with a halo, at least at the upper end, r ,«;
spores ellipsoid-spindle shaped, or slipper-shaped, blunt, straight,
4 — locular, the third cell usually the largest, surrounded by a;
6-9f-ii ''6-7"
broad gelatinous halo, -— /^; — //, Stirton. Sper-
20 — 28 20—28
mogonia numerous, forming minute black specks over the thallus
1 _ i
"5 — 2
spermatia straight, very slender, — P-
42 — 73
On twigs and trunks of Alnus, Gazos Creek, alt. 75 feet, and on dead twigs of Lupinus arbor eus at Pescadero Point, 10 to 25 feet alt. ; on Quercus agrifolia at Laguna Creek, and on bark of Pseudo- tsuga taxijolia at Santa Cruz. Dr. Hasse has also collected it on Jug- lans in the Santa Monica Range near Los Angeles.
The reference of our material to Stirton's plant may be incorrect, but it agrees with no other described in the accessible literature.
Specimens in the Anderson Herbarium at Santa Cruz, marked Opegrapha varia seem to be this same species.
XIV. Phaeographis Miill. Arg.
Phceographis Miill. Arg. Mem. Soc. Phys. et Hist. Nat. Geneve, 29: 1887.
Thallus uniform crustaceous, with Trentet>ohlia alga. Apothecia innate, appressed, or sessile, usually linear, simple or branched; disk usually narrow, fissure-like; proper margin and hypothecium black to colorless; hymenium gelatinizing, not turning blue with I; spores dark, spindle-shaped or caterpillar-like, bi-locular to many-celled.
About ICO species, mostly on bark in the warmer parts of the earth, represented with us by but one species.
I. PH^OGRAPHIS INUSTA (Ach.) Mull. Arg.
Graphis inusla Ach. Syn. Meth. Lich. 85. 1814; Canada. Graphis imista Leighton, Lich. Flora Grt. Brit. cd. 3, 431. 1879.
Thallus determinate, thin, uniform, smooth or slightly wrinkled, yellowish white to white; with KOH yellowish brown.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 69
Apothecia numerous, rather large, slightly sunken, variously and
irregularly shaped, curved, wavy, often branching, much resembling
characters in Oriental script; disk broad, plane, brownish-black to
black, more or less pruinose; thalline margin white, mostly entire,
somewhat overhanging, seemingly burst out from below; proper
margin blackish brown, merging into the clear or very pale yellow
hypothecium; paraphyses not very abundant, slender, simple, free,
pale blue with I; asci elongate-clavate or sub-cylindrical; spores
brown, 8-celled or rarely 6-or 7-celled; mature spores at first pale
brown, broadly ellipsoid or oblong, often pointed at one end, viola-
II - 12^ ceous with I, the septa rather indistinct, ; n :; " /<; later be-
31-^ — 51-4
coming darker, thick- walled with heavy septa, caterpillar-like in 7* - 9f
A single specimen collected on Alnus, on the Gazos Creek, with Opegrapha prosiliens.
Widespread in the tropics; occurring also in Great Britain, France, Germany, Japan, and temperate America.
DIRINACE^.
Thallus uniform crustaceous, corticated; alga Trentepohlia. Apo- thecia circular or elongate, with both proper and thalline margin; h^-pothecium thick, black; spores spindle-shaped, 4 to 8-locular, colorless or brown.
XV. Dirina E. Fr.
Dirina E. Fries, Syst. Orb. Veg. 244. 1825.
Cortex of unseptate hyphae arranged perpendicularly to the cor- tex. The proper margin thin, the thalline margin thick; paraphyses simple, unbranched; spores 8, colorless.
A genus of few species, on maritime rocks and on bark of trees near the sea.
I. DIRINA FRANCISCANA A. Zahlbr.
Dirina frajiciscana A. Zahlbr., in Herre Botanical Gazette, 43: 270, 1907.
Thallus effuse, of thick, rounded, irregular tuberculate areoles, uniform crustaceous, sub-cartilaginous, forming heaped patches.
yo HERRE
Color varying from yellowish or brownish yellow to a dingy ashy gray, the last most common; KOH — ; CaCl202 — .
Apothecia numerous, of medium size, rounded elevated, sessile; surface of disk minutely granular, ashy-gray pruinose; the thalline margin thick, prominent, white, obtuse, soon flexuous, often intri- cately so; epithecium dark, 56/^ high; hypothecium black, thick, 140/^ high, blue or bluish with I; paraphyses typical, thecium wine-red
70 — 134.5 with I; asci clavate, straight or curved, long stalked, 15 g _ 22 a '"'
spores fusiform, straight or slightly curved, quadrilocular,
5-8.5
23-8 - 33-5 '"
On rocks 50 to 75 feet above the sea at Point Lobos, San Fran- cisco, growing with Dendrographa minor.
Near Dirina repanda of Europe and Northern Africa, but with a thicker and differently colored thallus and with different spores.
ROCCELLACE^E.
Thallus fruticose, erect or decumbent, attached to the substratum by a holdfast; with distinct cortex and medulla; alga Trentepohlia. Apothecia circular and linear, innate or sessile.
A maritime family, on trees and rocks, mostly found on tropical coasts. Some species furnish a valuable dye.
XVI. Dendrographa Darb.
Dendrographa~Ddir:h\s)\\xt, Ber. der Deutsch. Bot. Gesellsch. 13: 313. 1895; Monograph. RoccelL, Bibliotheca Botanica, 45: 1898.
Thallus erect or decumbent, tufted or matted; filaments terete or compressed basally, branched, with infrequent lateral soredia; color gray. Apothecia lateral, circular, the disk black, white pruinose; spores quadrilocular, colorless, spindle-shaped or shghtly curved.
On maritime shrubs or on rocks and earth, on the coast of Cali- fornia.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 7 I
I. DENDROGRAPHA MINOR (Tuck.) Darb.
Roccella leucophcea var. minor Tuck.,
Dendrographa minor Darbishire, Ber. der Deutsch. Bot. Gesellsch.
16: 13. 1898. Dendrographa minor Herre, Proc. Wash. Acad. Sci. 7: 393. 1906.
Thallus erect or more often lax and decumbent, tufted; terete and hair-like or slightly flattened below, much and intricately branched, forming dense tangled clumps; color gray, or basally blackening. Sterile. Large globose lateral soredia sparingly pres- ent, these apparently taking the place of apothecia.
Abundant on rocks and earth 50 to 100 feet above the sea near Golden Gate, San Francisco; collected on rocks at Mission Dolores by Bolander, but now extinct there. Fertile specimens collected on rocks at Monterey by Dr. W. G. Farlow, in 1885. On high bluffs at Tomales Bay, Marin County, Bolander in Tuck. Herb. Re- corded from the islands of Lower CaHfornia by Dr. Hasse.
Specimens in the Tuck. Herbarium are marked "A R. leucophcea vix diver sa."
CYCLOCARPINEiE.
Thallus from the simplest uniform crust to the highest foliaceous or fruticose form; in the crustaceous forms fastened to the substra- tum by the hj^Dhse of the h^^othallus or the medulla, in the other forms usually by rhizoids, holdfasts, or an umbiHcus. Cortex absent in most crustaceous forms, or variously developed, on the upper side or on both sides. Algas of various families, Protococcus, Pleiiro- coccus, Palmella, Trenlepohlia, Gloeocapsa, Nostoc, Scytonema, Stig- onema, Calothrix and Rivularia.
The apothecia are usually disk, shield, or plate-like; sometimes they are urn-like or globose with a very narrow or minute disk and immersed in the thallus so that they resemble the Pyrenocarpeae. The apothecia vary from innate and sessile to stalked, in some forms the stalk resembling a fruticose thallus, the podetia of authors. A proper margin usually evident, sometimes lacking; when formed of hyphai which enclose no algae, soft and nearly or quite colorless, it is biatorine; when formed of the thallus, black and coal-Hke, it is lecideine; when formed of the thallus and enclosing algas, it is lecan- orine. Hjpothecium variously colored, clear to black; paraphyses
72 HERRE
very diverse, but no mazaedium is formed ; asci permanent, one to many spored. Spores simple, or two to many celled, polar-bilocular, and muriform, colorless to dark brown, sometimes with a halo. Soredia often greatly developed and in some families of the highest importance.
KEY TO FAMILIES.
A. Spores typically bi-locular or polar-bilocular, with much thickened walls, the cells often connected by a thin tube or canal. B. Spores colorless, polar-bilocular.
C. Thallus uniform crustaceous or with a radiately plicate margin,
without cortex Caloplacacece
CC. Thallus f ohaceous or f ruticose, corticated Theloschistacecs
BB. Spores brown, polar-bilocular or with a septum.
D. Thallus crustaceous, uniform or lobed at the circumference,
without cortex BuelliacecB
DD. Thallus foliaceous or fruticose, corticated Physciacece
A A. Spores simple, multilocular, or muriform, colorless or rarely brown. E. Algae belonging to the Cyanophycea; thallus more or less gelatinous when wet.
F. Algse Nostoc; apothecia sessile, dish-like CollemacecB
FF. Algae not Nostoc. G. Algae Scytonema or Stigonema; apothecia urn or dish-like.
EphehacecB GG. Algsd Gloeocapsa; apothecia often not visible, apparently pyre-
nocarpous, urn or dish-like PyrenopsidacecB
EE. Thallus not gelatinous when wet. H. Thallus crustaceous, uniform or marginally lobed, attached by the hyphas of the hypothallus or medulla.
/. Thallus with Trentepohlia algae Lecanactidacece.
II. Thallus with Pleurococcus or Palmella algae. /. Asci many-spored; apothecia lecideine, biatorine, or lecanor-
ine AcarosporacecE
JJ . Asci 1-8 spored, or rarely 16-32 spored.
K. Apothecia with a proper margin, not enclosing algae.
LecidacecB KK. Apothecia with a thalline margin, enclosing algae. L. Proper margin well developed, black or dark, usually entire; apothecia innate; thalline margin poorly devel- oped DiploschistaceoB
LL. Proper margin wanting or poorly developed and clear; thalline margin well developed.
M. Apothecia sessile, the disk large LecanoracecB
MM. Apothecia sohtary or several, enclosed in thalline
warts; disk usually very small PertusariacetB
HH. Thallus usually foliaceous or fruticose, or occasionally reduced to squamules.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 73
iV. Thallus fruticose, erect or decumbent, structure radial, corti- cated, with a holdfast Usneacea
NN. Thallus foliaceous or squamulose. 0. Thallus large, attached by a central umbilicus . .Gyrophoracea
00. Thallus not umbilicate.
P. Apothecia long-stalked, the podetia simple or branched, naked or covered with leafy squamules. , . Cladoniacecs PP. Apothecia not long-stalked.
Q. Apothecia adnate by their entire under surface to the
foliaceous thallus PeltigeracecB
QQ. Apothecia sessile or elevated sessile.
R. Medulla lacking or feebly developed; thallus mostly of pseudoparenchyma; aXg^e Scytonema. Ileppiacece RR. Medulla well developed.
S. Spores spindle-shaped, multilocular; under side of
thallus usually with cyphellae SlictacecB
SS. Spores oval or ellipsoid, simple or rarely 2-4 locu- lar; thallus never with cyphellae.
T. Alga Scytonema PannariacecB
TT. Alga Pleurococcus or Palmella; apothecia lecan- orine Parmeliacea
LECANACTIDACEiE.
Thallus uniform crustaceous, without cortex, alga Trentepohlia. Apothecia circular, sessile or innate ; proper margin lacking, or rudi- mentary, or occasionally well developed; with or without thalline margin. Paraphyses branched, more or less entangled and twining. Represented with us by only one genus.
XVII. Lecanactis (Eschw.) Wainio.
Lecanactis Eschweiler, 4, Syst. Lich. Gen., 14. 1824, in part. Lecanactis Wainio.
Apothecia circular, lecideine, with black proper margin; a thalline margin wanting; hypothecium black, merging into the margin; asci 4 — 8 spored; spores colorless, elHpsoid, spindle or needle-shaped, 2 — 16 locular.
Rock and bark lichens, mostly of the warmer regions.
1. LECANACTIS ZAHLBRUCKNERI Herre.
Lecanactis zahlhrnckneri Herre, Botanical Gazette, 43: 270. 1907.
Thallus effuse, of small, irregular (sometines plicate) squamules,
which at first are scattered but soon become a thick, uniform, tarta-
74 HERRE
reous crust. Color a more or less evident rose-pink which soon fades out in herbarium specimens, leaving them whitish or ashy gray. Apothecia small, round, sessile, becoming convex; black, the disk gray pruinose, but eventually naked; the proper margin prominent but finally excluded. Epithecium dark or black, thick, 45 // to 50 // high, blue with I; hypothecium black, broad, 42 to 60 /^ high; para- physes typical; thecium brick- or vinous-red with I; asci clavate,
straight or curved, sometimes pointed at tip, '- fi; spores 8,
78 — 106
5 ~7
colorless, fusiform, straight or slightly curved, 4 — locular, — f^-
19.6 — 28
Rare on maritime rocks, 50 to 75 feet above the sea, at Point
Lobos, San Francisco; associated with Dendrographa minor, Artho-
pyrenia halodytes, and some species of Trentepohlia.
2. LECANACTIS CHLOROCONIA Tuck.
Lecanaciis chloroconia Tuck. Trans. Am. Phil. Soc, p.
Lecanactis premnea h. chloroconia Tuck. Syn. N. Am. Lich. II:
115. 1888.
Thallus small, thin, uniform, smooth to granulose; definite and limited by a black hypothalline line, or this obsolete, and the thallus diffuse; yellowish-greenish to yellowish-ashen; KOH yellow; CaCl202 — .
Apothecia small to medium size, circular, sessile, black; the disk pruinose or finally naked; the proper margin erect, rather thin, mostly entire, becoming somewhat angulose or wavy; epithecium granulose, greenish-blackish; hypothecium broad, black, continuous with the broad black margin; paraphyses branching or simple, free, their tips thickened and dark green; thecium colorless, turning wine- red with I; spores mostly 4 — locular, very rarely 3 — or 5 — locular, finger-shaped or broadly spindle-shaped, straight or slightly curved,
3¥ - 4.9 II — 15 and rarely 17
On bark oiAlnus, in tiny patches mixed with Opegrapha prosiliens, on Gazos Creek, near the Pacific.
A tropical lichen of wide distribution and occurring also in Europe and over a great part of North America.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 75
DIPLOSCHISTACEiE.
Thallus uniform crustaceous, corticated, attached to the substra- tum by the hyphai of the hypothallus or of the medulla; alga Proto- coccus. Apothecia circular, sunken in the thallus or appressed, with crater-like, or finally flat disk; proper margin well developed, entire, or only laterally developed; spores two- to many- celled or muriform,
XVIII. Diploschistes Norman.
Diploschistes Norman, Con. Prae. Nov. Gen. Lich. 20. 1852.
Thallus as above, as are the apothecia. Proper margin well developed, black or clear; paraphyses simple or branched at the tips; asci thin-walled, with 4 — 8 spores; these muriform, dark.
KEY TO THE SPECIES.
Apothecia medium-sized, from pit-like becoming open and plane with broad disk, black or gray pruinose i. scruposus
Apothecia minute, immersed, opening by a pore surrounded by a radiately striate margin 2. actinostomus <
I. DIPLOSCHISTES SCRUPOSUS (L.) Norman.
Z,/c/?e« 5cm/?05«5 Linne, INIantissa, 2: 131. 1771.
Diploschistes scruposus Norman, Con. 1852.
Urceolaria scriiposa Ach. Meth. Lich. 147. 1803.
Urceolaria scniposa Tuck. Syn. North Am. Lich. I: 222. 1882.
Thallus determinate to effuse, of rough, irregularly shaped, some- times contorted and pUcate, chinky or fissured areoles, forming a thick and hard or more or less crumbly and mealy crust; KOH — ; CaCl.Oj reddish; color various shades of gray, whitish, and ashen; rarely gray-brown and rusty.
Apothecia numerous, from small and immersed becoming superficial and medium-sized or large; from deep and pit-like finally open, plane, with broad black disk, often gray pruinose; the thalline margin thick, swollen, from entire to more or less rugose ; the proper margin arched, blackish, more or less denticulate; sometimes the thalline margin disappears, when the proper margin becomes prominent, thickened, and exceedingly fine, intricately entwined, transverse striae become visible; paraphyses slender, branching,
Proc. Wash. Acad. Sci., May, 19 10.
76 HERRE
their tips enlarged, brown; spores 4 — 8 in the asci, variously
1- J 10 ~ 15-5 shaped, ^-^ n,
26 — 41.5
Abundant throughout; occurring on various kinds of rocks, earth, the bases of old tree trunks, and the thallus of Cladonia pyxidata.
When the thallus is thick, soft, and crumbly or mealy, it forms the variety or species gypsacea of various authors. When it is on Cla- donias and mosses it is the subspecies bryophila; sometimes in this habitat it grows without a thallus and is then variety parasitica Sommerf.
All of these variations occur with us, as well as some others, but none of them depart widely enough from the average form to merit special description. Several varietal forms may be secured from one extensively spreading patch covering an irregular rock mass, where different conditions of light and moisture may affect the growth of different portions of the thallus.
Widely distributed in both the north and the south temperate zones.
2. DIPLOSCHISTES ACTINOSTOMUS (Pers.) A.Zahlbr.
Urceolaria actinostoma Persoon, in Ach. Lich. Univ. 288. 18 10. Urceolaria actinostoma Tuck. Syn. North Am. Lich. I: 222. 1882. Diploschistes actinostomus A. Zahlbr. Ascolichenes, 122. 1907.
Thallus of smooth, thick, closely compacted, angular or difform areoles separated by deep cracks and fissures; the whole forming a dense, determinate, suborbicular crust; rarely the crust is thin and indeterminate; color whitish, gray, mouse-colored, dusky, and in one specimen collected, black; the margin usually much paler and a white hypo thallus more or less evident; KOH — ; CaCl202 faint reddish.
Apothecia numerous, immersed, very small, opening at the surface by a minute pore, which is surrounded by an at length fully visible, radiately striate or stellate proper margin; said by authors to be gray pruinose, but not so with us; epithecium deep blackish-brown; para- physes thread-like, densely entangled; spores 3, 4, and 8 in the asci, variously disposed, ovoid to broadly ellipsoid, from colorless turning dusky with age, then dark brown and much shrunken and misshapen; 12 — It;
• ^~ PL.
18 — 26.9
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 77
A handsome lichen common on rocks in the foothills. The fruit superficially resembles that of the Verrucariaceod. Found in the temperate parts of Europe and North America.
LECIDEACE^.
Thallus crustaceous, horizontally outspread, uniform or margi- nally lobed, fissured, areolate to scale-hke or squamulose, without rhizoids; alga Protococcus; with or without a poorly developed cor- tex. Apothecia circular, sessile upon the thallus or rarely innate or elevated; the proper margin clear or black, the apothecia without thalline margin and without algae; hypothecium colorless to black; paraphyses rarely branched, their tips usually more or less thickened; asci usually with 8 spores, in some genera i — 8, rarely i6 — 32; spores colorless or brown in one genus, simple, to plurilocular or muriform.
KEY TO GENERA.
Thallus with a cortex; spores 2-8 locular XXII. Toninia
Thallus without cortex.
Spores simple, colorless XIX. Lecidea
Spores not simple.
Spores muriform, brown XXIII. Rhizocarpon
Spores not muriform, colorless.
Spores bilocular XX. Catillaria
Spores 4-16 locular XXI. Bacidia
XrX. Lecidea (Ach.) Th. Fr.
Lecidea Ach. Meth. Lich. 32. 1803.
Lecidea Th. Fries, Gen. Heterol. Europ. 88. 1861.
Lecidea A. Zahlbr. AscoUchenes, 130. 1907.
Thallus crustaceous, varying from uniform to lobate at the cir- cumference, and from small areoles to warty, scale-like, or more or less leafy squamules, without rhizoids, not corticated or with a thin cortex, naked or sorediose; alga Protococcus. Apothecia circular or angulate or irregular from lateral pressure, innate, sessile, or ele- vated-sessile, the proper margin not enclosing algaj and varying from clear to black; hypothecium colorless to black; paraphyses un- branched; spores 8 or rarely 16, small, simple, colorless, usually ellip- soid, ovoid, or narrow-oblong, straight or slightly curved, j
78 HERRE
About 500 described species, principally inhabitants of cold or temperate regions. The typical representatives of the genus are especially abundant on igneous or metamorphic rocks in alpine or cold regions, and are among the most difficult of plants to satisfac- torily determine.
KEY TO THE SPECIES.
I. Section Psora : — Thallus squamulose, more or less lobed.
A. Squamules brick-red or reddish-flesh color; KOH-I-. .1. decipiens A A. Notred;KOH— .
B. Thallus light brown to chestnut; of small to medium-sized squa- mules 2. gloUfera
BB. Squamules minute; dull brown-black 3. scotopholis
II. Section Biatora: — Thallus uniform, granulose or much reduced;
apothecial margin clear or colored, but never black. a. Scales large, more or less lobate ; apothecia large . 4. granulosa phyllizans aa. Areoles quite small; apothecia small or minute 5. coardata
III. Section Eulecidea : — Thallus uniform ; apothecial margin black.
A. Thallus absent 17. auriculata diducens
A A . Thallus usually well developed. B, Thallus brown or blackening. C. Thallus forming small isolated patches among other Hchens
7. intumescens CC. Thallus indeterminate, spreading.
D. Hypothecium colorless or slightly brownish; no reaction
with 1 8. manni
DD. Hypothecium dark; I-f
E. Thallus brown to dark brown or nearly black; squamules
flat to concave 6. fumosa
EE. Thallus fawn-colored; squamules convex to sub-globose
9. atro-lulescens BB. Thallus yellow, gray, whitish, or white. F. White or nearly so.
G. On bark; KOH yellow; CaCl202— 18. melancheima
GG. On rocks.
H. Thallus thin or very thin, marked by tortuous black lines
II. criiciaria HE. Thallus of crumbs or scales, without tortuous black
lines 16. goniophila
FF. Thallus gray, grayish, or yellowish. /. On bark or dead wood.
/. Thallus gray, ashen, or whitish; CaCUOj— ... 20. parasema J J. Thallus olive or yellow to gray and whitish; CaCLOj
brick-red 19. olivacea
II. On rock.
K. Hypothecium black or black-brown.
Thallus ashy or dusky gray 10. grisella
KK. Hypothecium pale to brown.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 79
L. Thallus yellow or sulfur -colored; hypothecium dark
brown 23. enter oleuca theioplaca
LL. Thallus whitish gray, ashen, or bluish-gray. M. Yellow with KOH.
N. Hypothecium thick, brown. Hymenium blue, then
brown with 1 21. latypcea
NN. Hypothecium colorless to dusky; hymenium
blue with 1 22. enteroleuca
MM. Thallus without reaction to KOH.
O. Without evident h>pothallus, ash-colored or leaden
gray; apothecia small 15. platycarpa
00. A black hypothallus more or less evident. Apo- thecia from small or medium to very large. P. Apothecia more or less gray pruinose; thallus
ash-colored 12. lithophila
PP. Apothecia not pruinose. Q. Thallus ashy-gray or whitish. .. . 13. tessellata QQ. Thallus bluish-gray or gray ... 14. lapicida
I. LECIDEA DECIPIENS (Ehrh.) Ach.
Lichen decipiens Ehrhart, Hedwig Stirpes Crypt. 2:7. 1789.
Lecidea decipiens Ach. Meth. Lich. 80. 1803.
Biatora decipiens Tuck. Syn. N. Am. Lich. II: 13. 1888.
Thallus of scattered to crowded, rather thin, appressed, smooth scales of medium size; often shield-shaped or round-lobed and cre- nate; more or less concave or furrowed, sometimes plicate; of a bright reddish flesh-color, brick-red, or darkening; the margin more or less white-edged; beneath white; with KOH the thallus is first rose pink, than a plum color; CaCl202 — .
Apothecia small to rather above medium size, usually marginal but also occurring in the middle of a scale, closely adnate, from cir- cular becoming angular in shape; the black disk soon convex; gener- ally the small, paler margin is hardly visible and soon entirely disap- pears; occasionally it is white and persistent; paraphyses conglu- tinate, their tips umber; hypothecium clear to pale brown; asci narrowly clavate; thecium blue with I; spores ellipsoid or ovoid,
S - 7
-/I.
9.5 - 16
On earth in lime rock crevices, near the summit of Black Moun- tain, at an altitude of 2700 feet, mingled with Toninia caeruleo-nigri- cans and Dcrmatocarpon hepaticum.
So HERRE
A common xerophyte of calcareous earth throughout Europe and North America; in this country most abundant in the western half.
2. LECIDEA GLOBIFERA Ach.
Lecidea glohifera Ach. Lich. Univ. 213. 1810; Switzerland. Biatora glohifera Tuck. Syn. N. Am. Lich. II: 10. 1888.
Thallus of small or medium size, thickish, rigid, rounded and cre- nate lobate, ascendant and imbricate scales; their surface usually polished and often concave; color pale or whitish brown and greenish to various shades of chestnut, the under side white; KOH — ; CaCl202 -.
Apothecia numerous, small to medium size, marginal, elevated- sessile, black or brown-black, moderately convex to sub-globose, soH- tary and circular or often confluent and irregular; margin visible only on young apothecia; white within to the naked eye; paraphyses conglutinate and difficult to make out, the epithecium a broad dark brown (reddish-brown?) band; hypothecium very pale brownish; thecium very pale reddish or becoming colorless, blue with I, the asci narrow, clavate; spores rare, eUipsoid and ovoid,
9.75 - 14.7
On dry rocks in sunny places in the foothill canons; the only local- ity from which I have specimens obtained within our territory is Stevens Creek Canon, alt. 900 feet. It is common in Alum Rock Park, Mt. Hamilton Range, near San Jose, at an altitude of about 500 or 600 feet. In our plant the scales are smaller than given by Tuckerman, and smaller and darker than in a specimen from the Sprague Herbarium, collected in Washington.
A hchen of calcareous earth, occurring over the greater part of Europe and North America; in the latter region most abundant on the Pacific Coast.
3. LECIDEA SCOTOPHOLIS (Tuck.) Herre.
Biatora scotopholis Tuck. Lich. Cahf. 24. 1866.
Biatora scotopholis Tuck. Syn. N. Am. Lich. II: 11. 1888.
Thallus effuse, of minute, rather thin, rounded, areolate squam- ules with rugulose surface; their borders finely crenate and slightly
THE LICHEN FIORA OF THE SANTA CRUZ PENINSULA 8l
elevated; fissured or crowded and sub-imbricate; dull brown-black, the irregular thallus dull black to the naked eye, by which the minute scales are hardly visible; upon a black, fringing hypothallus; KOH - ;CaCl202 -.
Apothecia small, innate-sessile or adnate; disk flat, red-black to black, finally becoming convex, and the stout, shghtly lighter colored margin then disappearing; epithecium granulose, pale brown; paraphyses strict, coherent; hymenium 44 to 56/1 high, blue with I; hypothecium colorless; asci clavate; spores ovoid-ellipsoid,
3 -5
— I.
8 -II
On sandstone and other rocks at 1000 feet alt., on Mt. San Bruno, also in the foothills and mountains, probably throughout, up to an altitude of 1800 feet. Recorded by Tuckerman from the coast of California and the Dalles of the Columbia, Oregon.
Often intermingled with the thallus of Rhizocarpon holanderi (Tuck.), to which it bears a curious resemblance.
4. LECIDEA GRANULOSA PHYLLIZANS A. Zahlbr.
Lecidea granulosa Schaer var. phyllizans A. Zahlbr. Beih. Bot. Cen-
tralbl. 13: 159. 1902. Biatora glebulosa Tuck. Syn. N. Am. Lich. II: 16. 1888, in part.
Thallus determinate to effuse, often spreading extensively, of wavy, undulate, sub-cartilaginous, rounded squamules, incised or crenate, imbricate, at the circumference becoming radiate and lobate; whitish ash-colored to pale buff; KOH distinct yellow; CaCl202 red.
Apothecia scattered, or grouped, and then becoming conglom- erate, sessile, large, 1.75 mm. wide, from plane to turgid convex; at first with a thin proper margin which is finally excluded; disk dull reddish or yellowish brown, or blackening, papillate, with a faint bloom. In the field the disk is reddish flesh-color. Epithecium granulose, of a sordid yellow-brown color; paraphyses coherent, indistinct; hymenium pale sordid yellowish; h^-pothecium colorless
3-6
or nearly so; asci narrowly clavate; spores /^; sterigma simple,
9-14
1.2 — 1.5
straight; spermatia acicular, /^.
7-9
82 HERRE
Common in the foothills on sandstone and occasional in the moun- tains on the same substratum, up to 3000 feet. Specimens collected by Bolander at Mission Dolores, San Francisco, and at Ukiah, were named Biatora glehiilosa by Tuckerman.
Described by Dr. Zahlbruckner from specimens collected by Dr. Hasse on Mt. San Gabriel, in southern California.
5. LECIDEA COARCTATA (Sm.) Nyl.
Lichen coarctatus Smith, English Bot. 5:8. t. 534. 1795. Lecidea coardata Nyl. Lich. Scand. 196. 1861. Biatora coardata Tuck. Syn. N. Am. Lich. II: 15, 1888. Parmelia elacista Ach. Meth. Lich. 159. pi. 4./. 4. 1803. Lecanora ornata Sommerf. Suppl. Fl. Lapp, 92. 1826.
Thallus whitish or gray, areolate, scattered; thin or disappearing or forming a continuous, fissured crust; areolae quite small, the largest scarcely exceeding f mm. in width, furfuraceous, convex, and mostly approximate; the red reaction of authors with CaCl202 is not seen in our plant.
Apothecia sessile and sub-innate, small or minute; disk reddish brown, flat, fi:cly pap Hate, with a thin, slightly devated margin, which is concolorous or a little darker, persistent; frequently with a coarctate, spurious, pulverulent thalline margin; paraphyses indis- tinct, thread-like and twining; hypothecium and epithecium color- less; thecium blue or brown with I; asci oblong saccate; spores ovoid-
„. . , 7-5 - II ellipsoid, — ■ fi.
17-25
On sandstone and earth in the foothills and mountains. The forma elacista (Ach.) characterized by the effuse, very thin, scurfy, or almost entirely disappearing thallus, occurs on clay, along the sum- mit of the first ridge east of Los Gatos, at about 1500 feet elevation, and on sandstone at 2000 feet elevation on Castle Rock Ridge.
The forma ornata of authors, characterized by a more luxuriant development of the thallus, with marginally crenate, flat to convex squamulcs, occurs on sandstone in the foothills.
A variable and quite common lichen of Europe and North Amer- ica. (coardata, narrow or appressed, from the appearance of the false
thalline margin.)
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 83
6. LECIDEA FUMOSA (HofTm.) Ach.
Patellaria fumosa Hoffmann, Deutsch. Fl. 2: igo. 1791. Lecidea fumosa Acharius, Meth. Lich. 41. 1803. Lecidea fusco-atra Tuck. Syn. N. Am. Lich. II: 75. 1888.
Thallus spreading extensively, areolate squamulose, fissured, or the small squamules crowded and uniform, concave, rarely flat or convex, with a distinct, often paler margin, which from entire becomes intricately flexed; from shining brown varying to dark brown and nearly black, with a fringing black hypothallus; KOH — ; CaCloOs - .
Apothecia sessile, not very numerous, scattered or sometimes conglomerate, i to 1.25 mm. wide; disk smooth, flat, black, with a thin, grayish-black, at first entire, finally flexuous or lobate margin; becoming strongly convex and finally immarginate; hymenium color- less, blue with I; paraphyses coherent; epithecium bluish black; hypothecium horny, dark brown, about half the thickness of the
5 - 7-5 hymenium; asci narrowly clavate; spores ellipsoid, ^ /^.
On various rocks at all elevations, in the foothills and mountains, from 200 feet to the summit of the range.
Generally distributed in the mountainous regions of Europe and North America,
7. LECIDEA INTUMESCENS (Flot.) Nyl.
Lecidea hadia var. intumescens Flotow, Lich. Siles. no. 175, 1829.
Lecidea intumescens Nyl. Lich. Paris, no. 58, 1854.
Lecidea insularis Nyl. Bot. Not. 177. 1852.
Lecidea insularis Tuck. Syn. N. Am. Lich. II : 76. 1888.
Thallus a tartareous, determinate, warty, plicate or broken crust, of convex, cervine-brown squamules, several grouped to form small islands scattered among other crustaceous lichens, especially Lec- anora sordida and Rhizocarpon geographicum; hypothallus not dis- tinct.
Apothecia small or minute, innate to sessile; disk black; margin thin, persistent; epithecium brown; paraphyses coherent; h}pothe- cium brown; asci clavate; thecium blue with I; spores broadly
5—7
ellipsoid, /^.
10 - 13
84 HERRE
On sandstone in the mountains, at 3000 feet altitude, and also rarely at lower elevations. Occurring in the Oakland Hills (Bo- lander), and probably elsewhere in CaKfornia. Generally distrib- uted in Europe.
8. LECIDEA MANNI Tuck.
Lecidea manni Tuck. Syn. N. Am. Lich. II: 75. 1888. Z,ga(/m wawm Hasse,The Bryologist 11:6. 1908.
Thallus indeterminate, of rather large and conspicuous, thick, convex scales or areoles, from scattered becoming crowded and im- bricate; their surface smooth, with rounded or occasionally crenate or finely toothed edge, often with a gray, dusky, or blackish mar- gin; color buff to yellow brown; the dusky or blackish hypo thallus indistinct; KOH-f CaClsOs reddish.
Apothecia not numerous, innate, sessile, of medium size, circular or irregular, the black disk flat, soon plano-convex or moderately convex; bordered by a paler, erect, rather thick, entire, and sinu- ous margin; hypothecium colorless or sHghtly brownish; I-; spores
4.8 - 5 "5-7
ellipsoid, /^; — /<" Tuck.
12 — 19.5 II — 16
A single specimen collected on felspathic rock, at the summit of
Loma Prieta, at an elevation of 3793 feet. A comparison of this
with Tuckerman's type specimen shows them to be identical in
every respect except the paler color of my specimen. Mt. Diablo,
the type locality, is about 75 miles north of Loma Prieta, and is
the terminal peak of the Inner Coast Range east of San Francisco
Bay. The plant has also been found by Dr. Hasse in Ventura
County. (Named for Horace Mann, Jr. who collected hchens in
California and the Hawaiian Islands in the '60s.)
9. LECIDEA ATROLUTESCENS Nyl.
Lecidea atrolutescens Nyl. in lilt., 1896.
Thallus cartilaginous, indeterminate, composed of convex to sub-globose squamules, from ^ to 2 mm. wide; fawn-colored or buff, paling toward the margins, often crenulate and lobulate, either scattered or approximate; hypothallus indistinct.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 85
Apothecia sessile, becoming large, i to 2 mm. wide, circular, numerous, often crowded, and then irregular or distorted; disk black, usually with a white or gray bloom; at first moderately con- vex, with a turgid and lighter colored margin, becoming subglobular, the margin persistent and in larger apothecia sinuate or distorted; epthecium granulose, brown; paraphyses conglutinated; hymenium colorless to light brownish, deep blue with I, 80 to 100 /« high;
5-6
hypothecium very dark brown; spores oblong ellipsoid, <«.
12 — 14
8 and — [i. Spermatia not seen.
On sandstone at Grizzly Peak, at an altitude of 2700 feet. Here- tofore known only from Southern California, where it was discovered by Dr. Hasse.
10. LECIDEA GRISELLA (Elk.) Nyl.
Lecidea fumosa var. grisella Eloerke, in litt.
Lecidea fumosa var. grisella Schaerer, Enum. Crit. Lich. Europ..
no. 1850. Lecidea grisella Nylander, Lich. Lapp. Or. 160. 1867,
Thallus indeterminate, of minute, then scattered or barely con- tiguous areoles, plane or moderately convex, dull ashy gray or dusky gray in color; KOH- or faintly yellowish; CaCl202 red.
Apothecia small to medium, numerous, innate-sessile, closely appressed, black; the disk plane or soon moderately convex, bordered by a thin entire margin, which becomes angulose and is long per- sistent, finally disappearing; epithecium dark brown; paraphyses conglutinate, thecium deep blue with I; hypothecium blackish brown, broad; spores not observable in my specimens, the asci
6-7
poorly developed or their contents not differentiated; fi
II - 15
according to Hue.
On rocks in the foothills near Stanford University. A European
lichen apparently not distinguished by American authors.
86 HERRE
II. LECIDEA CRUCIARIA Tuck. Lecidea cruciaria Tuck. Syn. N. Am. Lich. II; 67. 1888 ; Santa Cruz.
Thallus efifuse, thin to very thin, of minute scurfy scales, or closely areolate; white, more or less plainly marked by tortuous black hypo- thalline lines, best seen when wet; medulla with I-; hypo thallus black.
Apothecia numerous, small to medium, sessile, flat; disk black, opaque, from smooth becoming minutely roughened; from flat becoming moderately convex and the originally stout, wrinkled, at length flexuous margin disappearing; epithecium bluish or green- ish black, with KOH becoming sooty brownish black; paraphyses coherent, capitate, with bluish black apices; asci clavate and in-
flated-clavate, — /'; hypothecium pale greenish brown and dark-
4.8 - 7J
ening; spores ellipsoid, -i^ 7 /<; hymenium blue with I.
9i - i7i
Tuckerman's specimens were from Santa Cruz, on sandstone. I have found it, however, only on Monterey shale, along the coast for 50 miles north of Santa Cruz and extending back into the ''chalk hills" ten miles or more from the coast, at altitudes from 50 to 1400 feet.
12. LECIDEA LITHOPHILA (Ach.) Th. Fr.
Lecidea lapicida i. Uthophila Ach. Lich. Univ. 160. 18 10. Lecidea Uthophila Th. Fr. Lich. Scand. II: 495. 1874. Lecidea pruinosa Tuck. Syn. N. Am. Lich. II: 66. 1888. Lecidea priiinosa Macoun, Cat. Canadian Plants, VII: 154. 1902.
Thallus thin, tartareous, of small ash-colored squamules, loosely approximate; the rimose-areolate character of the thallus mentioned by various authors not marked; the black hypothallus but little evident; KOH- ; CaClaOs-.
Apothecia sessile, i to f mm. wide; disk concave to flat, black, more or less hght gray pruinose, the thin black margin finally dis- appearing; epithecium brown; paraphyses simple, erect, coherent; hypothecium almost colorless; asci clavate; spores rarely seen,
6-7
ellipsoid, n-
Q - 15
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 87
On sandstone at Grizzly Peak at an altitude of 2775 feet. A lichen of northern and alpine Europe; in America reported from Greenland, Newfoundland, a number of localities in Canada, and in Texas.
13. LECIDEA TESSELLATA Flk.
Lecidea tessellata Floerke, Deutsch. Lich. no. 64. 181 5. Lecidea tessellata Tuck. Syn. N. Am. Lich. II: 68. 1888.
Thallus usually determinate and more or less orbiculate, limited by a black hypothalline band or line which is rarely obsolete; uni- form crustaceous, thick, sub-tartareous, of flat areoles, from deli- cately rimose becoming plainly fissured; pale ashy gray or whitish with a faint blue tinge; KOH — ; CaCl202— ; medulla without reac- tion with I.
Apothecia numerous, scattered or occasionally thickly grouped, from small to medium and very large (2.5 mm. broad), innate to sessile; disk flat to moderately convex, black, occasionally with a faint bloorfi; margin thick, black, erect; persistent, sometimes crisped or flexuous; a spurious thalhne margin is seen with some apothecia; epithecium bluish-black, paling downward; paraphyses, coherent, strict; hymenium colorless or very pale blue, 80 /i high, blue with I; hypothecium colorless to pale ash-color, as high as the hymenium; asci narrowly spatulate; spores rarely to be seen,
^Z'; / ~ ."/' Tuck. 6 6—10
A handsome and conspicuous lichen on igneous rocks in the foot- hills, at elevations of a few hundred feet. Generally distributed over Europe and North America.
{tessellata, checkered, like a mosaic pavement, alluding to the contrasting thallus and apothecia.)
14. LECIDEA LAPICIDA (Ach.) Am.
Lichen lapicida Ach. Lich. Suec. Prodr. 61. 1798, exclud. syn- onymy. Lecidea lapicida Ach. Meth. Lich. 37. 1803. Lecidea lapicida Am.,
Lecidea pantherina v. lapicida Th. Fr. Lich. Scand. 2 : 493. 1874. Lecidea polycarpa Tuck. Syn. N. Am. Lich. II : 69. 1888.
88 HERRE
Thallus thick to moderately thin, determinate, hmited by a more or less evident black hypothallus, uniform crustaceous, of flat areoles separated by minute fissures which later become broad and conspicuous cracks; KOH -; CaCl202-; medulla without reaction with I.
Apothecia at first small and innate, then appressed and large to very large, numerous, single and circular or usually grouped and then angular, i to 2.5 mm. wide; disk at first flat, soon sHghtly' convex, black; margin thin, persistent, slightly elevated; epithecium brown; thecium colorless, 80 /^ high, blue with I; hypothecium almost colorless or faint yellowish-gray or yellowish-brown; asci inflated
oblong-clavate; spores oblong ellipsoid, //.
ID — i4
On sandstone; at Castle Rock, altitude 3000 feet, and elsewhere along the summit of the ridge.
A lichen of alpine regions and the cooler parts of Europe and North America.
15. LECIDEA PLATYCARPA Ach.
Lecidea platycarpa Ach. Lich. Univ. 173. 1810.
Lecidea platycarpa Tuck. Syn. N. Am. Lich. II: 73. 1888.
Thallus ash-colored or leaden gray, indeterminate, uniform, thick and tartareous, becoming more or less fissured, or thin and granulose to finely pulverulent; no hypothallus evident; KOH — ; CaCl202 — .
Apothecia numerous, small, i to i mm. wide, appressed, scattered ; disk black, sHghtly papillate, flat; margin thin, becoming obsolete; the younger apothecia show a spurious, thin thalline margin now and then; margin shghtly horny; epithecium brown, much narrower than the hypothecium; paraphyses conglutinate ; hypothecium brownish-black or blackish-brown; asci ventricose; spores ovoid-
6 — 7.1;
ellipsoid, -^fJ-
^ '15-20
On sandstone in Santa Cruz, altitude about 50 feet. A lichen of the subarctic and temperate regions of Europe and America.
In our plant the thallus is much more developed and the apothe- cia smaller than in specimens gathered by me in the Alps.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 89
16. LECIDEA GONIOPHILA (Flk.) Schaer.
Lecidea immersa var. goniophila Floerke, Berl. Mag. 311. 1809. Lecidea goniophila Schaerer, Lich. Helvet. Exsicc. no. 531. Lecidea goniopliila Schaerer, Enum. Lich. Europ. 127. 1850.
Thallus effuse, of small, dingy white crumbs or minute scales, or quite obsolete; KOH pale yellow; CaCl202 very pale reddish.
Apothecia sessile, medium to small, J to i mm. wide, scattered; disk from a little concave to flat, finally convex, dull black, when wet suggesting red-black; margin concolorous, entire, somewhat turgid, glistening, finally obsolete; epithecium dark bluish-black; paraphyses coherent, hair-like; hymenium about 60 « high, pale blue with I; hypothecium pale dingy yellow; asci saccate, 20 x 50 //; spores ovoid ellipsoid, one end often pointed, the episporium dis-
7 — 10
tmctly double, 7- 1^-
■' '14 — 16
On comparison with authentic specimens in the museum at Vienna, our plant seems to be closest to the variety atro-sanguinea Hoffm., distinguished by the nearly obsolete thallus and dark red-black scattered apothecia.
On sandstone near Devils Canon, at an altitude of 2500 feet. Described originally from Germany and found also in France, Switzerland, Austria, and Italy.
17. LECIDEA AURICULATA DIDUCENS Th. Fr.
Lecidea auriculata didiicens Th.. Fries, Lich. Scand. 2: 499. 1874. Lecidea auriculata dediicens Jatta, Syll. Lich. Ital. 347. 1900.
Thallus absent. Apothecia medium to large; disk flat or slightly convex, black; margin persistent, thin to almost turgid, black, at first regular, at last sinuate; epithecium brownish black; paraphyses coherent, their capitate tips brownish black; hymenium colorless; hypothecium dusky; asci clavate, thecium intensely blue with I;
spores oblong ellipsoid, — — [j.] according to Fries .- _ ,«•
On various rocks and at various elevations, from near sea level ascending to 3000 feet at Castle Rock.
90 HERRE
i8. LECIDEA MELANCHEIMA Tuck.
Lecidea melancheima Tuckerman, Proc. Am. Acad. Arts, and Sci.
260. 1847. Lecidea melancheima Tuck. Syn. N. Am. Lich. II: 81. 1888.
Thallus creamy white, indeterminate, moderately thick, of irreg- ularly shaped, conjointed and rimose, rugulose, sub-lobate squam- ules or warts; KOH yellow; CaCl202 — .
Apothecia numerous, 4 to i mm. wide, appressed to sessile; disk glistening and very black, fiat to strongly convex, and then at times tuberculate, often wavy; margin very thin, becoming fiexuous, finally excluded; epithecium dark brown, gradually paling down- ward; paraphyses loosely coherent; h3^othecium pale or colorless; thecium 60 ,« high, colorless to light brown, blue with I; asci inflated
clavate; spores ellipsoid, //; spermatia not seen.
7 — 12
On dead wood, fences, roofs, limbs of Pseudotsuga taxifolia, etc., from sea level up to 3000 feet.
Common in New England; Colorado; Central Europe.
Lecidea elabens E. Fries, Act. Stock. 256, 1822, a similar plant from northern and Alpine Europe, is different. Schaerer, Enum. Crit. Lich. Europ. 131, says: "Apothecia atra, sub lamina cornea strato inferiore carbonaceo, innata, immarginata; disco exasperate, papillato. "
19. LECIDEA OLIVACEA (Hoffm.) Mass.
Verrucaria olivacea Hoffmann, Deutsch. Fl. 2: 192. 1791. Lecidea olivacea Mass. Ric. Aut. Lich. Crost. 71. 1852. Lecidea enteroleuca i.flavida Tuck. Syn. N. Am. Lich. II: 80. 1888. Lecidea enteroleuca Hasse, Pacific Slope Lichens distrib. C. F. Baker, no. 628. 1902.
Thallus thin, effuse, of scattered, minute, scale-like granules, or uniform crustaceous, of tiny areolate granules or crumbs, or becom- ing warty and uneven; color from an olive-brown or yellowish to greenish gray and whitish; KOH — ; CaCl202 brick-red or clay- red.
The forma geographica BagHetto is distinguished by the small, thin to very thin, determinate thallus, sharply limited by the con-
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 9 1
spicuous black hypothalline lines, and the yellowish or greenish- brown color.
Apothecia small or minute, numerous, scattered, black or rusty- black, sessile or sub-immersed, the disk concave to plane, finally moderately convex, with a thin, erect, entire margin which is finally excluded; epithecium dusky greenish; thecium blue with I; hypo-
7 - 8i thecium brown ; spores ellipsoid or ovoid ■ !J-
Common on the bark of various trees from the foothills to the summit of the highest peaks. A European lichen particularly abundant in the Mediterranean region. We have both the typical plant and the forma geographica.
20. LECIDEA PARASEMA Ach.
Lichen parasemus Ach. Lich. Suec. Prodr. 64. 1798.
Lecidea parasema Ach. Meth. Lich. 35. 1803.
Lecidea enteroleuca e. achrista Tuck. Syn. N. Am. Lich. II: 80. 1888.
Thallus effuse, thin, contiguous and rather smooth, or tartareous, becoming chinky or dispersed and made up of minute scurfy or warty areoles; whitish, gray, ashen, to brownish ash color; KOH — or sometimes yellowish; CaCl202 — ; hypo thallus indistinct or ab- sent.
Apothecia small, sessile, black; disk at first flat and often more or less tuberculate, with an evident entire margin which is some- times flexuous; soon convex and tumid, rugulose or papillate, the margin finally obsolete; epithecium bluish black; paraphyses free, their bluish black tips abruptly thickened; hypothecium faintly colored to brown; asci clavate, thecium blue with I; spores oblong
6-8 ellipsoid, ^^-3^ /..
A variable bark lichen occurring throughout our territory and found all over Europe and North America; one of the commonest species in most temperate regions, but with us less abundant than the closely related Lecidea olivacea.
21. LECIDEA LATYP^A Ach.
Lecidea latypcBa Ach. Meth. Lich. SuppL 10. 1803.
Lecidea enteroleuca Tuck. Syn. N. Am. Lich. II: 79. 1888, in part.
Proc, Wash, Acad. Sci., May, 1910.
92
HERRE
Thallus indeterminate, of thickish, unequal, whitish, gray, or yellowish brown warts, more or less dispersed or continuous and granulate-areolate; hypothallus indistinct, black; KOH yellow; CaCl202 faintly reddish yellowish.
Apothecia numerous, scattered or conglomerate, from innate to
sessile, i to i i mm. wide; disk black, long remaining flat, but finally
convex and tuberculate or rugulose; margin at first elevated, entire
or crenulate and sinuate, later disappearing; epithecium bluish
black; paraphyses loosely coherent; hymenium colorless or pale
gray, with I blue, soon turning brown; hypothecium brown, thick;
5 — 10 asci inflated clavate; spores broadly elHpsoid, — — — r -"•
Common on various rocks in the maritime region and in the foothills, at no very great elevation. Originally described from Sweden and not rare in the mountains of Europe.
22. LECIDEA ENTEROLEUCA Ach.
Lecidea enteroleuca Ach. Lich. Univ. 177, 1810.
Lecidea enteroleuca Tuck. Syn. N.Am. Lich. II: 79. 1888, in part.
Thallus a thin, effuse, granulose or minutely areolate or warty crust, or now and then disappearing; the small areoles or warts scattered, loosely approximate, or becoming crowded and even heaped; grayish white to dark ashy gray; KOH yellow; CaCl202 — or faintly reddish; hypothallus black.
Apothecia from. 5 to 1. 5 mm. wide; disk black, at first flat, soon con- vex, becoming subglobose; the thin, black, horny margin finally disappearing; paraphyses loosely coherent; epithecium bluish to brownish black; asci clavate, thecium pale reddish to colorless, becoming blue with I; hypothecium colorless to dusky; spores
5 — 10
ellipsoid to broadly ellipsoid, sometimes falsely bilocular, — — — ^/^;
spermatia long, needle-like, curved.
Common on various rocks in the foothills and widely distributed both as to latitude and altitude; a variable plant.
According to the character of the thallus and color of the hypo- thecium several forms are recognized, of which we have the follow- ing:
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 93
var. AEQUATA (Flk.)
Lecidea sahuletorum coquala Floerke, in Schaer. Spicil. 152. 1828. Lecidea enter oleuca cequala Tuck. Syn. N, Am. Lich, II: 80. 1888.
Thallus of whitish or gray squamules, irregularly distributed or crowded and then rimose areola te; KOH yellow; CaCl202 red.
Apothecia at first innate, then subsessile to superficial, often crowded but retaining their regular circular form; disk black, from flattish to convex; margin regular, entire, but eventually disap- pearing as the disk becomes more convex; tips of the loosely coherent paraphyses bluish black; hypothecium pale or colorless;
6 - 8.5 asci inflated clavate or wedge-shaped ; spores — — Z^-
On various rocks in the foothills and mountains. A Hchen of Central and Northern Europe, also occurring along the Atlantic coast of America.
var. THEIOPLACA Tuck.
Lecidea enteroleuca var. theioplaca Tuck. Genera Lichenum, 179.
1872. Lecidea enteroleuca var. theioplaca Tuck. Syn. N. Am. Lich. II: 79.
1888.
Thallus of pale yellow or sulfur-colored, globose or crenulate warts, mostly irregularly distributed and areolate, or closely com- pacted into a thin, uniform crust; KOH — ; CaCl202 vermilion; the color of the thallus precludes a reaction with KOH.
Apothecia small to medium, numerous, irregular, concave or plane, the thin, entire, greenish margin paler than the black disk, and finally excluded; hymenium colorless or brownish; hypothecium dark brown; spores as in the type.
On cliffs bordering the sea, at Point Lobos, San Francisco, and southward along the coast, at Point San Pedro and Pescadero.
Described by Tuckerman from about San Francisco and also determined by him from South Carolina and New Jersey.
In the author's opinion this is a species rather than a variety, distinguished by the different hypothecium, the different chemical reactions, and other minor distinctions.
94 HERRE
XX. Catillaria (Mass.) Th. Fr.
Catillaria Massalongo, Ric. sul. Aut. Lich. Crost. 78. 1852. Catillaria Th. Fries, Gen. Heterol. Europe. 88. 1861.
Thallus crustaceous, uniform or marginally lobed, without cor- tex. Apothecia circular, innate or sessile, with clear to black proper margin but no thalline margin; disk concave to convex, variously colored; hypothecium clear to black; paraphyses simple, free or coherent, capitate; spores 8, rather small, colorless, ovoid or ellipsoid, elongate or short, straight or curved, bilocular, with thin walls and without a halo.
A large genus, representatives occurring in all parts of the world and upon all kinds of substrata.
KEY TO THE SPECIES.
A. On rocks.
B. Thallus purplish black; apothecia black i. suhnigrata
BB. Thallus ash-colored; apothecia dark brown and blackening;
white pruinose 2. franciscana
A A. On bark.
C. Thallus whitish ash-color or gray; becoming yellow with KOH
3 . tricolor CC. Thallus greenish white; no reaction with KOH 4. globulosa
I. CATILLARIA SUBNIGRATA (Nyl.)
Lecidea suhnigrata Nyl. Flora, 370. 1866.
Lecidea suhnigrata Leighton, Lich. Fl. Grt. Brit. ed. 3. 331. 1888.
Thallus indeterminate, of purplish black squamules, imbricate, lobed and crenulate, rugulose; KOH — ; CaCl202 — .
Apothecia sessile, .5 to i mm. wide; disk flat, black, finely papil- late, at last markedly convex; margin at first thick but becoming partly or wholly obsolete; hymenium 68 M thick, pale purpHsh gray, paHng downward, blue with I; paraphyses subcoherent, the
5 — 7
tips clavate; hypothecium colorless; spores ellipsoid, fx,
10 — 12
On rocks near Stanford University, at an elevation of 500 feet. A lichen of the British Isles.
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 95
2. CATILLARIA FRANCISCANA (Tuck.) Herre. Biatora franciscana Tuck. Syn. N. Am. Lich. II: 32. 1888.
Thallus effuse, of small but thick and coarse squamules; these numerous and close together, concave, rugose, undulating, often crenate and lobulate, or sometimes closely appressed, few in number, or nearly disappearing; occasionally passing into warty areolate conditions; ash-colored with lighter colored margin, hardly darken- ing; hypothallus indistinct or absent; KOH — ; CaCl202 — .
Apothecia of medium size, i to 1.25 mm. wide; disk from slightly to strongly convex, dark dull brown to blackening, with a white bloom; the rather stout, lighter colored margin finally excluded; epithecium brown; hypothecium colorless or faintly colored; paraphyses strict, not septate, their slightly thickened tips light brown; asci elongate clavate, about as high as paraphyses; hymenium colorless, blue
3 — 5
with I; spores narrowly oblong-ellipsoid, //.
i4 — 22
On rocks all along the Pacific coast, from near the Cliff House, San Francisco, southward. Recorded by Tuckerman from the Oak- land Hills and by Dr. Hasse from the coast of Southern California.
3. CATILLARIA TRICOLOR (With.) Th. Fr.
Lichen tricolor Withering, Arrang. 4: 20. 1796. Catillaria tricolor Th. Fr. Lich. Scand. 2: 574. 1874. Biatora mixta E. Fries, Vet. Ak. Handl. 267. 1822. Biatora mixta Tuckerm. Syn. N. Am. Lich. II: 30. 1888.
Thallus whitish ash-color to dull gray, and from nearly smooth to granulate, becoming almost chinky, areolate, the small areolae rugu- lose; usually limited by a black hypothalline line; KOH yellow; CaClzOz-.
Apothecia small to minute, appressed sessile; disk at first flat, soon becoming convex, flesh-color to reddish brown and blackish, pruinose; the thin margin usually persistent; epithecium and hypo- thecium colorless; paraphyses simple, free, thread-like, slightly knob- like at the apex; thecium blue with I; asci inflated clavate, — fx;
40
96 HERRE
spores ellipsoid to spindle-shaped, straight or slightly curved, 2.8-5.6
9.75-18
Abundant on the bark of various living trees in the foothills and along the coast; rarely on old fences in the mountains.
The var. pacifica of Tuck., distinguished by a black, Kmiting hypo- thallus and more distinct septum in the spores, and the var. atlantica of the same author, are both found on this coast.
Common in Europe, on bark and dead wood, in New England, and on the Pacific coast.
4. CATILLARIA GLOBULOSA (Flk.) Th. Fr.
Lecidea globulosa Floerke, Deutsch. Lich. 181. 1815. Catillaria globulosa Th. Fr. Lich. Scand. 2: 575. 1874. Biatora globulosa Tuck. Syn. N. A. Lich. II: 32. 1888.
Thallus greenish white, thin, effuse, of minute, crowded warts or granules, or these now and then scattered; KOH— ; CaC]202 — .
Apothecia small to very small, sessile or semi-immersed in the thalline warts, soon convex and sub-globose, immarginate ; very young and small apothecia are fiat with a thin margin; disk dark brownish black to black, opaque; epithecium pale sordid yellowish; paraphy- ses conglutinated, indistinct, their tips thickened; hymenium color- less, blue with I; hypothecium colorless; asci clavate; spores nar-
3~3-5 rowly oblong, faintly septate, at times a Httle curved, 7 f^-
On old fences near Los Gatos, at an altitude of 450 feet. A European lichen reported in this country from the White Moun- tains and from British America.
XXI. Bacidia (DeNotaris) A. Zahlbr.
Bacidia DeNotaris, Giorn. Bot. It. an. 2, tom. I: 189. 1846. Bacidia A. Zahlbruckner, Ascohchenes, 135. 1907.
Thallus uniform crustaceous, without cortex. Apothecia circu- lar, sessile or rarely innate or elevated, the disk plane or strongly convex; the proper margin colorless or dark; paraphyses simple, free or coherent, their ends often thickened; hypothecium clear to dark;
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 97
asci with 8, rarely i6 spores; these from 3-to multilocular, spindle to needle-shaped, with both ends ahke or one end prolonged into a tail, straight, curved, or spiral, without a halo.
A large genus, of more than 200 species, found all over the world, on stones, bark, w^ood, moss, etc.
KEY TO THE SPECIES.
A. Thallus usually pale yellow, and yellow with KOH; apothecia blue,
then violet with KOH i. herrei
AA. Thallus never yellow and not changed by KOH.
B. On rocks; thallus black with greenish or grayish cast 2. ioessa
BB. On trees; thallus not blackish.
C. Apothecia clouded flesh-color and darker; thecium blue with
I 3. ncBgelii
CC. Apothecia black; thecium not blue with 1 4. akompsa
I. BACIDIA HERREI A. Zahlbr.
Bacidia herrei A. Zahlbruckner, Annales Mycologoci, 6: 130. 1908.
Thallus sub-orbiculate, becoming effuse, more or less chinky, of granulose, densely imbricated and crowded, thickish squamules, or thin and reduced to mere granules; without soredia or isidia, but the granules sometimes almost coralloid; usually of a pale yellow color, sometimes whitish, rarely greenish gray; KOH yellow; CaCl202 — .
Apothecia sessile, small to medium, sparsely distributed to approxi- mate, sometimes forming a heap of several together, basally con- stricted, circular, or sub-angulose when crowded; disk plane to con- vex, red, not pruinose; the proper margin thin, concolorous, entire, from prominent finally excluded; with KOH an apothecial section turns an intense deep blue, soon changing to violet, the epithecium losing the blue last; epithecium granular, broad, dark red; hypothe- cium colorless; hymenium more or less reddish, 90 — iio/i high, bluish with I; paraphyses close together, free, simple, not septate; asci short, oblong-clavate; spores arranged lengthwise in the asci, colorless, needle-like to narrowly spindle-shaped, usually much attenuated at one end, straight to slightly curved, indistinctly pluri-
2 — 3 < septate (5 — 8), ^/^> 35 ~ 40." longiseti.y — i.8«latis", A.
Zahlbr.
On sandstone and the bark of Pseudotsuga taxifolia and on dead wood of Pseudotsuga taxifolia and Adenostomafasciculatum, at Devil's
pS HERRE
Canon, altitude 2300 feet, Castle Rock, 3000 feet, and Grizzly Peak, 2700 feet. Probably found all along the summit of the range in similar localities.
The above description somewhat altered from Dr. Zahlbruckner's excellent diagnosis. While the specimen he described was found on dead Adenostoma, I regard the typical plant to be the one with yel- low, orbiculate, thickish thallus, growing on sandstone.
A handsome, conspicuous, but not very abundant plant.
Strongly characterized by the color of the thallus and apothecia, as well as by the beautiful apothecial reactions with KOH.
2. BACIDIA lOESSA Herre, new species.
Thallus effuse, thin, of scattered, minute to small, thick, rounded or sub-globose, sometimes sub-pHcate or difiform granules or crumb- like squamules, which are occasionally aggregate; on a thick, promi- nent, often scurfy hypo thallus; color black with a greenish or grayish cast; dark olive-green when wet; KOH — : CaC]202 — .
Apothecia numerous, small, sessile, black; the flat disk bordered by a small, entire, sometimes paler margin, but soon becoming con- vex, when the margin is excluded finally; epithecium blackish, with KOH becoming purplish or rosy violet, the color suffusing the the- cium; the latter blue with I; paraphyses free, thread-like, rather lax, with sub-globose tips which are violaceous dusky to blackish ; hypo- thecium colorless to pale brownish; asci subcylindrical to clavate
10 — 15
/<; spores 4 — locular, spindle-, finger-, and sickle-shaped,
3-5-6
14 . 5 — 20
On igneous rocks on a dry liill side. Hidden Villa Canon, at an alti- tude of 800 feet, and probably in similar situations all through the foothills. The specimens scanty; apparently close to Tuckerman's Bialora arlyla, Synopsis, 11: 37, but I cannot bring the two together. {ioessa, from coeoaa violet colored, from the epithecial reaction with KOH.)
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 99
3. BACIDIA N^GELII (Hepp.) A. Zahlbr.
Bialora nagelii Hepp. Exiscc. no. 19. 1853.
Biatora ncEgelii Tuck. Syn. N. Am. Lich. II: 36. 1888.
Bacidia ncogelii A. Zahlbr. Ascolichenes, 135. 1907.
Thallus of minute, thickish, scale-like granules, forming a more or less chinky crust, or occasionally thin; color ashy gray or greenish ashen; KOH-;CaCl202-.
Apothecia numerous, minute to small, sessile, circular, at first
plane but very soon becoming strongly convex and excluding the
thin, entire, scarcely evident margin; color a clouded flesh-color as
nearly as can be defined, soon darkening and then blackening; para-
physes distinct but coherent, slender; hypothecium clear; thecium
5 — v.c blue with I; spores spindle-shaped to ellipsoid, — /^;with
I, 2, 3, 4 septa, mostly 4-locular. Dr. Zahlbruckner states in Ascolichenes "sporen bis 8 zellig" but I find none with more than 5. Th. Fries states, Lich. Scand. II: p. 379, "sporae primitus simplices, dein 2-4, raro 6-8 blastae."
On bark of Umbellularia and other trees, mixed with Lecania dimera, Catillaria tricolor, and other lichens.
A bark lichen of both Europe and North America. (Named for Karl Wilhelm von Naegeli, botanist and philosopher, professor at Munich from 1858 to 1891.)
4. BACIDIA AKOMPSA (Tuck.) Herre.
Biatora akompsa Tuck. Syn. N. Am. Lich. II: 47. 1888.
Thallus pale ash-colofed to dusky greenish ash-colored, effuse, interruptedly granulose or scurfy, as if poorly developed; no chemi- cal reaction evident.
Apothecia scattered, small to minute, sessile; disk dull black, more or less convex; margin thin, indistinct; epithecium pale grayish brown; paraphyses coherent, hair-like; hypothecium colorless; asci clavate or narrowly spatulate; spores needle-shaped, 4 to 5 locular,
2 — 3 1.5 — 2.5
— /^: according to Tuck., — 7 u; thecium not colored
18 - 20 ' ^ ' 18-24
by iodine.
lOO HERRE
On bark of Pinus radiata near Stanford University, at an altitude of 200 feet. Given by Tuck, as collected by Bolander on the bark of Pinus insignis, coast of California. The Monterey pine, Pinus radiata {Pinus insignis) occurs wild only about Monterey and on the southwestern coast of the Santa Cruz peninsula. Bolander's speci- mens were undoubtedly collected on the coast between Pescadero and Santa Cruz, a locality where he collected a number of lichens.
XXII. Toninia (Mass.) Th. Fr.
Toninia Mass. Ric. sul. Aut. Lich. Crost. 107. 1852. Toninia Th. Fries, Gen. Heterol. Europ. 80. 1861.
Thallus crustaceous-squamulose to sub-foliaceous, swollen or inflated and sub-pedicellate, marginally lobed ; without true rhizoids; upper side with a firm cortex.
Apothecia circular, sessile, the proper margin variously colored, horny, of radiately arranged, thickened hyphae; paraphyses simple, free or confluent, often capitate; hypothecium clear or dark; spores 8, elongate or ellipsoid, 2 to 8-locular, without halo.
About 80 species, on rocks and earth, mostly xerophytes of the cool temperate and alpine regions.
KEY TO THE SPECIES.
A . Spores bilocular.
B. Thallus compact crustaceous; usually white pruinose; with KOH
dusky brown i. caruleo-nigricans
BB. Thallus scattered; not pruinose 5. massata
A A. Spores 4-plurilocular.
C. Thallus of livid brown and blackening squamules, slightly reddish
with KOH 2. squalida
CC. Thallus not affected by KOH.
D. Thallus tawny brown, extending downward in stout brownish
stems 3. caulescens
DD. Thallus of minute squamules forming a dark greenish black or dusky gray crust 4. aromatica
I. TONINIA C.ERULEO-NIGRICANS (Lightf.) Th. Fr.
Lichen caruleo-nigricans Lightf oot. Flora Scot. 805. 1777. Toninia cceruleo-nigricans Th. Fr. Lich. Scand. 2 : 336. 1874. Lecidea cceruleo-nigricans Tuck. Syn. N. Am. Lich. II: 61. 1888.
Thallus indeterminate, of thick, gyrosely plicate, turgid, medium sized or small squamules crowded into a compact crust, appressed,
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA lOI
or sometimes extended downward into stipes; mostly dusky greenish
but varying also from whitish to brown-green and black-green,
usually white pruinose; with KOH turning dusky brown.
Apothecia medium to large, sub-innate to sessile, at first concave
but soon plane or plano-convex, finally strongly convex; the black
disk with a thick, whitish or pruinose margin which soon disappears;
epithecium granulose, dark; paraphyses thick, free, their enlarged or
spatulate and blunt tips dark greenish; thecium colorless, blue with
I; hypothecium brownish; spores bi-locular, spindle-shaped to nearly
2.5 — 3.5 ^, 2-4
needle-like, /«; /<," Tuck.
20 — 30 14 — 27
On earth in rock crevices on Black Mt., at from 2400 to 2700 feet elevation. An earth and Hme-rock lichen of Europe; in America only in cold mountains or the far north.
In our specimens the disk is naked, but in specimens collected by me in Styria the apothecia are mostly white pruinose, the whole plant often seemingly covered with hoar frost.
2. TONINIA SQUALIDA (Schleich.) Mass.
Lichen squalidus Schleicher, PI. Crypt. Helvet. Cent. Ill, no. 75.
1807. Toninia squalida Mass., Ric. Aut. Lich. Crost. 108. 1852. Lecidea squalida Ach. Lich. Univ. 169. 1810. Lecidea squalida Tuck. Syn. N. Am. Lich. 64. 1888.
Thallus a close, uneven crust of small, rather thick and closely appressed, rugose-pHcate squamules, often sub-Iobate; livid brown and blackening; shghtly reddish with KOH.
Apothecia numerous, small, closely adnate, the black disk plane, bordered by a thickish, regular margin; apothecia also larger, becom- ing irregular, confluent, and convex, when the margin is excluded; pale wdthin; paraphyses free, slender, their blackish tips enlarged and rounded; hypothecium pale reddish brownish; epithecium granulose, violaceous or purplish with KOH, the entire internal structure becoming more or less suffused with the same tint; thecium blue with
12 — 15
I, the asci clavate, -r /^; spores 4-6 locular, finger or needle-
48 - 52
2.5 — 4
shaped, straight or curved, — Z^-
^ '28-37
I02 HERRE
Rare; on earth in rock crevices near the summit of Black Moun- tain, at an elevation of 2700 feet. A lichen of alpine and arctic Europe; in North America recorded from Greenland and the moun- tains of the Pacific Coast.
3. TONINIA CAULESCENS Anzi.
Toninia caulescens Anzi, Cat. Lich. Prov. Sondr. 67. i860. Lecidea squalida caulescens Tuck. Syn. N. Am. Lich. II: 64. 1888.
Thallus tawny brown, squamulose, the turgid squamules convo- lute, scattered or usually crowded and imbricate, extending down- ward in stout brownish stems; KOH — ; CaCl202 — .
Apothecia closely sessile, deeply concave, from small and round to large and lobulate; disk dull black, papillate; the prominent turgid margin persistent, at first regular, round, becoming at length sinuate on the larger apothecia; epithecium brown, with KOH violaceous brown; hypothecium dark reddish brown; paraphyses separate, the brown tips abruptly capitate; hymenium pale yellowish, intense blue with I, soon changing to a sordid vinous red; asci spatulate, almost equahng the hymenium in height; spores fusiform, 4 to 8 (10?) locu-
2-5 — 5 lar, ^ ^ [i-
24 - 50
On earth and rocks at 50 to 100 feet above the sea. Point Lobos, San Francisco, and on earth above the sea a few miles south of Point San Pedro.
A lichen of alpine and arctic Europe and of the Pacific coast of the United States.
4. TONINIA AROMATICA (Sm.) Mass.
Lichen aromaticus Smith, Eng. Bot. pi. 25, f. 1777. 1807.
Toninia aroniaticaMa.ss . Symm. 54. 1855.
Lecidea aromatica Ach. Lich. Univ. 168. 1810.
Lecidea aromalica Tuck. Syn. N. Am. Lich. II: 131. 1888.
Thallus effuse, of minute squamules or crumb-like granules, mostly contiguous, imbricate, irregularly and confusedly rugose, forming a dark greenish black or dusky gray crust.
Apothecia sessile, often clustered, f to i mm. wide; disk black, at first slightly concave with a thin, entire or crenulated margin, soon
THE LICHEN FLORA. OF THE SANTA CRUZ PENINSULA I03
flat, the margin not elevated, and finally convex and distorted, the
margin disappearing; epithecium dark, brown-black and purplish
black, with KOH violet; hypothecium dark yellowish brown; para-
physes separate, some with grayish violet, globose tips;hymenium
pallid or pale violaceous gray, intensely blue with I; spores quadrilo-
3—4 cular, narrow fusiform with obtuse ends, [i\ spores straight
or sometimes slightly curved.
On sandstone near Mayfield, at an altitude of 400 feet.
This plant of Europe and Northern Africa has been reported only from Ontario and California in North America.
5. TONINIA MASSATA (Tuck.) Herre.
Lecidea massata Tuck. Lich. Calif. 25. 1866.
Lecidea massata Tuck. Syn. N. Am. Lich. II: 63. 1888.
"Thallus of small, scattered, turgid, glebous squamules becoming at length plicate, pale greenish and glaucescent; apothecia small to middling-sized (0 mm., 5—1 mm, 5 in width) peltate, flat, but the t in uneven margin at length disappearing, finally convex and irregu- lar, pale within, the hypothecium rufous-brown. Spores cymbiform, bilocular, 9 — 16 by 3 — 5/^-
On the earth in gravelly soil, San Francisco, California (Bolander, Tuckerman, /. c. 1866). Colorado, Brandegee in Herb. Sprague. Spermogones not observed."
I have not been able to find the above lichen and give the descrip- tion written by Tuckerman.
6. TONINIA RUGINOSA (Tuck.) Herre.
Lecidea ruginosa Tuck. Lich. Calif. 25. 1866.
Lecidea ruginosa Tuck. Syn. N, Am. Lich. II: 64. 1888.
"Thallus of rounded, turgid, glebous squamules which become more or less crowded together, wavy, and rugose-plicate, and are finally cancellated, from greenish to at length tawny brown; apothecia ample to large (i mm., 5 to 3 mm. in width,) flat, at length flexuous- lobate, scarcely excluding the stout margin, pale within, the hypo- thecium brownish. Spores acicular, 4 — plurilocular, 25 — 40 by 2 — 3 mic. Spermatia filiform, i:owcd, on sub-simple stcrigmas.
I04 HERRE
Serpentine rocks on the coast of California, (Bolander). Squa- mules less developed than in the last preceding, scarcely lobed. Apothecia originally rufous."
The above copied from Tuckerman's description in the Synopsis, II: 64. "The last preceding" refers to "Lecidea squalida."
As yet I have been unable to find the above described lichen, though Bolander 's specimens undoubtedly came from about San Francisco.
XXIII. Rhizocarpon (Ram.) Th. Fr.
Rhizocarpon Ramond, in DC. Fl. Fr. 2: 365. 1805, in part. Rhizocarpon Th. Fr. Gen. Heterol. Europ. 92. 1861.
Thallus uniform crustaceous, without cortex; often with a strongly developed hypothallus.
Apothecia circular, sessile upon the thallus, innate, or between the areoles, with a black or brown proper margin and a dark hypothe- cium; paraphyses lax, branched and twining; asci with from i to 8 spores which are colorless to dark, bilocular to multilocular, or muri- form, with a plainly visible halo.
Species numerous, on rocks in arctic and temperate regions.
KEY TO THE SPECIES.
A . Spores colorless, muriform i, distinctum
AA. Spores brown, 4-locular to muriform, B. Thallus yellow.
C. Thallus sulfur-yellow; medulla with I- 2. viridi-atrum
CC. Thallus lemon-yellow; medulla blue with I.. 3. geographicum
BE. Thallus not yellow.
D. Thallus of dark brown or reddish black squamules . .4. bolanderi DD. Thallus more or less gray.
E. Asci with 2 spores; thallus dark gray and blackening.
5. geminata EE. Asci with 8 spores; thallus brownish, bluish-gray or black- ish gray 6. petrmum
I. RHIZOCARPON DISTINCTUM Th. Fr.
Rhizocarpon distinctum Th. Fr. Falk. Bleck, 16 (nomen) ; Lich. Scand. 2: 625. 1874.
Thallus sub-determinate to effuse, thin, rimose-areolate, the areo- lae minute to small, sHghtly concave or flat; hypothallus black; color of plant whitish to leaden gray and darkening; KOH — ; CaCl20:. — .
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 105
Apothecia small, .5 to .75 mm. wide; disk black, papillate, with a
thin grayish, at first slightly elevated, margin, continuing plane with
the finally convex disk, but not wholly excluded; epithecium brown
black, gradually paling downward, with KOH pale violaceous; hypo-
thecium pale brown and darkening; hymenium palUd, with I turning
intensely blue; paraphyses conglutinated; asci saccate; spores color-
10 — 16
less, muriform, //.
24 - 36
On rocks in the mountains; New Almaden, 1200 feet; Castle Rock, 3000 feet; and on maritime rocks near Pescadero. A lichen of northern, alpine, and southern Europe.
2. RHIZOCARPON VIRIDI-ATRUM (Flk.) Korb.
Lecidea viridi-atnim Floerke,
Rhizocarpon viridi-atrum Korber, Syst. Lich. Germ. 262. 1855.
Diplotomma viridi-atrum Jatta, Syll. Lich. Ital. 432. 1900.
Thallus greenish or sulfur-yellow, of minute, thickish, tartareous, fiat or rounded granules or squamules; the h^-pothallus but httle, or not at all evident; KOH — ; CaCl202 — ; medulla not affected by I.
Apothecia numerous, of medium size, innate or closely appressed, dull black, not pruinose; the disk more or less minutely roughened, at first plane, with a thin, entire or irregular margin; soon moderately convex and the margin disappearing ; paraphyses indistinct, coherent; epithecium broad, black, purplish red with KOH; hypothecium blackish-brown; thecium deep blue with I; spores quadrilocular, elhp-
7-5 — i3-5
sold or oblong, dark brown, becoming nearly black, /^;
17-28.5
perhaps murilocular, but too dark to determine positively.
A distinct species, very rare with us; collected but once, on sand- stone in the foothills 4 miles west of Stanford University, at an alti- tude of 400 or 500 feet. Not rare in Europe, but apparently not dis- tinguished by American collectors.
3. RHIZOCARPON GEOGRAPHICUM (L.) DC.
Lichen geographicus Linne, Spec. Plant. 1140. 1753. Rhizocarpon geographicum DC. Fl. Fr. 2: 365. 1805. Buellia geographica Tuck. Syn. N. Am. Lich. II: 103. 1888.
Io6 HERRE
Thallus greenish yellow to bright lemon-yellow, determinate, rimose areolate, the flat areolae crowded into a chinky crust, or else the areolae are scattered and tumid, forming small clumps; hypothal- lus distinct, black; medullary hyphae blue with I; not affected by other reagents.
Apothecia immersed or between the areolae and on the same level
as the thallus, mostly angular from pressure of adjoining areolae or
other apothecia, small, numerous and often grouped; disk always
flat, black, opaque, the margin thin, black, indistinct; epithecium
brownish-black; paraphyses loosely coherent, the brown tips scarcely
thickened ; hypothecium brownish black; thecium colorless, blue with
16 — 20
I; spores dark brown, 2 to 4 locular and muriform, -^ > fi-
20—40
The forms contigua and lecanorina of authors are not rarely found mixed with the type, on the same specimen.
A beautiful and conspicuous lichen. Abundant on various rocks at 2500 feet and above, and in the cold and foggy San Francisco region descending as low as 500 feet. Found in nearly all moun- tainous regions of the world and characteristic of all very high peaks.
4. RHIZOCARPON BOLANDERI (Tuck.) Herre.
Buellia holanderi Tuck. Gen. Lich. 189. 1872.
Buellia holanderi Tuck. Syn. N. Am. Lich. II: 103. 1888.
Thallus indeterminate and spreading extensively, cartilaginous, of small to very small, brown or reddish-black, sometimes shiny, flat or slightly concave, round or sinuate squamules; these with a shghtly elevated black border, scattered or approximate; in the latter case forming an areolate-diffract crust upon a conspicuous black hypo- thallus that to the naked eye gives the predominant color to the thal- lus; no chemical reactions of thallus or medulla.
Apothecia small, dispersed, from partially innate to sessile; disk flat or shghtly convex, naked, black; margin quite thin, becoming finally obscure; epithecium dark brownish violaceous black; thecium pale, deep blue with I; paraphyses conglutinate; hypothecium of same color as epithecium; asci saccate and inflated saccate, about as high as thecium ; spores colorless to dark smoky gray and dark brown, solitary or in twos, with a thick gelatinous halo, muriform, oblong
THE LICHEN FLORA OF THE SANTA CRUZ PENINSULA 107
20 — 36
ovoid or broadly ellipsoid, — -^^ — !i\ according to Tuckerman,
20 — 25
solitary, in twos or in fours, fi-
^ 30-50
On igneous rocks and sandstone throughout, but most abundant on the higher peaks. Often intermingled with Lecidea scotopholis Tuck., which is very similar in appearance.
Type locahty, <