r ! 5 !j; ( iii^i ?i^';■ii^,i' •- ■■ ; ) 1 . ;''■''''•, !*}>;•!' M ;•: ^ f ; 5 ^ I'll- ''v. .J );:•'!;} ■. -)';'J:' ■': :■ n)5^'i;'''';':{'fr^ -■--■: ■■■■■ i^mm\m^>>-. ' ■ ' ■ .:,.',;,.■: i.'' ■■■■ •','-:■ ■ ■'■'■'. •■- :'■ Im-mm 1 i :;lii PROCEEDINGS OF THE California Academy of Sciences FOURTH SERIES Vol. X 1920 SAN FRANCISCO Published by the Academy 1920 COMMITTEE ON PUBLICATION George C. Edwards, Chairman C. E. Grunsky Barton Warren Evermann, Editor CONTENTS OF VOLUME X Plates 1-9 PAGE Title-page i Contents Hi A Further Study of Variation in the Gopher-Snakes of Western North America, by John Van Denburgh 1 (Published August 6, 1920) Description of a New Species of Rattlesnake (Crotalus lucasensis) from Lower California, by John Van Denburgh 29 (Published August 6, 1920) Description of a New Subspecies of Boa (Charina bottce utaliensis) from Utah, by John Van Denburgh 31 (Published August 6, 1920) Description of a New Lizard (Dipsosaurus dorsalis lucasensis) from Lower California, by John Van Denburgh : 33 (Published August 6, 1920) Undescribed Tipulidae (Diptera) from Western North America, by Charles P. Alexander 35 (Published August 6, 1920) Three New Species of Dolichopodidae (Diptera) from California and Nevada, by M. C. Van Duzee _ 47 (Published August 6, 1920) Two New Species of Syrphidse (Diptera), by A. L. Lovett 51 (Published August 6, 19^) Notes on Some Undescribed Calif ornian Helices, by S. Stillman Berry.... 53 (Published August 11, 1920) A New Genus and Species of Grasshopper from California, by Morgan Hebard _ 71 (Published December 29, 1920) Color Changes and Structure of the Skin of Anolis carolinensis, by Charles E. von Geldern 77 (Published February 12, 1921) Report of the President of the Academy for the Year 1920, by C. E. Grunsky 119 (Published July 2, 1921) Report of the Director of the Museum for the Year 1920, by Barton Warren Evermann 128 (Published July 2, 1921) Index 165 JUNE 11, 1923 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. X, No. 1, pp. 1-27, pis. 1-2 August 6, 1920 A FURTHER STUDY OF VARIATION IN THE GOPHER-SNAKES OF WESTERN NORTH AMERICA BY JOHN VAN DENBURGH Curator, Department of Herpetology In August of last year the writer and Joseph R. Slevin pub- lished the results of a study of some three hundred specimens of Pitnophis from western North America^ Probably no one who has not undertaken such a study can realize the amount of time and effort necessary for gathering, analyzing, digesting, and formulating the data derived from such a mass of material as was involved in our earlier study of the garter-snakes of the genus Thamnophis. The time and effort expended were so great that in the subsequent study of Pitnophis the attempt was made to reach conclusions without so much attention to detail. The result was that, while the conclusions reached were valu- able and probably correct, as far as they went, they never have been satisfactory as a solution of the problems in variation and distribution offered by this genus. Further study of the data published in that paper indicated the probability tliat the snakes 1 The Gopher-Snakes of Western North America. M/. . i» .itt *p! j.t» ui/t-jii^ jm iir.M'-UJ''^i^Mt.jj*..x3a'.xij..;a^.Mt 4jg'.ji«l3M...iW-2H..^»~JfB.Mhi:jff.ia-J^',jit»Mt-^trt^ Fig. 1 As one proceeds south and east from the cool north coast regions toward the wami south coast and interior desert regions, the average counts gradually increase. Although the transition is gradual, these gastrostege counts are of great use in the separation of these snakes into the two groups which we then regarded as two subspecies. This same difference and relationship are shown when the combined gastrostege and urostege counts are charted for the two groups, as has been done in figure 2. In this chart the VOL.X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES Fig. 2 upper curve (1) represents the counts in the snakes we called P. catenifer catenifer, the middle curve (2) those for P. caten- ifer annectens, and the lower curve (3) the same counts for P. vertebralis. That there is a very real difference between the first group and the others is evident. Turning now to a consideration of ^he number of dorsal blotches between the head and tail (at a point over the anus), we find that the snakes which have fewer gastrosteges are not all alike. Those from near the coast have more blotches than those from the interior. The same is true of the snakes with more numerous gastrosteges. The coast snakes have more numerous blotches than those from the interior. Figure 3 shows the number of blotches on the body in gopher-snakes from several geographical areas. The upper two curves (1 and 2) represent the snakes with fewer gastrosteges, which we called Pituophis catenifer cataiifer. It will be seen that there is a marked difference between these two curves. The upper curve (1) represents the snakes from the coast valleys and ranges from Oregon to Santa Barbara County, California. The second curve (2) shows the number of blotches in the snakes of this group from the Klamath-Modoc region, the Sacramento Valley, the western slope of the Sierra Nevada, CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Seh. and the northern portions of the San Joaquin. The difference between these two curves points to the necessity of recognizing two subspecies here, a coast race, P. catenifer catenifer, and an interior race, P. catenifer heermanni. The third, fourth, fifth, and sixth curves charted in this figure 3 represent respectively snakes (3) from the San Diegan Fauna, (4) from the desert regions of CaHfornia, the southern San Joaquin Valley and Nevada, (5) from Utah, and (6) fo f-t- 'ff Fig. 3 from Arizona. Curve 3 shows that the snakes from the coast of southern California have more blotches than those of any of the other regions where the gastrosteges are numerous. In this respect these snakes are like those from the northern coast, P. catenifer catenifer, from which, however, they differ in other respects. The difference betw'een these snakes and those repre- sented by the other curves necessitates recognition as a distinct subspecies, P. catenifer anncctens, using this name in a re- stricted sense. Vol. X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES 5 The sixth curve, representing the snakes of Arizona, also shows marked difference from all of the others of this group. The number of blotches is much reduced and is similar to that found in Pituophis vertehralis, shown in the seventh curve. It would seem that the Arizona snakes also are entitled to sub- specific rank. This leaves curves 4 and 5, which agree with each other and with curve 2. Bearing in mind the fact that the snakes of curve 2 {P. c. hcermanni) belong to the group having fewer gastrosteges, it seems necessary to regard those represented by curves 4 and 5 as deserving separate standing. It will be shown later that the snakes of Utah (curve 5) differ from those represented by curve 4. For the latter, the name P. catenifer deserticola would seem to be available. The number of blotches on the tail in these various races has been charted in figure 4. The same differences and relation- ships as are indicated by the blotches on the body are shown by these data. Turning now to a consideration of the scale-rows we find that the greatest number present on the body, although subject to great individual variation, also shows geographic variation. This is brought out in figure 5. Curves 1 and 2, representing P. c. catenifer and P. c. heermanni, agree in showing 31 rows as the most frequent number. The Ari^na specimens (curve 6) and those of P. c. amiectens (3), and P. c. deserticola (4) agree, and show 33 as the most frequent number. P. verte- hralis (curve 7) stands by itself in having 35 rows as the usual number. At the other extreme stand the Utah (curve 5) speci- mens, of which only 25 per cent have as many as 31 rows, while the majority have only 29. These Utah snakes are thus quite different from typical P. c. deserticola. In figure 6 are charted the number of urosteges in specimens of these several races. Curves 1 and 3 are of interest as show- ing that P. c. annectens (3) usually has more urosteges than P. c. catenifer or any of the other races. Curve 4 indicates a reduction in the number of urosteges in P. c. deserticola as com- pared with Utah specimens and P. c. heermanni. In work upon groups such as this, where individual variation is so great as almost to conceal geographic differences, I have found that the latter may be thrown into greater relief by what CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Se». /o /J, If- /c /r J.0 jf J.V- j^i. j.r iSL^ •♦■••■•^ f-—: » •*• i \ •••f:-4— i : : : : •f- ■f ■f i •♦- •; i I • ! • ■!■ — * f- ••*■-■■?—.*■■ ♦ . — •♦.••y-..* t : ■ -« •• ■■■• I ••• •I •♦•• ■\ f- "t — T-— *■-? ♦•■ •h-4- •I *- i : \ ■»• — I • ■f- • \ ♦••• "i — r-— |— ■< :• ^. ■•■;.■..• ■;.. • [••■ i i 1 4 1 \ ; : ;.. 4 T *•■•■•• ■* i-f-l f..i....i... .:.. ] ■ 1 ^ ' J ^ ^ : i • 1 *••■•>••••• i-. ♦•■■■•■■ -^ i.y\ j.-i I.- ..,.„.. ■••?••••■?■••• ■f- ...i i : ■-♦ 1 1 » i- -, ...\ •■••i t ! : ..i..-l M : •t — I ; ■ —4 t-—- ! •■■» : ; 1 i :- ■i—f- ■i— •:■■ .♦....!.. -t v > ■ .;....4. 1 : ^^^^ *■■-■♦ ' *■ r----t— f-t ■ t -v t .♦■■i - r- Vou X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES jur M. 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Individual variation in any one specimen rarely affects greatly all of the characters involved in geographic variation. Therefore, by adding to- gether, or subtracting various characters showing geographic differences, the factors of individual variation are proportion- ally reduced or buried, and geographical variations are less concealed. Figure 7 represents this principle as applied to certain charac- ters of the gopher-snakes. The number of dorsal and caudal blotches together has been subtracted from the number of gas- trosteges for each specimen. Certain differences between the subspecies become at once evident. P. c. annectens (curve 3) stands out as very distinct from P. c. deserticola (4) and the snakes of Arizona and Utah (5 and 6). P. c. heermanni (2) is also clearly different from P. c. deserticola (4). The Arizona snakes (6) plainly are not like those of Utah (5), nor like the Lower Californian P. vertehralis (curve 7). Still other differences between various subspecies exist but have not been charted. Thus, the snakes of Utah and Arizona most frequently have but one preocular, vr'hile the other races usually have two. In P. vertehralis the supralabials usually are nine, while eight is the number most frequent in the other races. It seems, then, that instead of the three kinds of gopher- snakes which have been recognized by authors for many years, the facts will be best set forth by according recognition to seven kinds from western North America. The chief differences be- tween these may be briefly set forth in the following table of averages. The complete scale-counts and localities are given in the earlier paper. The counts of the blotches are given here under the head of each subspecies with the numbers of the specimens and of the localities as in the former paper. Vol. X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES 11 § w o w w Q w > < fa o w -3 •^ •«* 1 8 10 CN 0 *i* 1 o\ ^ *^ ^0 fl, <=^ -^f ^ "C Vi a t— t^ rO lO «o «N ^ VD 0 0 '-^ ^ - ^ so "^ a ^-H 0 ~ •<* CN CN ^ ^ •O 0 *^ rr> so i! CN 6< 0 0 «o V> Ov Ov ^M r, 00 ■ 1 "5^ CN fO 0 CN JL, "^ «o 0 1 fO "O '^ ^ OS OS a CO »o •i. cs ■■a 00 0 0 1 d^ ^ 5^ CN ^ JS fN ^ 'b ^ »-H 0 (o PO t^ K (N %j o»- 0 ■* ,v. 00 p o> • ^ -^ ■ I 8 5; 0 00 <^ CN ~" ' t^ CN CN 0 'b 'b ■ ■>► ■ CN -<: 'b -b 0 ■* CN 0 es ^> rc 0 0 '-' CN •^ 8 0 1 <^ ^ • v-H <5 ^ «^ " Cs| M HJ ••-> % 0 (U 2 < £ 0 s ) a 1 s a 1 2 > pq 0 2 12 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Where individual variation is so great in races which inte- grade and are so closely related, it probably is impossible to make a "key" which will properly refer all specimens. How- ever, it is believed that the following synopsis will serve this purpose for most individual specimens, and perhaps for all series of specimens from one locality. Synopsis of Species and Subspecies a. Coloration on anterior half of body not largely red. b. The number of gastrosteges less the number of dorsal blotches on body and tail rarely exceeds 151. c. Gastrosteges fewer (200 to 230), average fewer than 225; urosteges fewer (51 to 80), average in males 70, females 63; scale-rows usually (70 to 75%) not more than 31. d. Dorsal blotches more numerous on body (56 to 93, average 70) and tail (14 to 30, average 21.4). P. c. catenifer. — p. 13 dd. Dorsal blotches fewer on body (48 to 70, average 57.6) and tail (13 to 19, average 15). P. c. heermanni. — p. 16 CO. Gastrosteges more numerous (217 to 243), average more than 225; urosteges more numerous (62 to 85), average in males 76, females 70; scale-rows usually (63%) more than 31. Dorsal blotches numerous on body (54 to 89, average 74) and tail (14 to 29, average 22.8). P. c. annectens. — p. 17 bb. The number of gastrosteges less the number of dorsal blotches on body and tail usually exceeds 151 ; gastrosteges more than 220. e. The sum of the number of scale-rows and of preoculars on both sides of head rarely exceeds 2)Z\ usually one preocular. P. c. stejnegeri. — p. 21 ee. The sum of the number of scale-rows and of pre- oculars on both sides of head usually exceeds Z2>. f. The sum of the number of caudal blotches and preoculars of both sides of head usually exceeds 16; usually two preoculars; posterior dorsal blotches not distinctly reddish. P. c. dcscrticola. — p. 19 ff. The sum of the number of caudal blotches and pre- oculars of both sides of head rarely exceeds 16; usually one preocular; posterior dorsal blotches often distinctly reddish or red-brown. P. c. rutilus. — p. 24 aa. Coloration on anterior half of body largely red. Gastrosteges 233 to 257; scales usually in 35 or 2i2> rows; dorsal blotches few (average 44 on body, 11 on tail) ; supralabials usually nine or more. P. vertebralis. — p. 27 Vou X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES 13 Pituophis catenifer catenifer (Blainville) Gsast Gopher-Snake Diagnosis. — Gastrosteges average fewer than 225 ; urosteges average 64 to 70; scale- rows most frequently 31; preoculars usually two ; supralabials most frequently eight ; dorsal blotches numerous, average on body 70, on tail 21.4; no red in colora- tion. Type locality. — California. Distribution. — The Coast Gopher-Snake occupies a rather narrow strip of territory along the Pacific Coast of the United States from Puget Sound to Santa Barbara County, California. The eastern limit of its range in the far north is not known, but does not include eastern Washington. In southern Oregon, it occurs near Roseburg and in the Camas Mountains, in Douglas County, but not near Klamath Falls, Klamath County, where it is replaced by P. catenifer heermanni. In California, it occupies the coast ranges and valleys east to the western edges of the Sacramento and San Joaquin valleys, where it intergrades with and, farther east, is replaced by, P. catenifer heermanni in the north and P. catenifer deserticola in the south. Still farther south, it probably intergrades with P. catenifer annectens in Santa Barbara or Ventura County. It has been taken in Siskiyou (Ft. Jones, Callahan, Mt. Shasta), Humboldt (Garberville), Trinity (Yolla Bolly Mountain), Mendocino (ten miles south from Willits), Lake (Middletown, Kelseyville, Lower Lake), Sonoma (Petaluma, Duncan Mills, Guerneville, Monte Rio), Napa (Napa), Solano (Buddha Can- yon), Marin (Inverness, Point Reyes Station, Mailliard, Mt. Tamalpais, Lagunitas, Manzanita, San Anselmo), Contra Costa (Walnut Creek, Antioch, Contra Costa, San Pablo Val- ley, Mt. Diablo, Moraga Valley), Alameda (Berkeley, Oak- land, Hayward), San Francisco, Santa Clara (Palo Alto, Stanford University, Sunnyvale, San Jose, Los Gatos, Alma, Coyote, Coyote Creek), Santa Cruz (Glenwood, Corralitos, Soquel), San Benito (San Juan), Monterey (Monterey, Car- mel, Bradley, Soledad, Coburn, Welby, Metz), San Luis Obispo (San Miguel, Pismo, Edna, Indian Creek, San Juan 14 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. River, source of the Salinas River), and Santa Barbara (Santa Cruz Island) counties. Variation. — One specimen has no loreal plates ; the other one hundred and three all have the normal 1-1. The preoculars are 2-2 in seventy-nine, or 78% ; 1-1 in seventeen, or 16% ; and 1-2 in six, or 6%. The postoculars are 3-3 in seventy-one, or 69% ; 3-4 in eighteen, or 17% ; 4-4 in eleven, or ll'^*' ; 4-5 in one, or 1% ; 2-3 in one, or 1% ; and 2-2 in one, or 1%. The temporals are 4—4 in thirty-nine, or 38% ; 3-3 in twenty-six, or 25% ; 3-4 in twenty-one, or 20% ; 4-5 in nine, or 9% ; 2-3 in three, or 3% ; 2-2 in three, or 3% ; 5-5 in one, or 1% ; and 2-4 in one, or 1%. The supralabials are 8-8 in fifty-six, or 54% ; 8-9 in twenty-seven, or 26% ; 9-9 in fourteen, or 14% ; 9-10 in three, or 3% ; 7-8 in two, or 2% ; 10-10 in one, or 1%. The infralabials are 13-13 in thirty-two, or 31% ; 12-12 in twenty- two, or 21% ; 12-13 in nineteen, or 18% ; 11-12 in nine, or 9% ; 13-14 in seven, or 7%; 11-11 in six, or 6%; 11-13 in three, or 3% ; 10-10 in three, or 3% ; 14-14 in one, or 1% ; and 10-11 in one, or 1%. The scale-rows are 31 in sixty-nine, or 68% ; 33 in twenty, or 20% ; and 29 in twelve, or 12% ; the average is 31.1 rows. The gastrosteges vary in number from 200 to 230, males having from 207 to 230, females from 200 to 230; the average in fifty-four males is 217, in forty-six females is 220. The urosteges vary from 53 to 79 ; males having from 59 to 79, females from 53 to 78 ; the average in fifty-three males is 69, in forty-five females, 64. The dark blotches between head and anus in seventy-five specimens vary from 56 to 93, the average being 70. On the tail, in eighty-three specimens, they vary from 14 to 30, and average 21.4. The counts of blotches in the various specimens are shown below. Vol. X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES 15 Blotches on Blotches on mJ im*k r\A«> Locality Number ■ «««h«ki-x_ta 1^ UlIlL>Cr Body Tail Body 1 L^ocajity ail C 1589 1 43434 64 f i 23 52 C 2314 1 S 1165 53 C 2434 1 40413 63 JO 54 C 1626 66 22 1 S 4026 55 C 848 79 22 1 S 1799 71 A ?7 55 43452 57 18 3 S 1798 69 A J4 55 C 849 56 17 4 S 1808 87 r A 1% 55 43519 70 21 5 S 4017 71 ] 19 55 C 4019 67 20 6 S 1150 63 / A W 55 C 4018 68 21 6 41667 66 f A n 56 C 5614 64 18 7 S 1119 71 19 57 C 4017 59 21 8 S 1118 63 A >1 57 S 4220 15 S 1773 63 15 57 S 1697 66 21 16 S 1168 67 A JO 57 S 1741 70 18 16 S 1117 71 A 21 57 30888 23 17 S 5806 73 17 57 C 5285 74 23 19 S 1749 59 18 57 27326 77 24 20 S 1748 64 A 23 57 C 975 74 22 21 S 1752 67 24 57 C 4845 68 20 22 S 7195 57 C 5283 77 28 23 S 1167 57 C 5282 93 28 23 S 4042 57 43377 67 19 33 S 4047 57 13766 80 23 34 S 4044 57 13767 85 25 34 S 4045 57 13768 78 20 34 S 4043 57 13769 75 22 34 S 1747 57 13770 84 22 34 S 1171 57 17858 68 27 34 S 1169 57 f A 22 57 43375 76 22 35 S 1751 58 ] 14 57 43376 66 19 35 41671 61 A 21 57 43321 72 18 36 43440 69 r A 21 58 43322 69 20 36 S 409?* 73 f A 25 59 43379 69 20 37 S 1681 82 r A 27 60 43373 67 19 38 S 1772 84 r A 26 60 43374 70 24 38 36061 66 1 21 61 C 4312 39 S 1740 62 r A 20 62 43412 71 23 41 43523 71 r A 23 63 39261 62 20 42 27942 74 A 27 64 43274 78 25 44 C 4911 84 \. 50 65 43418 62 22 45 C 4131 74 f A 23 66 43419 72 20 45 C 5284 75 c A 23 68 43364 85 22 46 29492 71 A 21 70 43365 66 22 46 S 4062 71 43413 49 16 47 44194 69 f A 20 73 43414 74 19 48 44943 77 A 23 76 43382 73 22 49 45119 63 41 36120 66 18 50 45120 88 r A 24 77 36121 56 17 50 45121 72 \ 19 77 45131 70 23 50 45127 71 f A 24 78 C 6166 51 16 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Se». Pituophis catenifer heermanni (Hallowell) Valley Gopher-Snake (Plate 1, fig. 1) Diagnosis. — Gastrosteges average fewer than 225 ; urosteges average 60 to 68; scale-rows most frequently 31; preoculars usually two; supralabials most frequently eight; dorsal blotches fewer, average on body 58, on tail 15 ; no red in coloration. Type locality. — Cosumnes River, California. Distribution. — The Klamath region, Oregon, and in Cali- fornia, Modoc County, the Sacramento Valley, the northern part of the San Joaquin Valley, and the western slope of the Sierra Nevada. In Oregon it has been taken near Klamath Falls, Klamath County. Californian specimens have been col- lected in Modoc (Canby, Goose Lake Meadows, Sugar Hill, between Alturas and Davis Creek, Dry Creek in the Warner Mts.), probably Shasta (McCloud River), Tehama (Tehama), Glenn (Fruto, Winslow), Butte (between Live Oak and Grid- ley), Yolo (Grand Island Landing), Placer (Lander near Col- fax), El Dorado (Fyffe, Riverton), San Joaquin (Tracy), Merced (Los Banos, Snelling), Mariposa (between Kinsley and Maculey's Stage Station, Coulterville, Pleasant Valley), Madera (vie. Madera), and Fresno (King's River, Dunlaps, Clovis) counties. Variation. — The loreal is 1-1 in all of the twenty-eight speci- mens. The preoculars are 2-2 in twenty-six, or 93% ; and 1-1 in two, or 7%. The postoculars are 3-3 in twenty-three, or 85% ; 4—4 in three, or 11% ; and 4-5 in one, or 4%. The tem- porals are 4-4 in eleven, or 45% ; 3-4 in five, or 21% ; 5-5 in three, or 13% ; 3-3 in three, or 13% ; 4-6 in one, or 4% ; and 2-3 in one, or 4%. The supralabials are 8-8 in twelve, or 48% ; 8-9 in eight, or 32% ; and 7-7, 7-8, 9-9, 9-10, and 10-10, each rn one, or 4%. The infralabials are 13-13 in twelve, or 50% ; 12-12 in six, or 25% ; 14-14 in two, or 8% ; 12-13 in two, or 8% ; and 12-14 and 11-14 each in one, or 4%. The scale rows are 31 in thirteen, or 46% ; 33 in eight, or 29% ; 29 in six, or 21%; and 35 in one, or 4%; the average is 31.3 rows. The gastrosteges vary in number from 209 to 231, males having from 209 to 231, females from 218 to 231 ; the average in six- Vou X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES 17 teen males is 219, in twelve females, 224. The urosteges vary from 55 to 74, males having from 61 to 74, females from 55 to 66 ; the average in fifteen males is 68, in eleven females is 60. The dark blotches between head and anus in twenty-eight specimens vary from 48 to 70, the average being 57.6. On the tail, in thirty-two specimens, they vary from 13 to 19, and average 15. The counts of the blotches are shown below. Blotches on Blotches on Number Locality Number Locality Body Tail Body Tail C4012 70 23 2 C3608 54 14 27 S 5631 61 19 9 C5595 59 15 28 S 5633 59 23 9 C2080 63 24 29 39637 62 19 10 C2081 60 17 30 27333 63 19 11 C2082 56 19 31 C6264 12 C 2083 56 20 32 C4016 55 16 13 S 6500 40 C4015 57 14 14 43521 48 15 43 C4014 63 21 14 43522 56 15 43 41670 50 16 18 C2759 57 43 C5883 54 24 C4013 63 20 67 C5884 55 17 24 C4011 58 19 69 C5885 53 14 25 20413 65 16 72 C5886 51 17 26 44161 58 18 74 41699 50 15 27 44241 57 18 75 Pituophis catenifer annectens (^aird & Girard) San Diegan Gopher-Snake (Plate 1, fig. 2) Diagnosis. — Gastrosteges average more than 225 ; urosteges numerous, average 70 to 76 ; scale-rows most frequently 33 ; preoculars usually two ; supralabials most frequently eight ; dorsal blotches many, average on body 74, on tail 22.8; no red in coloration. Type locality. — San Diego, California. Distribution. — This subspecies occupies the coast region of southern California and northern Lower California, and has been found on some of the islands off the coast. We have ex- amined specimens from Santa Barbara (Santa Barbara), Ven- tura (Pine Creek), Los Angeles (Charter Oak, Cold Water Canyon, La Crescenta, Pasadena, Mt. Wilson, Sierra Madre), 18 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. San Bernardino (Ontario), Riverside (Colton, San Bernar- dino Mts., Riverside, San Jacinto, San Jacinto Mts.), and San Diego (Warner Pass, Agua Caliente, Cahuilla Valley, Julian, Cuyamaca Mts., Campo), counties, California, and from En- senada, San Martin Island, and South Coronado Island, Lower California. It is probably this subspecies of gopher-snake which has been observed, but not captured, on Santa Catalina Island. Those of Santa Cruz Island, however, are Pitiiophis catenifer catenifer. Variation. — Sixty-nine specimens all have loreals 1-1. The preoculars are 2-2 in forty-four, or 64% ; 1-1 in twenty, or 29% ; and 1-2 in five, or 7%. The postoculars are 3-3 in forty, or 59% ; 4-4 in thirteen, or 19% ; 3-4 in eleven, or 16% ; and 2-3, 2-A, 4—5, and 5-5, each in one. The temporals are 4-4 in twenty-three, or 34% ; 3-3 in fifteen, or 22% ; 3-4 in twelve, or 18% ; 4—5 in five, or 7% ; 5-5 in five, or 7% ; 3-5 in two, or 3% ; and 2-2, 2-3, 2—4, 4-6, and 5-6, each in one. The supra- labials are 8-8 in thirty-three, or 48% ; 9-9 in nineteen, or 27% ; and 8-9 in seventeen, or 25%. The infralabials are 13-13 in thirty-four, or 49% ; 13-14 in ten, or 14%) : 12-12 in eight, or 12% ; 12-13 in seven, or 10% ; 14-14 in three, or 4% ; 11-13 in two, or 3%; and 10-11, 11-11, 11-12, 12-14, and 14-15, each in one. The scale-rows are 33 in thirty-seven, or 54% ; 31 in twenty-five, or 36% ; 35 in six, or 9% ; and 29 in one, or 1% ; the average is 32.4 rows. The gastrosteges vary in num- ber from 217 to 243, males having from 217 to243, females from 218 to 240; the average in forty-three males is 228, in twenty-five females, 231. The urosteges vary from 62 to 85, males having from 62 to 85, females from 62 to 83; the aver- age in thirty-nine males is 76, in twenty-four females, 70. The dark blotches between head and anus in seventy-one specimens vary from 54 to 89, the average being 74. On the tail in sixty-five specimens they vary from 14 to 29, and aver- age 22.8. The counts of the blotches are shown in full below. Vol. X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES 19 Blotches on Blotches on Number Locality Number Locality Body TaU Body Tail 8575 85 23 1 S 1163 37 8678 69 19 2 S 1162 79 27 37 13588 63 19 3 S 4050 71 21 38 13589 63 21 3 40060 69 25 39 S 5163 65 20 17 40061 82 24 39 38918 73 22 18 40062 75 22 39 40003 83 22 19 40063 84 27 39 C4311 73 21 20 40064 82 24 39 C 749 74 23 21 40065 79 25 39 C 750 67 20 21 40066 75 19 39 C4313 78 21 22 40067 77 20 39 C4310 69 21 22 40068 81 28 39 27534 65 20 22 40069 73 23 39 27774 71 18 22 40070 77 24 39 27806 76 25 22 40071 73 19 39 C 75 73 20 24 40072 69 21 39 S 1197 81 24 25 40073 69 22 39 S 1164 71 26 25 40074 79 25 39 S 1784 61 18 25 40075 71 21 39 S 1166 72 22 25 40076 80 26 39 S 1122 65 24 25 40077 78 26 39 S 4008 74 26 25 40078 72 20 39 S 1146 75 26 25 40079 81 39 S 1135 78 29 25 40080 76 24 39 S 1750 68 22 25 40081 79 24 39 C 104 89 29 26 40082 76 19 39 C 553 75 20 26 C 623 75 24 40 C 343 72 24 26 C 622 72 19 41 C 551 87 26 26 S 1160 79 26 42 C 552 78 23 26 vS 1149 77 26 42 S 5240 28 S 1155 73 21 42 S 6464 78 20 29 C1Q40 C38l9 79 28 43 S 4291 81 21 31 71 44 S 4268 80 24 31 43520 80 24 45 Pituophis catenifer deserticola Stejneger Desert Gopher-Snake Diagnosis. — Gastrosteges average more than 230 ; urosteges average 59 to 64; scale-rows usually 31 or 33; preoculars usually two ; supralabials most frequently eight ; dorsal blotches fewer, average on body 56, on tail 15.4; sometimes some orange, but not distinct red, in coloration. Type locality. — The Great Basin and the southwestern deserts. Distribution. — The Colorado and Mohave deserts, the south- ern part of the San Joaquin Valley, eastern San Luis Obispo 20 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Seb. County, the valleys east of the Sierra Nevada in California, and probably nearly all of Nevada; possibly Idaho and eastern Washington. From Nevada, I have examined specimens of this subspecies from Humboldt (Thousand Creek Flat, Virgin Valley, Big Creek Pine Forest Mountains), Washoe (Pyramid Lake, Nixon), Ormsby (Carson), Lander (Austin), Elko (Carlin) and Esmeralda (Palmetto Mountains), counties. Californian specimens examined have been collected in Im- perial (Silsbee), Riverside (Mecca), San Bernardino (Victor- ville, Hesperia), Mono (Benton), Kern (Walker Pass, Tehachapi Mountains, Isabella, Delano, Bakersfield, Button- willow), and San Luis Obispo (Simmler, Pozo, Palo Prieto, Shandon) counties. Variation. — Twenty-eight specimens from California and western Nevada show the following variations : The loreals are 1-1 in all. The preoculars are 2-2 in twenty-four, or 86% ; 1-2 in two, or 7% ; and 1-1 in two, or 7%. The postoculars are 3-3 in twenty-two, or 79% ; 4-4 in five, or 18% ; and 3-4 in one, or 3%. The temporals are 3-4 in eleven, or 39% ; 4-4 in five, or 18% ; 4-5 in four, or 14% ; 5-5 in three, or 11% ; and 2-3, 3-3, 3-5, 5-6, and 6-6, each in one, or 37o. The supra- labials are 8-8 in seventeen, or 61% ; 8-9 in six, or 21% ; 9-9 in four, or 14% ; and 8-10 in one, or 4%. The infralabials are 13-13 in thirteen, or 46% ; 12-12 in six, or 21% ; 12-13 in four, or 14% ; 12-14 in two, or 7% ; 13-14 in two, or 7% ; and 14—14 in one, or 4%. The scale-rows are 33 in eleven, or 39% ; 31 in eleven, or 397o ; 35 in three, or 11% ; 29 in two, or 7% ; and 37 in one, or 4% ; the average is 32.3 rows. The gastro- steges vary in number from 223 to 263, males having from 224 to 252, females from 223 to 263 ; the average in twelve males is 234, in sixteen females, 239. The urosteges vary from 50 to 72, males having from 58 to 72, females from 50 to 67; the average in twelve males is 64, in fifteen females, 59. The dark blotches between head and anus in twenty-eight specimens vary from 46 to 66, the average being 55. On the tail they vary from 12 to 21, and average 15.4. The counts of these blotches are shown in full below. Vol. X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES 21 Blotches on Blotches on Number Locality Number Locality Body Tail Body 1 Tail C1003 52 12 10 43383 66 18 33 C2761 54 15 11 43347 60 18 34 C2760 52 16 11 43429 50 15 35 39553 55 17 12 C2763 53 17 36 43381 56 16 13 C2764 56 17 36 39595 62 21 14 S 5649 50 14 48 38958 56 19 15 C 1529 53 13 49 38959 55 16 15 C1528 51 14 49 C2798 47 13 16 C1274 51 15 50 C3716 46 12 23 C1275 52 14 51 C3715 63 15 23 C1276 54 16 51 C 469 62 16 27 S 53 14 53 C 470 56 15 27 S 54 14 54 C 471 54 12 27 S 6406 61 15 55 36285 64 30 40504 58 17 55 C5365 52 14 32 Pituophis catenifer stejnegeri, new subspecies Utah Gopher-Snake (Plate 2, fig. 1) Diagnosis. — Gastrosteges average more than 230; urosteges average 60 to 66; scale-rows most frequently 29; preocular most often single; supralabials usually^eight ; dorsal blotches fewer, average on body 58, on tail 16.5; no distinct red in coloration. Type. — Cal. Acad. Sci. No. 14203, adult male, collected by Joseph C. Clemens, at Fort Douglas, Salt Lake County, Utah, June to July 4, 1908. Description. — Head somewhat flat-topped, with snout pro- jecting and rather narrow. Temporal regions not swollen. Rostral plate very large, prominent, not very narrow, often re- curved between internasals on top of snout ; bounded behind by internasal, anterior nasal, and first labial plates. Plates on top of head are a pair of internasals, a variable number of pre- frontals (normally four), a frontal, supraocular of each side, and a pair of parietals. Anterior and posterior nasals usually distinct. Loreal usually elongate. Preoculars usually one, oc- casionally two. Postoculars usually three, often two. Supra- 22 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. labials usually eight, often nine. Infralabials usually 13, often 12, sometimes 11 or 14. Temporals of first row varying from two to five, usually four. Genials in two pairs, anterior larger. Scales on body in 27 to 33 rows, usually 29, keeled except in a varying number of rows on each side. Anal plate not divided. Gastrosteges varying in number from 223 to 241, males having from 227 to 241, females from 223 to 240. Urosteges in two series of from 55 to 71, males having from 58 to 71, females from 55 to 62. The ground color is pale brownish or grayish-yellow, some- times more or less obscured by the spreading of the blotches or the presence of black or dark browh marks along the keels of its scales. Along the middle of the back, from the head to a point over the anus, is a series of from 50 to 68 (average 58) dark blotches. These blotches are brown on the central part of the body but are black anteriorly and posteriorly. On the upper surface of the tail are from fourteen to twenty (aver- age 16.5) blackish blotches. On the anterior portion of the body the blotches are more or less rounded, but posteriorly they tend to become quadrate. There are several series of alternating, often more or less confluent, dark blotclies or spots on the sides. Across the top of the head, between the preocular plates, is a moderately narrow brown band, very definite and well-defined. There are similar bands or spots below the cen- ter of the eye and running back and down from the upper post- ocular plate. The top of the head posteriorly has a few, small, scattered, dark spots. The spaces between the dark dorsal blotches on the posterior portion of the body are yellow or orange-yellow, usually somewhat obscured by dark brown streaks along the keels of the scales. The lower surfaces are yellow or yellowish-white with irregular spots or blotches of dark brown or black on the gastrosteges and urosteges. There is no definite median subcaudal black band. Length to anus . . 758 800 863 1028 1125 1125 Length of tail . . . 126 148 168 179 190 207 Variation. — Twenty-nine specimens from Utah show the following variations : The loreals are 1-1 in all. The preocu- lars are 1-1 in twenty-one, or 87% ; and 2-2 in three, or 13% VouX] VAN DENBURGH— VARIATION IN GOPHER-SNAKES 23 of those undamaged. The postoculars are 3-3 in twelve, or 52% ; 2-2 in ten, or 43% ; and 2-3 in one, or 4%. The tem- porals are 3-4 in ten, or 43% ; 4—4 in seven, or 31% ; 4—5 in three, or 13% ; 3-3 in one, or 4% ; 2-3 in one, or 4% ; and 3-5 in one, or 4%. The supralabials are 8-8 in thirteen, or 56% ; 8-9 in five, or 22% ; and 9-9 in five, or 22%. The infralabials are 13-13 in eight, or 38% ; 12-13 in five, or 24% ; 12-12 in four, or 18% ; 11-11 in two, or 9% ; 11-12 in one, or 5% ; and 13-14 in one, or 5%. The scale-rows are 29 in fifteen, or 56% ; 31 in seven, or 26% ; and 27 in five, or 18% ; the average is 29 rows. The gastrosteges vary in number from 223 to 241, males having from 227 to 241, females from 223 to 240; the average in twenty-two males is 233, in five females, 235. The urosteges vary from 55 to 71, males having from 58 to 71, females from 55 to 62; the average in twenty-three males is 66, in five females, 60. The dark blotches between head and anus in twenty-nine specimens vary from 50 to 68, the average being 58. On the tail they vary from 14 to 20, and average 16.5. The counts of the blotches are given in full below. Blotches on Blotches on Number Locality Number Locality Body TaU Body Tail 40961 66 16 56 14194^ 56 14 57 14207 54 14 57 14195 50 14 57 27198 64 17 57 14196 58 15 57 27199 52 15 57 14197 63 19 57 30913 51 16 57 14198 59 16 57 30914 60 18 57 14199 50 16 57 30915 61 20 57 14200 54 19 57 30916 61 17 57 14201 57 19 57 30917 54 16 57 14202 56 18 57 30918 61 16 57 14203 61 16 57 30919 63 16 57 14204 54 14 57 30920 53 18 57 14205 58 16 57 38756 63 18 57 14206 54 16 57 38757 60 14 57 38755 68 17 58 14193 66 18 57 Distribution. — Specimens of this subspecies are at hand from Thompson, Grand County, Wasatch Mountains, Wasatch County, and Fort Douglas, Salt Lake County, Utah. Remarks. — Three snakes from Boise, Ada County, and Blue Lakes, Twin Falls County, Idaho, and one from Wallula, 24 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Walla Walla County, Washington, may belong here rather than with P. catenifer deserticola, but their final disposition must await additional material. They have twenty-nine and thirty-one scale-rows and one or two preoculars (50% each), and gastrosteges from 23 1 to 244. Pituophis catenifer rutilus, new subspecies Arizona Gopher-Snake (Plate 2, fig. 2) Diagnosis. — Gastrosteges numerous, average more than 225 ; urosteges average 57 to 63 ; scale-rows most frequently 33 ; preocular normally single ; supralabials usually eight ; dor- sal blotches very few, average on body 46, on tail 12.5 ; colora- tion often somewhat reddish posteriorly. Type. — Cal. Acad. Sci. No. 33869, adult female, collected by J. R. Slevin at Tucson, Pima Co., Arizona, April 11, 1912. Description. — Head somewhat flat-topped, with snout pro- jecting and rather narrow. Temporal regions not swollen. Rostral plate very large, prominent, not very narrow, often re- curved between internasals on top of snout ; bounded behind by internasal, anterior nasal, and first labial plates. Plates on top of head are a pair of internasals, a variable number of pre- frontals (normally four), a frontal, supraocular of each side, and a pair of parietals. Anterior and posterior nasals usually distinct. Loreal usually elongate. Preocular usually one, oc- casionally two. Postoculars usually three, often four, some- times five. Supralabials usually eight, often nine, rarely 10. Infralabials usually 12, often 13, sometimes 11 or 14. Tem- porals of first row varying from two to five, usually four. Genials in two pairs, anterior larger. Scales on body in 29 to 35 rows, usually 33, keeled except in a varying number of rows on each side. Anal plate not divided. Gastrosteges varying in number from 222 to 258, males having from 222 to 237, fe- males from 227 to 258. Urosteges in two series of from 52 to 68, males having from 57 to 68, females from 52 to 60. The ground color is pale yellow or grayish-yellow, some- times more or less obscured by the spreading of the blotches Vol. X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES 25 or the presence of dark keels on the scales, especially laterally and anteriorly. Along the middle of the back, from the head to a point over the anus, is a series of from 37 to 55 (average 46) reddish-brown blotches. The brown of these blotches be- comes darker and redder posteriorly. These blotches often are margined with black. On the upper surface of the tail are from ten to fourteen (average 12.5) dark reddish-brown blotches or cross bands. On the anterior part of the body the blotches tend to be more or less rounded, posteriorly they are more quadrate, or are wider with concave anterior and posterior bor- ders. There are two or three series of alternating, often more or less confluent, dark blotches or spots on the sides. Across the top of the head between the preocular plates is a narrow brown band, more or less obsolete in adults. There are similar bands or spots below the center of the eye and running back and down from the upper postocular plate. The top of the head is light brownish-yellow, speckled with black. The spaces be- tween the dark dorsal blotches on the posterior part of the body are light yellowish or grayish-orange, usually without dark marks on the keels of the scales. The lower surfaces are yel- low or yellowish-white, with irregular spots or blotches of light or dark brown on the gastrosteges and urosteges. There is no definite median subcaudal dark band. Length to anus 456 1050 "*1115 1130 1140 Length of tail 68 154 153 183 165 Variation. — Sixteen specimens from Arizona show the fol- lowing variations : The loreals are 1-1 in all. The preoculars are 1-1 in ten, or 62% ; 2-2 in five, or 31% ; and 1-2 in one, or 6%. The postoculars are 3-3 in six, or 37% ; 3—i in five, or 31% ; 4—4 in four, or 25% ; and 4-5 in one, or 6%. The tem- porals are 4-4 in seven, or 47% ; 3-3 in three, or 20% ; 3-4 in two, or 13% ; 4—5 in two, or 13% ; and 2-3 in one, or 7%. The supralabials are 8-8 in eight, or 50% ; 8-9 in three, or 19% ; 9-9 in three, or 19% ; 9-10 in one, or 6% ; and 8-10 in one, or 6%. The infralabials are 12-12 in nine, or 60%; 13-13 in three, or 20% ; 13-14 in one, or 6% ; 14—14 in one, or 6% ; and 1 1-1 1 in one, or 6%. The scale rows are 33 in eight, or 50% ; 31 in six, or 37% ; 29 in one, or 6% ; and 35 in one, or 6% ; the average is 32.1 rows. The gastrosteges vary in number from 26 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Sek. 222 to 258, males having from 222 to 237, females from 227 to 258; the average in six males is 227, in nine females, 237. The urosteges vary from 52 to 68, males having from 57 to 68, females from 52 to 60; the average in six males is 63.5, in nine females, 57. The dark blotches between head and anus in 16 specimens vary from 37 to 55, the average being 46. On the tail they vary from 10 to 14, and average 12.5. The counts of these blotches are given in full below. Blotches on Locality Number Blotches on Locality Body TaU Body TaU 34755 17541 17546 17547 C 1824 S 1131 S 1705 S 1714 39 50 40 45 54 55 42 42 10 12 11 13 14 13 13 11 4 5 5 5 6 7 7 7 33869 33870 33447 C C C C C 48 44 54 43 50 37 51 46 13 13 13 12 14 13 14 8 8 9 Distribution. — I have examined specimens of the Arizona Gopher-Snake taken at Yuma, Yuma County, the Colorado River above Bill Williams River, Mohave County, Cave Creek, Maricopa County, Fort Lowell and the Santa Cruz River near Tucson, Pima County, and the Huachuca Mountains, Cochise County, Arizona. Specimens of Pituophis from Ari- zona have been recorded as collected at Oak Orchard, Camp Grant, Wilton Springs, Tucson, Gila River, White River Can- yon, Fort Whipple, Grand Canyon, and at Las Gijas in Pima County. Remarks. — The specimens from Mohave and Yuma counties, and a specimen from Silsbee, Imperial County, California (re- ferred to P. c. deserticola) show more or less intergradation between the Arizona and the Desert Gopher-Snakes. These specimens have very many gastrosteges, while those from ex- treme southeastern Arizona have fewer. Indeed, some of the latter have so few as to indicate intergradation with the gopher- snakes of New Mexico which Ruthven has referred to P. c. sayi, but which may possibly require recognition as a distinct subspecies, P. sayi dlinis (Hallowell). Vol. X] VAN DENBURGH— VARIATION IN GOPHER-SNAKES 27 Habits. — Ruthven states that a large specimen, secured near Tucson about sun-down on August 22, had recently swallowed an adult ground squirrel. These snakes live both on the desert plains near sea level and in mountain canons to an altitude of at least 5,300 feet. Pituophis vertebralis (Blainville) San Lucan Gopher-Snake Diagnosis. — Gastrosteges average moi-e than 240; urosteges average 62 ; scale rows most frequently 35 ; preoculars usually two; supralabials usually nine or 10; dorsal blotches very few, average on body 44, on tail 1 1 ; much red in coloration. Type locality. — "California." Distribution. — The southern half of the peninsula of Lx)wer California, Mexico. Variation. — Fourteen specimens show the following varia- tions: The loreal is 1-1 in all counted (six). The preoculars are 2-2 in thirteen, or 93% ; 1-1 in one, or 7%. The postocu- lars are 3-3 in fourteen, or 100%. The temporals in five speci- mens are 4-4 in three, or 60% ; 3—4 in on«, or 20%, and 4-5 in one, or 20%. The supralabials are 9-9 in seven, or 50% ; 9-10 in five, or 36% ; 8-9 in one, or 7% ; and 8-10 in one, or 7%. The infralabials are 12-12 in seven, or 50% ; 13-13 in three, or 22% ; 13-14 in two, or 14% ; 12-13 in one, or 7% ; and 14-16 in one, or 7%. The scale-rows are 35 in seven, or 54% ; 33 in four, or 31% ; 34 in one, or 7% ; and 31 in one, or 7% ; the average is 34 rows. The gastrosteges in fifteen specimens vary from 233 to 257, the average being 244; two males average 242, four females average 250. The urosteges in fourteen specimens vary from 57 to 67 ; the average being 62 ; two males average 63, and three females, 60. The dark blotches between head and anus in six specimens vary from 39 to 48, the average being 44. On the tail in six specimens they vary from 10 to 12, and average 11. Counts of the dorsal blotches on the body and tail in six specimensare45, 10;39, 11;48, 11; 43, 10; 44, 11; and 45, 12. PROC. CAL. ACAD. SCI., 4th Series, Vol. X VAN DENBURGH] Plate 1 Figure 1. — Pitiiopliis cati'iiifcr hccniiaiiui (Hallowell). Valley Gopher- Snake. Photograph from a living specimen collected five miles south from Madera, Madera County, California, Alay 10, 1920. Figure 2. — Pituophis catciiifcr anncctcns (Baird & Girard). San Diegan Gopher-Snake. Photograph from a living specimen collected near Campo, San Diego County, California, about May 12, 1920. PROC. CAL. ACAD. SCI., 4th Series, Vol. X [VAN DENBURGH] Plate 2 Figure 1. — Piliiophis catcuifcr stcjncgcri \'an Denburgli. Utah Gopher- Snake. Photograph of a Hving adult male from Provo Canyon, Wasatch Mountains, Wasatch County. Utah. r it Figure 2. — Pituophis cafcnifcr rutUus Van Denburgh. Arizona Gopher- Snake. Photograph of a living adult from Huachuca Mountains, Cochise County, Arizona. PROCEEDINGS OP THK CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. X, No. 2, pp. 29-30, pi. 3 August 6. 1920 II DESCRIPTION OF A NEW SPECIES OF RATTLE- SNAKE (CROTALUS LUCASENSIS) FROM LOWER CALIFORNIA BY JOHN VAN DENBURGH Curator, Department of Herpetology The first specimens of the diamond rattlesnake of the Cape Region of Lower California were sent from Cape San Lucas by John Xantus. Cope recorded them, in^861, as Caudisona atrox sonoraensis, but remarked that they were more delicately tinted than Sonoran specimens, the dorsal rhombs being more perfect and their yellow borders brighter. A considerable number of specimens have been collected in southern Lower California in more recent years, and have been recorded usually as C. atrox. The differences in coloration, however, seem to be of such constancy as to make it desirable to regard the San Lucan snakes as a species distinct from both C. atrox and the reddish C. exsul. I, therefore, propose for this snake the name Crotalus lucasensis, new species (Plate 3, fig. 1) Diagnosis. — Similar to C. atrox (Plate 3, fig. 2) but colora- tion brighter, much less punctulate, and with dorsal rhombs more completely enclosed in light borders. 30 CALIFORNIA ACADEMY OF SCIENCES [Paoc. 4th Ser. Type. — Cal. Acad. Sci. No. 45888, collected by Joseph R. Slevin at Agua Caliente, Cape Region of Lower California, July 26, 1919. Distribution. — The southern portion of Lower California, Mexico. Description of type. — Large. Head broad, fiat-topped ; ros- tral higher than wide, in contact with anterior nasal. Two nasals ; two preoculars ; three postoculars ; two loreals. Supra- ocular large, not raised into a horn-like process, separated from its fellow by about six or seven irregular rows of scales. Six- teen and seventeen superior and nineteen inferior labials, the first pair of the latter divided horizontally as in C. exsul. A single pair of genials. About five rows of scales between supralabials and eye. Scales in twenty-seven rows, keeled ex- cept in one or two rows on each side. Gastrosteges 186; urosteges 26. The general color is yellowish-brown, or brownish-yellow, with a series of large, darker brown blotches along the back. These blotches are well defined, are usually enclosed in con- tinuous light borders laterally as well as dorsally, and show little of the punctulate or pepper-and-salt style of coloration so characteristic of C. atrox. The sides are clouded or blotched with brown, more or less indefinitely outlined with light yellow or white. The head is somewhat mottled above. A yellow or white stripe runs across the side of the face from the preocular plates to the mouth. The scales behind and above this light stripe are darker than the ground color and are set off pos- teriorly by a light streak which runs down and back from the corner of the mouth. The tail is grayish with about four to six black cross-bands. The lower surfaces are yellowish-white. Length to anus 1070 mm. Length of tail to base of rattle 90 " PROC. CAL. ACAD. SCI., 4th Series, Vol. X [VAN DENBURGH] Plate 3 '^ %^'% m .' Epiphragmophora traskii willetti, new subspecies (Plate 4, figs. 3a-3c) Diagnosis: Shell depressed-conic, conspicuously umbilicate, the umbilicus deep, permeable to the apex, and having a diame- ter about one-ninth to one-eighth the greater diameter of the shell. Whorls about 6>4 or a trifle less, convex, the last de- scending somewhat in front. Aperture ample and very oblique (45°). Edges of peristome converging and connected by a very thin, transparent parietal callus. Lip but little thick- ened, everted somewhat throughout, but especially at the pillar where it is sufficiently reflected to indent somewhat the other- wise circular outline of the umbilicus. Color varying from near Prout's brown to tawny-olive, be- coming a little paler and yellower in the umbilical region, and with a clear-cut, deep, liver brown band of a width of about 2.2 mm. on the shoulder, bordered below by a light yellowish band (near naphthalene yellow of Ridgway) of about equal width and above by a much narrower, slightly less clear-cut band of the same color. Periostracum somewhat glossy and with a peculiar sheen. Lines of growth very numerous and quite regular. First half whorl delicately hyaline and nearly smooth except for a few weak incremental waves, with a fine weak papillation sometimes superimposed; next whorl and a half very finely and closely granose or wrinkly granose, with numerous, large, elongate, rather distantly spaced papillae superimposed, the latter ar- ranged fairly definitely in retractively curved, very obliquely slanting series; papillae on later whorls nearly, or quite, obso- lete; spiral sculpture consisting of a weakly developed series of Vol. X] BERRY—SOME UNDESCRIBED CALIFORNIAN HELICES 59 incised lines, barely to be detected on the third whorl, then gradually increasing in strength to the penultimate whorl, but again becoming very weak on the upper portion of the body whorl, and nearly, though not quite, obsolete below. Measurements: Maximum Minimum Altitude Diameter Number of diameter diameter umbilicus whorls mm. mm. mm. mm. WillettColl 31.6 26.2 18.8 3.8 6V2 " 31.4 26.6 19.0 3.7 6V2 " 30.3 25.1 18.4 3.3 6V3 " 30.1 25.2 18.5 3.8 6V2 " 29.7 24.5 18.2 3.2 6V3 , " " 29.3 24.2 17.3 Z.7 6V3 " 28.4 23.5 16.1 2>.7 6V2 Berry Coll. 4498... 31.4 26.3 19.1 3.8 6V2 Type 21.1 24.8 18.4 3.8 6V2 Berry Coll. 4498... 29.4 24.6 18.0 3.2 6V3 "... 29.+ 24.3 18.3 3.2 6V3 Type: Cat. No. 4497, Berry Collection. Paratypes in pri- vate collection of George Willett. Type Locality: Alt. 3500 ft.. Pine Canyon, Sespe Creek, Ventura County, California; Geor§;e Willett, March 24, 1919; dead but fresh shells of 10 adults, and 14 juvenals of various stages, both living and dead, found in rock slides. One living and three dead adults taken at same locality in 1916 or 1917. Remarks: The shell of this beautiful snail is easily the largest, finest, and most richly colored of any of the traskii- group. In some respects it reminds one of the large mountain species, petricola, and like it has suffered a considerable reduc- tion in the spiral sculpture usually so characteristic of the snails of this group. Nevertheless I believe its relationships are rather with the true traskii, an opinion which is fortified by the color and general texture of the shell and periostracum, quite different from the light, SonercUa-Wkt gloss of petricola. The majority of the specimens are in an excellent state of preservation and are remarkably constant in their characters. They appear to represent an unusually well marked race, not especially near to any of the previously described subspecies. 60 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Its chief taxonomic features are the large size, depressed spire, wide umbilicus, weakly developed spiral sculpture (especially on the base), rich brown color, and prominent spiral banding. Dead shells of what appears to be the same subspecies are before me from Sespe Canyon above the mouth of Tar Creek, collected by Harold Hannibal, and from Matilija Canyon, col- lected by H. N. Lowe, in May, 1919. Both localities are in Ventura County. Epiphragmophora petricola orotes, new subspecies (Plate 4, figs. 5a-5d; plate 6) Diagnosis: Shell thin, translucent, depressed-conic, con- spicuously umbilicate, the umbilicus deep, permeable to the apex, and having a diameter about one-ninth the greater diame- ter of the shell. Whorls about 5^, convex, the last descending somewhat in front. Aperture oval and very oblique (45°). Edges of peristome converging and connected by a thin, very delicate, parietal callus. Lip but little thickened and only very slightly reflected save at the pillar, where it tends to cover the edge of the umbilicus. Periostracum more or less glossy, often showing quite a high polish. Lines of growth fine and numerous. First half turn weakly radially costate, with a few scattered papillse ; next three-fourths of a whorl finely, closely granulose, with fine, weak incremental costations, and, over all, traces of larger papillae; granulation present to some extent on all remaining whorls, but of diminished importance as compared with the suddenly much increased incremental lines and the papillae ; latter now seen to be ranked, at least primarily, in the usual obliquely retractively slanting series, almost quincuncially ar- ranged, but the appearance of regularity often lost ; maximum development of these papillae attained on the upper surface of the third whorl, still strong on the penultimate whorl, but prac- tically absent from the body whorl except along the suture and within the umbilicus; spiral sculpture very poorly developed, only a few interrupted traces of incised threading being dis- tinguishable on the upper third of the last two whorls, even these becoming entirely obsolete below. 17.8 12.1 2.3 573 17.0 11.4 2.3 57. 20.5 14.0 . . • • • • 18.3 12.5 2.4 57. 17.5 12.+ 2.3 57. 17.2 11.8 2.4 57. Vol. X] BERRY— SOME UNDESCRIBED CALIFORNIAN HELICES 61 Color a warm golden brown, running fairly near a tawny- olive, becoming a little paler and yellower on the base, and with a conspicuous dark (liver brown) band of a width of about 1.5 mm. on the shoulder, bordered above and below by a rather narrower band of a few tints lighter than the body of the shell. Measurements: Maximum Minimmn Altitude Diameter Number of diameter diameter umbilicus whorls mm. mm. mm. mm. Smith Coll.(3700ft.)21.6 Type 20.4 Chace Coll 24.5 Berry Coll. 3988.... 22.1 " ....21.0 Type: Cat. No. 3905, Berry Collection. Type Locality: Altitude 2500 feet, near trail, south fork of Warm Spring Canyon, San Bernardino Mountains, California ; under logs; Allyn G. Smith, December 26, 1917; one specimen. Additional Localities: Alt. 3700 feet, near trail just south- east of summit. Warm Spring Canyon, San Bernardino Moun- tains, California; Allyn G. Smith, "December 26, 1917; two specimens. Alt. 6500 feet, west wall of Bridal Veil Falls Canyon near mouth, above Forest Home, San Bernardino Mountains, Cali- fornia, in talus; E. P. Chace, May 24, 1918; nine fully mature living specimens, several dead and young. (No. 3988 above are part of this lot.) Remarks: This neat little helicoid is practically a miniature race of the large E. petricola Berry ( :16, p. 107), with which alone it would seem to require any special comparison. From this it differs not only in its much smaller size, but also in its thinner shell, more polished periostracum, and still further re- duction of the spiral sculpture. It occurs in the same general region of the San Bernardino Mountains as the typical form, but has only been discovered at localities farther into the moun- tains, at all of which it appears relatively constant and quite sharply separable from petricola. 52 CALIFORNIA ACADEMY OF SCIENCES IP roc. 4th Ser. For purposes of comparison figures of the shell and micro- scopic sculpturing of the type specimen of pctricola are here appended (Plate 4, figs. 4a-4c; plate 5), especially as this species has recently been strangely misunderstood by Bartsch (: 16, p. 612), who referred it without qualification to E. traskii troskii (Newcomb). As a matter of fact very few of the numerous races described by him in the same paper have nearly the claim to separate recognition that petricola has, although I believe with him that most of them will stand. E. petricola is in fact the earliest described prototype of a whole series of southern Californian mountain snails, the exact relation of which to true traskii still remains to be determined. E. ::cchco Pilsbry ( :16), on the other hand, seems quite close to petricola. Both petricola and orotes are distinctly papillose over much of the upper surface. A very young petricola now at hand from the type locality (Cat. No. 3950, Berry Collection) shows that, when perfect, each papilla bears a minute, stubby, hair-like periostracal process. Epiphragmophora petricola sangabrielis, new subspecies (Plate 4, figs. 6a-6c) Diagnosis: Shell low-conic, thin, fragile, rather tumid, um- bilicate; the umbilicus rather narrow, barely permeable to the apex, and with a diameter about one-twelfth the greater diam- eter of the shell. Whorls 5>4, convex, the last swollen and slightly descending in front. Aperture rounded, sometimes slightly flaring, oblique (40°). Edges of peristome slightly converging and connected by a very thin, transparent parietal callus. Lip only slightly thickened ; everted near the pillar so as to indent the circular outline of the umbilicus. Periostracum somewhat glossy, often with a strong satiny sheen or semi-iridescence. Lines of growth numerous and fairly strong, though somewhat irregular. First half whorl when unworn showing rather strong, more or less interrupted, incremental wrinkles, and traces of a strong, coarse, overlying papillation; succeeding turns very finely wrinkly-granulose be- neath the retractively slanting lines of small and at first often nearly obsolete papillae, the latter increasing in strength to the \'0L. X] BERRY— SOME UNDESCRIBED CALIFORNIAN HELICES 63 penultimate whorl where they are always strongly evident as well as within the umbilicus and to a less degree over the region just behind the aperture on the body whorl ; papillae elsewhere on the last whorl more weakly developed. Spiral sculpture obsolete, a few weak traces persisting on the upper surface and peripheral region of the body whorl only. Color light golden brown near buffy citrine, paler and with more of a yellow tone below, with a dark, liver brown band of a width of about 1.0-1.5 mm. on the shoulder, bordered above and below by a rather narrower band slightly lighter in tone than the body of the shell. Measurements: Maximum Minimum Altitude Diameter Number of diameter diameter umbilicus whorls mm. mm. mm. mm. Type 26.3 21.0 15.7 2.2 5V., Paratype, Willett Coll 23.8 19.1 13.8 2.3 5V3 Type: Cat. No. 4848, Berry Collection. Paratypes in the collection of George Willett. (Neither specimen quite fully mature. ) Type Locality: Monrovia Canyon, San Gabriel Mountains, California; George Willett, March, r5l9; 14 specimens, for the most part not quite mature. Additional Localities: In addition to the lot from which the type was selected, the following specimens before me are pos- sibly referable to the same subspecies. They are at any rate very close, although the material is still inadequate for entire certainty. Millard's Canyon, north of Pasadena, San Gabriel Moun- tains, California; E. P. Chace, March 11 and June 3, 1917; one living adult, one living juvenal, seven dead of various ages. Eaton's Canyon, north of Pasadena, San Gabriel Mountains, California; E. P. Chace, September 3, 1917; two living adults, six dead of various ages. West fork San Gabriel River "just below the divide", San Gabriel Mountains, California; E. P. Chace, September 1, 1918; five dead shells. 64 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Remarks: This mountain race appears to be somewhat similar to Bartsch's avus in shape, size, and the narrow um- bilicus, but differs in the zvcak papillation of the upper surface, and the presence of a weak spiral sculpture. From zcchce Pils- bry, it is distinguishable by its thinner, more tumid shell, much narrower umbilicus, and the better developed papillation of the upper whorls. None of the other described races appears to require any special comparison. LITERATURE Bartsch, P. :16. The Californian land shells of the Epiphragmophora traskii group. Proceedings United States National Museum, v. 51, pp. 609-619, pis. 114-117, December, 1916. Berry, S. S. :16. Three new Helices from California. University California Pub- lications, Zoology, V. 16, pp. 107-111, January 5, 1916. :19. Three new alpine Vertigos from California. Nautilus, v. ZZ, pp. 48-52, text f. 1-8, October, 1919. Binney, A. '43. Descriptions of two undescribed species of North American Helices. Boston Journal Natural History, v. 4, pp. 360-362, pi. 16, f. 17, pi. 20, 1843. Newcomb, W. '61. Descriptions of new shells. Proceedings of the California Academy of Natural Sciences, v. 2, p. 91, February 4, 1861. Pilsbry, H. A. :13. Note on a new variety of Epiphragmophora tudiculata. Nau- tilus, V. 27, pp. 49-50, pi. 3, f. 15-17, August-September, 1913. :16. A new Californian land snail. Nautilus, v. 29, pp. 104-105, pi. 3, lower figs., January, 1916. 66 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Sek. EXPLANATION OF PLATES (Note: Figs. 4a-Sd on Plate I\', and Plates \' and VI, are from photographs by John Howard Paine. The remaining figures on Plate IV are from photographs by Berton W. Crandall.) Explanation, Plate IV (All figures natural size.) Figs, la-lc. Epiphragmophora tudiculata allyniana, new subspecies. Anterior, apical, and basal views of type specimen, from Jasper Point, Mariposa County, California. Figs. 2a-2c. Epiphragmophora traskii chrysoderma, new subspecies. Anterior, apical, and basal views of type specimen, from South Coronado Island, Lower California. Figs. 3a-3c. Epiphragmophora traskii ivilletti, new subspecies. Anterior, apical, and basal views of type specimen, from Pine Canyon, Sespe Creek, Ventura County, California. Figs. 4a-4c. Epiphragmophora petricola Berry. Anterior, apical, and basal views of type specimen, from Mill Creek Canyon, San Bernardino Mountains, California. Figs. 5a"5d. Epiphragmophora petricola orotes, new subspecies. An- terior, apical, basal, and lateral views of type specimen, from Warm Spring Canyon, San Bernardino Mountains, California. Figs. 6a-6c. Epiphragmophora petricola sangabrielis, new subspecies. Anterior, apical, and basal views of type specimen, from Monrovia Can- yon, San Gabriel Mountains, California. PROC. CAL. ACAD. SCI., 4th Series, Vol. X ['.BERRY] Plate 4 2.d 5a E-b 4-b 5b 0" 54 eb 68 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Explanation, Plate V Fig. 1. Epiphragnwphora petricola Berry, type. Portion of upper sur- face of last whorl about a quarter of a turn behind the aperture, showing periostracal sculpturing magnified about 15 diameters. Fig. 2. Epiphragviophora petricola Berry, type. Portion of basal sur- face of last whorl about a quarter of a turn behind the aperture, magnified about 15 diameters. PROC. CAL. ACAD. SCI., 4th Series, Vol. X [BERRY] Plate 5 70 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Explanation, Plate VI Fig. 1. Epiphragmophora petricola orotes, new subspecies, type. Por- tion of upper surface of last two whorls a short distance behind the aper- ture, magnified about 20 diameters to show the periostracal sculpturing. Fig. 2. Epiphragmophora petricola orotes, new subspecies, type. Por- tion of basal surface of last whorl just behind the aperture, magnified about 20 diameters. Fig. 3. Epiphragmophora petricola orotes, new subspecies, type. Por- tion of apical region, magnified about 20 diameters to show the perios- tracal sculpturing. (All photographs used on this plate reversed.) PROC. CAL. ACAD. SCI., 4th Series, Vol. X [BERRY] Plate 6 F'ROCEEDINOS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. X, No. 9, pp. 71-75, 1 text figure. December 29, 1920 A NEW GENUS AND SPECIES OF GRASSHOPPER FROM CALIFORNIA BY Morgan Hebard In the spring of 1917 (April 22-27), Mr. E. P. Van Duzee, curator of Entomology, Museum California Academy of Sciences, collected at Bryson in Monterey County, California. Among the material taken was a single specimen of grasshop- per, which seemed to represent an undescribed genus and species. Upon being informed of this fact, Mr. Van Duzee again visited Bryson, May 16-23, 1920, in order to obtain, if possible, additional material of the species. After considerable effort he was able to locate the habitat, and secured three additional specimens, all females. Esselenia,^ new genus The present genus shows a combination of characters which makes it most difificult to place. The form of the insect is more robust than that of any other North American Acridid ; in fact, it is of a type C|uite similar to that usual in the Ommexechinse. The head is as full, with face as perpendicular, as in Stira- pleura, the pronotum showing a generally similar construction of the lateral carinae. The lateral foveolfe of the vertex are, however, not visible from above, in this feature agreeing with Mesochloa and Phliho stroma, the vertex itself being as broad as in the latter genus and intermediate between them in show- ing a subobsolete medio-longitudinal carinula. The antennse ^ The small Esselenian tribe of Indians once inhabited the region from which this genus is known. 72 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th. Ser. are relatively heavy and flattened, much as in Mesochloa and Cordillacris, though not showing the slightly greater flattening proximad found in those genera. The pronotum, with caudal portion of disk produced mesad and laterad and margin between concave, is distinctive. In Mesochloa and Psoloessa alone do we find mere traces of con- cavity of the lateral portions of the caudal margin of the pro- notal disk. The inner spurs of the caudal tibiae are almost equal in length, in this agreeing with Cordillacris, but not as long and slender as in that genus. The ovipositor valves are very short, as in all the other genera referred to above. Taking all into consideration, we believe that Esselenia should be placed after Phlibostroma, though further study of the proper order of the related genera may lead to its being placed near Stiraplciira. The system generally in use for the linear arrangement of the genera of the Acridinae is clearly un- satisfactory, but can not be rectified until extensive studies of the subfamily are made. Genotype. — Esselenia vanduzeei, new species. Figure 1. — Esselenia vanduzeei, new species. Female. (X3) Dorsal view of type. Vol. X] HEBARD—A NEW GENUS AND SPECIES OF GRASSHOPPER 73 Generic description : Size medium small, form extremely ro- bust for the Acridinae. Head large ; face vertical ; vertex wide, its surface weakly concave and showing a subobsolete medio- longitudinal carinula, its lateral margins slightly raised and carinate; lateral foveolas not visible from above; frontal costa decidedly concave throughout between the strong lateral carinae. Eye rounded, scarcely deeper than broad, more nearly circular in outline than in Stirapleura and much less elongate than in any of the other genera here discussed. Antennae short, moderately heavy, flattened. Pronotum with a distinct and percurrent medio-longitudinal carina, cut only by the principal sulcus; disk bounded laterad by distinct but irregular carinae which are weakly convex-convergent to first sulcus, thence straight and strongly divergent. Caudal margin of lateral lobes and of each half of disk broadly concave, so that the pronotum is roundly produced at each shoulder and more strongly so mesad on the disk. Tegmina represented by large lateral pads. Ovipositor very short. Interspace between metasternal lobes of female slightly over twice as broad as long. Dorsal surface of caudal tibiae with eight to ten spines on external margin. Internal spurs of caudal tibiae heavy, the ventral spur being only slightly longer than the dorsal. Esselenia vanduzeei, new species We refer to the figure of this distinctive insect, as we know of no other species showing sufficiently close agreement to enable us to make a brief comparative analysis. Type: Female, No. 736, Museum California Academy of Sciences; Bryson, Monterey County, California, April 27, 1917; E. P. Van Duzee, collector. In addition to the characters described in the generic treat- ment, we would note the following : Lateral bounding carinae of vertex straight, very narrow and diverging from the sharply rounded apex, forming nearly a right angle, to a point above the lateral ocelli ; thence, for an equal distance, straight, broader and moderately converging to the point where they disappear into the rounded occiput. Cephalic margin of pronotal disk weakly convex, appreciably less than one-half the distance between the extremities of the caudal margin ; least width of disk about four-fifths the cephalic width. Surface of pronotal disk weakly tectate, moderately 74 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th. Ser. wrinkled, this decided toward the lateral carinse. Lateral lobes of pronotum with greatest depth equal to dorsal length; cephalic margin weakly convex and scarcely oblique to the ob- tuse-angulate, sharply rounded ventro-cephalic angle, ventral margin broadly convex to the rather broadly rounded, rect- angulate, ventro-caudal angle; caudal margin broadly concave; sulci deeply impressed on lateral lobes. Tegmina represented by broadly lanceolate pads with rounded apices, which are attingent (or weakly overlap), nearly as long as combined length of head and pronotum. Veins and cross-veinlets of tegmina heavy and distinct, forming a coarse and irregular network, in which, however, the principal veins, and particularly the humeral vein, are clearly defined. Short ovipositor valves with heavy apices sharply curved. Caudal femora heavy, the ventral and particularly the dorsal portions evenly and broadly lamellate, the lamellation of the dorsal margin being over half as wide as the external pagina. Measure»ients (in miUimeters) Female Length of body Length of pronotum Cau of idal width pronotal disk Length of tegmen Width of tegmen Length of Width of caudal caudal femur femur Type Paratype Paratype Paratype 20.2 22.2 20.2 20.8 5.6 6. 5.6 5. 4.8 5.3 5. 4.4 8.1 8.1 8. 7.6 3.8 4. 3.8 4. 12. 3.5 12.1 3.9 11.7 3.7 11.8 3.7 General coloration (type, intensive) dark chestnut brown. Head paling to hazel on occiput, cheeks tawny, this continued as a U-shaped band from eyes, its lower portion crossing the labrum, frontal costa and ventral portion of infra-ocular sulcus suffused with black. Antenn?e hazel, deepening to chestnut brown distad. Pronotum with median carina russet, bordered by a band of mars brown, lateral carinae buffy tinged with rus- set, this broadening into a band on the metazona, the triangular areas between these and the medio-longitudinal band velvety blackish brown. Lateral lobes of pronotum irregularly paling to tawny meso-proximad and with a buffy callosity mesad which extends as a narrow line ventro-cauded to the caudal margin above the ventro-caudal angle. Tegmina pecan brown, the veins of the dorsal field suffused with black. Ventral sur- face of abdomen argus brown. Cephalic and median limbs hazel, mottled with chestnut brown. Caudal femora light ochraceous-tawny in proximal two-fifths, this terminated in external face of dorsal surface by a large triangular patch of Vol. X] HEBARD—A NEW GENUS AND SPECIES OF GRASSHOPPER 75 velvety blackish brown, this surface flecked proximo-dorsad with this color also ; remaining distal portions pecan brown, suffused in genicular areas with blackish. Caudal tibise oppo- site genicular areas of caudal femora black, beyond this show- ing a broad buffy annulus, particularly distinct on the inner surface, remaining portions buckthorn brown, suffused distad with mummy brown ; proximal internal spines mummy brown, other spines and spurs buckthorn brown, tipped with mummy brown. One of the paratypes is similarly colored, except that the general coloration is much paler, walnut brown. Another has the color pattern much more strikingly defined, the blackish areas being as dark, but the medio-longitudinal band of head and pro- notum light ochraceous-tawny, the tegmina ochraceous-tawny except in the sutural half of the dorsal section and the proximal portions of the caudal femora light ochraceous-salmon. The remaining paratype is almost uniformly sayal brown in colora- tion, the dark triangle dorsad on the caudal femora remaining as the only trace of color pattern. In addition to the type, three female paratypes are before us, bearing the same data, but taken May 18, 1920. One of these is the property of the California Academy of Sciences, the other two are in the author's collection. In securing these paratypes, Mr. Van Duzee had some diffi- culty in locating the proper environment in which to search for the species. They were taken in the same canyon as the type, on the grassy or sandy borders of a small mountain stream. At the time there was no running water in the stream, but little pools with level sandy flats between and it was on one of these flats that all were located. The exact spot is in the canyon back of the Bryson school house, about two miles east of the Nacimiento River and the same distance north of the San Luis Obispo County line. The species probably reaches its greatest adult abundance in the early spring. We take pleasure in naming this singular species in honor of the ardent student and collector who discovered it and who has subsequently, not without considerable difficulty, secured addi- 'tional material and valuable data as to the immediate environ- ment in which it occurs. F'ROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. X, No. 10, pp. 77-117, pis. 7-9 February 12, 1921 X COLOR CHANGES AND STRUCTURE OF THE SKIN OF ANOLIS CAROLINENSIS BY CHARLES E. VON GELDERN The mechanism of the color changes in the lacertilia has been the subject of much investigation, especially in that of the African Chameleon. One need only refer to the works of Briicke, Keller, and Fuchs for historical resumes. Briicke, Pouchet, and Keller have attempted to solve this problem by a close study of the histological structure of the skin and Keller was able to demonstrate clearly the various elements essential for the production of the various color states. The histological structure of the skin of Anolis has, as far as I have been able to determine, been studied only by Carlton, who attempted to correlate the findings of Keller in the chameleon with those in Anolis. The Anolis carolinensis , or so-called Florida Chameleon, belongs to the family Iguanidae and is in no way related to the true chame- leons. Its habitat is the southeastern part of the United States and Cuba. It may be distinguished from all other North American lizards except the Geckos, according to Ditmars, by the expanded and flattened adhesive pads on the middle four phalanges of each foot. It is entirely insectivorous, subsisting mainly on flies and meal worms, which it is able to capture with almost unerring accuracy. Water in the form of scattered droplets is lapped by means of its thick tongue and these lizards soon become dehydrated and die if water be not supplied in this form. A characteristic flattened, semicircular projection of the skin, known as the throat-fan (Ditmars), dewlap or gular appendage 78 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. (Gadow), is produced at will in the mid- ventral region of the head and neck. This fan is produced by the hyoidean apparatus. The mechanism of this apparatus consists of a double, tapering car- tilage lying in the mid-ventral line and attached to the body of the hyoid just anterior to a line through the center of the eye. When erected, it carries the loose skin of the cervical region with it. This throat-fan is, according to Ditmars, purely ornamental and produced only by the males, being accompanied by a vigorous nodding of the head and neck. It is produced when a male spies a female or when it prepares for combat with another male. When the males are captured and held in the hand, this fan becomes prominent. Often when two males meet, each one will erect a ridge along the mid-dorsal line extending from the base of the occiput to a variable distance along the back. This may be accompanied by a marked lateral com- pression of the thorax and abdomen so that the lateral diameter is smaller than the dorso-ventral, whereas under ordinary condi- tions the reverse is true. The throat-fan is usually very prominent during this state and there is present a characteristic coloration which will be described later. The Anolis lives well in captivity when supplied with water in the form of scattered droplets and flies or meal worms. It soon becomes apparently very tame and will take insects from the fingers of its captor. Observations of Color Changes in the Living Anolis A general fact impresses itself after one has captured and observed many of these lizards in captivity, namely, that the range of variation in the color of the skin is by far greater in those animals which have been freshly captured. After a few weeks of captivity, although seemingly in perfect health, the color changes become less complicated and less brilliant. This observation may tend to explain the differences noted by different observers as to the color changes. Ditmars states that the color varies from different shades of brown to emerald green and that although these are the common hues of Anolis, other hues are striking, namely, golden yellow and slaty gray with the peppering of white spots over the back. These colors he believes occur during the transition from the two extremes, namely, brown and green. Carlton states that Vol. X] VON GELDERN—SKIN OF ANOLIS CAROUNENSIS 79 he has never seen any changes other than the different shades of brown to emerald green and bases his assirmption that the color changes are much more simple in Anolis than in the true chameleon, on the observations of Lockwood. To those observing these lizards in their natural environment, it is evident that the variation is not so simple as has just been noted. A slaty gray with no element of brown or green is of fairly common occurrence, as is also straw yellow. Furthermore over certain areas of the body the colors undergo even a greater variation than is ascribed to them by Ditmars. Over the mid-dorsal region, for instance, there is present in many of these lizards a narrow stripe of two or three millimeters in •width, extending from the cervical region to the sacrtim or even along the tail for a variable distance, in which further variations may be noted. That this stripe varies in different individuals is probable, and it may even be absent. Nevertheless, it is so char- acteristic of many specimens that a description of its changes should not be neglected. A bright pink color is often present along the stripe which may become darker until it assumes a brick-red color conveying the impression that there is placed there a thick pigment which does not belong to the skin. This stripe may also show a cream color or white, containing a faint suggestion of yellow or brown. This light color is most often observed in those lizards which have assumed the brown hue else- where on the body. When the mid-dorsal stripe assumes either a pink or a white stripe it is irregular at the edges and these irregular edges are dark brown. A less irregular, black stripe is often observed in the green state. I have been unable to discover any rule for the appearance of the dorsal stripe in its various states. It may be present or absent, brick-red, pink, white, or black in different lizards living under the same conditions, nor does the color state of the rest of the body influence the appearance to any extent. I believe that this stripe may appear at some time in all the animals in which it is absent, but that its appearance entails considerable change in the structiire of the skin so that its production must necessarily be a slow one. On the sides of the maxilla, posterior to the eye there may be present a black, quadrangular patch, measuring in large males about two by three millimeters. When present, this patch is of shiny black appearance and differs markedly from the rest of the 80 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. body even in the dark brown state. This patch appears during the time when two males are preparing for combat and is asso- ciated with the appearance of the mid-dorsal ridge, the lateral flattening of the abdomen and thorax, the extension of the legs so that the body is raised off the ground, and by a peculiar greenish mottling of the skin. The altered appearance of this animal pre- paring for combat, in color, form and action, shows such marked changes, that it is difficult to associate it with the animal under ordinary conditions. Even the eyes, which ordinarily are fairly prominent, recede so that the palpebral fissures show only the pupils. The movements are slow and awkward and the body may sway from side to side in a most fantastic manner. In general, the peppering of white spots on the dorso-lateral aspect of the body is characteristic, especially during the brown state, and these spots may be confluent on the lateral aspect of the cervical region. The lower border of the maxilla, the entire man- dible, and the ventral aspect of the entire body varies from a snowy white to dirty brown, gray, or peppered with black dots. The throat -fan when extended is a brilliant pink or vermilion and over it are scattered many white spots. When relaxed, the skin of the throat region is somewhat cream colored or white with reddish streaks at times. The following table indicates the various colors noted in different specimens at the same time and in the same specimen at different times: Table Showing Range of Colors in Various Areas OF THE Body I Dorso-lateral aspect: A Diffuse: — a. Golden yellow to straw yellow. b. Emerald green to dirty bluish green. c. Slaty gray of various shades. d. Light brown to dark mahogany brown. B Mottled:— a. Yellow with irregular patches of green. b. Yellow with irregular patches of brown. c. Emerald green or pea green with irregular areas of darker green to brown. d. Green or brown (usually the latter) peppered with white or hght turquoise blue spots. Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 81 II Mid-dorsal stripe: a. Pink with irregular brownish border. b. Brick-red with irregular brownish border. c. White or cream color with irregular brownish border, d. Black with fairly regular border. e. Brown or green matching surrounding skin. III Post-orbital patch: a. Black. b. Various shades matching the rest of the body. IV Ventral surface of the body : a. White. b. White with scattered black spots. c. Dirty brown or gray. On comparing this table with that of Briicke for Chamceleon vul- garis, one notices certain differences. For instance, pale flesh colors, lilac gray, steel blue and purple are not present in Anolis, but the greater simplicity which Carlton would attribute to the latter is questionable. Undoubtedly differences exist but the changes must be just as complicated in one as in the other. Causes of Color Changes The most evident general factors influencing the color states are light, absence of light, temperature, and various external stimuli. Carlton found that with few exceptions the brown state was brought about from the green state in four minutes by exposure to sunlight. Absence of light changed the brown to green in twenty- five minutes. With specially constructed containers, he found that, with few exceptions, if part of the body of a green lizard, either head or body, be exposed to direct sunlight, while the rest remained in the dark, the entire body became brown. Carlton concludes that there are nerve terminals in the skin which are directly sensitive to light and which, when stimulated in one area, send impulses which reach the efferent nerve endings of the skin over the entire body. Spinal cord section in no way changed the results, indicating that the action is either wholly reflex in character, or that spinal nerves do not necessarily influence the changes. Carlton was able to induce the green state by in- jections of .001% nicotin solution in small quantities and this change from brown to green was brought about in one minute. This suggested to him that this change is under the control of the S3anpathetic nerves. 82 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. He also found that the green state could be produced in three ways, namely, by subjecting the animals to the absence of light, by inhibiting the blood circulation, and by cutting off the nerve supply. Inhibiting the circulation, he found, was a more important factor than cutting off the nerve supply, in that it brought about more rapidly the green state and, furthermore, when both factors acted simultaneously, still greater rapidity in change occurred than when either one acted alone. Carlton believes that the green state represents the unstimulated state of the skin, which is suggested by the fact that ether narcosis, nicotin poisoning, and death are associated with the green state. The brown state, he believes, is brought about by stimulation of the nerve endings and represents "the state maintained through tonus established by the s>Tn pathetic nerves and dependent upon stimulation of the nervous end organs in the skin by light." Parker and Starratt, repeating Carlton's experiments on the rapidity of change from one color state to another, obtained results that were not uniform and found that changes would occur more rapidly on one day than on another and even at different times during the same day. By means of a constant temperature apparatus which could be illuminated at 115 candle-meters and at the same time brought from io°C. to so°C., they found the average length of time at various temperatures at which either the brown or green state could be produced from the opposite color state. They found that at io°C., the skin remained brown in either light or dark, but as the temperature was raised to 2o°C., the animals placed in the dark became green in 19.66 minutes. At 25°C., under the same conditions, the change took place in 13.23 minutes; at 3o°C., it took 10.93 minutes; at 35°C., 15.48 minutes. At 4o°C. to 45°C., the skin remained greenish gray to green in both light and darkness. On the other hand, when green lizards were placed in the light at 2o°C., the brown state was brought about in 4.23 minutes; at 25°C., 3.52 minutes; at 3o°C., 3.13 minutes, and at 35°C., 2.8 minutes. These investigators believe that at intermediate temperatures, namely, between 2o°C. and 3S°C., light is the controlling factor but that temperature is effective over this range is evident in that it may influence the rate of change. Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 83 Parker, in experimenting with Phrynosoma regale, found that the claw-like scales which fringe the lateral edge of the body be- came white when the animal was subjected to a temperature of 32°C. and placed in the dark, and when placed in the light these claw-like scales became almost black. At i9°C., these scales became black in 15 minutes when the animal was placed in the light, and they became white in 30 minutes in the dark. At i5°C. light again caused black, while darkness brought about a light color, but not white. From these results he concluded that a low temperature favored the production of the black state, whereas a higher temperature, the white state. Thus light produces in this animal, as in Anolis, a dark state, while its absence brings about the light state. Parker further believes that even in Stellio, Uromastix, and Veranus, which have always been considered to have a reversal of the light reaction, in that light causes just the reverse effect as in Anolis, namely, the production of light coloration in the light and dark coloration in the dark, that the apparent reversal is really a temperature effect and not a true reversal of the effect of light. Parker and Starratt mention the observations of Doctor Caswell Graves who stated that in the neighborhood of Beaufort, N. C, on hot, sunny days about as many green lizards as brown ones may be captured. These results are explained by Parker and Starratt by considering that some of tlfe animals are more sensitive to light than to heat and thus become brown, while others are more sensitive to heat than to light and become green. I do not believe that this explanation suffices, for if one notices individual lizards for a considerable length of time, one is struck by the frequent and rapid changes from green to brown and back again to green, apparently regardless of temperature and light. How much influence the otherwise varying nervous conditions exert on these color changes, it is difficult to say for it would seem almost impossible to control them. I have watched animals which were sunning themselves and apparently undisturbed and quiet undergo these changes in a rhythmic manner. Rapid changes are also frequently noticed when an Anolis changes slowly from one object to another, the change occurring while the transfer is being made. Redfield, after nimierous carefully checked experiments on Phrynosoma cornutum, was able to verify the conclusions of Parker, 84 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Carlton, Starratt, and others that the daily rhythmic changes of color are produced by the direct action of light and heat upon the melanaphores. He further states that Phrynosoma adapts itself to its surroundings, namely, if placed on a substratum of white sand it slowly assumes a light coloration irrespective of light or heat and if placed on a substratum of cinders it slowly takes on a dark coloration. He concludes that the color adaptation depends upon stimuli received through the eyes. He was able to bring about a pale coloration in various ways, such as forcibly opening the mouth or by the application of a weak faradic current to the mucous membranes of the mouth or cloaca. He believes that this proximal migration of the pigment may be brought about in two ways, namely, by nervous impulses which stimulate the melanophores through the sympathetic nervous system or by secretion of a hormone (adrenin) from the adrenals. The impulses are carried from the mouth or cloaca along the spinal cord to a center situated between eighth and thirteenth vertebrae and thence by sympathetic fibres to the adrenals. The stimulated glands secrete adrenin which is taken up by the blood stream and acts directly on the melanophores causing a proximal migration of the pigment. Redfield concludes from the fact that adrenalin produces proximal migration of the pigment in Anolis and from the work of Carlton, that impulses through the autonomic nervous system cause a distal migration of the pigment, that the melanophores of Anolis must possess a double innervation from two divisions of the autonomic nervous system. That this is possible he shows by analogous tissue, namely, the smooth muscle, the latter one "known to be innervated by antagonistic fibres belonging to two morpho- logically distinct parts of the autonomic nervous system." The explanation for emotional manifestations in Phrynosoma, Anolis and other animals is readily explained by his conclusions regarding the secretion of adrenin. When first placed in captivity the brown and green lizards in the same cage are about equally divided, but after remaining in captivity for a few weeks the greater proportion become brown in the daylight and the green produced by the absence of light has lost its former brilliance. Much has been written about the true chameleon and its adap- tation of color to its surroundings. Keller found, after placing speci- mens of Chamcsleon vtdgaris in a green house, that in a short space Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 85 of time he was able to find them only after a most careful search, in spite of the fact that when found they were often in plain view. However, he does not believe that the surrounding color plays any role but that other factors, which he did not attempt to explain, bring about these changes. Ditmars states that there is no rela- tion between the color of Anolis and its siuroundings. One must have great temerity to deny such a statement, but I have noted adaptations to the surroundings in Anolis which seem to be more than accidental. For instance, I have noticed that on dark brown fence rails which contained small areas of green lichen, some of the lizards resting on them assimie a dark brown color with irregular patches of brilliant green. In other words, a mixed state is often evident and the effect produced resembles fairly closely the surroundings. Almost invariably the lizards seen on the tnmks of the palm trees in New Orleans are brown and are often detected with great difficulty. The table below represents the findings on May i6, 191 7, from 11.30 A. M. to 1.45 p. M., during which time the temperature was 2 5.5°C. The environment is stated, as well as the number of lizards noted thereon, and the intensity of their color state. G repre- sents green and B brown. The sign + + + represents the greatest intensity of either green or brown, namely, either emerald green or mahogany brown, -f + represents a less intense color but still quite marked, while + indicates the least degree of intensity but one in which one is able to definitely st^e the color as being either green or brown. TABLE 2 Number of green lizards Total G G G G +++ ++ + Green foliage 0 2 6 8 Dark green foliage 1 0 1 2 Concrete 2 0 13 Brown tree trunk 0 0 2 2 Fence rail 2 3 6 11 Totals 5 5 16 26 86 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Number of brown lizards Total Total B B B B G & B + + + Green foliage 0 0 0 0 8 B B B + + + 0 0 0 1 0 2 2 0 3 1 0 1 Dark green foliage 1 1 0 2 4 Concrete 1 2 0 3 6 Brown tree trunk 0 1 0 1 3 Fence rail 1 3 7 11 22 Totals 3 7 7 17 43 Except the lizards seen on green foliage, the ntunber of green animals equals the brown ones. No brown ones were noted on green foliage but the observations here recorded are by far too limited to permit definite conclusions to be drawn. There was some difficulty in deciding whether an animal resting on a brown fence rail amid a mass of green foliage should be classed as one resting on a fence rail or on green foliage, but it was decided to place these with the former. One is justified, however, in drawing one conclusion from the table, namely, that under approximately the same conditions of temperature and light both green and brown lizards may be found and, even on sunshiny days with a moderate temperatiu-e, the green ones may even outnumber the brown ones. According to Parker, brown should be the prevailing color. It does not appear that in their natural environment the reason for the greater num- ber of green lizards can be accounted for on the ground that these animals reacted more strongly to temperature than to light. According to Parker and Starratt, the Anolis remains brown at io°C. and remains green at 4o°C., regardless of light. One would expect then, that at a temperature of 2 5.5°C., if there were a greater susceptibility to temperattire, the brown state would prevail for at this temperature there is active both the light and medium temperature influence. Evidently a factor which is of extreme importance in influencing the color state is the emotional or nervous condition which can not be easily controlled. The effect of the organs of internal secretion which are under the control, directly or indirectly, of the nervous system probably also influences the color states. Ditmars states that the sleeping Anolis is invariably green and that the same color is present during anger or fear. He states that if a cage containing a nimiber of these lizards be shaken, all Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 87 take on the green state, but after allowing them to rest for a short time, most of them assume the brown state. I have found this to be true in general. Also, if a brown Anolis is taken out of its cage and held in the hand it becomes green in a few minutes. This characteristic change occurs quickly even in animals which have been kept in captivity for a ntunber of weeks and have apparently become tame. It is evident that various factors influence color states and the problem becomes even more complex when one considers that certain areas of the skin may be light colored, as in the case of the mid-dorsal stripe, while the rest of the skin may be dark. One must admit that the skin is influenced by three factors, tem- perature, light and emotional or nervous conditions, induced, no doubt, by way of sense organ stimuli. Yet we get opposite effects in two areas of the skin of the same animal. One would hardly expect light and temperature to have a selective action on the skin. Structure of the Skin of Anolis The chief object of this paper is to present a review of the his- tological structure of the skin of Anolis and to add some observa- tions with the hope that the further investigation of the color changes may be enhanced and some of the factors governing the color states explained. It is not claimed that a knowledge of the minute structures and their relations will offer a full explanation of these changes, but without suclt* a knowledge, physiological experiments must fail to accomplish this end. It is only by keeping in mind the structure of the skin that the actual processes involved may be surmized and physiological data be applied in actual explanation of the problems. The skin of Anolis is comparatively thin and loosely attached except at the sides and dorsum of the head and tail. On closer observation it is seen that it is not smooth but is thickly studded with small, closely-packed scales which vary in shape, color, and size in different parts of the body. These small scales, designated as scutes by Carlton, are smooth and shiny. The scales situated along the mid-dorsal line are irregular in shape and size with slight tendency toward a hexagonal outline (Fig. i). They are well separated and measure 0.29 mm. length- wise to the body and 0.305 mm. in the transverse direction. A thickened, linear, longitudinal keel may be present in the mid-line of the scale. The scales on the dorso-lateral aspect of the body are 88 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. less irregular than those along the mid-dorsal line and none shows overlapping. They measure 0.248 mm. x 0.265 rnm- (Fig- 2). Those scales on the lateral aspect of the body are almost circular in outline and are arranged in two rows, vertical and horizontal, the intersection of the two rows making an angle of about 58°, and they measure 0.255 rnm. x 0.248 mm. (Fig. 3). The scales on the ventral aspect are larger and more closely arranged, the caudal edge of each scale overlapping slightly the cephalic end of the scale just behind it. The outline of the scale is somewhat oval or circular and there is present a homy ridge or keel extending down the middle which becomes more prominent as the caudal border is approached. The scales of the region measure 0.35 mm. x 0.36 mm. (Fig. 4). The scales on the tail differ markedly from those previously described. They are hexagonal in outline and closely packed and overlap each other to a greater extent than those on the ventral surface. There is present a median ridge and the surfaces on either side slope away from it. These scales measure 0.45 mm. X 0.27 mm. (Fig. 5). The dorsal aspect of the head is made up of bilaterally symmetrical plates of a more or less hexagonal form. Just posterior to the inter- section of the mid-dorsal line and the posterior edge of the orbital ridge is a marked pineal eye. The scales on the distended throat- fan are widely separated and are flattened and somewhat conical in shape. Those of the eyelids are so minute as to be barely visible. The color of the individual scales is the same as that described for the various areas of the body. When a general color state is assumed, for instance brown, isolated scales may be white, tur- quoise blue, lighter or darker brown than the general hue, or even green. Some extremely interesting features are noted in scales on the various parts of the body when observed under the low power of the microscope, by reflected and by transmitted light. The char- acteristics exhibited by the scales present points of similarity, but also some very marked differences. For these observations bits of fresh skin were taken from different parts of the body and mounted in glycerin and the appearance of the scales was studied both from the external surface and from the internal surface. The external siu-face of the lateral scales appears yellow by transmitted light. Scattered evenly throughout are somewhat indistinct, pale brown, stellate-shaped bodies, the melanophores, Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 89 which average about fifty in number for each scale. The spaces between the scales are transparent and contain many branching pigment cells (Fig. 6 A). By reflected light these scales appear emerald green and the interspaces black (Fig. 6 B). The internal surface appears blue by reflected light and the branching melano- phores appear distinct and dark brown or black (Fig. 6 C). The external surface of the scales of the ventral surface appears a pale straw color by transmitted light and contains isolated indis- tinct pale brown melanophores of from four to twelve in number. The branches of these melanophores become darker near their termination (Fig. 7 A). By reflected light the scales appear some- what as inverted glass cups containing crushed ice and the melano- phores are cobalt blue except at the termination of their branches which are brown (Fig. 7 B). The internal surface by transmitted light appears a pale straw color and the melanophores are distinct and black (Fig. 7 C). The scales along the mid-dorsal stripe differ from those just described in that, irrespective of other colors, they have a peculiar pinkish cast while the melanophores are much fewer in nvunber in many scales and lighter in color than those previously described. The scales of the throat-fan are pale straw color by reflected light and show the blue appearing melanophores. The wide spaces between the scales present many bright red linear streaks of varying sizes which branch and anastomose. The underlying color is pink and of granular appearance. -^By transmitted light the melanophores appear brown and more distinct and the spaces between the scales take on an orange color. Injections of india ink into the circulation would tend to show that the pink color is not entirely due to the vascularity of this structure but to some other coloring matter present. It would appear from the differences noted by reflected and transmitted light that the melanophores are separated from the surface of the scale by some substance which gives them a bluish cast by reflected light and pale brown by transmitted light. That they themselves are black is evident by viewing them from the internal surface of the scale where they present a sharp clear out- line (Fig. 6 A, B and C). In order to explain the green color of the scales of the lateral aspect of the body it is necessary to eliminate the yellow coloring matter which is noted by transmitted light. This may easily be accomplished by subjecting the skin to alcohol and ether in which this yellow substance readily dissolves. When 90 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. the scales are now viewed externally by reflected light, they no longer have the green appearance, but appear deep blue just as do the previously described scales viewed from the internal surface by reflected light (Fig. 6 C). The melanophores, however, are blue and indistinct, indicating, as was later found, that a semi- transparent reflecting layer exists between them and the surface and that this layer is evidently not affected by ether or alcohol. The explanation for the green color is now quite apparent, for blue rays are reflected and these in passing through a clear yellow medium present to the eye the green color. The semi-transparent reflecting layer lying between the melano- phores and the external surface reflects bluish-white light as is evident in those scales in which little or no coloring matter exists and in which the melanophores are sufficiently separated so as not to influence the reflected light to any extent. This is true of the scales on the ventral aspect of the body, in individual lateral scales, and the white ones often found along the mid-dorsal line. Histology That the tissue relationship of the skin might be more carefully analyzed with the purpose of attempting an explanation for the changes of color, a nimiber of lizards in various color states were killed and segments of the body fixed in different fluids. It was found that segments placed for about six weeks in a fluid consisting of 3-5% potassium bichromate loo parts, formalin 4 parts, and glacial acetic acid 5 parts, gave the best results. Tissue fixed in this fluid maintained the cellular relationship and the osseous structures were sufficiently decalcified so that entire sections of the body could be made. Formalin was found to act too slowly with segments of the body. Lizards in the green state became brown during its action. It was possible, however, to overcome this difficulty by injecting formalin quickly with a hypodermic needle under the green skin, thus obtaining almost immediate contact of the fluid with the entire internal surface of the skin. This method has a distinct advantage in that the animals may be studied on the table for a considerable length of time after fixation and the color state carefully recorded. Unfortunately, however, after a longer time the green color be- comes changed to a slaty gray after the use of formalin, due most probably to a change in the yellow coloring matter. Removing the skin and washing it in water soon after fixing prevents this Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 91 bleaching and bits of skin may then be held between pieces of hard- ened celloidin and sectioned. The solubility of the yellow sub- stance forbids imbedding in either celloidin or paraffin. These sections should then be mounted in glycerine. For examination of the cellular elements, tissues imbedded in both celloidin and paraffin were sectioned at 1 5 microns and stained with haematoxylin and eosin. Frozen sections treated with gold chloride for the purpose of showing the nerve endings have, so far, not been successful. In order that the histological picture be made as complete as possible, the elements of a single lateral scale will be described (Figs. 10, II, 12, 13 and 14). Where differences exist in the scales of other parts of the skin these will be mentioned. The layers comprising the scale will be taken up in their order, beginning from without inward. Epidermis. — The outermost, transparent layer or epidermis may be divided into an outer, horny layer or stratum comeum and an inner, stratum germinativum. The epidermis is consider- ably thicker near the siunmit of the scale than at the periphery where it becomes continuous with the thin epidermis of the space between the scales. The stratum corneum may usually be divided into two layers, an outer one which is separated by an interspace from an inner. This outer layer represents that portion which is ready to be cast off in moulting (Figs. 8 and 12). The inner layer does not stain with eosin, being straw-colored. The squamous cells may show clear, non-staining, round bodies, representing the degenerated nuclei. The layer undergoes marked keratinization at the apex of the scale with the formation of a homogeneous, homy ridge or keel. This thickening is more marked in the scales on the ventral aspect and those along the mid-dorsal line. Keratinization of the other cells of the stratum comeiun, other than in the keel, is not so marked and, due to the dehydrating effect of the air, become fairly well separated so that their outlines may be distinguished (Fig. 8). Briicke has described "interference cells" in the outer layer of the stratimi corneum in Chamcsleon vulgaris. These cells, he be- lieves, modify the color of the skin by reflected light to a very marked extent. Keller believes that the outer cells of the outer layer of the stratum comeum contain minute closely placed columns arranged at right angles to the surface of the cell. He calls this outer layer 92 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. the "Relief Schicht," and the inner surface of this outer layer the "Negative Relief Schicht." The latter presents the negative picture of the former in that instead of minute columns there are toothlike incisions corresponding to the columns of the "Relief Schicht." The layer between these two he terms the stratiun comeum. He further claims that the outer cells of the inner layer of the stratum corneum are similar to those of the outer layer and terms them the "Second . Relief Schicht." He believes that separation occurs along a line corresponding to the boundary between the "Negative Relief Schicht" and the second "Relief Schicht" and that when the outer layer is cast off the second "Relief Schicht" becomes the first and then a second line of cleavage occtus making a second "Relief Schicht" and a "Negative Relief Schicht." This cleavage goes on at regular intervals. He found that the fine colimm-like structures of the cells were very much more pronounced in the foot pads and at the apices of the scales. In Anolis the first or outer "Relief Schicht" of Keller is very prominent on the under surface of the adhesive pads of the second phalanges. The second "Relief Schicht" is also present when the outer layer of the stratum comeum is well separated from the inner layer, but no "Negative Relief Schicht" was noted. On the outer cells of the outer layer of the stratum comeimi of the scales of the general body, occasionally minute spicules resembling short cilia may be seen in stained preparations but such occurrence seemed very rare. In dried scrapings, the outer cells seen on the flat contain numerous dots giving them a stippled appearance when examined with the high dry or oil immersion lens. These probably represent the spicules that Keller has described for the chameleon. (See Fig. 9.) The stratimi germinativum takes the haematoxylin and eosin well. The cells are polygonal with fairly large vesicular nuclei. In the scales this layer is from two to three cells thick but in the epidermis between the scales it is at miost only two cells thick. The basal layer is composed of cuboidal and columnar cells with large vesicular nuclei. Their proximal borders, attached to an ill- defined basement membrane, are frayed and brush-like (Fig. 8). That the epidermis, through phenomena of interference, exerts some modifying influence on the color of the skin in Anolis is with- out doubt, but that it plays the important role which Briicke ascribes to it for the chameleon is doubtful. Keller, in fact, dis- agrees with Briicke as to the importance of this layer even in the Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 93 chameleon. The small, transparent spicules in the outer layer possibly cause some diffraction of light but this must be slight for the underlying cellular outlines are markedly clear vv^hen viewed through the epidermis. The Oil Droplet Layer. — A considerable amount of confusion exists in regard to the layer underlying the epidermis. Keller has described a layer in the chameleon which he designates the ochrophore layer. He does not believe that it is cellular for the elements composing it have no nuclei, but he believes that it is made up of bits of cytoplasm cast off from an underlying layer of cells which he ternis the leucophore layer. This ochrophore layer is found on the dorso-lateral aspect of the body but is almost entirely absent on the ventral aspect and entirely so on the foot- pads and spaces between the scales. Keller found that the ele- ments of the layer were brownish yellow by transmitted light and a bluish white by reflected light, had a granular appearance, and that they disappeared under the influence of mineral acids. He described these elements as more or less spindle shaped and verti- cally arranged, the ends in contact with the epidermis being more pointed than those of the opposite end. The elements farther removed from the epidermis had both ends rounded. Pouchet called this layer "Iridocytes" and believed the ele- ments to be cells although he could not make out the cellular structirre. In Anolis, Carlton describes what tie believes to be the ochro- phore layer of Keller. He admits that its structure is not similar to that found in the chameleon. I do not believe that Carlton saw a layer corresponding to that in the chameleon, but that he described the leucophore layer which he mistook for the ochrophore layer. I will take up my reasons for this asstmiption in the dis- cussion of the leucophore layer. In sections stained with haematoxylin and eosin, I was unable to make out any layer corresponding in either position, structure, or color to the ochrophore layer. In certain scales, more often those situated along the mid-dorsal line, a clear space or a space filled with large clear cells with large vesicular nuclei could be noted where the ochrophore layer should lie. Knowing that it was possible to dissolve out the yellow coloring matter in the scales with alcohol and ether, it did not seem improbable that in the preparation of the stained sections practically all trace of this layer had been lost. 94 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. For this reason formalin fixed skin was sectioned between blocks of celloidin very soon after fixing and then mounted in glycerine. In these sections, there is situated just beneath the epidermis a thin layer made up of bright yellow droplets of varying sizes (Fig. lo). These droplets appear beautifully refractive, do not contain any granular material, nor are they arranged like the elements in the ochrophore layer of Keller. Some of these drop- lets may even be found at times between the basal cells of the epidermis where they have probably migrated. By transmitted light this layer of droplets appears bright yellow (Fig. lo A), while by reflected light it disappears almost entirely, in other words it does not reflect light to any marked extent (Fig. lo B). In this respect it differs from the ochrophore layer of Keller, which its describer found to be bluish white by reflected light. The sections treated with Scharlach R caused this layer to be- come brownish red, indicating that the droplets composing it are lipochromes. This layer of oil droplets is practically absent on the ventral surface of the body and entirely so on the adhesive pads of the feet. To what extent it is present along the mid-dorsal stripe, I am unwilling to say. That it occurs here in individual lizards to the same extent as on the lateral scales is true; but whether it is greatly diminished in those lizards showing a white dorsal stripe, or whether it is present to the same extent but can be displaced to the periphery of the scale by a special mechanism and there does not effect the color, it is difficult to say. Keller believes that this layer can be removed from the field of action by special cells. One may conclude, then, that beneath the epidermis of the dorso-lateral scales there is a thin layer made up of transparent yellow oil droplets and that this disappears in stained preparations due to its solubility in alcohol, ether and clearing oils. Zanihophores. — Lying just beneath the epidermis of the lat- eral aspect of the body and in the oil droplet layer, are cells of varying sizes. The larger cells have a clear, round, un- stained cytoplasm in the sections stained with haematoxylin and eosin. The nuclei are large and vesicular and the chromatic ele- ments stain deeply (Figs, ii and 14). The nimiber of these cells varies in the scales of the different parts of the body, being prob- ably more numerous along the mid-dorsal stripe. In fresh speci- mens it was practically impossible to distinguish them with accu- Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 95 racy. These cells were not described by Carlton, but Keller has described similar cells for the chameleon and named them zan- thophores, and Pouchet also described similar cells for the chame- leon and believed that they contained fat droplets of 2.5 microns. He believed these cells to be analogous to the yellow cells of batra- chians and that they possessed the power of contractility. Keller also believed that they could expand or contract for he found them varying markedly in size. Doctor Irving Hardesty suggests that these cells secrete or control the accumulation of the oil droplet layer described above. If Keller and Pouchet be correct in their assumption that these large clear cells may expand, one might reason that during this state they practically fill the entire space between the epidermis and the underlying layer and force the yellow droplet layer towards the periphery of the scale so that it no longer influences the color states. For reasons which will be taken up later, I believe that the mechanism is not quite as Keller would have one believe, although undoubtedly these cells are more nimierous and almost replace the oil droplet layer in the white scales of the mid-dorsal stripe. If these large, spherical cells in Anolis are the zanthophores of Keller, and they resemble very closely those he figures and de- scribes, Carlton is wrong in stating that these cells do not exist in Anolis. The Leitcophore or Guanophore Layefr— The layer lying just be- neath the layer of oil droplets presents very marked differences from any of the structures previously described. In vertically sectioned scales, stained with haematoxylin and eosin, it is seen that this layer is thicker near the center of the scale and then gradually thins out until it disappears at the periphery. The layer forms then an inverted cup which thins out at the edges and fits into the hollow epidermal scale but does not come in immediate contact with it because of the intervening oil droplet layer. It is present in all the scales of the skin including those of the ventral aspect of the body (Figs. 8, 10, 11, 12, 13, 14 and 17). By reflected light it appears as a homogeneous bluish- white band (Fig. 10 B), and this appearance is not lost in those sections fixed in the fluid mentioned and stained with haematoxylin and eosin. Bits of the layer may be found isolated in the deeper fibrous layer, recogniz- able by the bluish-white color. 96 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. By allowing a minimal amount of light to come through the condenser of the microscope, the layer has a most brilliant opales- cent appearance. In unstained, freshly fixed formalin sections, by transmitted light, it has a pale brownish appearance (Fig. lo A), but in stained sections it appears darker and greenish brown (Figs. II, 12, 13 and 14). Fig. ID, A and B, represents the appearance of an unstained sec- tion of the layer by reflected and transmitted light. By reflected light this layer appears as a bluish-white cloud which obliterates the underlying structures, or at least makes them appear hazy and indistinct. In both the stained and unstained vertical sections the layer is seen to be composed of parallel rows of somewhat irregular blocks, their long axes being parallel to the outline of the epidermis. These blocks are of varying size and asymmetrical shape, and un- doubtedly possess small, deeply staining nuclei. Some sections show these nuclei better than others. The vertical section gives little idea of their morphology for when seen in tangentially cut sections they appear very irregular in outline and possess short pseudopodoid processes which may terminate in hooklike expan- sions or branches. Every conceivable shape exists and no simi- larity exists in these bodies except in their marked irregularity (Fig 14). In some sections these cells appear syncytial, for their processes are in juxtaposition, thus leaving numerous openings of various sizes between these apparently joined processes. Through these openings run the branches of the melanophores (Figs. 14 and 17). That really no syncytium exists appears likely, for in vertical sections no such connections between the processes can be made out. When viewed from above, the area around the nucleus has a bluish cast while the periphery is a pale greenish brown. One can conclude then that the cells of this layer are fairly thick, irregular plates of fairly uniform thickness throughout but with a marked irregular outline. Carlton describes a somewhat similar layer in the scale of Anolis which he calls the ochrophore layer and which he considers analo- gous to the ochrophore layer of Keller for the chameleon. He believes that this layer produces the green color and finds that by reflected light it appears bluish green and by transmitted light yellowish green. From the micro-photographs accompanying his paper, one cannot be mistaken as to the identity of the layer in question. He has noted the block-like, parallel arrangement in Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLTNENSIS 97 the vertical section and also the irregular appearance in the tan- gential section, stating that it had the appearance of a more or less homogeneous mass, irregular in outline, and penetrated in many places by the processes of the melenophores but he was unable to make out any cellular structure and denied the existence of nuclei except between the blocks. Why he should believe this layer to be the ochrophore layer of Keller, I am unable to say. He admits that its arrangement differs from Keller's ochrophore layer. If Carlton's ochrophore layer is responsible for the green state in Anolis, why is this layer present on the ventral aspect of the animal where no green color is ever present? Keller states that this ochrophore layer is almost entirely absent on the ventral aspect of the body of the chameleon, yet Carlton, in spite of these differences in structure and position, attempts to make these two layers analogous. Furthermore, Carlton's ochrophore layer closely fits the description of Keller's leucophore layer, which latter Carlton states does not exist in Anolis. Undoubtedly the layer in Anolis is the same as that in the chameleon except for possibly minor differences. Briicke described in the chameleon a white or yellow pigment which he finds separated into two layers, the inner being thicker and made up of closely packed colorless particles with rounded boundaries, which reflect light, resulting in the white appearance. He believes these reflecting granules^© be the product of cells whose processes force themselves between the dermal structures and lie between the epidermis and the underlying connective tissue. In these two layers he evidently includes both the ochro- phore layer and the leucophore layer of Keller. Pouchet has also described this layer and considers the white, dust-like material as the products of cells which, by the growth of the neighboring tissue, have been pressed into plates, and Keller describes these plates or blocks which he names leucophores. He considers their content similar to that found in the scales of cer- tain fish described by Klihne and which are said to be composed of guanine. The fact that both react positively to the murexide test leads to this assumption. He believes, as does Pouchet, that these leucophores have been pressed into plates by the pressure of the overlying and underlying tissues and that their edges adapt themselves to the neighboring structures due to the mechanical resistance of the latter, and, in consequence, assume very irregular 98 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. shapes except in those regions where no such pressure is exerted. In regions where no pressiire is exerted, they are rounded and He close together in the wide meshes of the connective tissue. I was unable to verify Keller's observations that the leucophores were rounded in those regions where the mechanical resistance is less, for those isolated cells which were noted by me showed in reality even greater irregularity, when seen in vertical section, than those in the leucophore layer proper. That the peculiar plate or block-like shape and the arrangement of these in parallel rows may be due to the pressure of the dense connective tissue from below seems likely, but only careful observations of the skin in various stages of development can determine this point. The white, dust-like material of Pouchet, the white pigment of Briicke, or the granules of Keller, which were described in the leucophores, were not noted by me in Anolis. The cytoplasm of these cells even in the fresh state was clear and apparently free from granules. This finding is interesting in that one would expect the reflecting power of these cells to be due to the denser granules, and probably such granules do exist but were invisible because of methods I employed in the study of these cells. No differences in shape or position were noted in the leucophore layer in the green and brown state. The cells of the layer seem to retain their characteristic appearance and relationship no mat- ter what color state of the skin existed. Carlton makes the same observation for his so-called ochrophore layer. The blue coloration of the melanophores by reflected light de- scribed for the scales of the ventral aspect of the body is undoubt- edly due to the leucophore layer as is also the white appearance of these scales. This will be touched upon later. The Melanophores. — Lying between the leucophore layer and the underlying connective tissue layer, and partially imbedded in both, are the melanophores described by Keller for the chameleon (Figs. 10, II, 12, 13, 14, 15, 16, 17). In Anolis three types of pigment cells are found, namely, those in the dorso-lateral scales which differ from those in the ventral scales by their smaller size and more delicate branching, those in the ventral scales, and a third type which is commonly situated just beneath the epidermis between the scales. The melanophores show a striking resemblance to the Purkinje cells as seen in Golgi preparations. A line passing through the cell bodies of the ma- jority of them would be more or less parallel to the epidermis except Vol. X] VON GELDERN—SKIN OF ANGUS CAROLINENSIS ' 99 at the periphery of the scale where they more closely approach the surface. A few cell bodies lie above or below this imaginary line, but in thick sections the bodies of the melanophores form a fairly thick, dark-brown layer of fairly regular width. The melanophores are best studied in vertical sections of the fixed material stained with haematoxylin and eosin. It was found unnecessary to use the methods adopted by Keller and others to bring out the finer branches of these cells, for in most of the prepa- rations these were clearly visible. The cell body is more or less rounded but considerable difference exists among them, some being much narrower than others (Figs. II and 12). The surface facing the epidermis is often concave but rarely it may be convex or apical. The nucleus may be round, oval, reniform, horseshoe-shaped or even double in rare instances. In some preparations it takes a fairly deep blue stain and has a vesicular appearance (Figs, ii, 15 and 17). The concavity when present is directed towards the epidermis. Coming off from the sides of the outer surface of the cell body are a varying number of permanent branches which run either vertically toward the inner surface of the epidermis or present a lateral curvature. The curvature may even be so marked that, at the proximal part, the branches may be directed first down- wards and laterally and then gradually curve laterally and up- wards (Figs. II, 12, 13 and 15). These^ranches run through the spaces among the leucophores and, as they approach the surface, lateral branches in turn give off further branches. This tree-like branching continues until beneath the under surface of the epi- dermis a layer of fine terminal branches exists. The contents of the melanophores consists of a varying amount of fine pigment granules imbedded in a mass of faintly brown, poorly staining cytoplasm. The arrangement and distribution of the pigment granules depends on the color state of the skin, being almost absent in the smaller branches in the green state but present even in the terminal branches in the brown state. Under the oil immersion lens the poorly staining cytoplasm may be followed even in the finest branches lying beneath the epidermis. Follow- ing them is, however, greatly facilitated by the presence of isolated pigment granules which have failed to migrate with the general mass of pigment. . The pigment granules are oval in shape and brown under magnification. Their number varies markedly, irrespective of the 100 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. distribution of the pigment. The number varies not only in melanophores in the same scale, but also melanophores of one scale may contain more pigment than those of another. Also the intensity of the color of the pigment may vary, individual or groups of melanophores containing a lighter brown pigment than others. This appears to be irrespective of the number of granules. These differences in amount and intensity in color are so striking that there is no doubt as to their occurrence (Figs. 11,12 and 13). In the green state of the skin of Anolis (Figs. 10 and 11) the pig- ment granules are present only in the bodies and proximal parts of the primar3^ branches of the melanophores. This proximal migration of the pigment is practically complete and the finer distal branches are clear and transparent. In some of these finer branches, however, a few scattered pigment granules may have failed to follow the mass of pigment and their presence allows one to detect more readily the finer branches. The bodies of the melan- ophores during the proximal migration of the pigment are neces- sarily darker than after distal migration. In proximal migration of the pigment it is noticed in the primary branches of the melano- phores that there is an area of gradation between the dense pig- ment on one side and the clear part on the other side where the pigment is much less dense. In this portion the pigment granules apparently arrange themselves in parallel rows (Figs. 11 and 15). This parallel arrangement has also been observed by Keller in the chameleon. In the brown state (Figs. 12 and 13) the finest branches lying immediately beneath the epidermis are filled with closely packed pigment. This gives the appearance of a thin, dark-brown layer lying just beneath the epidermis in vertical section. One might conclude that, in order that this appearance can be produced, the terminal branches must anastom.ose and fonn a plexus. That this is not the case, however, may be readily determined in tangentially cut sections where the terminal branches appear as separated but closely packed, dark-brown dots (Fig. 14). Keller makes this observation for the chameleon and Carlton for Anolis. It is probable that the green state of Anolis does not represent the maximimi degree of proximal migration of the pigment. In certain scales, pigment may be absent even in the primary branches and be confined entirely to the cell body which appears like a dark brown or black sphere. Furthemiore the pigment may be con- densed to such a degree that a clear broad halo of cytoplasm may Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 101 surround a central compacted mass of pigment (Fig. 15 A and B). These peculiar melanophores are often bilaterally arranged and present in a group of from two to three scales on each side of the body. This bilateral arrangement appears too marked to be acci- dental. Melanophores in this condition must explain the yellow- colors and white scales often to be observed in the living Anolis. Besides the distal and proximal migration of the pigment, any degree of migration may be present, namely, all but the terminal branches may be filled with pigment, or the terminal branches may contain scattered pigment, and so on. All of these conditions in- fluence the color state of the skin and must be associated with definite color states. The number of melanophores varies considerably in the various scales, but the average for those of the lateral aspect of the body is about fifty in number. This number is greatly reduced in the scales along the mid-dorsal line which are white and in which ten to fifteen pale brown melanophores appear to be the usual number. The latter are slightly smaller and their branches are more delicate and spread out more than the others. In the mid-dorsal line of other specimens where no white stripe exists but where color changes resemble those of the lateral aspect of the body, the me- lanophores cannot be distinguished from the others and appear in about the same number. The melanophores in the scales, on^the ventral aspect of the body are from five to twelve in nimiber. Their bodies are larger and more rounded and possess fewer primary branches. The terminal branches are followed with greater difficulty to the periphery. The pigment is usually thickly packed in the cell bodies giving the cells a dark-brown color. The nucleus due to this increased amount of pigment is rarely observed (Fig. 8). The pigment cells lying in the spaces between the scales vary markedly in number and position and, aside from their possession of branches and pigment content, show little resemblance to the true melanophores (Figs. 11, 12, 13 and 16). They more nearly resemble ordinary mesenchymal pigment corpuscles. The body resembles a flattened disc as is readily seen by comparing the verti- cal diameter as seen in vertical section (Fig. 13) with the horizontal diameter as seen from above (Fig. 16). The primary branches are thick and irregular and vary considerably in length. The terminal branches are short and terminate broadly in club-like ends. The cells are present just beneath the epidermis between the scales, 102 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. their branches spreading out and lying parallel to it. These cells may often be found in the deeper tissues of the body and give the impression of their being able to wander between the tissues (Fig. 13). That these cells may be converted into melanophores seems possible. That the pigment granules migrate in the fixed pseudopodic processes of the melanophores, instead of an amoeboid extension of and retraction of the processes themselves, is very probable. This migration has been clearly illustrated by Keller and Briicke for the chameleon, Carlton for Anolis, Degner ior Praunus flexuosus , Kahn and Lieben for Rana temporaria and Spaeth for Fundulus heteroclitus . Parker believes that the pigment migration is true for Phymosoma and, further, states that the migration of pigment in melanophores is influenced by light and temperature, either light or low temperature causing a distal migration and absence of light or high temperature causing a proximal migration. Although fully agreeing that the melanophores and their processes remain fixed and that their pigment undergoes migration, I am unable to see how any set of factors influence all melano- phores similarly. Under precisely similar conditions the melano- phores of the lateral aspect of the body may contain proximally migrated pigment, whereas the melanophores of the mid-dorsal stripe or melanophores of isolated scales may have the pigment in the terminal branches. In a single animal, in any color state, many exceptions may be found to the rule laid down by Parker. Gold chloride preparations repeatedly fail to reveal any nerve endings terminating on the bodies of the melanophores but that these exist seems most probable. Pouchet described a smaller pigment-bearing cell which he termed the erythrophore and which closely resembled the melanophore except that it contained a purplish-red pigment. Briicke overlooked these cells of Pouchet, but according to Keller, the cells only occur on the lateral scales of the chameleon in any great number and are not present in all indi- viduals. Keller described gradation forms, cells containing both brown and red granules in different proportions. Some cells may contain only a few red granules among brown ones while others may contain only a few brown ones, the greater proportion being red. Carlton was unable to find erythrophores in the skin of Anolis and denied their existence. I believe that Carlton is correct, for if these cells be present they must be extremely rare. No red pig- ment granules were observed in any melanophores of my sections. Vol. X] VON GELDERN—SKIN OF ANOUS CAROLINENSIS 103 It seems likely, however, that the pigment granules vary consid- erably in the intensity of their color as has already been stated. This conclusion is reached not only from a study of the pigment granules in the melanophores but also from their effect on the color of the skin. There is little doubt that a condensed mass of pigment will produce a darker brown than more scattered pigment, but the former will always be brown and can never be black or brick-red. If this be true, then the melanophores producing the post- orbital black patch must contain black pigment granules and those producing the brick-red stripe must contain reddish-brown granules. Furthermore, it is possible that an individual scale may contain melanophores of two or more kinds of pigment content and that these may act independently of one another. This is suggested by the microscopic appearance as well as the appearance of either a brick-red, brown, or black state in the scales of the mid- dorsal and the post-orbital stripes. Undoubtedly the amount of pigment present plays an important role, but many of these differ- ences cannot be satisfactorily explained. Partial distal migration may be responsible for a lighter brown color than maximal distal migration, but only up to a certain point. Any distal migration beyond this is not associated with a still lighter brown state but with a slaty or greenish-gray color. The Connective Tissue Layer. — Lyijjg beneath the leucophore layer, running into the concavity of the scale for a variable distance but approaching more closely the epidermis at the edges of the scale, is a fairly dense layer of white, fibrous connective tissue (Figs. 8, lo, II, 12 and 13). The fibres appear to run parallel but on closer inspection many vertical and oblique ones may be noted. The vertical fibres may be traced as they ascend among the cells of the leucophore layer where they break up into small fibril bundles which form a network beneath the epidermis. This layer takes on a bright pink color with eosin and contains many deeply staining stellate and spindle shaped nuclei. It is fairly vascular and nerves may be seen traversing it. Below the concavity of the scale are present fat corpuscles and large blood vessels (Figs. 11, 12 and 13). Beneath the dense connective tissue separating the skin from the underlying skeletal muscles, there is present a loose areolar connective tissue. Fine free pigment granules forming a fine line between the denser connective tissue and the looser 104 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. areolar tissue, may be seen in many sections. Free granules of red color aid in giving the red color to the extended throat-fan. The skin is extremely vascular but more so in some regions than in others. Larger vessels run beneath the denser connective tissue layer and run parallel with it, dipping into the scale. From these vessels branches are given off which run through the denser connective tissue and also through the leucophore layer to directly supply the epidermis (Fig. 13). It seems not improbable that a vaso-dilatation occurring under and in the leucophore layer, may exert a modifying influence on the color states. The pink color of the mid-dorsal stripe may be explained by the effect of a vaso-dilatation on a white stripe, and the red appearance of the throat-fan is no doubt in part due to blood, as well as to granules of red pigment in the subcutaneous connective tissue, which shows through the spaces between the scales. On the Mechanism of the Color Changes The essential structures present for the production of the various color states are the epidermis, the yellow oil droplet layer, the leucophore layer, the melanophores, and, possibly, the zantho- phores and the cutaneous blood supply. The skin of the scale is made up of four superimposed, inverted, hollow, cup-like layers, the outer being the epidermis. Next to this is the oil droplet layer, then the leucophore layer, and lastly, the connective tissue layer which, however, supports the integrity of the whole. The first and last named layers are continuous with those of the neighboring scales, but the second and third are limited to the scale. The epidermis is a transparent layer which acts largely as a protecting and supporting structure and, through interference phenomena, acts slightly, if at all, as a factor in the color states. The second or yellow oil droplet layer presents a thin trans- parent yellow mediimi which is extremely important in the pro- duction of many of the color states. In and superficial to it lie the fine terminals of the branches of the melanophores. The large zanthophores also lie in it and extend inward into the next layer. The oil droplets give a strong, bright color by transmitted light, but seem to reflect but little light (Fig. 10 A and B). It seems to act more as a filter than as a reflector. White light reflected from Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 105 the underlying layer, when passing through this yellow medium, must be so acted upon as to give the yellow. The leucophore layer, lying just internal to the oil droplet layer, acts essentially as a reflecting layer. It reflects a large proportion of the light which falls upon it, but it also screens the light to a great extent from the underhv'ing brown melanophores so that pigment granules, when only in the bodies of the melanophores, exert but little influence on the color states of the skin. On the other hand, however, if the primary branches of the melanophores, which pierce this layer, are filled with pigmient, the light which falls on it is reflected as blue light (Fig. loB). Further, cell bodies of the melanophores, lying internal to the leucophore layer, appear blue by reflected light in the scales of the ventral surface of the body in which the yellow oil droplet layer is very scant and in places absent (Fig. 7 B). Evidently then, the leucophore layer in part reflects all the rays of white light and also absorbs all but the blue raj'S from the light passing through it and reflected from the brown pigment within and internal to it. Traversing this leucophore layer are the large branches of the melanophores, con- nective tissue, and finer blood vessels. Partly imbedded in the lower stratum and beneath it are the melanophores. Fig. 17 is an attempt to show in perspective the various layers and their relation to one another. The only layer that remains fixed and present in all scales and not subject to variations is the leucophore layer. All the other elements may be either absent, increased, decreased, or subject to marked variations. All of these other elements function in con- junction with the leucophore layer and either by allowing it to be unobscured, partially obscured, or by entirely shutting it off from the light, produce the color phenomena. The appearance of white and pale blue, as found on the ventral aspect of the body or along the mid-dorsal stripe, may be explained as due to the oil droplet layer being either absent or that it has been forced to the edges of the scale. A purely white scale must mean that the mel- anophores are either absent or greatly diminished in nxunber, and very pale blue scales, that their pigment granules must have migrated entirely into the bodies of the cells. This allows the leucophore layer to act alone as a reflecting layer without the in- fluence of any other element. In addition, the stratum corneimi of the scales of the ventral surface is slightly thicker than in other regions, and this greater thickness, with the markedly developed 106 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. keel of the scale found here, no doubt results in more light being reflected from the outer surface of the scale and thus a whiter appearance. It seems probable that by a vaso-dilatation of the superficial capillaries a pink color may be imparted to the white scales. Further, in addition to the blood capillaries, red pigment is manifestly present in the subcutaneous connective tissue of the throat-fan. If, in the white scales, melanophores are present and send out pigment into the primary and larger branches, the light, which is now acted upon by the leucophore layer, is returned as blue. Decidedly blue scales are rare except in isolated scales on the lateral aspect of the body. Along the mid-dorsal stripe and on the ventral aspect, melanophores are too few in number to influence the color beyond a pale blue. I injected brown pigment (potassium bichromate solution) into the skin of the belly, and a blue color was readily produced. Higgins' brown ink furnishes the same result in the same way. The yellow or orange appearance may be readily explained by the presence of the yellow oil droplet layer through which light from the leucophore layer must be transmitted. The melano- phores, in the case of the yellow skin, must contain the pigment in their bodies, and the branches must be free of pigment. The degree of yellow color depends on the amount of oil droplets, the straw-yellow color being associated with a lesser amount than the deep yellow. Pale yellow is often present on the ventral aspect of some lizards due to the presence of a small amount of this substance. The emerald green is brought about by the migration of pig- ment into the primary and larger branches of the melanophores. Now the light which is reflected from the leucophore layer, due to the presence of pigment granules, is blue, and this blue, in pass- ing through the yellow oil droplet layer, mixes with yellow rays given by this layer and appears at the surface as green. By further distal migration of the pigment granules, light from the leucophore layer assumes a deeper blue which in turn produces the bluish-green color of the skin. By a still more distal migration of the pigment a muddy, greenish gray appears which, as the migration proceeds, becomes brownish gray, then light brown and, lastly, a deep mahogany brown is produced, which indicates that distal migration has pro- ceeded till the granules have accumulated immediately beneath Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 107 the epidermis sufficiently to block the action of both the yellow- oil droplets and the leucophore layer. The pigment granules now act entirely alone and produce the brown state (Figs. 12 and 13). Whether the yellow oil droplet layer can be dispersed under varying stimuli in those scales where it normally exists, that is, whether it is possible for a lateral green scale to assume the white or bluish state in a short space of time, is difficult to say. If, as is held by Keller for the chameleon, the zanthophores possess the power of dilating and can take up all the space held previously by the oil droplet layer and thus displace the latter to the edges of the scale, it would at least be possible for a fairly rapid change to occtir from green to white. Pouchet, however, believes the zan- thophores of the chameleon to be of yellow color and if this be so, Keller's explanation would not be valid. The zanthophores of Anolis appear transparent in all the conditions under which I could observe them. They seem to be absent in the skin of the ventral surface of the body. However, my observations are by far too meager for me to draw any definite conclusions. It is true that the zanthophores seem greatly increased in the white scales of the mid-dorsal stripe and that they form a transparent fairly thick layer between the epidermis and the leucophore layer. The objections to Keller's views are that if the zanthophores are able to disperse the oil droplet layer by dilating, they would probably also disturb the arrangement of the terminal branches of the melanophores, which we know'Soes not take place. One would also conclude that these cells must be under the control of the nervous system. No conclusions can be reached without observing the living animals for long periods of time and noting the changes of these white areas. Possibly the white areas are merely variations and are more or less permanent. Black or brick- red stripes may be also variations in the distally migrated pigment. The following table presents in a concise manner the mechanism involved in the various color states. Keller's theory in regard to the behavior of the zanthophores is included as a possibility, since their attributed function becomes necessary to explain white and blue changes in certain cutaneous areas, if these occur with any degree of rapidity. 108 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Table 3 Color State Oil Droplet Layer Zantho- phore Leuco- phore Melano- phore White absent or dispersed ® or absent ® Blue absent or dispersed ® or absent m , Pink absent or dispersed ® or absent ® vaso- dilatation Straw yellow partially dispersed © 2 9 Golden yellow 1 1 O or absent ® Emerald green .... n O or absent # 1. Bluish green 1 O or absent ® 2. Grayish green iiiiii I. O or absent ■Dili L ® 3. Brownish green. . . . ■iii 2. O or absent Iiiiii 2. ® 4. Light brown ■iillli 3. O or absent 3. ® 5. Mahogany brown. . — O or absent » O brown pig- ment granules Brick-red — O or absent « o brick-red pigment granules Black «. O or absent — o black pigment granules I I unobstructed, ■ill partially obscured, I^Bi obscured, ® dilatation of zanthophore, O contraction of zanthophore, O maximum distal migration of pigment, ® partial distal migration of pigment, ® maximum proximal migra- tion of pigment. No. 1-5 indicate comparative degrees of either obscuration or migration. For the final solution of this problem of the color changes in Anolis, three methods of attack m.ust be carried on and one must not lose sight of any one of them: Carefully controlled physio- logical experiments, histological studies of the skin, and careful Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 109 observations over long periods of time of the habits and color states of these lizards in their natural environment. The first method of attack is always open to criticism so long as the experi- ments are not carefully controlled; for instance in none of the experiments performed by either Parker and Starratt or Carlton were the factors of varying external stimuli taken into con- sideration. Lizards kept for long periods in confinement may give one set of results but one is not justified in drawing any general con- clusions as to the behavior of all lizards. I wish to thank Doctor Hardesty and Doctor Garey for their helpful suggestions. Summary 1. In its color changes, Anolis carolinensis shows a greater variety of colors than has been usually described for this animal. It may at times take on other colors than the emerald green, mahogany brown, and the variations intermediate between these. The variations, though apparently less frequent, correspond fairly 'closely with the variations described for Chamceleon vulgaris. 2. The color changes, in addition to general variations in Anolis carolinensis, as observed in its natural environment especially, seem to be induced by variations in external stimuli. Rhythmic changes of color may be observed with the animal in the same position with unchanged temperature ^d light, and emotional states interpreted as fear, sexual excitement, and anger (prelim- inary to and during combat) seem to more actively bring about color changes than temperature and light. Color changes in sympathy with environment (protective coloration) seem probable. 3. In structure, the skin of Anolis resembles that described by other investigators for Chamosleon vulgaris, except no "Negative Relief Schicht" could be distinguished in the stratum corneum of the epidermis, and no cells corresponding exactly to the eryth- rophores of Pouch et could be determined. Also the oil droplet layer described here for Anolis is not the same as the ochrophore layer described by Keller for Chameleon. 4. The observations of Carlton that the processes of the mel- anophores in Anolis are fixed or non-amceboid and that migration of the pigment granules occurs within them, is hereby confirmed. In this the nielanophores are similar to those described for 110 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Chairiceleon, Fundulus heierocUius , Rana temporaria, and other color-changing animals. 5. The color changes in Anolis depend upon the reciprocal physical action of four layers of the skin: the epidermis, the yellow oil droplet layer, the leucophore layer and the melano- phores. The physical characters making possible light inter- ference and absorption, and the mixing of transmitted and reflected rays, modified by the migration of pigment to different positions in these layers, result in the varieties of color apparent at the surface of the skin at different times and on different locali- ties of the body. The red coloration of the throat-fan is due to a rich capillary plexus and to the presence of a red coloring matter in the deeper layers. The effect of vaso-dilatation is also apparent in the pinlc stripe noted occasionally along the mid-dorsal line. 6. The oil droplet layer and the leucophore layer in general remain fixed and the various color states depend on the migration of the pigment granules in the fixed processes of the melanophores. Maximal proximal migration of the pigment is associated with yellow, while maximal distal migration produces dark mahogany brown. In the emerald green state the pigment lies in the primary and larger branches of the melanophores. Further distal migration is associated with bluish-green or slaty-gray color states depend- ing on the degree. Vol. X] VON GELDERN—SKIN OF ANOLIS CAROLINENSIS 111 BIBLIOGRAPHY Barbak, E. : ' i o. Zur chromatischen Hautfunktion der Amphibien. ^ iw*^.'- \ t s CO PROC. CAL. ACAD. SCI., 4th Series, Vol. X u CD PQ t^ U PQ ^ "S. *. -' > [VON GELDERN]Plate 8 ID CQ ID PROC. CAL. ACAD. SCI., 4th Series, Vol. X ) ^:^ - e ii^ t ■^^i^fcj' 14 •/-■v^ >•>■ 10 I [VON GELDERN ] Plate 9 <2.%^ Str. Corneum *^c©*^' Zanthophore Melanophore Conn Tissue Fat Connective Tissue Layer Leucophore Layer Fat — Str. Corneum Str. Germinat. Melanophore *ifS>VN.>^^. ,-:v<"- V. Str. Corneum Str Germinat. - Melanophore Leucophore Layer P Blood Vessel Connective Tissue P Fat Subcut Areolar Tissue 13 PROCEEDIIVG S OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. X, Nos. 11 and 12, pp. 119-163 July 2, 1921 XI REPORT OF THE PRESIDENT OF THE ACADEMY FOR THE YEAR 1920 By C. E. GRUNSKY President of the Academy In Academy affairs the year 1920 may be classed as rela- tively uneventful. Nevertheless some progress has been made. The membership campaign under the leadership of Mr. M. Hall McAllister has continued successful. The number of members has increased from 550 to 927. There were 403 new members admitted while the loss of members was 20 by death, 5 by resignation, and 1 dropped for arrearages. The present membership is made up of: Patrons 12 Honorary Members 30 Life Members 82 Fellows 20 Members 783 The Academy carries on its list of patrons the following names : LiT/ing Mr. William B. Bourn Mr. John W. Mailliard Mr. William H. Crocker Mr. Joseph Mailliard Mr. Peter F. Dunne Mr. M. Hall McAllister Mr. Herbert Fleishhacker Mr. Ogden Mills Mr. Joseph D. Grant Mr. Alexander F. Morrison Mr. A. Kingsley Macomber Mr. William C. Van Antwerp Deceased Mr. William Alvord Mrs. Oiarlotte Hosmer Mr. Charles Crocker Mr. James Lick Mr. John W. Hendrie Mr. Ignatz Steinhart 120 CALIFORNIA ACADEMY OF SCIENCES [Proo. 4th Ser. Those who were called by death are as follows : Mrs. Mary K. Brandegee Life April 3, 1920 Dr. John A. Brashear Honorary April 9, 1920 Dr. Frank S. Daggett Member April 5, 1920 Mr. I. W. Hellman Member April 9, 1920 Mrs. Charlotte Hosmer Patron November 30, 1920 Mr. G. Earle Kelly Member December 15. 1920 Mr. Curtis H. Lindley Member November 20, 1920 Mr. Bruce Martin Member 1920 Mr. W. W. Montague Life September 28, 1920 Mr. Percy T. Morgan Member April 16, 1920 Mr. Charles P. Punchard Member November, 1920 Dr. Pier Andrea Saccardo Honorary February 12, 1920 Mr. Leon Sloss Member May 5, 1920 Mr. L. E. Smith Life October 12, 1920 Prof. Dr. H. zu Solms-Laubach Honorary 1920 Dr. Franz Steindachner Honorary December 10, 1919 Mr. Sam B. Stoy Member August 18, 1920 Mr. Clement Tobin Member April 3, 1920 Dr. Ramsay H. Traquair Honorary November 22, 1912 Mr. Carlos Trover Life July 26, 1920 Mr. Raphael Weill Life December 9, 1920 The Academy has published during 1920 the following papers in continuation of the Fourth Series of the Pro- ceedings : Vol. H, Part H, No. XVH, pp. 311-345 Dermaptera and Orthoptera, by Morgan Hebard. Vol. IX, No. XHI, pp. 331-356 New Hemipterous Insects of the Genera Aradus, Phytocoris and Camptobrochys, by Edward P. Van Duzee. Vol. IX, No. XIV, pp. 357-365 Report of the President of the Academy for the Year 1919, by C. E. Grunsky. Vol. IX, No. XV, pp. 367-396 Report of the Director of the Museum for the Year 1919, by Barton Warren Evermann. Vol. X, No. I, pp. 1-27 A Further Study of Variation in the Gopher-Snakes of Western North America, by John Van Denburgh. Vol. X, No. II, pp. 29-30 Description of a New Species of Rattlesnake (Crotalus lucasensis) from Lower California, by John Van Denburgh. Vol. X, No. Ill, pp. 31-32 Description of a New Subspecies of Boa {Charina hotter utahensis) from Utah, by John Van Denburgh. Vol. X. No. IV, pp. 33-34 Description of a New Lizard {Dipsosaurus dorsalis lucasensis) from Lower California, by John Van Denburgh. Vol. X] GRUNSKY— PRESIDENT'S REPORT FOR 19^0 121 Vol. X, No. V, pp. 35-46 Undescribed Tipulid.t, (Diptera) from Western North America, by Charles P. Alexander, Vol. X, No. VI, pp. 47-49 Three New Species of Dolichopodid^ (Diptera) from California AND Nevada, by M. C. Van Duzee. Vol. X, No. VII, pp. 51-52 Two New Species of Syrphid^ (Diptera), by A. L. Lovett. Vol. X, No. VIII, pp. 53-70 Notes on Some Undescribed Californian Helices, by S. Stillman Berry. Vol. X, No. IX, pp. 71-75 A New Genus and Species of Grasshopper from California, by Morgan Hebard. During the year 1920, 11 free lectures have been delivered at the stated meetings of the Academy, as follows : January 7. "The Vegetation of New Zealand." Mr. A. H. Cockayne, Government Biologist of New Zealand. March 3. "American Aquariums." Dr. Barton Warren Evermann, Director of the Mu- seum, California Academy of Sciences. April 7. "A Commodity Unit to Supplement Money as a Standard of Value." Dr. C. E. Grunsky, President, California Academy of Sciences. May 5. "Some Bird Observations on the University Campus." Mr. Tracy I. Storer, Field Naturalist, Museum of Ver- tebrate Zoology, University of California. June 2. "The Work of the Audubon Society." Mr. C. B. Lastreto, Organizer and Ex-President of the Audubon Association of the Pacific. July 7. "The Birds and Mammals of the Stikine Region, British Columbia." Mr. H. S. Swarth, Curator of Birds, Museum of Ver- tebrate Zoology, University of California. August 4. "The Botanical Features of Mount Tamalpais." Miss Alice Eastwood, Curator of Botany, California Academy of Sciences. September 1. "Injurious Insects." Mr. E. O. Essig, Assistant Professor of Entomology, University of California. October 6. "Hunting Grizzlies with Bow and Arrow." Dr. Saxton Pope, Assistant Clinical Professor of Sur- gery, University of California. November 3. "The Pan-Pacific Scientific Congress and the Hawaiian Islands." Dr. Barton Warren Evermann, Director of the Mu- seum, California Academy of Sciences. December 1. "Birds, Fur Seals, Foxes and Other Animals of the Pribi- lof Islands, Alaska." Dr. G. Dallas Hanna, Curator of Invertebrate Paleon- tology, California Academy of Sciences. 122 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. The Sunday afternoon lectures delivered in the Museum building during 1920 have included the following: January 4. January 11. January 18. January 25. February 1. February 8. February IS. February 22. February 29. March 7. March 14. March 21. March 28. April 4. April 11. April 18. April 25. May 2. California's Fur-Bearers. Dr. H. C. Bryant, in charge Education and Publicity, California Fish and Game Commission. Life History of the Little Smelt or Grunion. Mr. W. F. Thompson, in charge State Fisheries Lab- oratory, California Fish and Game Commission. The Game Warden and His Work. Mr. J. S. Hunter, Assistant Executive Officer, Califor- nia Fish and Game Commission. The Commercial Fisheries of California. Mr. N. B. Scofield, in charge Commercial Fisheries Department, California Fish and Game Commission. The Ocean as an Abode of Life. Dr. W. K. Fisher, Director, Hopkins Marine Station of Stanford University. The Steinhart Aquarium. Dr. Barton Warren Evermann, Director of the Mu- seum, California Academy of Sciences. The Ocean Meadows, or the Microscopic Life of the Open Sea. Dr. C. A. Kofoid, Professor of Zoology, University of California. Fishes of the California Coast. Prof. E. C. Starks, Assistant Professor of Zoology, Stanford University. Marine Mammals. Dr. Harold Heath, Professor of Zoology, Stanford University. Alaska and the Fur Seals. Dr. Barton Warren Evermann, Director of the Mu- seum, California Academy of Sciences. Life of the Deep Sea. Prof. J. O. Snyder, Associate Professor of Zoology, Stanford University. Oceans of the Past. Dr. J. P. Smith, Professor of Paleontology, Stanford University. Systematic and Economic Phases of California Marine Algae. Dr. N. L. Gardner, Assistant Professor of Botany, University of California. Life Between Tides. Dr. W. K. Fisher, Director, Hopkins Marine Station of Stanford University. Plant Migrations. Dr. Douglas H. Campbell, Professor of Botany, Stan- ford University. The California Big Trees and History. Dr. L. L. Burlingame, Associate Professor of Botany, Stanford University. Some Plant Diseases. Prof. J. L W. McMurphy, Assistant Professor of Botany, Stanford University. The Trees of California. Dr. Leroy Abrams, Associate Professor of Botany, Stanford Universitv. Vol. X] GRUNSKY— PRESIDENT'S REPORT FOR ig^o 123 May 9. September 12. September 19. September 26. October 3. October 10. October 17. October 24. October 31. November 7. November 14. November 21. November 28. December 5. December 12. December 19. Plants and a Hungry World. Dr. George J. Peirce, Professor of Botany and Plant Physiology, Stanford University. The Origin of the Polynesian People. Dr. W. E. Safford, Bureau of Plant Industry, Depart- ment of Agriculture, Washington, D. C. A Recent Visit to the Hawaiian Islands. Dr. Barton Warren Evermann, Director of the Mu- seum, California Academy of Sciences. The Solar System. Dr. W. W. Campbell, Director, Lick Observatory, Mount Hamilton, California. Comets. Dr. A. O. Leuschner, Dean of the Graduate Division, University of California. The Binary Stars. Dr. R. G. Aitken, Astronomer, Lick Observatory, Mount Hamilton, California. The Nebulae. Dr. J. H. Moore, Astronomer, Lick Observatory, Mount Hamilton, California. What We Owe to Animal Experimentation. Dr. S. J. Holmes, Professor of Zoology, University of California. Early Ideas of the Earth. Prof. Earle G. Linsley, Professor of Geology and As- tronomy, Mills College. The Spectroscope — A Key to Celestial and Atomic Mys- teries. Dr. E. P. Lewis, Professor of Physics, University of California. Some Phases of Our Modern Conquest of the Air. Dr. W. F. Durand, Professor of Mechanical Engineer- ing, Stanford University. The Future of the Tropics a< a Factor in World Develop- ment. Dr. R. S. Holway, Professor of Geography, University of California. Cipher Messages from the Stars. Dr. J. H. Moore, Astronomer, Lick Observatory, Mount Hamilton, California. Earthquakes on the Pacific Coast of North America. — I. Dr. S. D. Townley, Professor of Applied Mathematics, Stanford University. Earthquakes on the Pacific Coast of North America. — II. Dr. S. D. Townley, Professor of Applied Mathematics, Stanford University. The United States in Haiti and Santo Dommgo. Mr. M. E. Beall, Berkeley, California. Ground has not yet been broken for the Steinhart Aquarium. This is not due to any lack of desire on the part of the Council and the Trustees to further its early con- struction. The trust which the Academy has accepted will be faithfully discharged. More than a year ago the bequest became available and the sum of $250,000 was paid to the 124 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. Academy. This sum was at once invested and interest has been accumulating. Dr. Evermann, accompanied by Civil Engineer T. Ronneberg, has made an inspection trip to all the notable aquariums in the United States in order that the Academy may have the advantage of the latest and best ideas on general arrangement, lighting and operation. The archi- tect, Mr. Lewis P. Hobart, thereupon made a number of pre- liminary studies from which it presently became apparent that the funds at our disposal were not adequate to provide a building of the size, dignity and architectural finish which would be required if the aquarium is given the place of honor in the Academy's building scheme. It was found im- practicable, in other words, to put the aquarium to the East of the unit in which our present exhibits and activities are housed, where it could be made, if funds permitted, an im- posing central feature at the rear of an aquatic court. When this fact became apparent the architect did the next best thing. He so arranged a design that only a small portion of the ex- terior of the building would require finishing in stone. But even with this arrangement under which only a narrow front of the building would be architecturally ornamental, there has been difiiculty in finding a site at once appropriate and ac- ceptable to the Board of Park Commissioners. The matter of selecting a site and of suggesting a building suited to the site is now in the hands of a joint committee of the Park Commission and of the Academy. It is regrettable that the funds placed by bequest at the disposal of the Academy are limited to an amount which will not srive to San Francisco all that is desired in connection with a first class, fully equipped aquarium in which the ma- rine life of the Pacific Ocean and the aquatic life of the streams which flow into the Pacific Ocean should be ade- quately represented. Even with extreme restriction of the exterior ornamentation of the aquarium building it will not be possible to provide more than about 50 tanks of moderate size. There will be no space provided in accomplishing this result for fishery and display exhibits, for research work, or for offices. The bare housing of the fish tanks, and of the machinery and appliances required to store, filter, aerate and cool or heat the water, is all that can be accomplished with the means at command. Vol. X] GRUNSKY— PRESIDENT'S REPORT FOR 1920 125 Some thought has been given to combining with the aquarium an auditorium or otherwise bringing under the same roof space that may be used for other purposes, with- out in any sense detracting from the distinctiveness of the Steinhart Aquarium. Nothing along this Hne seems possible of accomphshment. The aquarium once estabhshed will al- ways be the "Steinhart Aquarium." Enlargement or exten- sion by other bequests or endowments is not likely so long as the features added by such bequests or endowments are not sufficiently distinctive to perpetuate the names of those whose generosity prompts their making. Nor does it seem likely that the City, which is the beneficiary, could be pre- vailed on in the near future to increase the initial fund. Nevertheless, before it is too late, this thought is thrown out. It would indeed be a gracious thing if the City which has authorized the Academy to accept the bequest and to take charge of the erection and operation of an aquarium in Golden Gate Park, would add a like amount. It would thereby make possible the erection of a dignified structure adequate for immediate needs and arranged for expansion to meet any future requirements. The financial standing of the Academy will appear from the Treasurer's report. The endeavor of the Council and of the Trustees has been to do the utijaost that could be done within our resources as well in the matter of adding to the material in the Academy's collections as in research work and the publication of results. What has been done along these lines will appear more fully from the report of Dr. Evermann, the Director of the Museum, and the reports of the curators of the various departments. As you were advised a year ago our floating debt which had been incurred during the erection of the Museum build- ing, had been wiped out in 1919. This year we note with satisfaction that our indebtedness on the Academy's Market Street property has been reduced by $10,000, from $300,000 to $290,000. For a grizzly bear habitat group, as was noted in the President's last annual report, funds have been provided by Mr. Ogden Mills and the necessary specimens for the group have now been secured from the Yellowstone Park without 126 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. expense to the Academy, by Dr. Saxton Pope, assisted by Arthur Young and G. D. Pope. Permission was granted by the Park authorities to get four grizzly bear for this pur- pose. It is expected that before the close of this calendar year the group will be installed. It will fill the last available alcove and, thereafter, the need for more exhibit space will be felt more than ever. The Academy is a growing institution, as yet but inade- quately equipped to carry on the activities which should be its special concern. It should have more housing facilities for scientific and educational natural history material. It should be engaged in carrying natural history lessons into the school rooms of the city and it should have better facilities for popu- lar lectures on scientific subjects. Despite the present inade- quate facilities the weekly lectures (Sunday afternoons) have proved very successful. Their scope appears from the list submitted. They are attended to the limit of our small audi- torium's seating capacity. I am sure that I voice the senti- ments of all members and friends of the Academy who have attended any of these lectures, when I say that the kindness of those who have given the lectures, frequently at no little personal inconvenience, has been sincerely appreciated. In the President's last annual report note was made of the fact that on appeal to the Supreme Court, the will of Mr. S. F. Thorn under which the Academy was bequeathed some land near Santa Cruz and other property, had been sustained. It was subsequently learned that this decision was not rend- ered by the Court in banc. Upon further hearing, the Court in banc reversed the earlier decision, finding that the will was not holographic because the single word ''Cragthorn" had been inserted with a rubber stamp. The will was declared illegal and it now appears that the announcement of last year was at fault and that the Academy takes nothing under the will. Among the notable acquisitions of the Academy during the year is the W. Otto Emerson collection of bird skins, some 5300 in number. Most of the birds represented in this col- lection, which has great scientific value, are from Alameda County. It has come to us through the generosity of W. H. Crocker and John W. Mailliard. Vol. X] GRUNSKY— PRESIDENT'S REPORT FOR 1920 127 Private subscriptions by a number of friends of the Academy have made possible the purchase of the Albert Prager Herbarium, which contains many valuable plant speci- mens from various parts of the globe. Negotiations for the transfer of this herbarium from Leipzig, Germany, the home of Mr. Prager, to the Museum of the Academy, have been completed. Your officers again commend the zeal and ability with which the curators of the Academy's departments have con- ducted their work, which goes so far in maintaining the standing of the Academy as a scientific institution of real worth. Your officers are appreciative too of the interest taken by the membership in the activities of the Academy, and are gratified to note how popular the museum has be- come as evidenced by the large number of visitors reported by the Director of the Museum. The Academy stands ready to enlarge its usefulness. It can do so materially, however, only through outside help. As opportunity offers this fact should be made known to those who are so circumstanced that they can serve mankind by establishing endowments for useful purposes. They should know that the Academy is ready to serve. 128 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. XII REPORT OF THE DIRECTOR OF THE MUSEUM FOR THE YEAR 1920 BY BARTON WARREN EVERMANN Director of the Mtiseum The annual report of the Director for the year 1919 was presented to the Academy at the annual meeting February 18, 1920. At that time all except one of the spaces for large habitat groups in the mammal and bird halls had been filled. The one remaining space has been reserved for a Grizzly Bear Group. The grizzly bear is now extinct in California. None of us will ever see a living example of that magnificent animal which figured so prominently in the early history of the State; it is said there is no really good specimen of the species in any museum in the world. It has therefore been decided to put in a habitat group of a closely related species (Ursus imperator) which occurs in the Yellowstone Park, Dr. Saxton Pope (a member of the Academy) of San Fran- cisco, offered to secure the necessary animals, without ex- pense to the Academy, if the necessary permit were secured. Application was made to the National Park Service for a permit for Dr. Pope to kill the necessary animals, the pemiit was obtained, and Dr. Pope secured the animals. The taxi- dermists have begun the preparation of the group which, it is believed, will be completed by September, 1921. When Mr. Ogden Mills visited the Museum recently he was so pleased with what the Academy has already accom- plished in habitat group installation that he generously gave his check to the Academy for $5000 to cover the cost of the proposed Grizzly Bear Group. It is very gratifying to know that this Museum is so appreciated and the Academy feels grateful to Mr. Mills for this expression of his appreciation. Small Habitat Groups. — With the completion of the Grizzly Bear Group all the available spaces for large groups will have been utilized. There will remain only spaces for 24 small panel Vol. X] EVERMANN— DIRECTOR'S REPORT FOR 1920 129 groups — five in the bird hall and 19 in the mammal hall. It is hoped that all tliese may be completed within the next few years. PERSONNEL Only a few changes have taken place in the personnel of the Museum within the year. Mr. William Heim, who was employed as taxidemiist, took indefinite leave August 16, 1920, on account of ill health. It is hoped he may be able to return at an early date. Mr. Francis G. Gilchrist, who was employed May 10, 1920, as assistant in the department of Ornithology and Mammalogy, resigned July 30 to re-enter the University of California. Harvey R. Scott was employed as assistant taxidennist from July 28 to August 20. Chase Littlejohn has been employed as assistant curator, department of Ornithology and Mammalogy, since September 13. The employes of the Academy at this date are as follows : Dr. Barton Warren Evermann, Director and Executive Curator of the Museum, and Editor ; W. W. Sargeant, Secre- tary to the Board of Trustees ; Miss Susie Peers, Secretary to the Director; Joseph W. Hobson, Recording Secretary; Miss Alice Eastwood, curator, Department of Botany; Ed- ward P. Van Duzee, curator. Department of Entomology, and assistant librarian; Dr. John V&n Denburgh, curator, Department of Herpetology ; Dr. Roy E. Dickerson, honorary curator, Department of Invertebrate Paleontology; Dr. G. Dallas Hanna, curator. Department of Invertebrate Paleon- tology; Dr. Walter K. Fisher, curator, Department of Inverte- brate Zoology; Joseph Mailliard, curator, Department of Ornithology and Mammalogy; Joseph R. Slevin, assistant curator, Department of Herpetology; Chase Littlejohn, as- sistant curator, Department of Ornithology and Mammalogy; Mrs. Marian L. Campbell and Mrs. Kate E. Phelps, assist- ants, Department of Botany; Mrs. Helen Van Duzee, as- sistant, Department of Entomology and in the Library; Miss Mary E. McLellan, Library assistant; William Heim, as- sistant. Department of Exhibits (on leave) ; John I. Carlson, general assistant; Raymond Smith, general assistant; Georges Vorbe, Merle Israelsky, and William Barbat, temporary as- sistants, Department of Invertebrate Paleontology; William 130 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. C. Lewis, janitor; Fred Maag, carpenter and assistant jani- tor; George W. Edwards, assistant janitor; Frank W. Yale, night watchman; Mrs. Johanna E. Wilkens, janitress; Patrick J. O'Brien, day watch ; Archie McCarte, night watch. ACCESSIONS TO THE MUSEUM AND LIBRARY Accessions to the Museum and the Library have been un- usually numerous and valuable, as is shown by the detailed list in the appendix to this report (pp. 149-158). VISITORS TO THE MUSEUM In accordance with established policy, the Museum was open to the public every day in the year. The attendance by months for each of the years since the Museum was first opened to the public is shown in the following table: Month— 1916 1917 1918 ' 1919 1920 January 23170 25260 17241 27013 February 22058 23698 19586 23450 March 31606 26810 27397 25419 April 32175 23274 25994 32208 May 26154 26391 28369 37107 June 32123 29843 32248 36207 July 37193 31420 48028 52492 August 24619 31137 43730 53470 September 16448' 27866 29847 34007 42413 October 36933 20629 14743= 30463 33500 November 27718 21810 853^ 25246 19347 December 15002 21693 19588 21246 21340 Total 96101 321096 290542 351497 403566 A comparison of the number of visitors to the Museum of the California Academy of Sciences with those who visited the Smithsonian Institution and the National Museum at Washington in the year ending June 30, 1920, will prove very interesting. The visitors by months are given in the following table, which shows that the number visiting the Smithsonian Institution was only 21 per cent of those visiting the Academy Museum; the number visiting the Natural His- tory Building of the National Museum was only 4.7 per cent, greater than that at the Academy, while that at the Arts and Industries Building of the National Museum was only 62 per cent of that at the Academy, These figures should be very gratifying to us. 1 Attendance from September 22 to 30. .* Museum closed 29 days on account of the "flu." Vol. X] EVERMANN— DIRECTOR'S REPORT FOR igso 131 Number of Visitors by Months for the Year Ending June 30, 1920 AT THE Museum of the California Academy of Sciences The Smithsonian Institution AND National Museum in Washington, D. C. V(^nr California Academy of Sciences Smithsonian Institution United States National Museum and Month Arts and Industries Building Natural History Building 1919 Tulv 52,492 53,470 42,013 33,500 19,347 21,340 27,013 23,450 25,419 32,208 37,107 36,207 7,812 9,594 9,690 7,245 5,875 4,992 4,264 3,439 6,371 8,121 9,978 ^ 8,632 24,755 29,501 29,697 21,401 18,971 13,149 11,491 10,168 15,815 23,207 27,556 25,271 33,631 Aueust 45,392 September 44,974 October 36,906 November 38,420 December 23,751 1920 January 22,914 February 21,740 March 32,204 April 38,954 May 46,089 June 38,009 Total 403,566 86,013 250,982 422,984 COOPERATION WITH SCHOOLS The Museum continues to cooperate with the pubhc and private schools. Teachers and school officials are coming to realize more and more clearly that the Museum can be of real service to the schools. The number of schools visiting the Museum increases each year. During the year 1920 there were schools from San Francisco, Berkeley, Oakland, Ala- meda, Piedmont and Richmond, and classes from San Jose, San Rafael, San Anselmo, San Mateo, and Stockton. When- ever possible, the Director or some one of the Museum stafif accompanies the school through the Museum and explains briefly the general features of the installations, calling at- 132 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. tention to the educational value of the various exhibits, and then, when time permits, the class is taken into the lecture hall and shown moving pictures or stereopticon slides il- lustrative of some of the exhibits. The visits by schools in the year have been as follows: Schools of San Francisco Number of classes 280 Number of teachers with the classes 270 Number of pupils 19920 Schools outside of San Francisco Number of classes 26 Number of teachers with the classes 18 Number of pupils 444 Total number of classes 306 Total number of teachers 288 Total number of pupils 10364 THE PAN-PACIFIC SCIENTIFIC CONGRESS Under the auspices of the Pan-Pacific Union, the First Pan-Pacific Scientific Congress met at Honolulu x\ugust 2 to 21. The Director of the Museum attended the Congress, as a member of the Pacific Explorations Committee of the National Research Council and representing also the Califor- nia Academy of Sciences and the Pacific Division of the American Association for the Advancement of Science. He sailed from San Francisco July 28 and returned September 8. August 3 to September 1 were spent attending the ses- sions of the Congress and visiting places of interest on the islands of Oahu and Hawaii. The number of delegates and others in attendance upon the sessions of the Congress was more than 100. There were present delegates from New Zealand, Australia, the Philip- pines, China, Japan, Canada, England, and the United States, many of those from the United States representing various scientific bureaus of the Government. The papers and discussions at the sessions of the Congress covered many phases of the geology, meteorology, and natural history of the Pacific and its contained islands, and it was clearly shown that much of importance remains to be learned Vol. X] EVERMANN— DIRECTOR'S REPORT FOR 1910 133 regarding the hydrography, geology, and natural history of the Pacific. At the close of the Congress the unanimous verdict was that the meeting had been a decided success, and that other meetings should be held every two or three years, FIELD WORK OF THE MUSEUM STAFF Within the year the Museum carried on a number of field investigations, as fully set forth in the reports of the curators. MEETING OF THE PACIFIC DIVISION OF THE AMERICAN ASSOCIATION FOR THE ADVANCEMENT OF SCIENCE AND ITS AFFILIATED SOCIETIES The 1920 meeting of the Pacific Division was held at Se- attle, June 17 to 19. All the sessions of the Association and of its affiliated societies were held in various halls of the University of Washington. The Academy membership was well represented, at least 25 of its members being present. The Director of the Museum was in attendance as vice-presi- dent and chairman of the executive committee of the Pacific Division, and Secretary W. W. Sargeant as secretary of the Pacific Division. The Academy was represented on the pro- grams by the following members: ^ Mr. W. F. Thompson Dr. E. L. Packard Dr. Barton Warren Evermann Dr. Chester Stock Prof. John N. Cobb Dr. Wm. E. Ritter Mr. W. E. Allen Prof. Tracy I. Storer Air. Willis H. Rich Dr. E. P. Meinecke Dr. John C. Merriam Dr. W. F. Durand Dr. E. P. Lewis Dr. J. R. Slonaker Dr. J. W. Moore Miss Alice Eastwood Dr. R. G. Aitken Dr. D. T. MacDougal Dr. Bruce L. Clark Several others were in attendance. USE OF THE academy's COLLECTIONS AND LIBRARY BY INVESTIGATORS AND STUDENTS Students and investigators continue to avail themselves of the facilities offered by the Academy for study and research. The Library, with its more than 50,000 books and pamphlets 134 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. in the various departments of science, is in almost daily use by students. This has been especially true of geologists and paleontologists, who are interested in oil investigations and prospecting. The large research collections in the different departments have been consulted by a number of investigators. Mr. L. M. Looniis continues to avail himself of the Acad- emy's collections and Library in his study of sea birds. Messrs. Joseph Grinnell, Harry S. Swarth, Tracy I. Storer, and J. Eugene Law, of the Museum of Vertebrate Zoology, have had occasion to consult our collections of birds, mammals and reptiles. Numerous entomologists of the west coast make frequent use of the rapidly growing collections in that de- partment, and the botanists of the coast find it to their in- terest to consult the herbarium. The Boy Scouts, under Scout Master Harold E. Hanson, have continued the Friday night meetings at the Museum throughout the year. The Academy is glad to cooperate in this way. PUBLICATIONS BY THE MUSEUM STAFF The curators and other members of the Museum staff have continued active in research work and in their contributions to the scientific literature of their respective fields of investiga- tion, as evidenced by the following list of titles of papers published by them in 1919-1920: Evermann, Barton Warren 1. Nesting of the Wilson Snipe in California.