caren sehen ate ining tonto a - a Ms SS yememen nee Reet rer ‘ " at 2 ney pe ae te! : : ies i) Rhy, Ge iy ( a oe ee] cs ‘ a : , eye ; | ' r . =, 5 } ; 7 : : ? ah: ty | , : a \ : So SPR EEDING SC | cpp “rve7 @ maomiorna Acadent (OL Screncs wen vie . f —< 5 Le ee see 5 4 | , . ° ‘ P ie. ‘ ; ‘ . ’ * y y . } a * ea - ‘ ‘ - + <s a —_ —— — _ —<— ~ . . dl . be : “ 7 - » * aa. ‘ - ‘ ) 4 + ‘ \ ' mi PROCEEDINGS OF THE California Academy of Sciences Msrine Ej-ios: — mus 4102-Cal Laboratory LIBRARY tory SEP 1 8 1967 WOODS HOLE, Mass, FOURTH SERIES Vol. XXIX SAN FRANCISCO PUBLISHED BY THE ACADEMY 1956-1960 COMMITTEE ON PUBLICATION George E. Lindsay, Chairman Leo G. Hertlein Edward L. Kessel, Editor No. No. No. No. No. No. 10% Hale aot2: CONTENTS OF VOLUME XXIX JoHNSON, Martin W. The Larval Development of the California Lobster, Panulirus interruptus (Randall), with notes on Panulirus gracilis Streets (22 text figures). Pub- lishedeNovember*26;1056' = 2 2 eS ee SMITH, ALLYN G. Snails from California Caves (2 plates). Ruplished! janvdEyags. 105) =e ee ee Haas, Georc. Some Amphibians and Reptiles from Arabia (12 text figures). Published November 5, 1957 SOULE, JoHN D., and Mary MarsH Durr. Fossil Bryozoa from the Pleistocene of Southern California. Published INORG CYS) G5, 98 SY | ea = EE Ne OR acd eee en Va : McCiintTock, ELizABETH. A Monograph of the Genus Hydrangea (6 plates; 10 maps). Published November 5, 1 Sy /eee e cae OEN rme S) a OY a e JoHNsoN, Martin W. Bathycalanus sverdrupi, n. sp., a Copepod Crustacean from Great Depths in the Pacific Ocean (15 text figures). Published July 10, 1958 Menzies, Ropert J. The Distribution of Wood-Boring Limnoria in California (3 text figures). Published July HOMO S Que nome PON oe ee ee ee Hupss, Cart L., and ANDREAS B. RECHNITZER. A New Fish Chaetodon falcifer, from Guadalupe Island, Baja Cali- fornia, with Notes on Related Species (3 plates; 1 map). Pumsmeds [itihyat O05 8, 28. oes ae ee ee eee nee RICHARDS, ADRIAN F., and Bayarp H. BrRattTsTRoM. Bibliography, Cartography, Discovery, and Exploration of the Islas Revillagigedo (4 text figures). Published January 5. GONE eet One. Cot ek ON aE RT TT Site LE ATE oye HowpENn, ANNE T. A Revision of the Species of Pan- deleteius Schonherr and Pandeleteinus Champion of America North of Mexico (Coleoptera: Curculionidae) (2 plates). PublishedManuaryco~ [050m eee ee ee q STOHLER, Rupotr. Two New Species of West North Ameri- can Marine Gastropods (10 text figures). Published Febru- DEV 22 pe LO 0 Wee seit ieee ee ese Se SR eo LEviton, ALAN E. Report on a Collection of Reptiles frem Afghanistan. Published February 27, 1959 PAGES 147-256 257-265 267-272 273-313 315-360 361-421 423-444 445-463 No. No. No. No. No. Wee HERALD, Eart S. From Pipefish to Seahorse—A Study of Phylogenetic Relationships (3 text figures). Published May DOs MO SOs So Sutels ool eA ms. Se ee a Leviton, ALAN E., and WALTER C. Brown. A Review of the Snakes of the Genus Pseudorabdion with Remarks on the Status of Genera Agrophis and Tvphlogeophis (Serpentes: Colubridae) (10 text figures). Published May 90 1050 wt ee SL LOUKASHKIN, ANATOLE S., and NoRMAN GRANT. Behavior and Reactions of the Pacific Sardine, Sardinops caerulea (Girard), Under the Influence of White and Colored Lights and Darkness (23 text figures). Published May 29, 1959 McCautey, James E. The Morphology of Phyllaplysia zostericola, New Species (6 text figures). Published May OWNGGO! cvek cteseel A ee ee EXLINE, Harriet. Rhoicinine Spiders (Pisauridae) of Western South America (5 plates; 1 map). Published May OO GOs oo alegre she mee ee ee Imdex'to: Volume. Xochk <2 = oe" ess eek ee Errata 465-473 475-508 509-548 549-576 621 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. XXIX, No. 1, pp. 1-19, 22 figs. November 26, 1956 THE LARVAL DEVELOPMENT OF THE CALIFORNIA SPINY LOBSTER, PANULIRIS INTERRUPTUS (RANDALL), WITH NOTES ON PANULIRUS GRACILIS STREETS’ BY MARTIN W. JOHNSON Scripps Institution of Oceanography University of California, La Jolla, California a INTRODUCTION The California spiny lobster, Panulirus interruptus (Randall), is the only lobster occurring on the coast of California. Its main distributional range is usual:y considered to be from slightly north of Point Conception, California, to the Gulf of Tehuantepec, Mexico. For many years it has been a highly prized item in the fisheries of California and is considered eleventh in importance by value among the fisheries of the state (Wilson, 1948). The amount of research accorded the lobster has, however, not been commensurate with this economic im- portanee or with its biological interest. During its life history, the spiny lobster passes through a series of eleven floating larval “phyllosoma” stages in which the body is highly transparent and exceedingly compressed dorsoventrally. The last phyllo- soma stage metamorphoses to a ‘“‘puerulus” stage in which the animal is still transparent but structurally similar to the adult. The puerulus of some species may be found in plankton, but evidently it soon deposits calcium in the carapace and seeks the bottom to live in concealment. 1 Contribution from Scripps Institution of Oceanography, New Series, No. 878. [1] 2 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Allen (1916) carried out certain fundamental observations on the adult animals particularly in the vicinity of Santa Barbara, the Channel Islands, and La Jolla. His findings are especially pertinent to the present study in showing that the berried females may carry 5,000 to 500,000 eges each, depending upon size, and that the spawning season extends from March to August with its maximum activity in May, June, and July. He con- cluded that a period of from nine to ten weeks is probably required for hatching. The young hatched readily in aquaria, but attempts at rearing Figure 1. Stage I of Panulirus interruptus phyllosoma hatched in aquarium. Labels: a1, first antenna; a2, second antenna; cs, coxal spine; en, endopod; ez, exopod; mxp 3, third maxilliped; p 1, p 2, p 3, first, second, and third legs. Vou. XXIX] JOHNSON: LARVAL DEVELOPMENT OF SPINY LOBSTER 3 the larvae were futile. Other investigators have had similar experiences in attempts at rearing the phyllosoma larvae of other species. In a recent study, Lindberg (1955) found a maximum of 800,000 eggs produced by large females. The spawning season was essentially the same as indicated by Allen (1916). With regard to our knowledge of the biology of phyllosoma larvae in nature, rather little is known because of their scarcity, coupled with the lack of complete and identified developmental series. Their offshore or oceanic distribution also adds to the difficulty. In 1919 Sehmitt published a brief note on phyllosoma larvae caught off the coast of southern California. Larvae of several sizes were observed. The details of structure and stages of development were, however, not worked out in any ease, but a photograph was included showing a large specimen which appears to represent either the tenth or the eleventh phyllo- soma stage. A practical understanding of the biology of the California spiny lobster will require information regarding the abundance and extent of dispersal of its planktonic larvae from various areas within the geographical range of the species. It is probable that many of the larvae produced in the northern range of adult distribution are lost by drifting away from the area with the California Current. Unless many of the northern larvae are returned by some circuitous route in large local eddies, the mainte- nance of the local stock would depend in considerable measure upon re- eruitment of larvae from more southern waters, probably with the Da- vidson Current. A knowledge of the duration of the planktonic larval stages and the rate and direction of ocean currents is highly important in this connection, but we have no direct information on how long the larvae live in the plankton, though studies now in progress of the seasonal distribution of the stages point to a planktonic period of several months. The series of phyllosomas examined from plankton collections along the California coast to south of Cedros Island have not revealed any sig- nificant irregularity in structure, shape, or size, of the various develop- mental stages, indicating that only one species is involved in the northern range. However, Panulirys gracilis (until recently known as Panulirus mflatus (Bouvier) ) does overlap the southern range of Panulirus inter- ruptus at least in small numbers and enters into the commercial catch of Mexico, (Chase and Dumont, 1949). In a review of the conflicting synonymy of P. inflatus, Holthuis (1954) has concluded that the species is a synonym of P. gracilis Streets. The latter, being the older name, must be used. The larvae of P. gracilis may be expected to drift northward should they be caught in the Davidson Current, especially during its maximum development along the coast. For this reason, among others, it is essential 4 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. that the larval structure be known in detail in order to facilitate definite identification of the planktonie stages of the two species. This identification can now be made as a result of a study of larvae caught within the range of P. gracilis south of Point Eugenia and within the Gulf of California. The larvae of the two species are distinguishable on small but definite structural characteristics, which will be briefly mentioned here and re- ported upon in detail later. A survey of the seasonal and spatial distribution of the larvae is in progress. THE PHYLLOSOMA STAGES OF P. INTERRUPTUS Large numbers of the first phyllosoma stage were obtained in 1940 by hatching of eggs from a berried female kept in an aquarium (figure 1). Subsequent stages (figures 2 to 18) were all obtained from plankton hauls, especially from collections made along the coasts of southern and Baja California in connection with the Marine Life Research Program of the California Cooperative Oceanic Fisheries Investigations. ‘areful anatomical analysis of large numbers of larvae reveals eleven rather poorly defined developmental stages as summarized in table I. There is a considerable range in size within each “stage,” especially in the later stages, and specimens are often observed in which structures such as newly added legs, exopods, gills, second maxillae, and first maxillipeds show noticeable range in size or shape or even degree of segmentation. Whether or not this signifies a molt within the stage is not known, but it may be significant that the more advanced occur in the upper-size range for the stage represented. The fore-body (cephalic shield) of Panulirus interruptus is pear-shaped in outline in the early stages, and is much wider than the hind-body (thorax) during the first and second stages. But in the third stage, the hind-body has attained the width of the fore-body and remains thus dur- ing the subsequent stages. The pear-shaped fore-body of the earlier stages later becomes more uniformly oval as shown in figure 18. In no stage is the posterior margin of the hind-body deeply indented. In stage I, the eyes are not stalked, but in stages II to XI, they are on definite slender stalks which increase steadily in leneth with each molt. In all stages there is present a median eye spot. The first antennae (antennules) exceed in length the second antennae (antennae) during the first four stages, after which the second antennae Figures 2-5. Stages of Panulirus interruptus phyllosoma (continued). Labels: p 3, p 4, third and fourth legs. The distal ends of lengthy appendages showing no significant changes from stage to stage have been omitted in these and subsequent figures. VoL. XXIX] JOHNSON: LARVAL DEVELOPMENT OF SPINY LOBSTER Figure 2. Stage II. Figure 4. Stage IV. < 4 Figure 3. Stage III. Figure 5. Stage V. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. p4 Ww i f- 4 {ZN \\ LS GEN WA \ AY SSSs8 Figure 6. Stage VI. Figure 6a. Stage VI, Tip of third maxilliped. Figure 6b. Stage VI, Tip of first leg. Figure 6c. Stage VI, Tip of third leg. Figure 7. Stage VII. Figures 6-7. Stages of Panulirus interruptus phyllosoma (continued). Labels: p 4, p 5, fourth and fifth legs. VoL. XXIX] JOHNSON: LARVAL DEVELOPMENT OF SPINY LOBSTER i progress rapidly with each successive stage until in stage XI they exceed in length the fore- and hind-body combined. In stage I, the distal end of the first antenna bears a tuft of 4 setae and in each subsequent stage additional tufts are added proximally as transverse rows of setae. The endopod is first evident as a low bud in stage IV. In following stages it elongates, becoming a true branch, but re- mains small throughout the development and is not a free segment until stage VIII. Beginning in stage VI (sometimes stage VII), there are three segments in the pedunele, the first segment being equal to the combined leneths of segments two and three which are subequal. These proportions remain about the same in the next two stages, but in stages VITI-XI the third segment is somewhat longer than the second though both combined still equal the length of the first segment. The second antennal peduncle becomes three segmented in the seventh or eighth stage. These segments remain about equal, the third being slightly the longest. The structure of the mandibles and the first maxillae in stage XI is shown in figure 16. These appendages are very short and strong. The mandibular blade, when viewed from the ventral side of the larva, 1s seen from the edge. In figure 16, it is tilted slightly and the labrum lifted to show the broad scoop-shaped blade with the stronger teeth at the outer and inner angles. The two branches of the first maxillae are armed re- spectively with three strong hooks and three flexible setae, but prior to stages VII and VIII there are only two hooks and setae. There is present a small vestigial palp bearing two setae. In stage I the palp is prominent but diminishes in intermediate stages and later again enlarges. Thus these mouthparts, together with the second maxillipeds which are armed at the tips with small claws and heavy setae (figure 15) indicate that the larva must feed upon particulate material gathered without the aid of a real filtering device. The second maxillae in the first stage are composed of two segments, the distal one being much narrower. Each segment bears four long setae which in intermediate, and later stages are lost or shortened. There is but little change in this size relationship (figure 11) until about the sixth stage and in ensuing stages when the segments become noticeably flattened and the distal segment increases greatly in breadth especially posteriorly (figures 12-15). Similarly the first maxillipeds are at first simple buds which in the later stages gradually increase to three-lobed appendages (figure 15). The second maxillipeds are functional feeding appendages in all stages. There is no change from the first until the eighth stage when an exopod bud appears. In stage IX, this bud becomes separated as a short segment which in the next stage elongates and bears 2 to 4 setae at the tip and 8 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. a gill bud appears on the distal portion of the coxa. In stage LX, the exopod has become setose with five or six pairs of setae, the gill has become leaf- like, and a second gill bud has appeared on the proximal portion of the coxa. The third maxillipeds are very long and the exopod is already present and setose in the first stage. With each successive stage there is, however, an increase in the number of segments in the exopods. The number of segments as shown by the number of pairs of setae appears to be regular for each of the earlier stages, but become irregular in the later. The range in numbers of exopod segments for the third maxillipeds and the first and second legs for stages I to XI is respectively: 4 to 25; 6 to 32; and 6 to 33. The principal features of the development of the legs, especially the third, fourth, and fifth pairs, are adequately covered in table I which, together with the figures, will show the changes taking place with each stage. In contrast to the rapid development of the fourth legs, the fifth pair remain as very small buds through stages IIT to VII, and never exceed the abdomen in length. In no stage of the development of the legs is there present a strong spine at the distal end of the second basipods as shown for Panulirus gracilis (figure 22). The inner branches of the legs are readily lost so that a complete attached set of legs has not been found beyond the sixth stage, but judging from loose appendages found in the jar with the later stages the end seg- ments do not change after stage VI (figures 6a—6d). Coral spines are present in the first to ninth stages, but are usually all lost in the tenth, and are never present in the eleventh. In stage X gills are present on the dorsal distal end of the coxal seg- ment of the maxillipeds and legs and in stage XI they occur also on the proximal portion of the coxa at the juncture of the carapace and on the dorsal margin of the thorax at the base of these thoracic appendages. In stages V to XI, there are present spines on the ventral surface of the thorax at the bases of the third maxillipeds and each of the legs. The abdomen, in stages I to V, remains about the same with two small points at the tip. Complete segmentation occurs in the ninth stage though earlier indications of segments may be seen as blocks of tissue under the skin. The wropods become evident as buds in the sixth stage. They increase in size and definition with each successive stage and in stage IX are well- formed biramous appendages, but the rami are not separated from the basipods and their outer edges are entire. Upon reaching the tenth stage, the uropods are completely segmented and a small notch with a marginal Figures 8-17. Stages of Panulirus interruptus phyllosoma (continued). Labels: ex, exopod; g, gill; mx 2, second maxilla; maxp 1, maxp 2, first and second maxilli- peds; p4, p95, fourth and fifth legs. VoL. XXIX] JOHNSON: LARVAL DEVELOPMENT OF SPINY LOBSTER 9 Figure 8. Stage VIII, anterior end. Figures 11-12. Stages IV and VIII, second Figure 8a. Stage VIII, posterior end. maxilla with first and second maxillipeds. Figure 9. Stage IX, anterior end. Figure 13. Stage IX, same, reduced scale. Figure $a. Stage IX, posterior end. Figure 14. Stage X, same. Figure 10. Stage X, anterior end. Figure 15. Stage XI, same. Figure 10a. Stage X, posterior end. Figure 16. Stage XI, mouth parts: labrum (Figures 8-10a drawn to same scale.) (1) ; mandible (m); first maxilla (mz 1). Figure 17. Stage XI, pleopod. [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 10 Te Uevy} Jasuo, = + 98S0}9S = ‘O}aS (MBO SUIPN[IXxe) JUNUIsOS = SUIJUeM = 0) ex0ood UO (S)[[Is = 3 ‘S]IeJEp LoyJINJ 10J soinsy pue xo} veg podoxs = xa podopus = usa jusseid oulds [Bxoo = so svUUS}JUe PUOIIS = 7 B svuusJUue JsIg = TP oF | A SNOW®BIIG MBIO puB Boz if 32% ‘so OU BZ euUlveS poJUsWUIs9S ‘“SosF oules aules ynq oules 4nq aUIeS ‘0J0S "xO aules eules Z1S- 92% IX pnq ‘s T SNOUIB.ILG png ‘3 [ png ‘3 [ ‘so OU pnuq‘s[T pnq'st aules ‘sosun ‘SOS fF aules aules ynq sues aules ‘SOS [ ‘xo oules oules €?2-06L xX ez [dw090 at) ) “39S T SnHOWBIIq Spnq suo, ‘SaSzZ oules eures eules oules 10 pnq*x9 oules oules G6I-0'9L XI 499 qJeTo “so OU spnq spnq "S398 (L)Z oules ynq sures oules aules pnq ‘xo oules aules 0°9T-6°Z1 IIIA (qyeTo Soul} ‘39S T ‘ua -9UI0S ) + ‘39S & spnq spnq ‘SOS T aules oules oules oules “SOS fF sTseq O'LI-¥'6 IIA ‘39S [ ‘Ua ‘09S "xo + ‘39S &-Z spnq MOT 0 pnq “SOS F oules aules eules "B08 Z STSeq G8 —6°) IA ‘SOS [ ‘Xo pnq ‘ua 0 0 pnq “30S & ames oules oules ‘SOS T "BOS Z 8°9 -0'9 A pnq ‘xo pnq ‘ue 0 0 pnq ‘398 (Z)T oules oules oules “SOS T ‘SOS T 7G -9'F AI ‘OJOS "xo 0 0 pnq ‘30S T “So oules oules ‘SOS T “SOS T 8°& -E'E III ‘S08 [ "xo 0 0 0 pnq "so oules aules ‘SOS T ‘SOS T 82 -PZ II “So ({[eUIS } "so 0 0 0 0 ‘so ‘0J9S*xXO ‘OJOS ‘xO ‘398 T ‘SOS T GL -Z'T T spodoin spodoajd G ba] i fa] & 6a] @‘T 621 Gg dru c dru ED [ D ‘Ww Yi6U2] 2a6D78 SOILSIUMLOVUVHO ONIHSIOONILSICG TNAASA AO AUVINNWOAS SIOVYS BVULOSOT[AYG—SsnNgdnisazur snuynuvnd J W1daviE JOHNSON: LARVAL DEVELOPMENT OF SPINY LOBSTER 11 VoL. XXITX] LL a <4 a, owe, ~~ WW S77 Wit 4 a = SM te 7 - Stage XI of Panulirus interruptus phyllosoma. Figure 18. 12 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. spine occurs on the outer margin of each branch (figure 10a). In the last stage, this notch and spine are more pronounced. The pleopods appear first in the eighth stage, but they may be indi- cated earlier under the skin. In stage IX, they may appear either as blunt notched projections or somewhat more advanced, more elongated and flat- tened structures. In the next stage, they are more leaf-like and a notch appears on the inner margin of the endopod. Complete segmentation occurs in the last stage and the notch on the inner border now sets off a well- defined blunt spine-like process (figure 17). THE PHYLLOSOMA OF PANULIRUS GRACILIS Phyllosoma larvae of another species of Panulirus have been collected on 10 separate cruises at a total of 19 stations lying off Baja California, and on two cruises at eight stations in the Gulf of California. These larvae are being referred to Panulirus gracilis, the only other species known from the West Coast. From figure 22, it will be seen that this larva resembles that of Panulirus interruptus but in distinction from that species, possesses a strong spine at the distal end of the basipod on the posterior dorsal side of the segment closely adjacent to the exopod of each of the first three pairs of legs (figure 22 sp.). The complete series of larvae have not yet been found, but the spine persists as a distinguishing characteristic up through at least the eighth stage. Other differences have also been observed, but more material is needed for further study. THE PUERULUS STAGE OF PANULIRUS INTERRUPTUS (Figures 19-21) - _ During the metamorphosis from the last phyllosoma stage to the puer- ulus, the cephalic shield of the phyllosoma is greatly reduced in length and width. The thoracie¢ region is also narrowed and fuses with the cephalic shield to form the cephalothorax. Thus the two erstwhile leaf-like distinet anterior regions of the larvae now appear as one region (figure 19). This, however, remains to some degree dorsoventrally compressed and adult spination is beginning to appear on the dorsal surface. The abdomen has become slightly broader and noticeably longer. Other conspicuous changes include loss of the exopods of the thoracic appendages, reduction in the Figure 19. Puerulus stage of Panulirus interruptus, dorsal view. Figure 20. Puerulus of P. interruptus, diagrammatic habit sketch. Figure 21. Puerulus of P. interruptus, first pleopods. 13 VoL. XXIX] JOHNSON: LARVAL DEVELOPMENT OF SPINY LOBSTER 14 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. leneth of the eye stalks, and a greatly increased length of the second an- tennae to three times the body length. The puerulus, while similar in appearance to the adult, is wholly trans- parent except for the eyes and pigment bands at intervals along the sec- ond antennae, and the walking legs are carried more laterally (figures 19 and 20). The larval median eye is still present and statocysts occur in the first segment of the first antennae. The larval peripodal exopods which served as swimming organs in the phyllosoma stage are wanting and the animal is now apparently in the process of deserting the planktonic habit, but before deserting this pelagic existence the pleopods have developed into efficient swimming appendages. They have become heavily setose and each endopod bears a strong coupling process provided with hooks to interlock with a similar process of the oppo- site endopod (figure 21). Thus, in swimming, the appendages lend support to each other for more efficient propulsion. Only a few specimens of the puerulus stage have been caught and these only from near-shore sampling at La Jolla, April, 1947; in Baja Cali- fornia at Bahia San Roque, September, 1949; and at Cedros Island, August, 1952, where John MeGowan of Seripps Institution dip-netted four speci- mens. The animals are attracted to a light at night and most specimens have been caught in this way. The transverse dorsal grooves characteristic of the abdominal segments of Panulirus interruptus in older stages were not yet formed in the specimens. The size ranged from 20 to 21 mm. This agrees with the size reported by Gilchrist (1916) for Jasus lalandw in which phyllosoma of 33 to 35 mm. produced puerulus stages ranging from 22 to 26 mm. in length. DISCUSSION It has generally been assumed that the phyllosoma larvae are planktonic up to the puerulus stage. However, in experimental cultures Gilchrist (1916) found that after about six days the first phyllosoma larvae of Jasus lalandiit avoid light and return to the bottom. But the evidence is not clear on this point for he mentions the possibility of larvae seeking the bottom following each molt. Von Bonde and Marchand (1935) make a similar statement regarding the first phyllosoma of that species, and Kino- shita (1934) indieates the same for Panulirys japonicus. In no case was the duration of the whole series of phyllosoma stages known. Studies made on Panulirus longipes in Australian waters led Sheard (1949) to conclude that early phyllosomas are found in surface waters; the mid and late stages in mid to surface waters (migrating diurnally) ; and the late phyllosoma and puerulus stages in surface water. The dura- tion of larval life was estimated to be about five months. The phyllosoma larvae of the California spiny lobster have not been VoL. XXIX] JOHNSON: LARVAL DEVELOPMENT OF SPINY LOBSTER 15 taken in large numbers per haul in the plankton off the southern California coast. Usually they are widely dispersed and when present occur as one or a few individuals at a station. Stage I may, however, be quite abun- dant during August and September at coastal stations, especially off Baja California. The maximum concentration observed (cruise 28 station 105.32) was 488 per 1000 cubic meters of water strained by a 1-meter net. Data from analysis of plankton collected monthly over a period of several years from north of Point Conception to the tip of Baja California, and compris- Figure 22. Stage I of Panulirus gracilis phyllosoma. Label: sp, spine. 16 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ing hundreds of stations with routine sampling from 140-0 meters with a 1-meter net, show that complete cruises over the whole area sometimes yield no larvae whatever or only one to five late-stage individuals during January through May. More complete analysis of these data is needed and will be given in a later report. The finding of later stages floating above the 140-meter level in waters of depth ranging to several hundred meters and over 300 miles from shallow water shows that at least many of the larvae of our species do not seek the bottom regularly in the phyllosoma stages. The scarcity of the larvae in general may, however, indicate in part a tendency of the larvae to seek levels so near the bottom that they are not adequately sampled by most of our plankton towing. That this may be so, at least in shallow water, is suggested by the finding of twenty specimens in a dredge haul made by Robert Bieri and John Bradshaw of Seripps Institution with a small specially constructed net dragged along the bot- tom for a distance of 17 meters at a depth of 13 meters. These larvae were, however, all in stage I and may have been just recently released in the area and not yet undergone dispersal by swimming or water currents. Midwater-trawl hauls of long duration taken at depths ranging to 1500 meters or more over very deep water have yielded later stage larvae. The depth of actual catch, however, is not known and the amount of water screened in these hauls is relatively enormous when compared with that screened during the 1-meter net hauls. In view of the many deep hauls that have not yielded larvae, no conclusions can be made from these instances relative to the depth of concentration or habit of seeking the bottom. Nakamura (1940) working with the Japanese species, Panulirus japon- icus, states that the embryonie period is passed through in about one month and he deduced from related data that the phyllosoma stages require about the same length of time to reach the puerulus stage. This seems much too short a period. The status of the systematics of phyllosoma larvae was reviewed by Gurney (1936) in his analysis of the Discovery collection which in some 400 specimens contained about a dozen different generic or specific forms. Owing to lack of complete series, however, characters used for recognition of several genera, including Panulirus, were given with considerable res- ervation. The present complete series of larvae of Panulirus interruptus is therefore of special use toward establishing the characters which delineate that genus. The larval structure of Panulirus interruptus agrees in general with that of larvae known or believed to belong to other species of the genus. There is, however, a notable difference, no doubt specific, in the relative widths of the main divisions of the body of the older larvae. On the basis of material available to him, Gurney (1936) concluded VoL. XXIX] JOHNSON: LARVAL DEVELOPMENT OF SPINY LOBSTER 17 provisionally that in the genus Panulirus the hind-body is wider, sometimes much wider, than the fore-body. This diagnostic characteristic was estab- lished for later stages on Gurney’s Panulirus larvae A, B (P. argus), and D. Later Lebour (1950) and Lewis (1951), dealing with Panulirus argus, verity that the hind-body of that species becomes noticeably broader in the later stages. In Panulirus interruptus, however, the character does not hold, for in this species the hind-body has attained the width of the fore-body in stage III and remains equal in width throughout the ensuing stages. Thus the diagnostic description must be amended to accommodate not only the young stages of Panulirus argus as pointed out by Lebour (1950) but also the later stages of the present species. This makes it a rather unreliable generic characteristic, but useful specifically. Just what larval details must be stressed in the diagnosis of the species of Panulirus are, of course, not fully established as yet and it is likely that useful characters have been overlooked or omitted in the descriptions of series or isolated stages of larvae. Among the characters that may have some specific use are the spines observed in Panulirus mterruptus on the ventral side of the thorax at the base of each leg. Such spines are not shown or mentioned for other species, but Lewis (1951) in figure 4 B. and C. does show a series of structures suggesting low knobs on the ventral mid-line of the thorax in later stages of Panulirus argus. The presence and degree of persistence of coxal spines on the legs and third maxilliped appear also to vary with species within the same genus. In Panulirus argus no eoxal spine develops on the third maxilliped and those on the legs persist only through the fifth stage, whereas in Panulirus mterruptus the third maxilliped and all but the fourth and fifth pairs of legs bear these spines up through the ninth stage. There may be some variability in the later stages. The presence of a spine at the distal end of the basipod adjacent to the exopod of the first three pairs of legs as mentioned in the text for P. gracilis serves to separate that species from P. interruptus, but other characters such as relatively wider fore-body may be needed to separate it from Gurney’s phyllosoma Panulirus A. An obvious difficulty in establishing certainty in the systematics of phyllosoma larvae arises either from a considerable variability in the larval structures of some species or from erroneous descriptions as shown by the following unexplained discrepancies. Kinoshita (1934) states that in stage I Panulirus japonicus has no exopod on the third maxilliped and yet Nakazawa (1917) in figure I shows the exopod to be present in this stage. Crawford and De Smidt (1922) in figure 273 show no exopod on this appendage in stage I of Panulirus argus yet subsequent workers agree that the exopod is present. 18 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. SUMMARY The spiny lobster, Panulires interruptus (Randall), passes through eleven rather poorly defined phyllosoma larval stages. The larvae are plank- tonic during these stages which require several.months for completion. Through metamorphosis, the last phyllosoma stage becomes the puer- ulus stage which resembles the adult and becomes benthonie. All of the phyllosoma stages and the puerulus stage are described and figured. A second species of phyllosoma referred to Panulirus gracilis Streets was found overlapping the range of P. interruptus off Baja California, and in the Gulf of California. A description is given of this larva enabling its separation from P. interruptus. LITERATURE CITED ALLEN, B. M. 1916. Notes on the spiny lobster (Panulirus interruptus) of the California Coast. University of California Publications in Zoology, vol. 16, no. 12, pp. 139-152. CHASE Jr., F. A., and W. H. DuMont 1949. Spiny lobsters —- identification, world distribution, and U. S. trade, Com- mercial Fisheries Review, vol. 11, no. 5, pp. 1-12. CRAWFORD, D. R., and W. J. J. DE Smipr 1922. The spiny lobster, Panulirus argus, of Southern Florida: Its natural history and utilization. Bulletin of the U. S. Bureau of Fisheries, vol. 38, pp. 282-309. GILCHRIST, J. D. F. 1916. Larval and post larval stages of Jasus lalandii (Milne Edw.) Ortman. Journal of the Linnean Society of Zoology, vol. 33, no. 223, pp. 101-125. GURNEY, ROBERT 1936. Larvae of decapod crustacea Part III. Phyllosoma. Discovery Reports, vol. 12, pp. 400-440. Ho.itTuuts, L. B. 1954. On a collection of decapod crustacea from the republic of El Salvador (Central America). Zoologische Verhandlingen, Rijksmuseum van Natuurlijke Historie, No. 23, pp. 2-48. KINOSHITA, TORAICHIRO 1934. On the artificial propagation of Panulirus japonicus (De Haan) and some account of its larvae. Proceedings of the Fifth Pacific Science Congress, 1933, vol. 5, pp. 4191-3. LEBouR, MARIE V. 1950. Notes on some larval decapods (Crustacea) from Bermuda. Proceedings of the Zoological Society of London, vol. 120, Part II, pp. 369-379. LEWIS, JOHN B. 1951. The Phyllosoma larvae of the spiny lobster Panulirus argus. Bulletin of Marine Sciences of the Gulf and Caribbean, vol. 1, no. 2, pp. 89-103. Vou. XXIX] JOHNSON: LARVAL DEVELOPMENT OF SPINY LOBSTER 19 LINDBERG, ROBERT G. 1955. Growth, population dynamics and field behavior in the spiny lobster, Panulirus interruptus (Randall). University of California Publications in Zoology, vol. 59, no. 6, pp. 157-248. NAKAMURA, SYUYA 1940. Ecological studies of the spiny lobster, Panulirus japonicus (v. Siebold), with special reference to its conservation. Journal of the Imperial Fish- eries Institute (Tokyo), vol. 34, no. 1, pp. 101-113. NAKAZAWA, K. 1917. On the metamorphosis of Panulirus japonicus (in Japanese). Dobutsugaku Zasshi Tokyo, vol. 29, pp. 259-267. ScuHMITT, WALDO L. 1919. Early stages of the spiny lobster taken by the boat “Albacore,” California Fish and Game, vol. 5, no. 1, pp. 24-25. SHEARD, KEITH 1949. The marine crayfishes (spiny lobsters), family Palinuridae, of western Australia, with particular reference to the fisheries on the western Australian crayfish (Panulirus longipes). Commonwealth Scientific and Industrial Research Organization, Australia, Bulletin No. 247, pp. 5—45. Von Bonpbk, CEcIL, and J. M. MARCHAND 1935. The natural history and utilization of the Cape Crawfish, kreef or spiny lobster, Jasus (Palinurus) lalandii (Milne Edwards) Ortman. Depart- ment of Commerce and Industry, Fisheries and Marine Biological Sur- vey, Fishery Bulletin No. 1, pp. 5—54. WILSON, ROBERT C. 1948. A review of the southern California spiny lobster fishery. California Fish and Game, vol. 34, no. 2, pp. 71-80. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. XXIX, No. 2, pp. 21-46, 2 plates January 28, 1957 SNAILS FROM CALIFORNIA CAVES BY ALLYN G. SMITH Research Malacologist, California Academy of Sciences Several years ago a systematic exploration of California caves was under- taken by members of the Stanford Grotto, National Speleological Society. More recently, this work has been extended to certain eaves in Nevada, Ari- zona, and Mexico and the entire work is now being carried on under the auspices of the Western Speleological Institute, located in Santa Barbara, California. Scientific collections of fauna, flora, minerals, and other ma- terials are being assembled and accurate records are being kept by the Institute. Among the fauna taken by the cave explorers are several species of land snails, some of which are of more than passing interest. The initial lot of these snails was made available for study through the courtesy of Dr. Myra Keen, Department of Mineral Sciences, Stanford University. Subsequently, more snails were collected during further explorations and were sent in by Edward A. Danehy, Research Chairman of the Stanford Grotto, and by Raymond deSaussure and Arthur L. Lange of the Western Speleological Institute. Most of the snails colleeted by these speleologists represent two relatively common species living in the Mother Lode area in the foothills of central California, where many limestone caves are located. These are Monadenia mormonum (Pfeiffer) and Helminthoglypta cypreophila (““Newecomb” Bin- ney & Bland). They are not true cave snails. In fact, so far as known there are no true cave mollusks in the strict sense anywhere on the west coast of North America. Normally, snails of these and other species of the California foothills are found in rock piles or in thick brush, briers, or nettles, usually [ 21] 22 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH Ser. along water courses where the right conditions of moisture, shade, and food exist. Generally they go below the surface during the day, coming out to feed at night or on dark, rainy days. The entrances of caves often provide these habitat requirements. This could easily account for the occurrence of land snails in eaves, plus the added probability that they might be washed in through erevices or other obscure cave entrances. One species of land snail, collected in two eaves in Amador County, Cali- fornia, by members of the Stanford Grotto, proves to be a most unusual and remarkable find as it represents not only a new species but a new genus as well. In one sense, this is a rediscovery because it makes comparison possible with certain similar small California snails, thought to be fossil, which have puzzled land-snail specialists for considerably more than half a century. Another species, Pristiloma subrupicola spelaeum (Dall), was collected in eaves in Calaveras and Amador counties. One of these caves was the original locality at Cave City where the indefatigable collector, Henry Hemphill, first found this snail about seventy years ago. As the specimens taken by the Stanford speleologists constitute a rediscovery of the species, and as shells from Hemphill’s original collecting have been found at Stanford, an oppor- tunity is afforded to redescribe and figure this little-known species. The species of land snails found so far during the cave explorations in California are covered more fully in the following paragraphs. Family HELMINTHOGLYPTIDAE Monadenia (Corynadenia) hillebrandi (Newcomb). Two dead shells, taken in Church Cave, Fresno County, probably should be so identified. This represents a considerable extension of the present known range of the species in Mariposa and Tuolumne counties. Shells of this race were collected first by the writer in 1940 in a rockslide of lime- PLATE 1 Fig. 1. Monadenia mormonum (Pfeiffer) from Music Hall Cave, Calaveras Co., California, imbedded in a subaqueous calcite deposit. (C.A.S. Paleo. Type Coll., Hypotype 10293.) Fig. 2. Monadenia hillebrandi mariposa A. G. Smith, n. ssp. Dorsal view of holo- type from McLean Cave, Mariposa Co., California. Max. diam. 23.1 mm. Fig.3. Same. Ventral view. Fig. 4. Same. Side view. Fig.5. Same. Detail of sculpture on dorsal surface, ca. X20. Fig. 6. Haplotrema costatum A. G. Smith, n. sp. Dorsal view of holotype, some- what enlarged, from Lost Soldier’s Cave, Tulare Co., California. Max. diam., 5.8 mm. Fig.7. Same. Ventral view. Fig. 8. Same. Side view. VoL. XXIX] he (Cy ISUUUTEL § SNAILS FROM CALIFORNIA CAVES a ea a Fathom 23 24 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. stone near Boyden’s Cave, and later by the Chaces of San Diego, California, in the same vicinity. No living snails of this race have been reported. As the available shells collected so far are poorly preserved, their relationship to M. hillebrandi from farther north must await further collecting before being finally settled. Monadenia (Corynadenia) tuolumneana Berry. Dead shells were collected in two Tuolumne County localities by Ray- mond deSaussure in 1954: Crystal Butterfly Cave, one specimen (C.A.S. 34679), and Crystal Tuolumne Cave, several specimens (C.A.S. 34678). They agree in essential particulars with Berry’s description of this rela- tively new species of Monadenia. Monadenia (Corynadenia) hillebrandi mariposa A. G. Smith, new sub- species. , 9 (PL, figs: 2; 3; 4. and {) Houotyre: Shell bi-convex, depressed, umbilicate, acutely carinate. Whorls 514, the first 1834 turns representing the nuclear stage. Umbilicus wide, vortex-shaped, contained about five times in the major diameter, and only shghtly impinged upon by the reflection of the peristome. Peripheral earination formed by narrow shallow channels, the upper one deeper and extending over the final two or three whorls above the suture in such a way as to expose the keel. Sculpture of the nuclear whorls consisting of an array of closely-set papillations typical for the genus. Succeeding whorls decorated with rough irregular lines of growth, with scattered rounded papillae that become somewhat elongated and arranged spirally on the body whorl, giving the shell a dull granular appearance. Aperture set at an angle of 45°, the peristome reflected but not recurved, the edge sharp, with the upper portion flared slightly above the dorsal plane of the body whorl, angled at the termination of the peripheral keel, connected basally by a well-developed wash of callus, and white inside. Color of the shell reddish brown above and lighter below, especially in the umbilical area, with a single dark red-brown revolving band about one millimeter wide, bordered above and below by light colored narrower bands, the lower one coinciding with the peripheral keel or situated just below it. Maximum diameter, 23.1; minimum diameter, 18.6; height, 9.2; umbilicus diam- eter, 4.4 mm. California Academy of Sciences Paleo. Type Coll. 10283, a dead but well-preserved shell taken from the floor of MeLean Cave, Mariposa County, California (See. 20, T2S, RISE, MDM); collected in the zone of twilight to total darkness by Raymond deSaussure, November 16, 1953. (Western Speleological Institute Locality C-146.) Vou. XXIX] A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 25 PARATYPES: 15 adult shells and 4 juvenile specimens, collected with the holotype, all dead, in from fair to poor condition, some covered with a tena- cious layer of hardened reddish brown mud. Paratypes to be placed in the mollusk collections of the California Academy of Sciences (Paleo. Type Coll. 10284, 10285), the U. S. National Museum, the Academy of Natural Sciences of Philadelphia, Stanford University, the Los Angeles County Museum, the San Diego Society of Natural History, and the private collections of S. S. Berry, W. O. Gregg, M. L. Walton, and Walter Miller. OTHER LOCALITIES: One dead adult and one badly smashed juvenile speci- men taken in Barber Cave, Mariposa County, California (See. 20, T2S, RISE, MDM, USGS Yosemite Quad.) in the same general vicinity as MeLean Cave, by Raymond deSaussure, November 16, 1953. (Western Spel. Inst. C-147). These are C.A.S. 34818. REMARKS: Measurements of 10 specimens from the type lot of MW. h. mari- posa vield the following data: Max. Diam. Min, Diam. Height Umbilicus No. Whorls Largest shell -............... 25.4 mm. 20.6 mm. 10.4 mm. 5.3 mm. 55% Smallest shell _............ 20.9 ed: 8.3 4.1 5 Average of 10 .............. 23.5 19.3 9.5 4.6 536 The ratio of average maximum diameter to average umbilicus diameter 15. This new race of California mountain snail is remarkable for its un- usually acute carination, resembling that of Monadenia circumcarinata (Stearns). It appears to be quite closely related to M. tuolumneana Berry recently described from “among limestone rocks at the top of cliff above Crystal Cave, Baker Ranch, near Tuolumne City, Tuolumne County, Cali- fornia,” which is also a carinate race. Monadenia h. mariposa differs from M. twolumneana in its pinehed-in carina whereas the earina of the latter, while acute, does not have this peculiar feature. In addition, M. h. mariposa has a somewhat larger and a slightly more flaring umbilicus. Except for its acute pinched-in carina, J. h. mariposa is also related to M. hillebrandi yosemitensis from the Vernal Falls region of Yosemite Valley, which has a subearinate tendeney and a low spire. It differs from M. h. yosemitensis, however, in having a smaller average size, a somewhat smaller umbilicus (though of the same vortex shape), and in the sparser papillate sculpture. Compared to typical M. hillebrandi from the Yosemite region it ranges smaller in size, is lower spired, lighter colored, and somewhat less widely umbilicate. A point worthy of note is the habitat of these races of M. hillebrandi in areas of limestone outcrops, as indicated by their occurrence in or near limestone caves in both Tuolumne and Mariposa counties. In Kings Canyon 26 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. National Park shells of JM. hillebrandi, indistinguishable from typical central California specimens, have been collected by the author and by Mr. E. P. Chace of San Diego, California, near the entrance to Boyden’s Cave. Mr. deSaussure has also taken this southern race in Church Cave in Fresno County. These last two caves are both in limestone. All central California material previously collected or reported has been found where the predominating rock is granite. Monadenia troglodytes Hanna & Smith. Described as a fossil of Pleistocene Age by Dr. G Dallas Hanna and the writer from shells collected in Samwel Cave, Shasta County, during an earlier exploration by paleontologists of the University of California. A couple of dead shells of this species found by the Stanford speleologists outside the cave on a slope some distance away from the cave entrance indicate the possibility that this species may be found living somewhere in the vicinity. Monadenia circumcarinata (Stearns). A single dead, bleached specimen, picked up on the ground near Cave Man Cave, south of Paper Cabin Ridge in Tuolumne County, proved to be this long lost and much sought for species. Its rediscovery, since con- firmed, provided the first real clue to its exact habitat in the limestone along portions of the rim of the Canyon of the Tuolumne River (Hanna and Smith, 1954). This find proves the incorrectness of previously published localities for the remarkable species, first said to be near Turlock, in Stan- islaus County, and later near Columbia, Tuolumne County. Monadenia mormonum (Pfeiffer). Taken from nearly all of the limestone caves in the Mother Lode area of Amador, Calaveras, Eldorado, Mariposa, and Tuolumne counties by the cave explorers. Of all snail species collected, this was the most abundant. Helminthoglypta allyniana (Berry). Several specimens taken in McLean and Barber Caves, Mariposa County. Helminthoglypta cypreophila (“Newcomb” Binney & Bland). This was the next most frequently occurring snail in caves and was often collected along with Monadenia mormonum (Pfeiffer). Helminthoglypta tularensis (Hemphill). Dead shells were collected deep in Haughton’s Cave, Balch Park, Tulare Vout. XXIX] A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 27 County. Other poorly preserved specimens, long dead, taken from Cave No. 12-19, about 14 miles nne. of Springville, Tulare County, undoubtedly belong to this species. Helminthoglypta ferrissi Pilsbry. A large dead and decorticated adult specimen was found by Mr. deSaus- sure in Church Cave, Fresno County, along with a living juvenile and a fragmentary specimen. This race of H. ferrissi averages appreciably larger than the size of typical shells from Tehipite Valley, middle fork of the Kings River, the adult shell above mentioned having a major diameter of 34.2 mm. Other examples, including one fine adult found alive, were ecol- lected by the writer in 1940 near Boyden’s Cave, along with dead specimens of the southern race of Monadenia hillebrandi (Neweomb). Helminthoglypta proles (Hemphill). One well preserved but dead shell and another broken specimen were taken from Juniper Cave, Plumas County. This represents a northward extension of range for the species, which is partially confirmed by specimens collected earlier in Bear Valley, near Emigrant Gap, at the northern edge of Placer County. The best shell from Juniper Cave fits the description of the species in all respects and is not referable to the smaller and thinner subspecies H. p. mariposa Pilsbry, which lives in the Yosemite Valley region. Family CAMAENIDAE Ammonitella yatesii Cooper. Collected at two new cave locations not hitherto reported—Cave of the Catacombs (Miller’s Cave) in Calaveras County and Pioneer Cave in Eldorado County. This curious species, the only living one of its genus, has a limited distribution in the Mother Lode region of California, having been described many years ago from specimens collected in and near the eave at Cave City, Calaveras County. A large series of topotypes were collected in February, 1954, by Mr. deSaussure several hundred feet inside the cave in the zone of total darkness. All were dead. Several years ago snails of this species were collected by KE. P. Chace in limestone talus near the mouth of Boyden’s Cave in Fresno County, thus materially extending its geographic range southward. These were named A. yatesti allyni Chace based on minor differences between them and typical specimens from farther north. Dead shells of A. y. allynt were collected in Chureh Cave, Fresno County. 28 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Family HAPLOTREMATIDAE Haplotrema alameda Pilsbry. A good-sized series of dead shells was taken in Sutter Creek Cave, Ama- dor County; another comes from Pine Log Cave, and five additional speci- mens were collected in Small Cave, both located in Tuolumne County. A single dead shell of the subspecies H. alameda fieldi Pilsbry was found in Cave No. 12-1, Fresno County. ee costatum A. G. Smith, new species. U (Pl. 1, figs. 3, @, and 7.) Ho.uotyPeE: A dead adult shell, small for the genus. Upper surface nearly flat, base with a wide and deep vortex-shaped umbilicus contained about two and one-half times in the major diameter. Color white underneath a thin, slightly yellowish periostracum. Whorls four and one-quarter, the nuclear turns about one and three-quarters, the latter smooth and tumid. Half of the first post-nuclear whorl decorated with closely spaced, subobso- lete, transverse riblets similar to normal lines of growth. These give way abruptly to regularly spaced, coarse, rounded, sigmoidal riblets with chan- neled interspaces slightly wider than the riblets themselves. No spiral sculpture. Whorls moderately convex, with well impressed sutures. Outer lip descending, anteriorly protracted, reflexed at the upper edge by a small dimple, underneath which is a prominent, rounded denticle inside the aperture. Peristome thick, slightly recurved posteriorly, the outer and inner lips joined by a well defined ridge of callus. Maximum diameter, 7.5; minimum diameter, 6.2; height, 2.9; umbilicus diameter, 3.0 mm. Holotype in California Academy of Sciences (Paleo. Type Collection, 10286) collected in total darkness at depth in Cave No. 12-19, Tulare County, California, by Raymond deSaussure, August 8, 1953. PaRATYPES: Two additional specimens taken with the holotype, the first a somewhat smaller dead adult shell, the second a dead but fresh juvenile. Paratypes are in the California Academy of Sciences (Paleo. Type Coll., 10286a, 10286b (A.G.S. 9451) ). REMARKS: This remarkable little species of Haplotrema does not appear to be related at all closely to any described species. Its major characters con- sist of the prominent ribbing in relation to size, the lack of spiral sculpture in the adult phase, and the strong denticle beneath the edge of the upper outer lip. In the somewhat better-preserved juvenile shell the nuclear whorls are translucent and shining. With a magnification of 40 some Vou. XXIX] A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 29 extremely fine, closely spaced, spiral sculpture is seen on the early post- nuclear whorls. Haplotrema costatum is nearest in size and sculpture to H. caelatum (Mazyeck). It differs from the latter, however, in having a somewhat wider umbilicus, less tumid body whorl, and particularly in the configura- tion of the aperture and lip. In H. costatum this latter feature is similar to the lip structure of H. sportella (Gould) and H. voyanum (Newcomb) but without the denticle, whereas in H. caelatum the lip is simple, with a sharp edge even in the adult shell. On the basis of shell structure alone H. costatum probably belongs in the H. sportella group (section Ancomena H. B. Baker) along with H. voyanwm. Family ZONITIDAE Pristiloma gabrielinum (Berry). Three dead adults and one fine living specimen were collected in Church Cave, Fresno County, California; and another dead shell was found by Mr. deSaussure in Windy Cliff Cave, also in Fresno County. Comparison of these shells with others from the original habitat in the San Gabriel Moun- tains of Los Angeles County has indicated no appreciable differences. Thus the current find represents a considerable range extension for the species. Pristiloma juniperum A. G. Smith, new species. (PlS2 hess t., 2, and 3.) Houtotyrpe: A dead adult shell with general characters of P. gabr- elinum (Berry) from which it differs principally in the somewhat more tumid body whorl, correspondingly more capacious aperture, and the smaller tubular umbilicus. Whorls almost five and one-quarter. Like P. ga- brielinum, this species exhibits a well defined “false suture” below the true suture where the top of the whorl is appressed and fused to the preceding one, a feature that is continuous, starting at the nucleus. The posterior portion of the outer lip is sharply angled just before joining the columella, the angle forming a carina that curves around the umbilicus for about half a turn, giving it its tubular shape. In the immediate area of this umbilical earina the shell material is roughened into several irregular, closely spaced, spirally arranged striae, a feature not present in P. gabrielinum. Maximum diameter, 5.8; minimum diameter, 4.9; height, 2.9; umbilicus diameter, 0.3 mm. Holotype in the California Academy of Sciences (Paleo. Type Collec- tion, 10287), collected on the floor and under rocks in the Cavern Room of 30 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Juniper Cave, Plumas County, California (Western Speleological Institute, C-96), by Raymond deSaussure, July 20, 1953. PARATYPES: Three specimens (two badly broken) collected with the holo- type, (C.A.S. Paleo. Type Coll., 10287a, 10287b, 10287c (A.G.S. 9447) ). REMARKS: These snails appear to be worthy of a new name as they are suf- ficiently different from P. gabrielinum, with which the species has been compared, and also because the locality where it was discovered lies several hundred miles north of the known range of P. gabrielinwm. It comes from an area that is not at all well known conchologieally. Pristiloma chersinella (Dall). A single dead specimen was taken in McLean Cave, Mariposa County. Pristiloma subrupicola spelaeum (Dall). (Pl. 2, figs. 4, 5, and 6.) ?V[itrea] subrupicola ... var. spelaea Dall, 1895, Nautilus, 9:27. 2? Pristiloma (Ogaridiscus?) subrupicola (2?) spelaeum (Dall), H. B. Baker, 1931, Proc. Acad. Nat. Sci. Philadelphia, 83:90. Pristiloma subrupicola spelaeum (Dall), Pilsbry, 1946, Acad. Nat. Sci. Philadelphia, Mon. No. 8, vol. II, pt. 1, p. 418. The original specimens on which this subspecies is based were unques- tionably sent to Dall for identification. Probably they are in the U. S. National Museum at Washington, D. C. However, as Hemphill was in the habit of sending out only part of any lot he collected, it seemed certain that some of the original lot ought to be either in the California Academy PLATE 2 Fig. 1. Pristiloma juniperum A. G. Smith, n. sp. Dorsal view of holotype, some- what eniarged, from Juniper Cave, Plumas Co., California. Max. Diam., 5.8 mm. Fig. 2. Same. Ventral view. Fig. 3. Same. Side view. Fig. 4. Pristiloma subrupicola spelaeum (Dall). Dorsal view of C.A.S. Lectotype 10288 trom Cave City, Calaveras Co., California, coll. by H. Hemphill. Somewhat enlarged. Max. diam., 5.9 mm. Fig.5. Same. Ventral view. Fig.6. Same. Side view. Fig. 7. Speleodiscoides spirellum A. G. Smith, n. gen., n. sp. Dorsal view of holo- type, somewhat enlarged, from Violin Cave, Amador Co., California. Max. diam., 6.8 mm. Fig.8. Same. Ventral view. Fig. 9. Same. Side view. VoL. XXIX J A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 31 32 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. of Sciences where Hemphill’s main collection is deposited, or in the Hemp- hill “duplicates” at Stanford University. A thorough search of the Cali- fornia Academy collection proved fruitless but Dr. Myra Keen discovered a lot of five specimens at Stanford labeled “Pristiloma subrupicola spelaea Dall, Calaveras Co., probably from Cave City, coll. H. Hemphill.” Agree- ment between these shells and those collected at Cave City by the Stanford speleologists leaves little room for doubt that they are part of the original lot. The largest of the Stanford specimens has been selected for illustration. Through the courtesy of Dr. Keen it has been deposited in the Paleo. Type Collection of the California Academy of Sciences as Lectotype 10288. The shell evidently contained the living animal when found. It has four and seven-eighths whorls and measures: max. diam., 5.9; min. diam., 5.2; height, 2.4; diam. of umbilicus, 0.5 mm. It is thin and pellueid, with a whitish luster that is more pronounced on the last whorl. Axial sculpture consists of some wrinkling near the sutures and in the vicinity of the umbilicus, but this does not cross the periphery of the whorls, which are smooth and without any traces of lines of growth. Spiral sculpture consists of extremely fine wavy lines that can be seen well only at magnifications of about 40 or higher, although some heavier spiral impressions occur in the umbilical region. The sutures are not deeply impressed but where the upper end of the peri- stome becomes fused with the preceding whorl during growth, another line shows through, giving the aspect of a false suture just above the true su- tural groove. In shape, the shell agrees well with Pilsbry’s figure of Pristi- loma subrupicola (1946. p. 415, fig. 226), except that the umbilicus is wider. The other four specimens in the Stanford lot agree in most particulars with the above account. Two also were collected living, are somewhat smaller, but have more whitish luster than the larger shell has. Measure- ments of them are as follows: No. Whorls Max. Diam. Min, Diam. Height Diam. of Umb. 4% 5.0 mm. 4.5 mm. 2.5mm. 0.5 mm. 4 LN h 42) ey OLD ian The remaining two are dead shells, both somewhat damaged. The Stanford cave explorers collected this same species at the following localities, to which another is added where the writer found it later: Cave at Cave City, Calaveras County, Stanford Grotto, 9-b; Edward Dan- ehy, coll., July 1, 1950; 13 dead specimens (topotypes), C.A.S. 34819. Black Chasm, near Voleano, Amador County, Stanford Grotto, 68, lot 2; Charles Moller, coll., January 15, 1951; a single dead and possibly somewhat fossilized specimen found about 120 feet down from the eave opening. (C.A.8. 34820.) VoL. XXIX] <A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 33 Violin Cave, Amador County, Stanford Grotto, 142; Edward Danehy and Arthur Lange, colls., November 25, 1951; two living specimens (pre- served in aleohol) and two dead shells (C.A.S. 34821). The living speci- mens were found estivating about 50 feet from the entrance of the cave, in total darkness. The others were taken at the farthest point inside the cave where they appeared to have been washed, possibly through small crevices. Rippled Cave, Amador County, Stanford Grotto, 144; Gil. Lang, coll., February 3, 1952; three living specimens (preserved in aleohol) and one additional juvenile preserved dry (C.A.S. 34822). Bank of San Antomo Creek, near Sheep Ranch, Calaveras County; A. G. Smith, coll., May 18, 1952; a single dead adult specimen in a shaded rock- slide (C.A.S. 34823). Family ENDODONTIDAE Speleodiscoides A. G. Smith, new genus. DeEscripTION: Shell helical, small (7 mm. in diameter), fragile, disk-shaped, with flat or slightly concave spire, and with a wide, shallow umbilicus. Color white, the whorls being covered by a very pale horn-colored perio- stracum. Nuclear whorls about one and one-half, smooth and glossy, tumid; postnuclear whorls about five and one-half, tightly coiled, rounded, without internal teeth or other processes; aperture subcireular; lip simple, sharp, not entire. Sculpture consisting of fine growth lines and closely spaced, transverse growth ridges. Animal unknown. Genotype, the following species. REMARKS: This genus does not seem to be closely related to any other from North America. Provisionally it is referred to the family Endodontidae but this must remain a problematical assignment until living specimens can be found and their anatomy investigated. It is closest in size, shape, and coiling of the whorls to Helicodiscus Morse, 1864, but differs from that genus in the lack of spiral sculpture and of any internal laminae or teeth. It may also be compared with the South American genus Polygyratia Gray, 1847, section Systrophia, as defined by Tryon & Pilsbry in the Manual of Conchologyt. Several species of the general shape and coiling of Speleo- discoides are reported from Colombia, Peru, and Ecuador. These are some- what larger in size and although several are described as having no bar- riers or teeth inside the whorls, some have spirally striate sculpture that is lacking in Speleodiscoides. 1 Thiele (1931, p. 597) gives Systrophia generic status in the family Systrophiidae. 34 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Speleodiscoides spirellum A. G. Smith, new species. (Pie figs: 7-8) and.9>) Houotryee: A dead but well-preserved shell with general characters as de- seribed for the genus. Total whorls six and three-quarters, consisting of about one and one-half nuclear turns and five and one-half postnuclear turns. The embryonic tip is small but enlarges rapidly during the second half-turn, so that this portion of the nuclear whorls projects shehtly above the plane of the first two postnuclear whorls. Spire very slightly concave except for the projecting nuclear area. Sutures well impressed. Umbilicus very wide and shallow. Maximum diameter, 6.8 mm.; minimum diameter, 6.4 mm.; height, 1.6 mm. Holotype in the California Academy of Sciences (Paleo. Type Coll., 10289), collected in Violin Cave on South Fork of Dry Creek, west bank, Amador County, California (See. 4, T7N, R12E, MDM, USGS Pine Grove Quad.), by Edward Danehy and Arthur Lange, November 22, 1951, Stan- ford Grotto, 142. PARATYPES: 33 specimens collected with the holotype, of which about 20 are full-grown though all are dead shells in a more or less bleached condi- tion. Paratypes are in the California Academy of Sciences (Paleo. Type Coll., 10290, 10291, and 10292), Stanford University, the Los Angeles Mu- seum, the San Diego Society of Natural History, the Academy of Natural Seiences of Philadelphia, the United States National Museum, and the pri- vate collections of Dr. S. S. Berry, Dr. W. O. Gregg, Walter Miller, M. L. Walton, and Leshe Hubricht. OTHER Locauity: Black Chasm, near Voleano, Amador County, California, on the east side of the south branch of Sutter Creek, at an elevation of 2040 feet. Two lots were taken: the first containing nine bleached shells collected by Charles Moller, January 138, 1951 (C.A.S. 34824); the second, ten shells collected by Harold Treacy, May 5, 1951 (C.A.S. 34825). All were dead shells found deep down in this almost vertical shaft-like cave. ReMARKS: This remarkable small, closely-coiled, flat snail looks like a large Helicodiscus but the lack of internal teeth and the absence of even the slightest vestige of spiral sculpture immediately set it apart from this genus. Similar snail shells from the San Francisco Bay region have been mentioned in the literature by W. G. Binney (1886; 1890), Hanna (1939), and Pilsbry (1948) under the names Helicodiscus lineatus Say, H. fimbriatus var. sal- monensis W. G. Binney, and H. salmonaceus W. G. Binney. Although speci- mens from these early collectings have been at hand for many years, the lack of sufficient specific information on them and the fact that they have not been taken by any collector since has caused hesitation in recognizing VoL. XXIX] A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 35 them as part of the snail fauna of California. Discovery of these snails in fair quantity in two Amador County limestone caves leaves no doubt about their authenticity as well as their novelty. Although not one was found alive, several specimens in the type lot are fresh enough to lead one to the supposition that living snails of this species may be found by diligent search in the vicinity of the caves where the dead shells were collected. Discussion: Violin Cave is about eight miles southeast of the old gold set- tlement of FiddJetown (hence the name of the cave) and is around three or four miles air-line from Voleano and the Black Chasm location. Fiddler’s Cave is nearby. The cave entrance is 30 feet above the South Fork of Dry Creek at the base of a massive marble outcrop. Mr. Danehy has furnished the following comments: Violin Cave was almost dry when visited in November, 1951, although in the low, narrow crawlway where the estivating snails were found the gravel floor was damp. This point is approximately 50 feet from the entrance and was the beginning of the zone of total darkness. The cave is nearly hori- zontal and the farthest point reached is estimated as being 250 to 300 feet from the entrance. Here, soil and pebbles from the surface are accumulated below crevices. ... It was in the crevices and upon the soil that the dead shells were found. Almost every cave I have seen has places where soil and debris come in from the surface, even though no openings are evident. The snails taken from this cave were selected as the type lot because of the number of specimens collected, which permits a liberal distribution of paratypes. Measurements of the ten largest specimens are as follows: Dimension Range Average Maximum diameter .............. 6.0 — 7.4mm. 6.6 mm. ATT yp ee a eR os one de e 1.4 — 1.6 mm. 1.5 mm. Number of whorls ................ 64-7 634 All the shells in the lot show marked similarity in characters and seulp- ture. They differ only in shght irregularities in the coiling of the whorls. Black Chasm, where Speleodiscoides was first rediscovered, is located on a hilltop in an area of massive outerops of marble, weathered smooth into fantastic shapes in places—a characteristic of several locations in the Mother Lode country. It is described as follows in the January, 1951, re- port of the Stanford Grotto: It is an L-shaped vertical fissure in limestone, first entered by E. Sammis with a party in 1854. At that time the opening was about 18” across but has since been enlarged to a diameter of several feet in subsequent attempts to commercialize the cavern. The entrance confronts a steeply inclined mud slope composed of material washed in from above. Several vertical pitches follow before the first landing is reached—an estimated 120 feet below. From the entrance down to this landing there are found no foot- or hand-holds. 36 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Fifty or sixty feet below the landing lies the bottom of the cavern—a series of interconnecting (?) lakes, the first of which has a length of about 70 feet. Depths were estimated to exceed 20 feet. Between the landing and the lake level occurs one mud platform. The entire downward route is mud-coated, but away from the range of debris the cavern walls appear unsullied. The snail shells were reported as being found in a side passage extend- ing off from and above the first landing about 120 feet down from the mouth. How they found their way into this side passage is hard to imagine unless it has had some sort of surface connection. The Black Chasm locality was visited by the writer in May, 1952. A fairly thorough search of likely-looking piles of marble and country rock in the vicinity was made on the theory that the snails in the cave had dropped or been washed in from the surface. No snails were found, however, either of this or any other species, possibly owing to the fact that over much of the entire area were evidences of surface alteration resulting from placer or other gold-seeking operations. Time was not available to devote a simi- lar search around Violin Cave. The freshness of specimens from this loca- tion would appear to make such a search more likely to be successful if the species is still living. Shells of Speleodiscoides from the San Francisco Bay region that are available for comparison consist of two from Sausalito from the Reverend Joseph Rowell Collection, and two others from the W. J. Raymond Col- lection. Of the latter, one was labeled as collected at Sausalito by Voy in 1872, the other being found by Professor Raymond himself at “Laundry Farm” in the vicinity of what is now Leona Heights, a suburb of Oakland. All these specimens were presumed to be fossil or at least semifossil, a view that now seems questionable in the light of recent finds. Two of the four shells are adult or nearly so. They do not have a fossil appearance. Of the other two, one is young; the second is badly worn and broken. Two of the good specimens have the following measurements : CAS No. AGS No. Locality Collection Max. Diam. Height No. Whorls 34826 1937 Sausalito, Calif. Rowell 5.5mm. 1.5mm. 5% 34827 8811 ? Raymond 6.1mm. 1.4mm. 536 The two Raymond specimens were together in the same bottle, making it impossible to tell which one Voy collected and which Raymond himself found. The principal account of these snails was furnished by J. G. Cooper and published in an early Bulletin of the California Academy of Sciences (Cooper, 1885). As copies of this Bulletin are not readily accessible, it is pertinent to this discussion to quote what Cooper has to say: The fourth species (or specimens) of discoid sub-fossils have been in my Vou. XXIX] A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 37 hands since 1871, when Mr. C. D. Voy brought them to me as discovered by him “in an Indian mound north of S. F. Bay.” They are of the size and nearly the form of Helicodiscus lineatus (a species said to occur in California), but without a trace of teeth or sculpture. I considered them, however, imperfect specimens of that form until 1883, when W. O. Emerson found a few of the same much larger but more fossilized, near Haywards, on a sliding bank, where they seemed to come from under the roots of a large tree that had been undermined. These also are toothless, and some of both sent to Mr. Bland and Mr. Binney, have been decided to be a new species if not a new genus. As with the specimens of Hyalina, mentioned before, these were where they could not have been washed down by mountain streams. Without living specimens it will be inadvisable to describe these, especially as their being found in an Indian mound is some indication that they may increase instead of decrease with the settlement of the country. The chances, however, seem to be in favor of their being cave dwellers or more northern species, introduced where found by birds, but not able to increase there. This account is interesting from the information it gives about the con- ditions under which these snails were found. But it is all the more notable because of Cooper’s supposition that they might be cave-dwellers. It is difficult to see the reasoning back of this unless Cooper, who was a keen observer, had information in addition to what he published. Whether it was a shrewd guess or was based on more tangible evidence, the fact remains it has been proved true after a span of nearly seventy years! Compared with specimens from the two caves in Amador County, the San Francisco Bay shells have a flat rather than a slightly coneave spire, less prominent transverse rib-sculpture, and the whorls appear to be not quite as tightly coiled. As these differences are so slight, and as only two good specimens are available for comparison, determination as to whether they constitute a recognizable geographic subspecies is left pending the col- lection of additional material from the San Francisco Bay area. List OF CALIFORNIA CAVES EXPLORED, WITH RECORDS OF THE SNAIL SPECIES COLLECTED SHasta County* 1. SAMWEL Cave. Sec. 5, T35N, R3W, MDM, Shasta Nat. Forest Map; Stanford Grotto, 23; Edward Danehy eoll., September 20, 1950. Picked up on the ground on a prominent limestone bluff known as Eagle Rock, several hundred yards west of the cave entrance. Monadenia troglodytes Hanna & Smith. Two bleached immature specimens. Discovery of this species outside of the cave indicates that it may be found living in the vicinity, though described origi- * The counties are cited in geographic order from north to south. 38 bo CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. nally as a Pleistocene fossil. The depressed spire of the specimens, though quite immature, gives credence to the correctness of the identification. PLuMAS CoUNTY JUNIPER CAvE. Sec. 17, T23N, RIOE, MDM, USGS Downieville Quad.; Western Speleological Inst., C—96, C-103, and C-105; Raymond de- Saussure coll., July 20, 1953. Taken on the floor and under rocks in the Cavern Room in the zone of total darkness. Helminthoglypta proles (Hemphill). One adult and one broken specimen; dead. Pristiloma juniperum A. G. Smith, new species. Four specimens; dead. AMADOR COUNTY Buack CHASM. 34 mi. south of Voleano, Stanford Grotto, 68 and 104; Charles Moller and Harold Treacy ecolls., January 13 and May 5, 1951. “Found in side passage about 120 feet from the surface, in total dark- ness.”’ Speleodiscoides spirellum A. G. Smith, new genus and species. A total of 19 dead specimens, a few with deposits of small calcite erystals on one surface. Pristiloma subrupicola spelaewm (Dall). A single dead specimen. ViouIn Cave. Sec. 4, T7N, RIZE, MDM, USGS Pine Grove Quad.; Stanford Grotto, 142, Edward Danehy and Arthur Lange eolls., No- vember 25, 1951. Named for its location near Fiddletown. Speleodiscoides spirellum A. G. Smith. Thirty-four dead speci- mens. The type lot. Pristiloma subrupicola spelaeum (Dall). Two living and two dead specimens. RIppLeD Cave. Sec. 28, T7N, RI1E, MDM, USGS Big Trees Quad.; Stanford Grotto, 144 and 145, Gil Lang eoll., February 3, 1952. Helninthoglypta cypreophila (Binney & Bland). One living small adult. Pristiloma subrupicola spelaeum (Dall). Three living specimens. HarE Cave. Sec. 31, T6N, RI1E, MDM, USGS Sutter Creek Quad.; VoL. XXIX] A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 39 10. 188 Western Spel. Inst., C-34, Raymond deSaussure ecoll., April 5, 1953. Taken near second entrance in the twilight zone. Helminthoglypta cypreophila (Binney & Bland). Three dead ju- veniles. SUTTER CREEK CAvE. Sec. 29, T7N, R12E, MDM, USGS Pine Grove Quad.; Western Spel. Inst., C-61, Raymond deSaussure coll., May 25, 1953. Collected in lower right passage below a fissure, on the floor in total darkness. Haplotrema alameda Pilsbry. Twenty-seven adult and three young specimens, all dead. ELDORADO COUNTY PIONEER CAVE. Sec. 18, T12N, R9E, MDM, USGS Auburn Quad.; Stanford Grotto, 31, Edward Danehy ecoll., October 7, 1950. “Cave open to daylight due to large entrance.” Monadenia mormonum (Pfeiffer). Collected in 1875 by J. G. Cooper but not found there by the Stanford speleologists. Ammonitella yatest J. G. Cooper. One dead adult specimen. CALAVERAS CoUNTY CAVE OF THE CATACOMBS (MILLER’S CAVE). Sec. 2, T4N, RI3E, MDM, USGS San Andreas Quad.; Stanford Grotto, 139, Edward Danehy eoll., Mareh 28, 1951, and November 23, 1951. “Taken in the zones of twilight and total darkness. Some appeared washed [in] and others 5 seemed to be estivating.”’ Monadenia mormonum (Pfeiffer). Six living adults; eight adult and nine immature specimens, all dead. Helminthoglypta cypreophila (Binney & Bland). Two adults and one immature specimen, all dead. Ammonitella yatesi J. G. Cooper. Forty-three adults and 11 im- mature specimens, all dead. CAVE OF THE SKULLS. North side of the Stanislaus River, about half way between Columbia and Vallecito; Stanford Grotto, 133, Edward aa Danehy coll., Mareh 25, 1951. “Dead shells taken in the twilight zone.” Monadenia mormonum (Pfeiffer). One dead adult shell. Cave City Cave. Sec. 14, T4N, RI3E, MDM, USGS San Andreas Quad.; Stanford Grotto, 9a and 9b. Some shells taken January 7, 1950, 40 13. 14. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. in total darkness several hundred feet from the nearest of the three cav- ern entrances. Others collected March 24, 1951, about 40 feet from the entrance in the twilight zone; Edward Danehy coll. Also, Western Spel. Inst., C-173 and C—174, several hundred feet into the cave in total darkness, probably washed in; Raymond deSaussure, coll., February 7, 1954. Monadenia mormonum (Pfeiffer). Several dead specimens. Also collected by Hanna & Rixford (1923). Helminthoglypta cypreophila (Binney & Bland). One dead adult. Ammonitella yatesi (J. G. Cooper). Original lot found by L. G. Yates about 100 feet within the mouth of the eave (Cooper, 1868). Not reported by the Stanford Grotto but later a series of 60 speci- mens (topotypes) were collected by Raymond deSaussure of the Western Speleological Institute. Pristiloma subrupicola spelaeum (Dall). Thirteen dead specimens (topotypes). SHaw’s Cave. Near Mercer’s Cave; Stanford Grotto, 36, Edward Danehy coll., October 8, 1950. “Taken in total darkness. Washed in by water.” Monadenia mormonum (Pfeiffer). One dead (broken) adult and two immature specimens. SnaiL Cave. Sec. 4, T3N, RI4E, MDM, USGS Columbia Quad.; Stan- ford Grotto, 73, Ray Bennett coll., March 10, 1951. ‘Taken in twilight zone.” This eave and a number of others discovered have been given names for the first time. Snails were the only form of life found in Snail Cave. Monadenia mormonum (Pfeiffer). One living and one dead adult. CRYSTAL STANISLAUS Cave. Sec. 23, T3N, RI4E, MDM, USGS Colum- bia Quad.; Stanford Grotto, 56, Edward Danehy coll., November 25, 1950. Snails found “30 ft. in from entrance (twilight zone).” Also Western Spel. Inst., C-64, C-166, and C—169, Raymond deSaussure, eoll., January 30, 1954. Found in various locations from the mouth of the cave to the twilight zone inside. Monadenia mormonum (Pfeiffer). Two living immature specimens and 10 dead adults, some of large size, plus five immature shells, also dead. GRAPEVINE GuLCH Cave. Sec. 22, T3N, R14E, MDM, USGS Columbia Quad.; Stanford Grotto, 95, Edward Danehy and Jack Jacobus colls., Vout. XXIX] A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 4] iG: at 18. 19! April 17, 1951. “Found in twilight zone.” Also, Western Spel. Inst., C-3, C-5, and C-—7, Raymond deSaussure, coll., February 2, 1953. Monadena mormonum (Pfeiffer). A total of 21 adult shells and six immature specimens from three collectings, all dead. Also one living juvenile specimen. Helminthoglypta cypreophila (Binney & Bland). Two adults and three juvenile dead specimens. BUCKEYE CAVE. One of the Grapevine Gulch caves; Western Spel. Inst., C-6, Raymond deSaussure coll., February 2, 1953. From the twilight zone. Monadenia mormonum (Pfeiffer). Six dead specimens. Woot Hoituow Cave. Another of the Grapevine Gulch group. Ray- mond deSaussure coll., May 30, 1954, Western Spel. Inst. C—212, C216, and C—217. Taken in several locations from the entrance to the zone of total darkness. Monadenia mormonum (Pfeiffer). One immature living snail and eight other dead shells, five of which are adults. One of the last, broken and heavily inerusted, was found deposited with subaerial cave coral (PI. 1, fig. 1). KENNY’s GROTTO. Grapevine Gulch group of caves. Western Spel. Inst., C-215, Raymond deSaussure coll., May 20, 1954. From the floor in the twilight zone. Monadema mormonum (Pfeiffer). Five adult and four imma- ture shells, all dead. Helminthoglypta cypreophila (Binney & Bland). Two dead im- mature specimens. Music Hatt Cave. Between Moaning Cave and Parrott’s Ferry; Western Spel. Inst., C-2, Raymond deSaussure coll., February 1, 1953. Taken in total darkness at the foot of a 150-foot fissure. Monadenia mormonum (Pfeiffer). Fifteen adult and three imma- ture specimens, several almost completely covered or imbedded in calcite crystal masses, a number of others clean or mud covered. SPELEOGEN Cave. Near Music Hall Cave; Western Spel. Inst., C-—4, C-32, and C—50, Raymond deSaussure ecoll., February 2, April 4, and Aprils, 1953: Monadenia mormonum (Pfeiffer). Six adults and one immature, all dead, one found 160 feet down in a fissure. Helminthoglypta cypreophila (Binney & Bland). A single living juvenile taken on the floor in the twilight zone. 42 bo w bho coda | CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. TUOLUMNE CouUNTY Rating Cave. See. 34, T3N, R14E, MDM, USGS Columbia Quad.; Western Spel. Inst., C-67, C—77, C-81, and C-—82, Raymond deSaus- sure coll., May 16, 1953. Taken in various locations in the eave. Monadenia mormonum (Pfeiffer). Three living immature snails, plus eight adult and four juvenile dead specimens. Helminthoglypta cypreophila (Binney & Bland). A single dead juvenile. Pine Loe Cave. See. 36, T3N, R14E, MDM, USGS Columbia Quad.; Western Spel. Inst., C-87, Raymond deSaussure ecoll., July 18, 1953. Snail shells collected throughout the cave in the zones of twilight to total darkness. Monadenia mormonum (Pfeiffer). Eight adult and four juvenile shells, dead. Haplotrema alameda Pilsbry. One broken adult. SMALL Cave. Sec. 36, T3N, R14E, MDM, USGS Columbia Quad.; Western Spel. Inst., C-88, Raymond deSaussure ecoll., July 13, 1953. From the twilight zone in the front part of the cave. Monadenia mormonum (Pfeiffer). One adult and one immature, dead. Helminthoglypta cypreophila (Binney & Bland). One dead ju- venile. Haplotrema alameda Pilsbry. Three adults and two juveniles, dead. Goutp- ToorH Caves. See. 14, T3N, R14E, MDM, USGS Columbia Quad.; Western Spel. Inst., C-161, Raymond deSaussure eoll., Janu- ary 30, 1954. On the ground, under rocks, daylight to twilight zones. Monadenia mormonum (Pfeiffer). One broken adult and eight im- mature dead specimens. Helminthoglypta cypreophila (Binney & Bland). One dead ju- venile. CRYSTAL TUOLUMNE Cave. Sec. 21, TIN, RI6E, MDM, USGS Sonora Quad.; Western Spel. Inst., C-183, Raymond deSaussure coll., Febru- ary 6, 1954. On the ground just outside the entrance. Monadenia tuolumneana Berry. Four adult and one immature, all dead specimens. (C.A.S. 34678.) Helminthoglypta cypreophila (Binney & Bland). One dead adult. Vou. XXIX] A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 43 30. 31. 32. CrystaL ButterRFLY Cave. Sec. 21, TIN, RI6E, MDM, USGS Sonora Quad.; Western Spel. Inst., C-184, Raymond deSaussure coll., Feb- ruary 6, 1954. Just inside the entrance in the twilight zone. Monadenia tuolumneana Berry. One dead adult. (C.A.S. 34679.) CRYSTAL PALACE Cave. Sec. 26, T3N, R14E, MDM, USGS Columbia Quad.; Western Spel. Inst., C-60 and C—64, Raymond deSaussure and Bill Wise ecolls., April 17 and May 24, 1953. In twilight zone. Monadenia mormonum (Pfeiffer). One living immature snail and four dead (two adult) examples. Cave Man Cave. Sec. 28, TIN, RI6E, MDM, USGS Tuolumne Quad.; Western Spel. Inst., C-380, Raymond deSaussure coll., April 2, 1953. Monadenia circumcarinata (Stearns). <A single dead, bleached shell found on the ground outside the cave. This find constitutes a rediscovery of the species. (AGS No. 9419.) PINNACLE Point Cave. See. 7, T3N, RI5E, MDM, USGS Stanislaus Quad.; Western Spel. Inst., C-160, Raymond deSaussure coll., Jan- uary 29, 1954. Found on the floor of the cave in the zones of twilight to total darkness. Monadenia mormonum (Pfeiffer). Two dead and broken juveniles. Helminthoglypta cypreophila (Binney & Bland). Two adults and two juveniles, dead. Marreosa County CAvE oF OrpHEUS. Sec. 4, T3S, RISE, MDM, USGS Yosemite Quad.; Western Spel. Inst., C-36, C-39, and C-41, Raymond deSaussure coll., March 8, 1953. Taken at several locations, including a lake at the bottom of the cave. Helminthoglypta cypreophila (Binney & Bland). Six adults and three juvenile shells, all dead. Minurrep Cave. USGS Sonora Quad., near Bower Cave; Western Spel. Inst., C-22, Raymond deSaussure coll., May 25, 1953. On the floor in the twilight zone. Monadenia mormonum (Pfeiffer). A single living juvenile speci- men. McLean Cave. See. 20, T2S, RISE, MDM, USGS Yosemite Quad.; Western Spel. Inst., C-146, Raymond deSaussure coll., November 16, 1953. On the floor in the twilight and total darkness zones. 44 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. Monadenia hillebrandi mariposa A. G. Smith, n. ssp. Sixteen adult shells and four juveniles, all dead. Type lot. Helminthoglypta allyniana (Berry). Four adults and one juve- nile, dead. Pristiloma chersinella (Dall). One dead adult. 33. BARBER Cave. Close to McLean Cave. Western Spel. Inst., C-147 and C-148, Raymond deSaussure coll., November 16, 1953. In the zone of total darkness. Monadenia hillebrandi mariposa A. G. Smith. One adult and one (broken) juvenile, dead. Helminthoglypta allyniana (Berry). One living adult and four dead shells (three immature). FRESNO CoUNTY 34. CHurcH Cave. Near Boyden’s Cave; Western Spel. Inst., C—132, C-133, and C-134, Raymond de Saussure ecoll., September 19, 1953. Various locations in the zones of twilight and total darkness. Monadenia hillebrandi (Neweomb). One adult and one juvenile (broken), dead. Helminthoglypta ferrissi Pilsbry. One very large adult, dead; one fragment; and one young living specimen. Ammonitella yatesi allyni Chace. Five adults, dead. Pristiloma gabrielinum (Berry). One living adult; three dead adult shells (one badly broken). 35. Cave No. 12-1. About 9 miles sse of Badger; Stanford Grotto, 9, Ar- thur Lange ecoll., July 2, 1952. Found on the floor. Haplotrema alameda fieldi Pilsbry. One dead adult shell. a : 36. WINpy Cuirr Cave. Across the river from Boyden’s Céve, Kings Canyon Nat. Park, near Cedar Grove. Taken from the floor in the back section of the eave in total darkness. Pristiloma gabrielinum (Berry). A single dead adult specimen. TULARE COUNTY 37. HAUuGHTON’s Cave. Balch State Park. Western Spel. Inst., A-18, Ray- mond deSaussure ecoll., July 8, 1952. Taken in ‘total darkness about 300 feet below ground [level].” VoL. XXIX] A. G. SMITH: SNAILS FROM CALIFORNIA CAVES 45 Helminthoglypta tularensis (Hemphill). Two dead bleached adults, one broken. 38. Cave No. 12-19. About 14 miles nne of Springville; Stanford Grotto, 115 and 116, Raymond deSaussure coll., August 8, 1953. Helminthoglypta tularensis (Hemphill). Two young shells and a fragment referable to this species. Haplotrema costatum A. G. Smith, n.sp. Two adults and one im- mature, dead. Type lot. REFERENCES BAKER, H. BURRINGTON 1931. Nearctic vitreine land snails. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 83, pp. 89-90. Berry, S. STILLMAN 1955. A new Sierran pulmonate of the genus Monadenia. Bulletin of the South- ern California Academy of Sciences, vol. 54, part 1, pp. 14-16, pl. 6, figs. 1-3. BINNEY, W. G. 1878. The terrestrial air-breathing mollusks of the United States and the adja- cent territories of North America, Vol. V. Bulletin, Museum of Com- parative Zoology at Harvard College, vol. 4, pp. 185-186. 1885. A manual of American land shells. United States National Museum, Bul- letin 28, p. 75. 1886. A second supplement to the fifth volume of the terrestrial air-breathing mollusks of the United States and adjacent territories. Bulletin, Mu- seum of Comparative Zoology, Harvard University, vol. 13, no. 2, p. 35. 1890. A third supplement .. ., ibid., vol. 19, no. 4, p. 220. BOONE, ANDREW R. 1954. They seek our past underground. Westways (Automobile Club of South- ern California), September 1954, pp. 6—7, 5 text figs. CHACE, E. P. 1951. A new subspecies of Ammonitella. Nautilus, vol. 64, no. 4, p. 122, pl. 9, figs. 5, 6. Cooper, J. G. 1868. On a new California terrestrial mollusc. American Journal of Conchol- ogy, vol. 4, part 4, pp. 209-210, pl. 18, figs. 1-3. 1875. On shells of the Pacific slope of North America, No. III. Proceedings of the California Academy of Sciences, vol. 6, pp. 18-19. 46 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 1885. On fossil and sub-fossil land shells of the United States. California Acad- emy of Sciences, Bulletin 4, pp. 248-250. 1887. West Coast Pulmonata; fossil and living. Proceedings of the California Academy of Sciences, ser. 2, vol. 1, no. 1, pp. 12-13. DALL, W. H. 1895. Description of a new Vitrea from Puget Sound. Nautilus, vol. 9, no. 3, pp. 27-28. HANNA, G DALLAS, and RIxForRD, EMMET 1923. Notes on some land snails of the Sierra Nevada, with description of a new species. Proceedings of the California Academy of Sciences, ser. 4, vol. 12, no. 4, pp. 43-47, pl. 4, figs. 4-7. HANnnNA, G DALLAS, and SMITH, A. G. 1933. Two new species of Monadenia from northern California. Nautilus, vol. 46, no. 3, pp. 79, 84-86. 1954. Rediscovery of two Californian land snails. Nautilus, vol. 67, no. 3, pp. 69-76, pl. 8, figs. 5-7, text fig. 1. HaAnnA, G DALLAS 1939. Exotic Mollusca in California. Bulletin of the Department of Agriculture, California, vol. 28, no. 5, pp. 301-302, pl. C, figs. 11-18. HEMPHILL, HENRY 1892. Note on Helix Yatesii Cooper. Zoe, vol. 3, no. 1, pp. 45—47. PILSBRY, HENRY A. 1939. Land Mollusca of North America. Academy of Natural Sciences of Phila- deiphia, mon. 3, vol. 1, part 1, pp. 54-57, 838-85, 562-563, figs. 22(6-8), 23, 40(1-6), 372. 1946. Ibid. Vol. 2, part 1, pp. 415, 418. 1948. Ibid. Vol. 2, part 2, pp. 633-634. THIELE, JOHANNES 1931. Handbuch der Systematischen Weichtierkunde. Erster band, zweiter teil, p. 597. Gustav Fischer, Jena. Tryon, G. W., and Piuspry, H. A. 1887. Manual of Conchology, 2nd ser., vol. 3, pp. 124-125, pls. 25, 26. Academy of Natural Sciences of Philadelphia. 1894. Ibid. Vol. 9, pp. 81-84, pl. 20. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. XXIX, No. 3, pp. 47-86, figs. 1-12 November 5, 1957 SOME AMPHIBIANS AND REPTILES FROM ARABIA By GEORG HAAS The Hebrew University, Jerusalem INTRODUCTION Much of Arabia has long been difficult of access to European and American zoological collectors, and the fauna of the vast region between the Red Sea and the Persian Gulf is still quite inadequately known. Explorations for oil have brought American geologists to the oil field regions of Saudi Arabia, and thanks to the avocational interest of some individuals, collections of amphibians and reptiles have come to the United States National Museum and to the California Academy of Sciences. A considerable collection of reptiles, with a few amphibians, was pre- sented to the California Academy of Sciences by Mr. John Gasperetti in 1947. Through the kindness of the late Mr. Joseph R. Slevin, curator of herpetology at the Academy, this material was sent on loan for study and report to Dr. Karl P. Schmidt at the Chicago Natural History Mu- seum, who has long maintained an interest in the fauna of southwest Asia. I was studying in the laboratory of the Division of Reptiles at the Museum when the Gasperetti collection arrived in Chicago, and Dr. Schmidt and I made a preliminary examination of it. When other duties promised to delay his report on this material, Dr. Schmidt kindly turned the project over to me for a formal report. This was especially gratifying as some collections from southern Arabia had meanwhile been presented to the Hebrew University in Jerusalem by the members of the British Anti- Petal 48 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Locust Expedition of 1944, under the direction of Dr. J. E. Hardy, then head of the Department of Entomology of the British Mandatory Goy- ernment. This material, together with specimens from the Trucial Coast collected by Mr. Desmond Vesey-Fitzgerald, and specimens from central Saiidi Arabia collected by Mr. A. R. Waterston, has been included in the following report. I have listed the localities below, and through the courtesy of Dr. Schmidt, a map, prepared by Miss Margaret G. Bradbury, has been provided. I wish to thank Dr. Schmidt for his aid and advice, and for his revi- sion of this manuscript. I also wish to thank Mr. Hymen Marx, assistant in the Division of Reptiles at the Chicago Natural History Museum, who has aided Dr. Schmidt by checking the revision. I am especially indebted to Mr. J. C. Battersby, in charge of the Division of Reptiles at the British Museum (Natural History), where I have been privileged to study on several oceasions. Finally, I wish to thank Mr. Alan E. Leviton of the California Academy of Sciences for carefully going over the whole manu- seript, checking the specimen numbers and the literature, and otherwise aid- ing in the final preparation for publication. Specimens in the collection of the California Academy of Sciences are designated by the abbreviations CAS; those in the Hebrew University of Jerusalem by HUJ. Representative specimens from the Academy’s col- lection have been deposited at the Chicago Natural History Museum; these specimens are designated CNHM. LOCALITIES (All localities are in Satidi Arabia. See map, figure 1.) Abqaiq Jebel Hajar Abu Dhabi Jebel Qara Abu Shaiba (Oil field region) Moreiwa Post (northwestern Al Khobar Arabia ) Buraimi Mudawara Dammam Qana, north Nejd Dhahran Qatif Doha Dhalum Ras at Tanura Dhufar Sakaka El Alat (Oil field region ) Sharja Dar-el-Hadj Shimal (Oil field region) Hail Shinas Bir Hirmas Tebuk Jebel Aja Tomayiah (not identified) Jebel Dam Trucial Coast at 23° 10’ N., 53° Hy VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 49 Among the localities at which specimens were collected, Abu Shaiba, El Alat, and Shimal are not to be found on any map available to me; they appear to be in the vicinity of the Dhahran-Abqaigq oil fields. Dar- el-Hadj and Bir Hirmas (not named on the accompanying map) are lo- eated, respectively, two stations south and two stations north of Tebuk, on the Hedjaz Railway. The locality “Tomayiah” is not identifiable; based upon the date of collection of the specimen, CAS 84626, Scincus philbyt, collected February 10, 1945, it does not seem to belong to the Gasperetti collection. Baghdad O Dy —! aR Ay @ \ ne Sakaka ~. rd Jebel Jauf Qana % Hail TAs Jebel Aja Pe Jebel Hajar”"* SAUDI ARA Ad Dam S Mukalla Figure 1. Map of Arabia, showing principal localities mentioned in the text. 50 CALIFORNIA ACADEMY OF SCIENCES | Proc. 4TH SER. Class AMPHIBIA Order SALIENTIA Family BUFONIDAE Genus Bufo Laurenti Bufo pentoni Anderson. Bufo pentoni ANDERSON, 1893, Ann. Mag. Nat. Hist., (6) 12:440 (Suakin). MATERIAL EXAMINED (2): HUJ 524, from west of Jebel Hajar (alt.: 500 feet), and HUJ 527, from Buraimi, Trucial Oman, collected in 1944, by Vesey-Fitzgerald. Bufo dhufarensis Parker. Bufo dhufarensis PARKER, 1931, Ann. Mag. Nat. Hist., (10) 8:518 (Milwah, Alaud, southern Arabia). MATERIAL EXAMINED (3): HUJ 528, from central Arabia; HUJ 525, from Dhufar, collected in 1944, by Waterston; HUJ 526, from Buraimi, collected in 1944, by Vesey-Fitzgerald. Family RANIDAE Genus Rana Linnaeus Rana ridibunda ridibunda Pallas. Rana ridibunda Pauuas, 1771, Reise Rus. Reich., 1:458 (Gurev, north coast of Cas- pian Sea). Rana ridibunda ridibunda MERTENS, 1925, Abh. Senck. Ges., 39:55. MATERIAL EXAMINED (9): CAS 84273, 84407, 84409-84413, and CNHM 74012, from Dhahran, collected in 1945, by Gasperetti; CAS 84415, from Qatif, collected in 1945, by Gasperetti. Class REPTILIA Order CHELONIA Family EMypDIDAE Genus Clemmys Ritgen Clemmys caspica caspica (Gmelin). Testudo caspica GMELIN, 1774, Reise durch Russland, 3:59: pls. 10-11 (Hircania [northern Persia] ). Clemmys caspica caspica ScuMmipT, 1939, Field Mus. Nat. Hist., Zool. Ser., 24:89. Vout. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 51 MATERIAL EXAMINED (2): CAS 84420, from Shimal, collected in 1946, by Gasperetti; CNHM 74009, from Qatif, collected in 1945, by Gasperetti. Order SAURIA Family GEKKONIDAE Trigonodactylus Haas, new genus TYPE SPECIES: T'rigonodactylus arabicus Haas. DEFINITION: Habitus very slender. Head and body strongly depressed. Digits one to five, strongly dilated and flattened dorso-ventrally at their base by fan-like extensions of their skin, which overlap between the digits, which are, however, free; laterally digits are fringed by a series of mar- ginal, projecting, triangular, flat scales. Claws thin and long, strongly compressed laterally. Fingers one to four gradually increasing in length; fifth finger ends at level of second finger. Minute, imbricate, triangular scales cover undersides of fingers; only a few transversely widened scales at their tips. Upper surface of hands covered with keeled, imbricate, rather elongate scales. Toes more elongate and slender; with strongly compressed, long claws; margins of toes fringed; length of toes increasing from first to fourth; fifth toe shorter than first; sealing as on hand. A prominent plantar fold extends from base of third toe to base of first toe; deep folds run proximally from outer sides of first, second, and third toes to con- verge to proximal end of fold between hallux and third toe. Dorsal seales flat, juxtaposed, slightly keeled, elliptical. Pupil a vertical slit. No anal or femoral pores. Moderate hemipenial swellings. Tail cireular in eross section, tapering uniformly. Trigonodactylus arabicus Haas, new species. {Figure 2.) HouotypE: CAS 84321, male, from Abqgaiq, Satidi Arabia, collected September 29, 1945, by John Gasperetti. PaRaTYPES (3): CAS 84318, 84323, females, from the type locality, collected in 1945, by Gasperetti; CNHM 73992, male, from Dhahran, col- lected in 1945, by Gasperetti. (All specimens rather poorly preserved.) DeEscripTION: Habitus slender and strongly depressed, especially the head; limbs very slender; head rather elongate; snout obtusely pointed, equaling length of orbit; length of orbit greater than its distance from ear opening, which has the form of an elliptical, slanting slit. Eye very large; pupil vertical. Upper eyelid bent vertically downward anteriorly, forming a fold at anterior margin of orbit; fold widest below. No lower 52 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. eyelid. Interorbital constriction, at narrowest spot, as wide as three head tubercles. Nostrils open at summit of a blunt prominence, followed posteri- orly by a depression. An internarial depression extends into rostral. Col- lar constriction very distinct. Adpressed forelimb reaches anterior border of eye; bent backward, it reaches two-thirds the distance between fore- and hindlimbs. Adpressed hindlimb extends forward beyond axilla. Ros- tral cleft, as seen from above, undivided at anterior vertical slope; posteri- orly, rostral concave; dorsal portion of shield twice as wide as long; posterior border transversely truncate, with median re-entrant niche for an unpaired granular scale. Rostral enters nostril; its deeply coneave ventro-lateral border is in contact with inner nasals, which are separated from each other by 2 flat polygonal seales. Nostril defined by rostral, 3 nasals (4 nasals on left side, but the third is abnormally excluded from nostril), and first supralabial. Supralabials 13/14, gradually diminishing in height. Head covered above by more or less hexagonal, flat, not imbri- cating scales, which are smallest at rectangular bend of eyelid; scales on upper side of snout are larger than those adjacent to anterior upper labials; all slightly rugose; temporal scales much smaller, almost granular, between eye and ear opening. Infralabials 9/11. Mental with convex posterior border, somewhat wider than deep, with a slight anterior median prominence. No chin shields. Gular scales minute; lateral ones, near infra- labials, elongate, keeled, and subimbricate; more median ones cycloid, imbricate, and keeled, and gradually diminish in size toward neck. Dorsal seales elongate at middle of back, but are more or less cycloid at sides; their diameter equals the long dimension of the mid-dorsals; all are juxtaposed, not imbricate. Limbs covered dorsally by keeled, cycloid, im- bricate scales, which are much larger than the dorsal body seales; the scales on the dorsal surface of the hindlimbs are much enlarged. Ventral scales are bluntly triangular, keeled, and slightly larger than gulars; ventro- medially, scales are imbricate; laterally, they are juxtaposed. On under- side of extremities, the eycloid, imbricate, keeled scales are arranged in oblique, regular series; they diminish in size toward the proximal end of limb. The largest ventrals, in the pelvic area, are much smaller than the ventral tibial scales. At first glance, the fingers seem to be webbed, but closer examination shows that the proximally enlarged skin folds of the fingers overlap (see diagnosis of the genus). The tail is covered dor- sally by elongate, transversely truncate, keeled scales (up to 3 keels on a scale), arranged in more or less transverse, rather irregular whorls. Ventrally, these transverse series are more distinct. A ventro-median raphé is conspicuous. The subeaudals are subimbricate, elongate, keeled, with rounded distal ends. Two distinct hemipenial swellings present; the two obliquely arranged openings for the hemipenes are placed some- what behind the vent. A cluster of enlarged scales is present on both sides VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 53 of the root of the tail, the innermost 2 to 3 of these seales are conical and much enlarged. A erescent-shaped transverse black marking crosses the snout (fig. 2), linking the anterior corners of the eyes; the concavity of the crescent is directed backward. A blackish streak extends from the posterior border of the eyes to a point above the aperture of the eye, curving slightly up- ward. A faint stripe on each flank follows the direction of the temporal streak. Upper side of tail with about 10 darker transverse bands, bordered in black and narrower fhan the yellowish intervals. Venter whitish. A few seattered dark markings are present on nape and on upper side of limbs. There are no color differences between the sexes. MEASUREMENT OF HOLOTYPE (in mm.): Snout to vent 31.5; tail 34; length of head 9.5; width of head 5.5; height of head 3.0; forelimb 12.5; hindlimb 17.5. (2 Z ay fy a SOYA) af SAN uae \. é { A Y VAT Lense @ Figure 2. Trigonodactylus arabicus Haas, new species. Left: Dorsal view of head of CAS 84321, holotype, showing color markings (stippled) and pholidosis of rostral region. Right: Plantar surface of right hand of CNHM 73986. 54 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Genus Stenodactylus Fitzinger Stenodactylus slevini Haas, new species. (Figure 3.) HouotyPe: CAS 84592, male, from Dhahran, Satidi Arabia, collected in 1946, by John Gasperetti. Paratypes (2): CAS 84540, allotype, from the type locality, collected in 1946, by Gasperetti; CNHM 74026, female, from Al Khobar, collected in 1946, by Gasperetti. Diacnosis: Habitus rather stout. Arrangement of subdigital scales similar to that in S. petri, from which S. slevini differs in having the nostril strongly swollen and bordered by 3 nasals and the rostral; the first upper labial reaches the nostril only at a point where the third nasal and the rostral touch each other at the lower periphery of the nostril. The tail does not show the sudden contraction immediately behind the basal swelling, as in SN. petri, but tapers gradually as in S. elegans. Inner nasals separated by one granule, or more or less in contact. DESCRIPTION: Snout rather short, longer than distance from eye to ear opening. Ear opening small (half the height of the eye), vertically elliptic. Rostral a little broader than high, with a median dorsal cleft in a mid- dorsal depression. Granules on snout polyhedral, closely juxtaposed, very rugose, larger dorsally, especially dorso-laterally, in front of the orbit, and much larger than occipital granules. Inner nasals in contact. Upper Jabials 14/14; lower labials 13/12. Mental with parallel lateral borders; posterior border convex, somewhat bulging beyond suture with first infra- labials; length to width about 3 to 2. Nostrils swollen; opening directed forward and upward. The nasal swelling is emphasized by a median dorsal depression entering the rostral and a postnarial depression between the third nasal and the labials. Pupil a vertical slit. Upper eyelid bent vertically downward at anterior border of eye; lower lid very narrow, almost invisible. Back covered by rather elongate, juxtaposed granules; similar granules, but broader and slightly larger, on flanks; they form juxtaposed, blunt, triangular tubercles whose apices point backward. All dorsal seales rugose, bluntly keeled on posterior part of back and on upper side of femur. Gular granules slightly smaller than ventrals; latter granules are triangular, subimbricate, and faintly keeled mid-ventrally, rugose and subtubereular in pectoral region. Pelvic ventrals flat and rugose. No preanal or femoral pores. Cycloid, rugose granules present on underside of limbs. A group of much enlarged conical seales oceur at each side of swollen base of tail; three of these form a sloping series, VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 55 extremely enlarged in male. Ventral side of hemipenial swellings cov- ered with cycloid, imbricate, keeled scales. The adpressed forelimb ex- tends beyond tip of snout; extended backward, forelimb falls short of groin. Adpressed hindlimb reaches axilla. Fingers are covered dorsally by 7 series of pointed, imbricate, keeled scales. The marginal fringes of fingers and toes of moderate size. Enlarged medial series of scales beneath digits twice as wide as the 2 intermediate series. Claws curved, laterally compressed, sharp. Fingers short and stumpy, less flattened dorso-ventrally than in Ceramodactylus. First finger shorter than fifth, third and fourth almost equal, second a little shorter. First and fifth toes end at about same level, first to fourth gradually increasing in length. Behind the basal swelling of the tail the scales form obscure rings; these seales are imbri- cate, keeled, and rugose dorsally and ventrally. Back, inclusive of head and upper side of limbs, with a black and dark brown reticulation on a pale brown ground color; reticulations denser on body, widely meshed on head. Nine dark brown eross-bars on tail, bor- dered with black seams, wider than the lighter intervals. Venter pure white. Dark spots on rostral, mental, and upper and lower labials. Figure 8. Stenodactylus slevini Haas, new species. Upper and lower sides of hand of CNHM 74026. 56 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. MEASUREMENTS OF HOLOTYPE AND ALLOTYPE (in mm.) : Holotype Allotype (CAS 84592) (CAS 84540) Total lene tie treaee we tytn pve 5 Ve sec. 0 aces 5.8 105.5 104.5 Wench oi head are rire ese cue casei une Oe s 17.5 16.5 Ten obi omstaleeses ea Arce le ks ye aie sae 40.0 36.0 REMARKS: Paratype CNHM 74026, a female, differs from the two other specimens in having a remarkably pointed and more flattened head; it otherwise resembles the allotype. The forelimb does not extend beyond the tip of the snout; it ends between the snout and the anterior border of the eye. This species differs from Stenodactylus elegans in having an acces- sory series of tricarinate scales between the lateral fringes and the trans- versely enlarged median series of scales beneath the fingers and toes. Corre- sponding in this respect with S. petri, S. slevini differs from S. petri in the eradually tapering tail, in habitus, and in lepidosis. Stenodactylus arabicus Haas, new species. (Figure 4.) Ho.totyPeE: HUJ 2678, female, from Lat. 24°10’ N., Long. 53° E., on the Trucial Coast, Satidi Arabia, collected December 31 (year un- known), by Desmond Vesey-Fitzgerald. DESCRIPTION: Snout rather pointed; nostrils open on top of almost tubular swellings; internasal depression slight; postnasal depression deep. Head much narrower than in Ceramodactylus major. Rostral deeply cleft. First labial widely excluded from nostril; latter shield defined by the rostral and 3 nasals. Inner nasals meet near the suture of the rostral; rostral forms antero-ventral circumference of nostril, and is as high as wide. Upper labials 15/15; lower labials 13/13. Mental slightly longer than wide, with lateral borders somewhat converging posteriorly; the slightly convex border does not bulge beyond first infralabials. No chin shields. Head scales (polyhedral tubereules) rugose, flattened, largest at base of snout in front of eyes. Dorsal scales subimbricate, elongate, cycloid, perfectly smooth, as are the scales on upper side of limbs. Seales on flanks more cycloid, tubercular, juxtaposed. Ventral scales clearly juxtaposed, slightly rugose, not keeled, much larger than gular scales. Caudal scales arranged dorsally in transverse whorls, with dorso-median irregularities corresponding to an irregular ventro-median seam; dorsally, scales keeled only on distal half of tail; subeaudal scales smooth, with parallel trans- verse borders, keeled at distal half of tail. The tail tapers very gradually Vou. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 57 from the insignificant basal swelling to an extremely slender, whip-like end. Limbs very slender; anterior limb, extended forward, reaches beyond tip of snout; adpressed backward, it reaches the groin. At both sides of root of tail a cluster of 3 to 4 conical, enlarged scales are present. Lateral digital fringes not prominent; the individual scales bicuspid. The medial, transversely enlarged series of subdigital scales bear 3 longitudinal keels; between these and the lateral fringes, a single row of triangular scales intervenes, as in S. petr. Color of upper side very pale; some irregular, disconnected, dark, wavy linear markings obscurely cross the dorsum, mostly interrupted and ir- regularly arranged. Some isolated dark spots present on head, but not on snout. Tail with about 10 brown transverse bars with dark borders against the pale ground color. No markings on limbs. Ventral side white. MEASUREMENTS OF HOLOTYPE (in mm.): Total length 104; length of tail 37; length of head 16; width of head 11; forelimb 21; hindlimb 28. Remarks: This species differs from the similar Stenodactylus petri in the arrangement of the scales around the nostril, and in coloration and body proportions. However, the two species have a similar arrangement of the subdigital scales. Figure 4. Stenodactylus arabicus Haas, new species. Upper side of hand of HUJ 2678. 58 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Genus Ceramodactylus Blanford Ceramodactylus major Parker. (Figure 5.) Ceramodactylus major PARKER, 1930, Ann. Mag. Nat. Hist., (10), 6:594 (Rub’al Khali, Hadramaut, Satdi Arabia). MATERIAL EXAMINED (27) : CAS 84266, 84509, 84514, 84522-84523, 84525, 84528, 84558, 84590, 84595, and CNHM 74013, from Dhahran, collected in 1946, by Gasperetti; CAS 84361-84368, 84370-84372, 84374, 84482, and CNHM 74014, from Abqaiq, collected in 1945-1946, by Gasperetti; HUS 2677, from Sharja, Trucial Coast, collected in 1944, by Vesey-Fitzgerald; HUJ 2676, from 23 miles north of Hail, collected in 1944, by Waterston. Remarks: The series from Dhahran and Abqaiq is rather homogene- ous, but there is variation in the number of transverse subdigital scales, which ranges from 5 to 8. The largest specimen, CAS 84522, a female, has a total length of 137 mm., and a tail length of 66 mm. This is smaller than the type which measured 152 mm. in total length and 72 mm. in tail length. The inner nasals may or may not be in contact. All of these specimens belong certainly to this species and cannot be = Figure 5. Ceramodactylus major Parker. Underside of fingers of CNHM 74014. Vout. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 59 referred to C. doriae, as they have well developed digital fringes and rugose scales. The two specimens in the Hebrew University collection are from lo- ealities more than 800 miles apart. The Sharja specimen corresponds fairly well with the series from the Bahrein area, but the specimen from Hail has a very pale coloration compared with the series from the Bah- rein area. This specimen from Hail, HUJ 2676, has a buff ground color; whitish ocelli are present on the flanks and head; the darker markings on the upper side are very faint. The first supralabial is excluded from the nostril; the rostral and third nasal are narrowly in contact. The nostrils are swollen as in typical specimens; five series of ventral scales are pres- ent under the third finger. It is best to consider this aberrant specimen from Hail as belonging to C. major. Pseudoceramodactylus Haas, new genus TYPE SPECIES: Pseudoceramodactylus khobarensis Haas. DEFINITION: Somewhat intermediate between T'eratoscincus Strauch and Ceramodactylus Blanford. Digits not dilated; furnished with a long, thin, curved claw; not depressed; ventral side bulging, wider than dorsal surface; covered inferiorly between the lateral fringes (formed by pointed seales) by regularly disposed diagonal series of uniform, sharply pointed, minute, juxtaposed seales (about 14 in one diagonal series). Body cov- ered dorsally with equal, eycloid to hexagonal, smooth, juxtaposed, flat scales. Head seales smooth, as are the triangular, elongate, deeply imbri- cate ventrals. No preanal or femoral pores in males. Pupil vertical. Nos- trils remarkably swollen, on a turret-like prominence. A pair of chin shields in contact with the pointed, very elongate mental; mental extends beyond the first infralabials. Pseudoceramodactylus khobarensis Haas, new species. (Figures 6 and 7.) Houotyrpe: CAS 84458, male, from Al Khobar, Satidi Arabia, col- lected June 5, 1946, by John Gasperetti. PaRATYPES (6): CAS 84459, allotype, collected June 5, 1946; CAS 84462, female, collected June 6, 1946; CAS 84461, female, collected June 6, 1946; CAS 84460, female, collected June 5, 1946; CNHM 73994, male, collected May 30, 1946; all specimens from the type locality, col- lected by Gasperetti. CAS 84431, male, from Dammam, collected May 30, 1946, by Gasperetti. 60 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Description: Habitus of a slender Stenodactylus, with remarkably swollen nasal area, strong neck constriction, and slender legs with rather short fingers. Head rather elongate; snout longer than distance between eye and ear opening; latter rounded, its vertical diameter contained three times in horizontal diameter of orbit. Rostral as broad as high, cleft dorsally for two-thirds of its length. Granules on snout small, largest dorso-medially behind rostral and in front of orbits, gradually diminishing in size toward frontal area. Nostrils directed somewhat upward, at top of a turret-like prominence, defined by rostral and 3 nasals. First labial widely excluded from nostril border. Inner nasals separated by 2 granules. Upper labials 9/10; lower labials 9/8. Mental one and two-thirds times as long as broad, with lateral borders strongly curved toward the first labials, and with a triangular continuation beyond them which is flanked on either side by elongate postmentals; postmentals meet medially behind mental. Gular tubercules largest near mental, postmentals, and infra- labials; they diminish in size posteriorly. The two series of largest gular tubercles, those bordering the infralabials and those bordering the post- mentals, meet behind the mental and postmentals, and form a belt as wide as the length of the mental. The back and upper side of the limbs covered by rather large, Juxta- C Figure 6. Pseudoceramodactylus khobarensis Haas, new species. a) Dorsal view of snout of CNHM 73994. b) Lateral view of snout of CAS 84462. c) Mental region of CAS 84458, holotype. VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 61 posed, flat, and smooth, rounded tubercles; these are polyhedral in shape on top of the head. Ventral scales form chevron-like, transverse, sub- imbricate series, with posteriorly open angle; the scales are cycloid and smooth, largest in the pelvic area. The mid-ventrals are bigger than the mid-dorsals; the lateral scales are smaller than the mid-dorsals. Be- tween the round gular tubercles and the flattened pectorals there is a region covered with elongate, juxtaposed tubercles. The scales in the pectoral region are much smaller than those in the pelvic area. On the tail the scales of the anterior third are irregularly arranged; be- yond this region they form annuli; this arrangement is somewhat irreg- ular mid-dorsally, and to a slight degree mid-ventrally. Very strong hemipenial swellings present in the male. In both sexes a cluster of en- larged scales is present at either side of the root of the tail (better devel- oped in the male); each cluster consists of about 5 triangular scales whose pointed ends project upward; in the male they are imbricate and are at least four times larger than the adjacent normal seales; in the female the scales of the cluster are only three times as large as the surrounding ones. Adpressed forelimb reaches the groin; adpressed hindlimb reaches the anterior border of the pectoral girdle; the outstretched forelimb reaches Figure 7. Pseudoceramodactylus khobarensis Haas, new species. Upper and lower sides of hand of CAS 84459. 62 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. the tip of the snout. Upper side of fingers covered by slanting series of approximately 7 flat, imbricate seales. At the base of the claws, at the tips of the fingers, a transversely enlarged scale is present ventrally; this is followed by another seale of similar shape, but much narrower. About 14 minute, pointed, thorn-like, juxtaposed seales occur in regular diagonal series from one lateral fringe to the other. Third and fourth fingers are of almost equal length; the first is shorter than the fifth. First and fifth toes end at about the same level; fourth toe slightly longer than the third. The tail tapers very gradually and does not form a whip-like end portion. MEASUREMENTS OF HOLOTYPE AND ALLOTYPE (in mm.) : Holotype Allotype (CAS 84458) (CAS 84459) Motallenethh ssh tlh. eesti tech er acce eee ene eee 89 95 Pail Wen othr: 2383s ie pgeerwcrns, ¢ eee. ahaa teeters OT 40 eneth-or torelanli-c.2 sich ee 21 24 bene thy ofehamdiinmpn eens seen ee 29 30 Weneth ot meade ee escent. ae ee aa ee 15 15 Wartelbliwost they eet vers esi Brecon Scones eee a ee 99 9.5 Remarks: In the allotype (female), CAS 84459, there is a black, irreg- ular, reticulate pattern, leaving more or less cireular bright ocelli; several dark cross-bands are present on the tail, the spaces between them more or less mottled; body white ventrally, with darker specks on the lateral and anterior gular region; anterior pectoral area with an ill-defined trans- verse dark marking; markings on ventral side of the tail resemble the dorsal ones, but are fainter; scattered single dark scales are present on the underside of the limbs. In the holotype (male), CAS 84458, the dark dorsal markings show some ‘“‘condensations” in the form of ill-defined transverse bands on the occiput, neck, and pectoral area, with two bands across the body followed by 6 to 7 cross-bands on the tail. Ventral surface as in the female, but dark areas are fainter. In both specimens small dark spots are present on the supra- and infra- labials; tubercles on the head are usually entirely either bright or dark. Genus Alsophylax Fitzinger Alsophylax blanfordi (Strauch). Bunopus blanfordii Straucu, 1887, Mém. Acad. Sci. St. Pétersbourg, (7) 35:61: pl. 1, figs. 13-14 (Egypt [in error]: designated as Arabia [Schmidt, 1939, Field Mus. Nat. Hist., Zool. ser., 24:55]). Alsophylax blanfordii PARKER, 1931, Ann. Mag. Nat. Hist., (10) 8:519. VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 63 MATERIAL EXAMINED (4): CAS 84341, from Abqaiq, collected in 1945, by Gasperetti; CNHM 73993, from Dhahran, collected in 1946, by Gas- peretti; HUJ 2686, from Tebuk, collected in 1942, by Hardy; HUJ 2687, from central Arabia, collected in 1942, by Hardy. Remarks: The single male, CAS 84341, has 15 preanal pores. In the specimen from Tebuk, HU.J 2686, a female, the dorsal scales are more strongly keeled and more closely juxtaposed, especially in the temporal region, than in the remaining specimens. Genus Gymnodactylus Spix Gymnodactylus scaber (Heyden). Stenodactylus scaber HrybdEN, 1827, in Ruppell, Atlas Reise nord. Afrika, Rept., p. 15, pl. 4, fig. 2 (vicinity of Tor, Sinai). Gymnodactylus scaber DuUMERIL and BrBron, 1836, Erp. Gén., 3:421. MATERIAL EXAMINED (14): CAS 84480, 84483, from Abqaiq, collected in 1946, by Gasperetti; CAS 84389-84390, from Al Khobar, collected in 1945, by Gasperetti; CAS 84439, from Ras at Tanura, collected in 1946, by Gasperetti; CAS 84275-84276, 84512, 84517, 84532-84533, 84538, and CNHM 74018-74019, from Dhahran, collected in 1945-1946, by Gasperetti. Remarks: The preanal pores vary from 4 to 6; three specimens, of the fourteen, have 3 pairs of chin shields instead of the usual 2 pairs. Genus Pristurus Ruppell Pristurus carteri carteri (Gray). Spatalura carteri Gray, 1863, Proc. Zool. Soc. London, 1863:236: pl. 20, fig. 2 (Makulla, southern Arabia; corrected to Masirah Island [Parker, 1931, p. 515]). Pristurus carteri carteri Parker, 1931, Ann. Mag. Nat. Hist., (10) 8:515. MATERIAL EXAMINED (1): HUJ 2694, from Dhufar (at sea level), col- lected in 1945, by Vesey-Fitzgerald. Remarks: This specimen is much paler than specimens of Pristurus cartert tuberculatus. The nostril is set between a large crescentic anterior nasal and the first labial, the gap being closed by a minute posterior nasal. The ear-opening is an oblique slit. The snout is more pointed in profile than in P. c. tuberculatus. 64 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Pristurus carteri tuberculatus Parker. Pristurus carteri tuberculatus PARKER, 1931, Ann. Mag. Nat. Hist., (10) 8:515 (Dhu- far, littoral [Ain al Rizat, Milwah Alaud, and Sahalnaut]). MATERIAL EXAMINED (3): HUJ 2685, 2695-2696, from Jebel Qara (2000 feet altitude), collected in 1943, by Vesey-Fitzgerald. Remarks: HUJ 2685 and 2696 have clearly enlarged lateral tubercles; in HUJ 2695 these are feebly developed and are restricted to the lumbar region. Pristurus rupestris rupestris Blanford. Pristurus rupestris BLANFoRD, 1874, Ann. Mag. Nat. Hist., (4) 13:454 (Muscat and Kharag Island; restricted to Muscat [Schmidt, 1952, Nat. Hist. Misc., no. 7, p. 2 and 1955, Vidensk. Medd. fra Dansk. naturh. Foren., 117:202]). Pristurus rupestris rupestris ScuMiptT, 1953, Fieldiana: Zoology, 34:256. MATERIAL EXAMINED (2): HUJ 2688, 2693, from Dhufar (at sea level). collected in 1945, by Vesey-Fitzgerald. Remarks: The female specimen has a light vertebral line, and the male has the normal dorsal ocellate pattern. Genus Hemidactylus Oken Hemidactylus persicus Anderson. Hemidactylus persicus ANDERSON, 1872, Proc. Zool. Soc. London, 1872:378: fig. 2 (Shiraz, Persia). MATERIAL EXAMINED (1): CAS 84388, from Al Khobar, collected in 1945, by Gasperetti. REMARKS: The 8 preanal pores are arranged in an angular series, with the apex directed forward; 9 lamellae are present beneath the inner finger, and 11 beneath the third finger; 8 lamellae are present beneath the inner toe, and 11 beneath the third toe (Boulenger [1885, p. 131] gives, re- spectively, 10 and 10, and 9 and 12). The rostral is not fused with the first labials. Genus Ptyodactylus Gray Ptyodactylus hasselquisti (Donndorff). Lacerta hasselquistii DonnporFF, 1789, Zool. Beytr., 3:133 (Egypt). Ptyodactylus hasselquistii DuMERIL and Bipron, 1836, Erp. Gén., 3:378. VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 65 MATERIAL EXAMINED (1): CAS 84591, from Dhahran, collected in 1946, by Gasperetti. Remarks: The variability in this species is astonishing. The pres- ent specimen differs conspicuously from Palestinian ones, but a definite subspecifie classification is not yet feasible. There are 15 supralabials in this specimen as compared with 14, the usual number in Palestinian specimens; also, 11 infralabials are present, compared with 13 to 15 in the Palestinian form. The inner chin shields are very elongate, three times longer than broad; they meet behind the mental for one-third of their total length (in specimens from Palestine they are always short and separated from one another). The inner en- larged pair of chin shields is followed by enlarged, posteriorly decreasing postmentals, 4 on the right side and 5 on the left. The head, seen from above, is much narrower than in Palestinian specimens. The dorsal tu- bercules are flatter and larger in the Arabian specimen. The ventral seales of the tail form about 6 transverse series in every annulus as com- pared with 4 in the typical form. MEASUREMENTS (in mm.): Snout-vent length 70.5; tail length 60. Family AGAMIDAE Genus Agama Daudin Agama sinaita Heyden. Agama sinaita Hrypen, 1827, in Riippell, Atlas Reise nord. Afrika, Rept., p. 10, pl. 3 (Sinai). MATERIAL EXAMINED (4): HUJ 2702-2703, from Tebuk, Jauf Moun- tains, collected in 1942, by Hardy; HUJ 2704, from north of Jebel Aja, collected in 1944, by Waterston; HUJ 2705, from Jebel Aja, west of Hail, collected in 1944, by Waterston. Remarks: The terrain north of Jebel Aja is designated as rock and shrub; at Jebel Aja, it is granite rock. The lizards were active in the heat of the afternoon. The 4 specimens agree closely with specimens from southern Palestine. Agama pallida Reuss. Agama pallida Reuss, 1834, Mus. Senck., 1:38: pl. 3, fig. 3 (upper Egypt). MATERIAL EXAMINED (3): CAS 84610, and CNHM 74017, from Dhahran, collected in 1946, by Gasperetti; HUJ 2701, from north of Jebel Aja, col- lected in 1944, by Waterston. 66 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Remarks: In CAS 84610, the enlarged dorsals are smooth, and are as wide as 4 small dorsals; there are 7 small dorsals in a longitudinal line between the enlarged dorsals. In CNHM 74017, the enlarged dorsals are keeled, smaller, and more numerous, with only 4 or 5 small dorsals be- tween them. Agama agilis Oliver. Agama agilis OLiver, 1804, Voy. Emp. Ottoman, 4:394: pl. 29, fig. 2 (Baghdad, Iraq). MATERIAL EXAMINED (1): CAS 84416, from Qatif, collected in 1945, by Gasperetti. MEASUREMENTS (in mm.): Total length 220; tail leneth 125. Agama persica Blanford. Agama persica BLANFORD, 1881, Proc. Zool. Soc. London, 1881:674: pl. 49 (Deh Bid and Kazerun, Iran). MATERIAL EXAMINED (11): CAS 84417, from Qatif, collected in 1945, by Gasperetti; CAS 84477, 84491-84492, from Abqaiq, collected in 1946, by Gasperetti; CAS 84516, 84541, 84611, 84613-84614, and CNHM 74015- 74016, from Dhahran, collected in 1946, by Gasperetti. Remarks: All the males in this series have a single row of false pre- anal pores, as indicated in Blanford’s original description, not two rows, as stated by Boulenger (1885, p. 345). Eight of the specimens at hand (of both sexes) have a vertebral band of rather small keeled scales clearly set off from the larger lateral scales. CAS 84611 does not exhibit the row of smaller mid-dorsal scales; CAS 84477, from Abqaig, and CNHM 74015, from Dhahran, are intermediate. Some specimens have remarkably long tails; CAS 84477, a male, has a total length of 264 mm., and a tail length of 167 mm. In specimen CAS 84477, four longitudinal dark brown stripes fuse with the dark area on the upper side of the head. A broad dark band links the posterior angle of the orbit with the ear opening. Above this there is another stripe at the upper border of the temporal region. The gular region is black. The sides of the back and belly are black; medio-ventrally, from the anterior border of the pectoral girdle to the anterior border of the pelvic girdle, there is a slightly paler dark stripe; the rest of the belly is lighter. On the back there are two indistinctly defined longitudinal series of rectangular dark spots. The tail is very light, with very faint transverse gray cross-bars, which are much narrower than the light inter- vals. The dark upper side of the body contrasts strikingly with the light tail. VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 67 Agama jayakari Anderson. Agama jayakari ANDERSON, 1896, Contri. herpet. Arabia, p. 65, (Maskat). MATERIAL EXAMINED (2): HUJ 2699-2700, from Shinas, Batinah Coast, collected in 1944, by Vesey-Fitzgerald. (HUJ 2700 is without a field label.) REMARKS: These specimens differ from the original description in hav- ing 77 seales around the body, instead of 90; in having the ear opening larger than the eye, instead of smaller; and in the considerably larger size. The male specimen, HUJ 2699, measures 360 mm. total length, and 207 mm. tail length; the female, HUJ 2700, measures 298 mm. total leneth, and 165 mm. tail length. In corresponding measurements, the largest male and female in Anderson’s series measured respectively 316 mm. and 178 mm., and 266 mm. and 151 mm. The male specimen has no preanal pores, and has a much larger gular pouch. Further description of this form is deferred for comparison with more extensive material of Agama jayakari. Genus Phrynocephalus Kaup Phrynocephalus arabicus Anderson. Phrynocephalus arabicus ANDERSON, 1894, Ann. Mag. Nat. Hist., (6) 14:377 (Had- ramout). MATERIAL EXAMINED (5): HUJ 2706-2710, from Abu Dhabi, collected in March, 1944, by Vesey-Fitzgerald. (HUJ 2710 is without a field label.) RemMARKS: This series agrees fairly well with the original description of P. arabicus. The dorsal surfaces of the Lmbs in HUJ 2709 and 2710 are covered with keeled scales. The adlabial series of scales (adjacent to the lower labials) does not reach the first lower labial or the mental; the next series extends to the mental and first labial. The adlabials of P. nejdensis are better developed and reach the first labial. There are 3 or 4 more or less distinet dark cross-bands beneath the tail; the distal third of the tail is invariably black. MEASUREMENTS oF HUJ 2710 (gravid female) (in mm.): Total length 72; tail length 38. Phrynocephalus maculatus longicaudatus Haas, new subspecies. HouotyrE: CAS 84449, male, from Doha Dhalum, Satdi Arabia, col- lected July 22, 1946, by John Gasperetti. 68 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Paratypes (14): CAS 84442-84448, 84450-84452, 84455, and CNHM 74021-74022, from the type locality, collected in 1946, by Gasperetti; CAS 84511, from Dhahran, collected in 1946, by Gasperetti. DiaGnosis: Differs from Phyrnocephalus maculatus maculatus in hav- ine the posterior supraorbital scales strikingly flattened and enlarged, longer than wide, and larger than the mid-dorsal scales; a few dorsal scales are keeled or with an indication of mucronation; nostrils are directed for- ward instead of upward; tail is longer than twice the distance from the gular fold to the vent; dark coloration of the distal part of the tail is pro- nounced ventrally. MEASUREMENTS (in mm.) : Holotype CAS 84449 2 CAS 844424 CAS 84443 ¢ CAS 844449 CNHM 74022 9 Total length.. 196 187 186 159 171 Tail length ... 124 114 114 96 105 Phrynocephalus nejdensis Haas, new species. Hotoryre: HUJ 2711, male, from Nafud, near Qana, North Nejd, WNW of Hail, Satidi Arabia, collected May 19, 1944, by A. R. Waterston. Dracnosis: Related to the P. luteoguttatus-arabicus group, differing from them in the small size of the gular scales, smaller ventrals and prox- imal subeaudals, higher numbers of upper and lower labials, and flatness of the dorsal head seales. DESCRIPTION OF TYPE: Scales on snout juxtaposed, somewhat imbricate in the supraocular, frontal, and temporal areas; each dorsal seale on the snout with a pit near its anterior border; these pits are absent in the upper labia! region; nostrils between 2 crescent-shaped nasals, directed forward, not upward; 2 scales, one behind the other, between the 2 pairs of nasals; 8 longitudinally ridged supraoculars on the left side, 9 on the right side; rostral not differentiated; 14 upper labials on each side, the posterior ones enlarged; 14 lower labials on each side, each with 2 or 3 pits in its lower border; 2 chin shields in contact with the lower labials (thus different from P. luteoguttatus) and 2 with the enlarged mental; 3 series of scales between the orbit and upper labials; 4 scales between the anteroventral border of the first nasal and the series of upper labials; 12 very prominent fringe-like scales on the lower eyelid; a group of about 4 longitudinally set conical scales present above the tympanic area; en- larged occipital shield equals about 3 of the adjacent scales; the wrist of the extended forelimb reaches the snout; extended backward, the finger- tips of the forelimb reach the middle of the backward extended thigh; hind VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 69 leg, stretched forward, extends well beyond the tip of the snout; claws long, strongly compressed laterally, weakly curved, 3 mm. long on the fourth finger. The subequal dorsals and upper nuchals are feebly keeled and imbri- cate; juxtaposed, flat tubercles present toward the root of the tail; scales narrower, but not shorter, at the sides, granular in the axilla and in the eroin; proximal upper ecaudals larger than largest dorsals; more distal eaudals sharply keeled and elongate; gulars imbricate and pointed, grad- ually diminishing in size posteriorly; ventrals mucronate, some of them indistinctly keeled; scales on upper sides of limbs smooth, larger than those on the back; subeaudals smaller than ventrals. Counted perpendicularly to the tiers of the seales, there are, in a leneth of 5 mm., 12 gular seales (gular region), 6 ventrals (pectoral area), 12 dorsals (mid-body), and 10 dorsals (at base of tail). There are about 115 seales around the middle of the body. Dorsally, there is a fine gray reticulum enclosing round yellow spots; upper sides of limbs with transverse bars; posterior half of tail blackish, with faint darker cross-bands; a series of black spots present on each side of the anterior portion of the tail; the ocellate pattern of the back continues onto the anterior part of the tail; the latter is uniformly gray distally. Ventral surface white; collar region yellow with longitudinal gray stripes. MEASUREMENTS OF HOLOTYPE (in mm.): Total length 116; tail length 62; forelimb 32; hindlimb 50. REMARKS: Phrynocephalus nejdensis is related to P. luteoguttatus; the latter has a longer tail and has the medial gular scales smaller than the ventrals. Phrynocephalus nejdensis macropeltis Haas, new subspecies. Houotyre: CAS 84619, male, from Dhahran, Satidi Arabia, collected in 1946, by John Gasperetti. PaRATYPEs (48): CAS 84287-84290, 84292, 84294-84295, 84297, 84379- 84387, 84479, 84484-84485, from Abqaiq, collected in 1945-1946, by Gas- peretti; CAS 84262-84263, 84277-84278, 84280-84281, 84283-84286, 84502, 84510, 84513, 84536, 84542-84543, 84545, 84547, 84617-84618, 84620, 84622- 84623, and CNHM 73998-73999, from Dhahran, collected in 1945-1946, by Gasperetti; CAS 84421, from Shimal, collected in 1946, by Gasperetti; CAS 84432, 84437, from I] Alat, collected May 31, 1946, by Gasperetti. DiaGcnosis: Differs from Phrynocephalus nejdensis nejdensis in having larger ventral scales, about 90 scales around the middle of the body (com- 70 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. pared with 115 in the typical subspecies), head scales more bulging, and some of the pectoral scales keeled. DescripTION: Habitus of Phrynocephalus; the adpressed forelimb ex- tends backward beyond the groin; the adpressed hindlimb extends forward to the anterior angle of the orbit; the tail is slightly longer than the head and body together. Dorsal seales are nearly uniform, wider than long, a very few shghtly enlarged, some feebly keeled; anterior ventral scales are keeled and mu- eronate, but not in all specimens; gular scales smooth, pointed, larger than the dorsals; caudal scales keeled from the first third of the tail to its tip. Fringes present on both sides of the third and fourth fingers and toes, and are longest on the outer side of the fourth toe; claws very long (in one large specimen, the claw on the fifth toe measured + mm. in length), shorter on the fingers than the toes. Two parallel transverse folds present on the neck; a lateral cervical fold extends to the groin. Color variable, some specimens nearly uniform, either dark or light, others with an ocellate pattern, or mottled; last third of the tail is black below, less uniformly so above. MEASUREMENTS OF HOLOTYPE (in mm.): Total length 114; tail length 58. Genus Uromastyx Merrem Uromastyx microlepis Blanford. Uromastix microlepis BLANFORD, 1874, Proc. Zool. Soc. London, 1874: 656: pl. 53 (Basra, Iraq). MATERIAL EXAMINED (6): CAS 84430, from Jebel Dam, collected in 1946, by Gasperetti; CAS 84575-84577, and CNHM 73989, from Dhahran, collected in 1946, by Gasperetti; HUJ 2698, from Sakaka, collected in 1945, by Hardy. Family VARANIDAE Genus Varanus Merrem Varanus griseus (Daudin). Tupinambis griseus DAUDIN, 1803, Hist. Nat. Rept., 8:352 (Egypt). Varanus griseus BOULENGER, 1885, Cat. Lizards Brit. Mus., 2:306. MATERIAL EXAMINED (8): CAS 84298, 84300-84302, 84473, and CNHM 74020, from Abqaiq, collected in 1945, by Gasperetti; CAS 84267, from Vou. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 71 Doha Dhalum, collected in 1945, by Gasperetti; CAS 84422, from Shimal, collected in 1946, by Gasperetti. REMARKS: Specimen CAS 84422 is a handsomely colored juvenile. The adult specimens are of moderate size, none attaining the maximum of 1200 mm. recorded for this species. Family AMPHISBAENIDAE Genus Diplometopon Nikolsky Diplometopon zarudnyi Nikolsky. (Figure 8.) Diplometopon zarudnyi Nixousky, 1907, Ann. Mus. Zool. Acad. Sci. Petrograd, 10:277: figs 1-3 (Nasrie, Arabistan, Iran). MATERIAL EXAMINED (29): CAS 84274, 84406, 84529, 84534, 84539, 84559, 84588, and CNHM 74007, from Dhahran, collected in 1945-1946, by Gasperetti; CAS 84343-84355, 84373, and CNHM 74008, from Abgqaiq, collected in 1945, by Gasperetti; CAS 84426, from Shimal, collected in 1946, by Gasperetti; CAS 84454, from Doha Dhalum, collected in 1946, by Gasperetti; HU.J 2690-2692, from between Hadj and Tebuk, collected in 1942, by Hardy; HUJ 2689, from Sharja, collected in 1944, by Vesey- Fitzgerald. REMARKS: Most of the specimens do not show a dorsal impressed line, but the annuli are interrupted on the mid-dorsal line by 2 triangular Figure 8. Diplometopon zarudnyi Nikolsky. Dorsal and lateral views of head of HUJ 2696. 72 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER. scales meeting at their apices. Ventrally, a deep median groove is always present. CAS 84345 has a completely divided frontal. Family LACERTIDAE Genus Acanthodactylus Wiegmann Acanthodactylus scutellatus hardyi Haas, new subspecies. Acanthodactylus scutellatus PARKER, 1931, Ann. Mag. Nat. Hist., (10) 8:521. Houotyre: HUJ 2682, from Hirmas Station, Satidi Arabia, collected December 15, 1942, by J. E. Hardy. DracNosts: Differs from Acanthodactylus scutellatus scutellatus, the form found in Egypt and Palestine, in having the dorsal scales entirely smooth, 12 ventral plates across the belly instead of 14, fourth supraocular broken up into granules, no enlarged anterior gular scales bordering the chin shields, prefrontals only slightly longer than broad, median suture of nasals very short, and the temporal granules completely smooth. Remarks: Parker (1931) records Acanthodactylus scutellatus from southern Arabia. In the conspicuous character of the number of scales across the belly, his specimen agrees with ours. He, also, suspected a dis- tinguishable geographic race in southern Arabia. Acanthodactylus fraseri Boulenger. Acanthodactylus fraseri BouLENGER, 1918, Jour. Bombay Nat. Hist. Soc., 25:373 (Zobeiya, Shariba, Iraq). MATERIAL EXAMINED (10): CAS 84271, 84596, from Dhahran, collected in 1945-1946 by Gasperetti; CAS 84333, 84338-84339, 84486, and CNHM 73987-73988, from Abqaiq, collected in 1945-1946, by Gasperetti; CAS 84433, 84436, from El Alat, collected in 1946, by Gasperetti. REMARKS: These specimens have 4 supraoculars in front of the sub- oculars; the subocular reaches the labial border. Acanthodactylus fraser is smaller than A. cantoris. Acanthodactylus cantoris schmidti Haas, new subspecies. Houotyrpe: CAS 84599, male, from Dhahran, Satidi Arabia, collected in 1946, by John Gasperetti. VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 73 PaRATYPES (24): CAS 84268-84270, 84530, 84598, 84600-84606, 84608— 84609, and CNHM 74010-74011, from Dhahran, collected in 1945-1946, by Gasperetti; CAS 84332, 84336-84337, 84340, 84342, from Abgqaiq, col- lected in 1945-1946, by Gasperetti; CAS 84419, from Qatif, collected in 1945, by Gasperetti; CAS 84423, from Shimal, collected in 1946, by Gas- peretti; HUJ 2691, from north of Tebuk, collected in 1942, by Hardy. Diacnosis: Larger than Acanthodactylus cantoris arabicus; dorsal pat- tern reticulate instead of lineate; in the dorsolateral band of strongly keeled seales, the scales are about twice the size of the dorsals between them; the difference in size of these scales is most pronounced in the adult males. MEASUREMENTS (in mm.) : Holotype CAS 84599 3 HUJ 2681 3 MMLC TING MG ayers tee cs es eters wid es 8 = 241 281 ‘TRL GTC 1 pee Senter Ee Coase cn ee ae ee 160 187 Genus Eremias Wiegmann Eremias brevirostris (Blanford). Mesalina brevirostris BLANKoRD, 1874, Ann. Mag. Nat. Hist., (4) 14:32 (Kalabagh, Punjab, and Tumb Island, Persian Gulf. Restricted by Schmidt, 1939, Field Mus. Nat. Hist., Zool. Ser., 24:66, to Kalabagh, Punjab). Eremias brevirostris BOULENGER, 1887, Cat. Lizards Brit. Mus., 3:89. MATERIAL EXAMINED (14): CAS 84391-84392, 84394-84395, 84397— 84402, 84404, and CNHM 73990-73991, from Dammam, collected in 1945, by Gasperetti; HUJ 2679, from central Arabia, collected in 1942, by Hardy. Remarks: The proposal of a Syrian subspecies, Hremias brevirostris microlepis, by Angel (1936, p. 112), based upon specimens lifted out of a series of supposedly typical FE. brevirostris, is certainly not adequately established. Eremias guttulata guttulata (Lichtenstein). Lacerta guttulata LICHTENSTEIN, 1823, Verz. Doubl. Mus. Berlin, p. 101 (Egypt). Eremias guttulata guttulata WETTSTEIN, 1928, Sitzber. Akad. wiss. Wien, (math.- natur:), L3%s Abt, 14782: MATERIAL EXAMINED (1): HUJ 2683, from central Arabia, collected in 1942, by Hardy. 74 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H SER. Eremias adramitana Boulenger. Eremias adramitana BouLeNGER, 1917, Ann. Mag. Nat. Hist., (8) 19:279 (Hadra- maut). MATERIAL EXAMINED (1): HUJ 2680, from Abu Dhabi, collected in 1944, by Vesey-Fitzgerald. Family ScINcIDAE Genus Eumeces Wiegmann Eumeces taeniolatus (Blyth). (Figure 9.) Eurylepis taeniolatus Biytu, 1854, Jour. Asiatic Soc. Bengal, 23:470 (Salt Range, Punjab, India). Eumeces taeniolatus SroticzKaA, 1872, Proc. Asiatic Soc. Bengal, 1872:75. cenlinmelers Figure 9. Humeces taeniolatus (Blyth). Dorsal view to show distinctive color pattern. VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 75 MATERIAL EXAMINED (1): HUJ 2670, from 23 miles north of Hail, col- lected in 1944, by Waterston. Remarks: This distinct species, described from India, is represented in the collections of the British Museum by specimens from Arabia. These eome from El Khubar, southwest Arabia, and from Museat. The remark- able color pattern is shown in the accompanying figure. Our specimen has 9/8 upper labials, 58 transversely widened mid-dorsal scales, and 19 seale rows around the body. Maleolm Smith (1935, p. 342) has shown that Kumeces scutatus (Theo- bald) is a strict svnonym of Eumeces taeniolatus (Blyth). Genus Scincus Gronovius Scincus gasperettii Haas, new species. (Figure 10.) HouotyrE: CAS 84519, male, from Dhahran, Satidi Arabia, collected July 27, 1946, by John Gasperetti. PaRATYPE (1): CNHM 73986, male, from Abqaiq, collected in July, 1945, by Gasperetti. DiaGcnNosis: A species of the S. meccensis-nutranus group, with 5 supra- oculars, snout rather short, and ear-opening visible as an oblique slit; dis- Figure 10. Scincus gasperettii Haas, new species. Dorsal view of head of CAS 84519, holotype. 76 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. tinguished from both S. meccensis and S. mitranus by having 26 scale rows around the body. DeEscrIpTION: Upper labials 8; lower labials 7; 3 loreals; 3 supracili- aries; 2 azygous post-mentals. Rostral not in contact with the median internasal; the 2 postnasals meet in a short sagittal suture; prefrontals paired; frontal normal in holotype, transversely divided in paratype, with anterior portion twice as long as posterior part; 4 triangularly pointed seales along the anterior border of the ear opening; nuchals 4/4. General color light brown, the posterior border of dorsal seales dark brown; patches of dorsal scales unicolor, so that a faint pattern of darker and lighter spotting is produced. No dark brown lateral spots present. MEASUREMENTS (in mm.): Holotype CAS 84519 4 CNHM 73986 ¢ OPAL CTI CUA. S.2cctcxe ee eepeteK ae oro bine eae 114 115 amen Clas os coe ce NR te. Ake sohciieue waeuare 44 49 Scincus meccensis Wiegmann. Scincus meccensis WIEGMANN, 1837, Arch. Natur., p. 127 (Arabia). MATERIAL EXAMINED (1): HU.J 2673, from Mudawara, central Arabia, collected in 1944, by Waterston. REMARKS: This specimen, with only 22 scale rows around the body, comes closest to Scincus meccensis. It has 6/5 supraoculars; 3 supracili- aries, the first much the longest; rostral with a short suture with the fronto- nasal; snout elongate; loreal depression slight; 8/7 upper labials; frontal one and two-thirds as lone as wide; prefrontals paired; 5/7 nuchals; 4/5 triangular scales in front of the oblique ear opening. Definitive allocation of this specimen is obviously impossible without additional material from central Arabia. Scincus deserti Haas, new species. (Figure 11.) Hototyee: HUJ 2674, from 23 miles north of Hail, Satidi Arabia, collected in 1944, by A. R. Waterston. PaRatTyPeE (1): HUJ 2752, from central Arabia, collected in 1944, by Waterston. Diacnosis: A Scincus with 26 seale rows around the body; ear open- ing visible, an oblique slit, with 4 triangular and somewhat fringed scales covering it; 6 supraoculars; frontonasal in contact with rostral; 2 pre- VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA fil frontals; snout longer and narrower than in S. gasperetti; frontal elongate, one and one-half times as long as wide, narrowed posteriorly. Four faint brownish lateral spots present on each side. RemMArRKs: In the paratype there are 27 scale rows around the body; also, there is a fifth pair of brown spots in front of the forelimbs, with a brown marking on each knee. This species differs from Scincus sconcus in the absence of transverse dorsal bars; in having 4 fringed scales in front of the ear opening instead of 2; and in the presence of the lateral spots. Scincus deserti differs from S. phailbyi in having the ear opening clearly discernible; the supraocular region does not bulge as in S. philbyi; the snout tapers gradually and does not have the spatulate form of the latter species; and the eanthus rostralis is feebly marked, whereas it is sharply defined in S. philbyt. MEASUREMENTS (in mm.) : Holotype HUJ 2674 HUJ 2752 TM estaill) Il ervaa ay “2 pee eee eee Ore enna 157 165 pl ITaaaiLtiy Se Os See eee eee eee 70 59 Present bn ae Sea tintc tenes Seely Bees cca KE ncad! Shama aie labs 28 29 [BUTE ITNT ak Slo RR es Co ene 29 oT Figure 11. Scincus deserti Haas, new species. Dorsal view of head of HUJ 2674, holotype. 78 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Scincus philbyi Schmidt Scincus philbyi Scumipt, 1941, Field Mus. Nat. Hist., Zool. Ser., 24:162: fig. 18 (Anaiza, Satdi Arabia). MATERIAL EXAMINED (45): CAS 84296, 84303-84306, 84308-84310, 84312-84317, 84319-84320, 84324-84330, 84487-84489, 84494, and CNHM 74004, from Abqaiq, collected in 1945-1946, by Gasperetti; CAS 84535, 84580-84587, and CNHM 74003, from Dhahran, collected in 1945-1946, by Gasperetti; CAS 84414, from Qatif, collected in 1945, by Gasperetti; CAS 84427, 84429, from Shimal, collected in 1946, by Gasperetti; CAS 84435, from El] Alat, collected in 1946, by Gasperetti; CAS 84626, from Tomayiah, collected in 1945, by Gasperetti; HUJ 2671-2672, from Sharja, collected in 1944, by Vesey-Fitzgerald. REMARKS: Of the 45 specimens in this series, 14 specimens have 2 prefrontals, which is normal for other species in the genus, and 31 indi- viduals have the prefrontal fused into a single scale, as in the type series of S. philbyx. The number of brown lateral spots is variable; in some specimens they were faint or absent. The color of the spots varies from yellow to very dark brown. A small third loreal scale may be present anterior to the 2 normal loreals. Figure 12. Scincus philbyi Schmidt. Dorsal view of head of CAS 84414, to show divided prefrontal. VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA ow Family CHAMAELEONIDAE Genus Chamaeleo Laurenti Chamaeleo calyptratus Duméril and Duméril. Chamaeleo calyptratus DuMrERIL and DumeErRIL, 1851, Cat. Méthod. Rept., p. 31 (no locality given).—Anderson, 1896, Contri. Herp. Arabia, p. 62 (Yemen).— Schmidt, 1953, Fieldiana: Zoology, 34:259. MATERIAL EXAMINED (1): HUJ 2675, from Jebel Qara, Dhufar, col- lected in 1948, by Vesey-Fitzgerald. Order SERPENTES Family Borpar Genus Eryx Daudin Eryx jayakari Boulenger. Eryx jayakari BOULENGER, 1880, Ann. Mag. Nat. Hist., (6) 2:508 (Muscat, Arabia). MATERIAL EXAMINED (15): CAS 84857-84358, 84476, 84493, 84499, and CNHM 74023, from Abgqaiq, collected in 1945-1946, by Gasperetti; CAS 845538, 84570-84572, 84574, and CNHM 74024, from Dhahran, col- lected in 1946, by Gasperetti; CAS 84434, 84438, from El Alat, collected in 1946, by Gasperetti; HUJ 3868, from Sharja, Trucial Coast, Oman, collected in 1944, by Vesey-Fitzgerald. REMARKS: Twelve specimens had 5 scales between the eyes; two speci- mens had 4 seales between the eyes. Genus Coluber Linnaeus Coluber ventromaculatus Gray. Coluber ventromaculatus Gray, 1834, Illus. Indian Zool., 2: pl. 80, fig. 1 (Bengal). MATERIAL EXAMINED (5): CAS 84544, 84554, 84565, and CNHM 74000, from Dhahran, collected in 1946, by Gasperetti; HUJ 3863, from Jebel Qara, collected in 1943, by Vesey-Fitzgerald. Remarks: In the single juvenile specimen, CAS 84565 (much dam- aged), the anterior dorsal spots are longitudinally elliptic, changing to circular at mid-body, and to transverse saddle-shaped markings posteriorly; toward the vent the markings disappear; they are absent on the tail. 80 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. In 3 adult males, the ventrals vary from 203 to 206, the caudals from 95 to 100. There are 214 ventrals and 119 caudals in the single female (HUJ 3863). Genus Lytorhynchus Peters Lytorhynchus diadema arabicus Haas, new subspecies. Hototype: CAS 84478, male, from Abqaiq, Satidi Arabia, collected May 17, 1946, by John Gasperetti. PARATYPES (8): CAS 84504, 84526, 84557, 84579, and CNHM 73955, from Dhahran, collected in 1946, by Gasperetti; HUJ 3859, from Satdi Arabia, collected in 1944, by Waterston; HUJ 3860, from Dhahran, collected in 1943, by Vesey-Fitzgerald; HUJ 3867, from Moreiwa Post (not located on map), collected in 1942, by Hardy. Diacnosis: Habitus more slender than the typical subspecies, with wider intervals between the dorsal spots, and a higher number of ventrals (177-193 in L. d. arabicus as compared with 161-172 in L. d. diadema). DescripTION: Loreal single; 1 to 3 preoculars; 2 postoculars; 2 temporals in the first row, 2 or 3 temporals in the second row. Rostral strongly pro- jecting; deeply excavated beneath. Upper labials 8, the fifth or fourth and fifth enter the orbit; lower labials 7 to 11, usually 9 or 10. Seale rows at mid-body 19; ventrals 177-193; caudals with maximum of 47. Thirty-five to 50 dark blotches present on the body, 9 to 16 blotches on the tail. Genus Malpolon Fitzinger Malpolon moilensis (Reuss). Coluber moilensis Reuss, 1834, Mus. Senck., 1:142; pl. 7, fig. 1 (Moilah, Arabia). Malpolon moilensis Parker, 1931, Ann. Mag. Nat. Hist., (10) 8:522. MATERIAL EXAMINED (9): CAS 84474, 84500, 84625, from Abqaiq, col- lected in 1946, by Gasperetti; CAS 84515, 84551, 84567-84568, and CNHM 74002, from Dhahran, collected in 1946, by Gasperetti; HUJ 3861, from Dhahran, collected in 1943, by Vesey-Fitzgerald. ReMaArRKS: This species has a wide, but discontinuous distribution, confined to widely scattered localities from Palmyra to the middle of the Arabian Peninsula. Vor. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 81 Genus Psammophis Boie Psammophis schokari (lorskal). Coluber schokari ForsKAL, 1775, Descr. Anim., p. 14 (Yemen). Psammophis schokari BouLENGER, 1896, Cat. Snakes Brit. Mus., 3:157. MATERIAL EXAMINED (4): CAS 84418, from Qatif, collected in 1945, by Gasperetti; CNHM 74001, from Dhahran, collected in 1946, by Gas- paretti; HUJ 3866, from 23 miles north of Hail, collected in 1944, by Wat- erston; HUJ 3865, from Jebel Qara, collected in 1943, by Waterston. Genus Naja Laurenti Naja haje arabica Scortecci. Naja haie arabica Scortecct, 1932, Atti Soc. Ital. Sci. Nat., 71:47 (San ’a Yemen). Naja haje arabica Parker, 1941, Brit. Mus. (Nat. Hist.), 1:5. MATERIAL EXAMINED (1): HUJ 3864, from Jebel Qara, collected in 1943, by Vesey-Fitzgerald. ReMarRKS: This specimen is a juvenile, 535 mm. in total length and 106 mm. in tail length. There are 214 ventrals and 77 caudals. The scales are in 19 rows; they do not increase on the neck. General color yellowish brown, darker on top of the head; the dorsal scales have narrow dark margins; white below, with a median interrupted dark brown band; each ventral with a dark posterior border. Genus Walterinnesia Lataste Walterinnesia aegypti Lataste. Walterinnesia aegyptia LATASTE, 1887, Le Naturaliste, 1887:411 (Egypt). MatTreRIAL EXAMINED (3): CAS 84498, and CNHM 74025, from Abqaiq, collected in 1946, by Gasperetti; CAS 84578, from Dhahran, collected in 1946, by Gasperetti. Remarks: Marx (1953) has shown that Naja morgani Mocquard is clearly a synonym of Walterinnesia aegyptia. 82 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Family HypRoPHIUDAE Genus Hydrophis Latreille Hydrophis cyanocinctus Daudin. Hydrophis cyanocinctus DAupIN, 1803, Hist. Nat. Rept., 7:383 (Coromandel). MATERIAL EXAMINED (10): CAS 84463-84464, 84466-84468, 84470- 84472, and CNHM 73996-73997, from Al Khobar, collected in 1946, by Gasperetti. Remarks: The longest specimen measures 673 mm. in total length, and 73 mm. in tail length. Family VIPERIDAE tenus Pseudocerastes Boulenger Pseudocerastes fieldi Schmidt. Pseudocerastes fieldi ScumipT, 1930, Field Mus. Nat. Hist., Zool. Ser., 17:227: pl. 2, text fig. 2 (Bair Wells, Trans-Jordan). MarTERIAL EXAMINED (1): HUJ 3883, from Sakaka, near Jauf, collected in 1942, by Hardy. Remarks: This specimen extends the known range of this species, which includes Sinai, southern Israel, and southern Jordan, into northern Arabia. Genus Cerastes Wagler Cerastes cerastes (Linnaeus). Coluber cerastes LINNAEUS, 1758, Syst. Nat., 10th Ed., 1:217 (‘“‘Habitat in Oriente’’). Cerastes cerastes ANDERSON, 1899, Bih. Svenska Vet. Akad. Handl., 24:29. MATERIAL EXAMINED (23) : CAS 84503, 84505-84506, 84520-84521, 84527, 84531, 84555, 84560-84564, 84566, and CNHM 74005, from Dhahran, eol- lected in 1946, by Gasperetti; CAS 84481, 84490, 84495-84496, from Abgqaiq, collected in 1946, by Gasperetti; CAS 84424-84425, and CNHM 74006, from Shimal, collected in 1946, by Gasperetti; CAS 84440, from Abu Shaiba, collected in 1946, by Gasperetti. VoL. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 83 REMARKS: Five specimens in this series of 23 are horned. I find no relation between the development of horns and body size. None of the specimens have dark tails. Genus Echis Merrem Echis coloratus Giinther. Echis colorata GUNTHER, 1878, Proc. Zool. Soc. London, 1878:978 (Jebel Sharr, Midian). MATERIAL EXAMINED (1): HUJ 3862, from Satidi Arabia (exact lo- eality unknown), collected in 1944, by Waterston. CONCLUSION In this paper 48 species of reptiles and 3 species of amphibians are enumerated. With the exclusion of Jordan and the Sinai Peninsula, about 115 species and subspecies of reptiles and 8 species and subspecies of amphibians are presently known from Arabia. The number of endemic forms in this region is considerable; the genera Stenodactylus, Phryno- cephalus, Agama, Uromastyx, and Scincus include a great number of species and subspecies known only from Arabia. In the wider relations of the herpetological fauna, the affinities of the elements in western and southwestern Arabia with the fauna of northeastern Africa are striking. These affinities are exemplified, at the generic level, by T'ropiocolotes, Pris- turus, Philochortus, Latastia, and Rhynchocalamus; at the species level, these affinities are exemplified by Agama sinaita, Agama cyanogaster, Uro- mastyx aegyptius, Acanthodactylus boskianus, Eremias guttulata, Scincus hemprichi, Boaedon lineatus, Coluber rhodorhachis, Dasypeltis scabra, Telescopus dhara, Naja haje, and Bitis arietans. Many species range over the whole peninsula, but those mentioned above are restricted to the west and south coasts and their hinterlands, and are found in northeastern Africa as well. A similar correspondence exists between the south Iran- ian and Iraqi herpetofauna and that of eastern and southeastern Arabia. In this southeastern part of the peninsula there are many forms that do not reach western Arabia. To this group belong Aslsophylax tuberculatus, Henudactylus persicus, Agama persica, all of the Arabian species of the genera Phrynocephalus and Diplometopon, and the species Acantho- dactylus frasert and Eumeces taeniolatus. The number of species with these eastern affinities is smaller than that of the western list. Many species, however, range from Sind, in northwestern India, across southern 84 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Iran and Arabia, into Syria and Egypt. Examples are afforded by Eremus guttulata and Eremias brevirostris, Echis carinatus, and Varanus griseus. Human ageney ean have played no more than a minor role in such dispersals. The last comprehensive work on the amphibians and reptiles of Arabia was A Contribution to the Herpetology of Arabia, by John An- derson, published in 1896. Every subsequent collection from this area has yielded a harvest of species previously unknown. The time seems ripe for a renewed comprehensive work on the rich Arabian herpetofauna, the example for which has been set by Colonel Richard Meinertzhagen’s recent account of the birds. BIBLIOGRAPHY In addition to the special papers referred to in the text, the follow- ing bibliography includes citations to all of the literature bearing on the herpetology of Arabia which has appeared subsequent to the pub- lhieation of Anderson’s comprehensive work of 1896, and which has been available to me. ANDERSON, JOHN 1896. A contribution to the herpetology of Arabia, with a preliminary list of the reptiles and batrachians of Egypt. London (privately printed), li + 122 pp. 1898. Reptilia and Batrachia. In: Zoology of Egypt, vol. 1. London (Bernard Quaritch), 436 pp., 50 pls., 14 text-figs. 1901. A list of the reptiles and batrachians obtained by Mr. A. Blayney Percival in southern Arabia. Proceedings of the Zoological Society of London, 1901, pp. 137-152, 2 pls. ANGEL, FERNAND 1936. Reptiles et batraciens de Syrie et de Mesopotamie recoltés par M. P. Pallary. Bulletin de VInstitut égyptien (d’iigypte), T. 18, pp. 107-116. BOULENGER, GEORGE ALBERT 1885. Catalogue of the lizards in the British Museum (Natural History). Lon- don (Taylor & Francis), 2nd Ed., vol. 1, xii + 436 pp., 32 pls. FLOWER, STANLEY 1933. Notes on the recent reptiles and amphibians of Egypt, with a list of the species recorded from that kingdom. Proceedings of the Zoological Society of London, 1933, pp. 735-851, 1 map, 1 text-fig. Marx, HyMEN 1953. The elapid genus of snakes Walterinnesia. Fieldiana: Zoology, vol. 34, pp. 289-296, text-figs. 38—40. Vout. XXIX] HAAS: AMPHIBIANS AND REPTILES FROM ARABIA 85 MEINERTZHAGEN, RICHARD 1954. Birds of Arabia. Edinburgh (Oliver & Boyd), 624 pp. ParRKER, HAMPTON WILDMAN 1930. 1931. 1932. 1933. 1938. 1941. Three new reptiles from southern Arabia. Annals and Magazine of Nat- ural History, ser. 10, vol. 6, pp. 594-598. Some reptiles and amphibians from S. E. Arabia. Annals and Magazine of Natural History, ser. 10, vol. 8, pp. 514-522. Reptiles and amphibians. In: Bertram Thomas, Arabia Felix. New York (Scribners & Sons), pp. 341-344. Reptiles and amphibians. In: St. John B. Philby, The Empty Quarter. London. Appendix G, pp. 397-398. Reptiles and amphibians of the southern Hejaz. Annals and Magazine of Natural History, ser. 11, vol. 1, pp. 481-492. British Museum (Natural History!) Expedition to south-west Arabia, 1937-8. London, vol. 1, no. 2, pp. 3-6. ScuHMIDT, KARL PATTERSON 1930. 1933. 1939. 1941. 1952. 1953. 1955. Reptiles of Marshall Field’ North Arabian Desert Expedition, 1927-1928. Field Museum of Natural History, Zoology Series, vol. 17, pp. 221-230, pl. 2, text-figs 1-2. A new snake from Arabia. Field Museum of Natural History, Zoology Series, vol. 20, pp. 9-10. Reptiles and amphibians from southwestern Asia. Field Museum of Natural History, Zoology Series, vol. 24, pp. 49-92, 1 map. Reptiles and amphibians from central Arabia. Field Museum of Natural History, Zoology Series, vol. 24, pp. 161-165, 1 map, 1 text-fig. Diagnoses of new amphibians and reptiles from Iran. Natural History Miscellany, no. 93, pp. 1-2. Amphibians and reptiles from Yemen. Fieldiana: Zoology, vol. 34, pp. 253-261, 1 map. Amphibians and reptiles from Iran. Videnskabelige Meddelelser fra Densk Naturhistorisk Forening i Kjgbenhaven, volume 117, pp. 193-207, 2 text-figs. ScHmipt, Kart P., and Hymen Marx 1956. The herpetology of Sinai. Fieldiana: Zoology, vol. 39, no. 4, pp. 21-40, 1 map, 1 text-fig. SCORTECCI, GIUSEPPE 1932. 1933. Rettile dello Yemen. Atti della Societa italiana di scienze naturali del Museo civile di storia naturale, Milano, vol. 71, pp. 39-49, text-figs. Leptotyphlops yemenicus sp. n. Atti deila Societa italiana di scienze naturali del Museo civile di storia naturale, Milano, vol. 72, pp. 165— 166, text-fig. 36 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. SmitH, MatcoLtm ARTHUR 1935. Reptilia and Amphibia. Vol. II, Sauria. In: Fauna of British India. London (Taylor & Francis), xiii + 440 pp., 1 map, 1 pl., 94 text-figs. STEINDACHNER, FRANZ 1900. Expedition S.M. Schiff ‘Pola’ in das rothe Meer, nordliche und sidliche HAlfte. 1895/96 und 1897/98. Zoologische Ergebnisse. XVII. Bericht iiber die herpetologischen Aufsammlungen. Denkschriften der k. K. Akademie der Wissenschaften, Wien, (math.-naturw. Klasse), Bd. 69, S. 325-335, 2 Taf. 1903. Batrachier und Reptilien von Siidarabien und Sokotra, gesammelt wahrend der siidarabischen Expedition der kaiserlichen Akademie der Wissenschaften. Sitzungberichte der k. K. Akademie der Wissen- schaften, Wien, (math.-naturw. Klasse), Bd. 112, Abt. 1, S. 7-14. VESEY-FITZGERALD, DESMOND 1944. From Hasa to Oman by car. Geographical Review, vol. 41, pp. 544-560, 5 figs., map. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. XXIX, No. 4, pp. 87-146 November 5, 1957 FOSSIL BRYOZOA FROM THE PLEISTOCENE OF SOUTHERN CALIFORNIA* BY JOHN D. SOULE! and MARY MARSH DUFF? Allan Hancock Foundation, University of Southern California INTRODUCTION Except for the work of Gabb and Horn (1862), that of Canu and Bassler (1923), and the species listed in a paper by Clark (1931), little has been published with reference to the fossil Bryozoa of the Pacific coast of North America. The Bryozoa reported in this paper have been assembled from various sources. Through the courtesy of Mr. George P. Kanakoff we were able to examine fossil material collected by the Section of Invertebrate Paleon- tology of the Los Angeles County Museum from a number of sites. Addi- tional specimens were collected by the authors in San Pedro, California, and in Santa Barbara, California. The survey disclosed 56 species of Bryozoa of Pleistocene age. Of this number, 19 apparently are new to the fossil record, 10 have not been previously reported as part of the fossil fauna of the Pacific coast, and 4 are reported only as fossil and have not as yet been numbered among the living Bryozoa of the eastern Pa- cifie waters. For each species recorded we have established the following format: * Contribution no. 197, Allan Hancock Foundation, University of Southern California, Los An- geles, California. 1 Allan Hancock Foundation. 2 Pittsburgh, Pennsylvania. Formerly with the Hancock Foundation [ 87 ] 88 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. (1) The name of the species, author, date. (2) A limited synonymy, re- stricted to the original name, followed by previous eastern Pacific citations. (3) Brief description, and where applicable, measurements. (4) Locality or localities from which examined specimens were collected. (5) Oceur- rence, fossil and Recent records gathered from as many sources as were available. (6) Bathymetric range when available, including the number of records obtained. The age assignments are those of the original authors. For additional data, excellent descriptions, and illustrations, the monu- mental work of Osburn, Bryozoa of the Pacific Coast of America, 1950, 1952, and 1953, should be consulted. ACKNOWLEDGMENTS The authors wish to acknowledge their deep debt of gratitude to the late Dr. Raymond C. Osburn for his aid and encouragement during the preparation of this paper, and for the time and effort he so willingly spent in verifying the identification of our specimens. Also we express our appre- ciation to Dr. William K. Emerson of the American Museum of Natural History for many valuable suggestions during the preparation of the manuscript. Grateful acknowledgment is made to Dr. William Easton, Department of Geology, University of Southern California, for his help in initiating this study. PHYLUM BRYOZOA CLass ECTOPROCTA Order GYMNOLAEMATA Sub-order CHEILOSTOMATA ANASCA Levinsen, 1909 DIVISION I, INOVICELLATA Jullien, 1888 Within this division we have no fossil representatives to report. DIVISION II, MALACOSTEGA Levinsen, 1909 Family MEMBRANIPORIDAE Busk, 1854 Genus Membranipora Blainville, 1830 Membranipora tuberculata (Bosc), 1802. Flustra tuberculata Bosc, 1802, partim, Hist. Nat. Vers., vol. 3, no. 1, p. 118. VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 89 Flustra tehuelcha bv’OrBigNy, 1847, Voy. Amer. Mer., vol. 5, pt. 4, p. 17, pl. 8, figs. 10-14. Membranipora tehuelcha, Ropertrson. 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, p. 265, pl. 15, figs. 16, 17, pl. 16, fig. 18. Membranipora tuberculata, CANU and BAsSsLerR, 1923, U. S. Nat. Mus. Bull. 125, pp. 22, 23, pl. 33, figs. 3-5. Nichtina tuberculata, Hastrines, 19380, Proc. Zool. Soc. London, 1929, p. 706, pl. 3, figs. 9, 10. Mamobranipora tuberculata, OspurN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 23, 24, pl. 2, figs. 4-6. DescripTION: Zoaria incrusting. Zooecia variable in proportions, large, elongate, alternate, surrounded by a calcareous border with 4-6 denticles. Opesia occupying about two-thirds of the front. Cryptoeyst bearing a pair of thick rounded ecaleareous tubercules, that may fuse in the mid-line to form a plate. Locauities: LACMIP (Los Angeles County Museum, Invertebrate Paleontology) loe. 66-2, Newport Harbor Mesa, Newport, California, Pleistocene. OCCURRENCE: Pleistocene: Santa Monica, California [Rustic Canyon] (Canu and Bassler, 1923). Pliocene: Red Crag of Walton-on-the-Naze, England (Kendall, 1931). Recent: World wide in warm waters of both Atlantic and Pacifie oceans. DeptH: Shallow water. Intertidal to 108 fathoms reported in the literature. Membranipora tenuis Desor, 1848. Membranipora tenuis Dresor, 1848, Proc. Boston Soc. Nat. Hist., vol. 3, p. 66. Membranipora denticulata Busk, 1856, Quart. Jour. Micro. Sci., vol. 4, p. 176, pl. 7, figs. 1, 2. Hemiseptella grandicella CANU and BASsLER, 1923, U. S. Nat. Mus. Bull. no. 125, 105 Wal, pol, By sake aA Membranipora tenuis, OSBURN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 26, 27, pl. 2, figs. 9, 10. Description: Zoaria inerusting. Zooecia elongate, quadrangular, dis- tinet, separated by a narrow groove in which may be a thin brown line. Mural rim distinctly tubereulated. Cryptocyst depressed, finely tubereu- lated. Free border of the eryptoeyst bearing spinules that project into the oval opesia. The spinules straight, curved, or branched. Opesia occupying most of the frontal area. MEASUREMENTS: Opesia 368/195 mp. Zooecia 769/333 muy. 90 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. Locauities: LACMIP loe. 86-b, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. West Second and Pacific Avenue, San Pedro, Cali- fornia, Pleistocene. OccURRENCE: Lower Miocene: Bowden Marl, Bowden Jamaica (Canu and Bassler, 1923). Recent: Pacific: southern California, Mexico, Costa Riea (Haneoek collections). Atlantic: Denmark (Levinsen, 1894); Massa- chusetts (Desor, 1848, Osburn, 1912); North Carolina (Canu and Bassler, 1928); Florida (Smitt, 1873); Puerto Rico (Osburn, 1940); Brazil (Mar- eus, 1937); Tunis (Canu and Bassler, 1930); Netherlands (Lacourt, 1949). DereptH: Shore (intertidal) to 36 fathoms. Based on 17 records. Remarks: Not previously known as fossil from the Pacifie coast. Membranipora savarti (Audouin), 1826. Flustra savarti Aupovuin, 1826, in Savigny, J. C., Descrip. l’Egypte, Hist. Nat., vol. 1, pt. 4, p. 240, pl. 10, figs. lla, b. Acanthodesia savartii, CANU and BASssLER, 1920, U. S. Nat. Mus. Bull. no. 106, p. 100, pl. 21, figs. 2-4. Acanthodesia savartii, CANU and BASSLER, 1923, U. S. Nat. Mus. Bull. no. 125, pp. 30, 31, pl. 11, figs. 1-9, pl. 2, figs. 2, 3, pl. 5, figs. 1-5, pl. 46, figs. 8, 9. Acanthodesia savartii, CANuU and Basser, 1930, Proc. U. S. Nat. Mus., no. 2810, vol. 76, art. 13, p. 4. Membranipora savarti, OsBuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 27, 28, pl. 2, fig. 7. DescrIPTION: Zoaria incrusting. Zooecia moderately large, with short denticle projecting into the opening from the center of the basal region. Ooecia and avicularia wanting. MEASUREMENTS: Opesia, 380/173 mp. Zooecia 518/300 mz. Locauitres: LACMIP loc. 66-2. Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OccuRRENCE: Pleistocene: Florida (Canu and Bassler, 1923). Plo- cene: Red Crag of Walton-on-the-Naze, London, England (Kendall, 1931). Miocene: Australia (Canu and Bassler, 1920); Virginia (Canu and Bass- ler, 1923); Torino, Italy (Scotti, 1936); New Jersey (Richards, 1942). Recent: Florida, Tortugas Island, Gulf of Mexico, Puerto Rico (Osburn, 1940); Venezuela (Oburn, 1947); Sulu Sea, Celebes Sea, Zamboanga, P. L., Indian Ocean, Zanzibar, Red Sea, Ceylon, Galapagos (Canu and Bassler, 1929); Brazil (Marcus, 1937); Philippine Islands (Harmer, 1926); Suez Canal (Hastings, 1927); Great Barrier Reef, Australia (Livingston, 1927) ; Japan, Java, New Guinea, Singapore, Malay Peninsula (Okada and Mata- VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 91 wari, 1938); San Diego, California, Panama, Baja California, Costa Riea, Galapagos, Colombia, Mexico (Hancock stations). DeptH: 5 to 75 fathoms (shallow water characteristically). Remarks: Not previously known as fossil on the Pacifie coast. Genus Conopeum Gray, 1848 Conopeum commensale Kirkpatrick and Metzelaar, 1922. Conopeum commensale KIRKPATRICK and MrtzeLAAr, 1922, Proc. Zool. Soc. London, p. 985, pl. 1, figs 1, 4-7, 9. Conopeum commensale, OspuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 30, 31, pl. 2, figs. 12-15. DeEscripTION: Zoaria incrusting, multilaminar. Zooecia distinct, oval, alternate. Zooecial margins, in protected portions of a zoarium, granular, serrated along the opesial border. Proximal to each zooecium are two low tubercules, often fused so as to appear as a single prominence. No avicu- laria or ovicells. MEASUREMENTS: Opesia, 403/208 mp. Zooecia, 530/299 mp. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. LACMIP loc. 136, Newport Bay Road, Newport, Cali- fornia, Pleistocene. OccuRRENCE: Recent: Atlantic: Brazil (Mareus, 1937, 1938, 1939); Argentina (Kirkpatrick and Metzelaar, 1922). Pacific: Baja California, Mexico, Guatemala, Costa Rica, Panama, Ecuador, Peru (Haneock sta- tions). DeptH: Shore collecting (intertidal) to 130 fathoms (21 records). ReMARKS: Not previously listed as fossil. Genus Antropora Norman, 1903 Antropora tincta (Hastings), 1930. Crassimarginatella tincta Hastines, 1930, Proc. Zool. Soc. London, 1929, pp. 708, 709, pl. 5, figs. 16a, p. 117, fig. 120. ‘‘Wembranipora ? denticulata (old)’’ B.M. 99.7 1.1048 Busk (fide Hastings). Membranipora lacroizi RoBertson, 1908, partim, Univ. Calif. Publ. Zool. vol. 4, no. 5, pp. 261, 262, pl. 14, fig. 5. Antropora tincta, OsBuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, p. 54, TO, 4h, male, (5 105 UA, aie 0G te 92 CALIFORNIA ACADEMY OF SCIENCES | Proc. 4TH SER. Description: Zoaria inecrusting. Zooecia with a narrow tuberculate eryptoeyst. Around each of the zooecia are 5 to 6 small triangular open- ings. Our fossil specimens give evidence of a few avicularia scattered among the zooecia. MEASUREMENTS: Opesia, 242/162 mp. Zooecia, 368/230 mu. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OccuRRENCE: Recent: Panama Canal, Galapagos Islands (Hastings, 1930); Mazatlan, Mexico (Busk 1856, fide Hastings); California, Gulf of California, Baja California, Mexico, Guatemala, Costa Rica, Panama, Eeua- dor, Peru, Galapagos Islands (Hancock stations). DeptH: Shore (intertidal) to 103 fathoms (51 records). Remarks: Not previously found as fossil. Genus Cauloramphus Norman, 1905 Cauloramphus spiniferum (Johnston), 1838. Flustra spinifera Jounston, 1838, Trans. Nat. Hist. Soc. Northumb. Durk. Newcle., vol. 2, p. 266, pl. 9, fig. 6. Membranipora spinifera, ROBERTSON, 1900, Proc. Wash. Acad. Sci., vol. 2, p. 324. Membranipora spinifera, Ropertson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, p. 265, pl. 15, fig. 15. Schizoporella spinifera, CANU and BAsster, 1920, U. S. Nat. Mus. Bull. 106, pp. DoOleOO Ss Cauloramphus porosus CANU and BaAsster, 1923, U. S. Nat. Mus. Bull. 125, p. 48, Die sb sstigs, 17. Membranipora spinifera, O’DoNoGHUE and O’DonocHur, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 168. Cauloramphus spinifer, O'DONOGHUE and O’DoNOGHUE, 1925, Univ. Wash. Pub. Puget Snd. Biol. Sta. Trans., vol. 5, pp. 97, 98. Cauloramphus spinifer, O’DoNoGHUE and O’DonoGHuE, 1926, Contr. Canad. Biol., n. S., Vol. 3, NO.70, Ds SD: Cauloramphus spiniferum, Hastines, 1930, Proc. Zool. Soc. London, 1929, p. 713. Cauloramphus spiniferum, OsBuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, DD. 55, 56, pla 5, ties 9: DEscRIPTION: Zoaria inerusting. Zooecia oval, alternate, margin wide, raised and roughened by minute tubercles. Zooecia having a variable number of spines in the recent Pacific material, 10-12, that have been eroded in the fossil material. This left shallow sockets which are considered to be “pores” by Canu and Bassler (1923). Recent species showing avicu- VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 93 laria tapering to small bases adjacent to the larger spines; thus the ‘“‘pores” seen on the fossil specimens are not just the bases of avicularia, but also of spines. Aperture occupying entire frontal area. MEASUREMENTS: Opesia, 621/264 mp. Zooecia 747/368 muy. Locauities: LACMIP loe. 130-7, opposite Berth 79, Los Angeles har- bor, Timm’s Point formation, Pleistocene. LACMIP loc. 66-2, Newport Harbor mesa, Newport, California, Pleistocene. Bath-house Beach, Santa Barbara, California, Pleistocene. W. Second and Pacific Avenue, San Pedro, California, Pleistocene. OCCURRENCE: Pleistocene: Santa Barbara, California (Canu and Bass- ler, 1923). Recent: Pacific: Alaska (Robertson, 1902); British Columbia (O’Donoghue and O'Donoghue, 1923, 1925, 1926); California, Mexico, Peru, Galapagos Islands (Hancock Foundation station records); Galapagos Islands (Hastings, 1930). Atlantic: Arctic, Greenland, Sweden (Borg, 1930, 1933); E. Finmark (Norman, 1903); England (Johnston, 1838, 1847; Alder, 1857; Hineks, 1862, 1880; Bassindale, 1941, 1943; Mar. Biol. Assoc., 1931) DeptH: Shore (intertidal) to 230 fathoms, based on 35 records. Family ALDERINIDAE Canu and Bassler, 1927 Genus Callopora Gray, 1848 Callopora circumclathrata (Hincks), 1881. Membranipora circumclathrata Hincxks, 1881, Ann. Mag. Nat. Hist., ser. 5, vol. 8, Ty JUHI, By sites ale Membranipora circumclathrata, Roperrson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 259, 260, pl. 14, figs. 1, 2. Membranipora circumclathrata, O’DONOGHUE and O’DONOGHUE, 1923, Contr. Canad. Biol., n. s., vol. 1, p. 166. Callopora circumclathrata, CANuU and BASsSLER, 1923, U. S. Nat. Mus. Bull. 125, p. 43, pl. 34, figs. 1-3. Callopora circumclathrata, O’DoNOGHUE and O’DoNnoGHuUE, 1925, Univ. Wash. Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 97. Callopora circumclathrata, O’DoONOGHUE and O’DoNnoGHUE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, p. 97. Callopora circumclathrata, OsBuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, jo, Hay, GOs pollk 75 sales ale DESCRIPTION: Zoaria inecrusting, due to erosion, lacking the spines prominent on the recent material. Zooecia elongate, oval, distinct. Mural rim of well preserved zooecia finely beaded, granular. Attachment points 94 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. of spines, both lateral and distal, indicated by small pits. Proximal avicu- laria abundant, variable in size. Few ovicells intact, smooth, raised, rounded. MEASUREMENTS: Opesia 276/185 mp. Zooecia 518/299 mu. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OccuRRENCE: Pleistocene: Santa Monica, California (Canu and Bass- ler, 1923). Recent: Pacific: British Columbia, Puget Sound region (O’Don- oghue 1923, 1925, 1926); California (Robertson, 1908, Hancock station ree- ords); Galapagos Islands (Hancock stations). DeptH: Shore (intertidal) to 131 fathoms (based on 11 records). Callopora corniculifera (Hincks), 1882. Membranipora corniculifera Hincxs, 1882, Ann. Mag. Nat. Hist., ser. 5, vol. 10, p. 468, pl. 20, figs. 4, 4a. Hincksina (Membranipora) corniculifera, CANU and BaAssteEr, 1920, U. S. Nat. Mus. Bull. 106, p. 112. Cauloramphus triangularis CaANu and BAssuter, 1923, U. S. Nat. Mus. Bull. 125, pp. 48, 49, pl. 33, figs. 14-16. Callopora corniculifera Ospurn, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, Dp. 166;.6%, pl. 7, fe. 1 DESCRIPTION: Zoaria inerusting. Zooecia distinct, large opesia ovoid, with narrow distal end. Sears of 6—8 spines on the opesial rim. Avicularia small, located on the mural rim of the distal half of the zooecia. Ovicells large, globose, some with a varied process immediately above the aperture. MEASUREMENTS: Zooecia 640/410 muy. Locauities: LACMIP loc. 130-7, opposite Berth 79, Los Angeles har- bor, Timm’s Point formation, Pleistocene. OCCURRENCE: Pleistocene: Santa Barbara, California (Canu and Bass- ler, 1923). Recent: British Columbia (Hincks, 1882); California (Osburn, 1950). DeptH: Two records (Hancock) 6 and 38 fathoms respectively. Hincks gave no depth data on the British Columbia specimens. Genus Copidozoum Harmer, 1926 Copidozoum tenuirostre (Hincks), 1880. Membranipora tenuirostris Hincxks, 1880, Ann. Mag. Nat. Hist., ser. 5, vol. 6, pp. (Os Te plo} fies 3: Vou. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 95 Membranipora tenuirostris, HincKs, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 10, p. 465. Callopora tenuirostris, CANU and BAssLErR, 1920, U. S. Nat. Mus. Bull. 106, p. 146, figs. F, G, pp. 147, 154, pl. 29, figs. 10, 11. Callopora tenuirostris, O’DoNoGHUE and O’DonoGHutr, 1926, Contr. Canad. Biol. Fish., vol. 3, no. 3, p. 79, pl. 3, fig. 24. Callopora tenuirostris, CANU and Basser, 1930, Proc. U. S. Nat. Mus., no. 2810, Wolk, 7h; era al}, To, 24 os Gy OE Copidozoum tenwirostre, OsBuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, 105 7s 1, te aiksy 2S DeEscrRIPTION : Zoaria inerusting. Zooecia oval, broader below than above, with wide border. Few small spines (spinules) projecting into orifice from the rim. Avicularia found in interzooecial spaces, irregularly spaced throughout the zoarium. MEASUREMENTS: Opesia 345/218 mp. Zooecia 437/276 muy. LocauitiEs: Bath-house Beach, Santa Barbara, California, Pleistocene. OCCURRENCE: Miocene: Touraine (Canu, fide Canu and Bassler, 1920). Eocene: Mississippi (Canu and Bassler, 1920). Recent: Pacific: British Columbia, Queen Charlotte Islands (Hincks, 1884, O’Donoguhe, 1926) ; Galapagos Islands (Canu and Bassler, 1930) ; California, Gulf of California, Channel Islands, Baja California, Mexico, Costa Rica, Panama, Colombia (Haneock stations, Osburn, 1950); Galapagos Islands (Hancock stations). Atlantic: Brazil (Marcus, 1937); Florida (Canu and Bassler, 1928); Gulf of Mexico (Canu and Bassler, 1928); Cape Verde Islands (Waters, 1918) ; Mediterranean (Waters, 1879, 1885); Madeira (Hincks, 1880). DeptH: Shore (intertidal) to 164 fathoms (37 records). ReMARKS: Not previously recorded as fossil on the Pacific coast of North America. Genus Tegella Levinsen, 1909 Tegella robertsonae (O’Donoghue and O’Donoghue), 1926. Membranipora unicornis, ROBERTSON, 1900, Proc. Wash. Acad. Sci., vol. 2, p. 324. Membranipora occultata Ropertson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 262, 263, pl. 14, figs. 6-9. (non) Membranipora occultata WATERS, 1887, Quart. Jour. Geol. Soc. London, vol. 43, p. 48, pl. 6, figs 12, 12a, pl. 8, fig. 40. Tegella (Membranipora) occuitata, CANU and BASSLER, 1920, U. S. Nat. Mus. Bull. 106, p. 166. Membranipora occultata, O'DONOGHUE and O’DoNOGHUE, 1923, Contr. Canad. Biol., nas volede no: 10s pe L6r. Tegella robertsoni O’DoNoGHUE and O’DonoGHUE, 1926, Contr. Canad. Biol. Fish., ie ih WO By 10), BS fob te 96 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Tegella robertsonae, Ospurn, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 81, 82, pl. 9, fig. 5. Description: Zoaria incrusting. Zooecia alternate, with large apertures occupying most of the frontal area. Inner rim of the margin finely notched. Avicularia large, frequent, raised, pointed obliquely forward. Occasional smaller broad avicularia may be found. Ovicells small, globose, may be sur- mounted by avicularia. MEASUREMENTS: Aperture, 125/162 mp. Zooecia, 690/483 muy. Locauities: LACMIP loc. 76, “lumber yard” San Pedro, California, eoll. Dr. A. T. Tieje. Pleistocene. Bath-house Beach, Santa Barbara, Calli- fornia, Pleistocene. OCCURRENCE: Pleistocene (early): Japan, Dizodo Beds, Boso Penin- sula (Sakakura, 1935). Recent: Pacific: Alaska (Robertson, 1900, 1908) ; British Columbia, (coll. E. F. Ricketts in Hancock collection) ; Vancouver Island (O’Donoghue and O’Donoghue, 19238, 1926). DeptH: 6 to 12 fathoms (one record). Remarks: Not previously recorded as fossil from the Pacific Coast of North America. Genus Chapperia Willey, 1900 Chapperia patula (Hincks), 1881. Membranipora patula Hincks, 1881, Ann. Mag. Nat. Hist., ser. 5, vol. 7, p. 150, pl. 9, fig. 4. Membranipora patula, Hincks, 1882, Ann. Mag. Nat. Hist., ser. 5, vol. 10, p. 465. Membranipora patula, Ropertson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, p. 263, pl. 15,+fig. 10. Membranipora patula, O’DoNoGHUr and O’DoNOGHUE, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 167. Chaperia galeata CANU and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, pp. 52-54, pl. 34, figs. 9, 10, but not 8. Amphiblestrum patulum, O’DonocHUrE and O’DonoGHuE, 1925, Univ. Wash. Pub. Puget Snd. Biol. Sta. Trans., vol. 5, p. 97. Amphlestrum patulum, O’DonoGHUE and O’DonoGHuE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, pp. 83, 84. Chapperia patula, OsBURN, 1950, Allan Hancock Pac. Expds. vol. 14, no. 1, p. 89, pl. 10, figs. 1, 2. Description: Zoaria inerusting. Zooecia distinct, large, broad, and short. Raised distal rim bearing the sears of 4-6 spines. Avicularia small, triangular, numerous, located in front of the spines on the border of the distal rim. Ovicells large, globular, smooth. VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 97 MEASUREMENTS: Opesia, 299/230 mp. Zooecia 690/356 mu. Locauities: Bath-house Beach, Santa Barbara, California, Pleistocene. OCCURRENCE: Pleistocene: Santa Barbara, Santa Moniea, California (Canu and Bassler, 1923) Pliocene: New Zealand (Waters, 1887); Italy (Manzoni, 1875). Miocene: Australia (Waters, 1882); Italy (Sequenza, 1882). Recent: Pacific: Queen Charlotte Islands (Hincks, 1882); Califor- nia (Hineks, 1881, Robertson, 1908); Vancouver Island region (O’Don- oghue and O’Donoghue, 1923, 1926); Puget Sound, Washington (O’Don- oghue and O’Donoghue, 1925); California, Mexico, Baja California, Ga- lapagos Islands (Hancock stations). DertH: Shore (intertidal) to 252 fathoms (44 records). DIVISION III, COILOSTEGA Levinsen, 1909 Family Microporipar Hincks, 1880 Genus Microporina Levinsen, 1909 Microporina borealis (Busk), 1855. Salicornaria borealis Busk, 1855, Quart. Jour. Micro. Sci., vol. 3, p. 254, pl. 1, figs. 1-3. Cellaria borealis, H1incks, 1882, Ann. Mag. Nat. Hist., ser. 5, vol. 10, p. 463. Cellaria borealis, RopertTson, 1900, Proc. Wash. Acad. Sci., vol. 2, p. 322. Cellaria borealis, RopertTsoNn, 1905, Univ. Calif. Pub. Zool., vol. 2, no. 5, pp. 287, 288, pl. 16, fig. 102. Cellaria borealis, O’DoNoGHUE and O’DoNoGHuUr, 1923, Contr. Canad. Biol., n. s., vol. l. no; 105 p: 1165. Microporina borealis, CANU and BaAsster, 1923, U. S. Nat. Mus. Bull. 125, p. 84. Microporina (Salicornaria) borealis, O'DONOGHUE and O’DoNOoGHUE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, p. 95. Microporina borealis, OsBuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, p. 106, pl. 11, fig. 2. DESCRIPTION: Zoaria erect. Zooecia elongate, alternate, distinet with lateral walls raised. Aperture rounded distally with a pair of teeth pro- jecting upward on the proximal rim. On some zooecia, avicularia occu- pying the area just distal to the aperture. No avicularian mandible pre- served. MEASUREMENTS: Aperture, 103/161 mp. Zooecia 690/264 muy. Locauities: LACMIP loc. 68—B, Newport Harbor mesa, lower north, Newport, California, Pleistocene. LACMIP loc. 130-7, opposite Berth 79, Los Angeles harbor, Timm’s Point formation, Pleistocene. 98 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 47TH SER. OccuRRENCE: Recent: Pacific: Alaska (Robertson, 1900, 1905); Queen Charlotte Islands, British Columbia (Hincks, 1882); Vancouver Island re- gion (O’Donoghue and O'Donoghue, 1923, 1926); Japan (Canu and Bassler, 1929, Sakakura, 1936, Silén, 1942); Alaska (Osburn, 1950). Atlantic: Greenland (Busk, 1855, Osburn, 1936); Spitzbergen (Nordgaard, 1917); Hudson Bay (Osburn, 1932). Depth: 6 to 69 fathoms (5 records). Remarks: This appears to be the first occurrence of this species in the fossil record. Genus Labioporella Harmer, 1926 Labioporella sinuosa Osburn, 1940. Labioporella sinuosa OsBuRN, 1940, N. Y. Acad. Sci., Sci. Sur. Porto Rico and Vir- gin Islands, vol. 16, pt. 3, p. 377, pl. 5, figs. 40, 41. Labioporella sinuosa, OspurN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 109, 110, pl. 11, fig. 12. Description: Zoaria incrusting. Zooecia regular, mural rim raised, beaded. Orifice large, arched anteriorly, almost straight across posteriorly. No avicularia. No ovicells. MEASUREMENTS: Aperture, 92/172 mp. Zooecia 494/195 muy. Locairtes: West Second and Pacific Avenue, San Pedro, California, Pleistocene. OccuRRENCE: Recent: Pacific: Gulf of California, Baja California, Mex- ico, Panama, Ecuador (Osburn, 1950). Atlantic: Puerto Rico, Virgin Islands (Osburn, 1940). DeptH: Shore (intertidal) to 55 fathoms (4 records). REMARKS: This species has not been previously recorded in the fossil record. Family THALAMOPORELLIDAE Levinsen, 1909 Genus Thalamoporella Hincks, 1887 Thalamoporella californica (Levinsen), 1909. (Non) Membranipora gothica Busk, 1856, Quart. Jour. Micro. Sci., vol. 4, p. 176, pl. 7, figs. 5-7. Steganoporella Rozieri form gothica H1ncks, 1880, Ann. Mag. Nat. Hist., ser. 5, vol. (6; p: 380) pl. 16; fizs93: VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 99 Thalamoporella rozieri, ROBERTSON, 1908, Univ. Calif. Publ. Zool., vol. 4, no. 5, pp. 277-279, pl. 17, figs. 27-29, pl. 18, fig. 30. Thalamoporella Rozieri var. E. (californica) LEVINSEN, 1909, Cheilo. Bryozoa, p. 184, pl. 66, figs. 2a-d, Copenhagen. Thalamoporella californica, HAstTinGs, 1930, Proc. Zool. Soc. Lond., 1929, no. 42, p. 716, pl. 10, figs. 47-53, pl. 11, figs. 56, 57. Thalamoporella californica, OSBURN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 111, 112, pl. 12, fig. 2. Description: Zoaria large, incrusting or erect. Zooecia oblong, quad- rangular, alternate. Margin broad, separating zooecial rows, and frequently rising anteriorly on each side of the orifice into blunt processes which may be continued in an are around the oral extremity. Orifice prominent, occu- pying about one-third of the front. Seattered avicularia, large, some occu- pying the place of a zooecium in the colony. Ovicells, when present, large, smooth, globose, with a bilobate appearance. MEASUREMENTS: Aperture, 118/69 mp. Zooecia 586/483 my. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OCCURRENCE: Recent: Pacific: California (Robertson, 1908, Osburn, 1950) ; Galapagos Islands (Hastings, 1930, Osburn, 1950); Baja California, Mexico, Costa Rica, Panama (Osburn, 1950, and Haneock stations). DeptH: Shore (intertidal) to 80 fathoms. Remarks: This species is new to the fossil record. Family LUNULARIDAE Levinsen, 1909 Genus Discoporella d’Orbigny, 1852 Discoporella umbellata (Defrance), 1825. Lunulites umbellata DEFRANCE, 1823, Dict. Sci. Nat., vol. 27, p. 361. Cupularia canariensis, Ropertson, 1908, Univ. Calif. Publ. Zool., vol. 4, no. 5, p. 314, pl. 24, figs. 90, 91. Cupularia umbellata, CANu- and BASsLER, 1923, U. S. Nat. Mus. Bull. 125, pp. 80-82, pl. 2, figs. 15-19. Cupularia robertsoniae, CANU and BASsLER, 1923, U. S. Nat. Mus. Bull. 125, p. 82, pl. 34, figs. 5-7. Discoporella umbellata, Hastines, 1930, Proc. Zool. Soc. Lond., 1929, no. 42, pp. 7H, GIS), ole alals ake Gye Cupularia umbellata, CANuU and Basser, 1930, Proc. U. S. Nat. Mus., no. 2810, vol. WG. ene, ies aoe Jak ay 100 CALIFORNIA ACADEMY OF SCIENCES | Proc. 4rH SER. Discoporella umbellata, OspuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 113, 114, pl. 11, figs. 7-10 DESCRIPTION: Zoaria incrusting, flattened, wider than high, shaped like an umbrella or coolie-hat. Zooecia spirally arranged, elongate, distinct. MEASUREMENTS: Opesia, 241/184 mp. Zooecia, 368/240 mp. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OCCURRENCE: Pleistocene: Santa Monica, California (Canu and Bass- ler, 1923). Pliocene: South Carolin, Florida (Canu and Bassler, 1923); Italy (Waters, 1921). Miocene: Florida, South Carolina, North Carolina, Costa Rica, Jamaica, Santo Domingo (Canu and Bassler, 1923); Franee, Austria, Hungary (Waters, 1921); Italy (Scotti, 1936); New Jersey, South Carolina (Gabb and Horn, 1862). Oligocene: France (Waters, 1921). Recent: Pacific: California, northern Channel Islands to Eeuador, Galapagos, Colombia, Panama, Costa Rica, Mexico, Gulf of California (Os- burn, 1950); Galapagos (Canu and Bassler, 1930, Hastings, 1930); Colom- bia (Hastings, 1930); Guatemala, Baja California (Haneock stations). At- lantic: Florida, North Carolina (Smitt, 1873, Osburn, 1914); Panama, Tortuga, Aruba (Osburn, 1947); Gulf of Mexico (Canu and Bassler, 1928) ; Brazil (Silén, 1942); Canary Islands (Calvet, 1907); Spain (Norman, 1909): DeptH: Shore (intertidal) to 160 fathoms (187 records). DIVISION IV, PSEUDOSTEGA Levinsen, 1909 Family CELLARIDAE Hincks, 1880 Genus Cellaria Solander, 1786 Cellaria mandibulata Hincks, 1882. Cellaria mandibulata Hincks, 1882, Ann. Mag. Nat. Hist., ser. 5, vol. 10, pp. 463, 464. Cellaria mandibulata, Hincxs, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 18, p. 209, oe Ge gies We Cellaria mandibulata, Ropertson, 1905, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 288, 289, pl. 15, figs. 87-89, pl. 16, fig. 103. Cellaria mandibulata, CANU and BAssurErR, 1923, U. S. Nat. Mus. Bull. 125, p. 86, pl. 34, figs. 11-14. Cellaria mandibulata, O’DoNnoGHUE and O’DoNnoGHurE, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 165. Cellaria mandibulata, OsBuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, DD 6) elena e hi ole VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 101 DESCRIPTION: Zoaria of the fossil material only fragments of the masses of irregular branches such as are found in the specimens of this species living in present waters. Zooecia large, elongate, narrowed at the proximal end, situated within short slender internodes. Frontal raised, convex, minutely porous. Avicularia long, resembling most an extra large zooecium. Ooecia, when present, circular or oblong, occupying the space upon each zooecia above the orifice. MEASUREMENTS: Aperture, 103/138 mp. Zooecia 598/276 mp. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, south location, Newport, California, Plestiocene. Bath-house Beach, Santa Barbara, Cali- fornia, Pleistocene. Packard’s Hill, Santa Barbara, California, Pleistocene. West Second and Pacifie Avenue, San Pedro, California, Pleistocene. OccURRENCE: Pleistocene: Los Angeles, California (Canu and Bassler, 1923). Recent: Pacific: Queen Charlotte Islands (Hincks, 1882, 1884), Vancouver Island region (O’Donoghue and O'Donoghue, 1923); California (Robertson, 1905, Osburn, 1950); Mexico (Osburn, 1950); Channel Islands (Haneoek stations). Atlantic: Aruba Island Caribbean Sea (Osburn, 1947). DerptH: Shore (intertidal) to 131 fathoms (75 records). Cellaria diffusa Robertson, 1905. Cellaria diffusa RoBEeRTSON, 1905, Univ. Calif. Pub. Zool., vol. 2, no. 5, pp. 289, 290, pl. 15, fig. 90, pl. 16, fig. 104. Cellaria diffusa, CANuU and Basser, 1923, U. S. Nat. Mus. Bull. 125, p. 86, pl. 34, figs. 19, 20. Cellaria fissurifera CANU and BASsLeER, 1923, U. S. Nat. Mus. Bull. 125, pp. 85, 86, pl. 34, figs. 15-18. Cellaria diffusa, O’DONOGHUE and O’DoNoGHUE, 1923, Contr. Canad. Biol., n. s., Volos enon l0. ps 165: Cellularia diffusa, O‘DONoGHUE and O’DonoGHur, 1925, Univ. Wash. Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 100. Cellularia diffusa, O’DoNoGHUE and O’DonoGHuE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, p. 96. Cellaria diffusa, OsBuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 117, 118, pl. 12, fig. 9. DeEscrIPTION: Zoaria consisting of fragments of elongate cylindrical internodes, connected by joints. Internode variable in length. Zooecia depressed, narrower below and above than in the middle, truncated at each end. Avicularia occurring in place of zooecia, small, almost square. MEASUREMENTS: Aperture, 80/103 mp. Zooecia, 540/312 mp. Localities: West Second and Pacific Avenue, San Pedro, California, 102 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Pleistocene. LACMIP loc. 64, Hill-top quarry, Pleistocene. LACMIP loe. 130-7, opposite Berth 79, Los Angeles harbor, Timm’s Point formation, Pleistocene. OccURRENCE: Pleistocene: Santa Monica [Rustic Canyon], California (Canu and Bassler, 1923); Los Angeles, California (Canu and Bassler, 1923.) Recent: California, Channel Islands, Baja California, Galapagos, Mexico (Osburn, 1950, and Hancock station records); California (Robert- son, 1905); Puget Sound (Robertson, 1905, O’Donoghue and O’Donoghue, 1925); Vancouver Island (O’Donoghue and O’Donoghue, 1923, 1926). DeptH: Shore (intertidal) to 118 fathoms (27 records). DIVISION V, CELLULARINA Smitt, 1867 Family ScRUPOCELLARIIDAE Levinsen, 1909 Genus Amastigia Busk, 1852 Amastigia rudis (Busk), 1852. Caberea rudis BuskK, 1852, Cat. Mar. Polyzoa, Brit. Mus., Cheilo. pt. 1, p. 38, pl. 46, figs. 1-3. Amastigia rudis, OsBuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, p. 127, pl. 16, figs. 3-5. DESCRIPTION: Zoaria erect, chitinous, multiserial. Zooecia crowded, ovoid. Avicularia of two types; frontal, small, triangular, dorsal, elongated, vibracula-like, pointing proximally. Marginal zooecia may have greatly enlarged frontal avicularia, gigantic, pointing proximally in contrast to the other smaller avicularia directed distally. Ovicells prominent, low, globose, with a heavily chitinized distal area well demarked from the proximal. MEASUREMENTS: Zooecia range between 420 mp and 490 my. Locauities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. OCCURRENCE: Recent: Pacific: Australia (Busk, 1852, MacGillivray, 1887, Haswell, 1880, Hincks, 1884); Netherlands East Indies (Harmer, 1926); Japan (Ortmann, 1889. Yanagi and Okada, 1918, Okada, 1934, Sakakura, 1935; Okada and Mawatari, 1935, Okada and Mawatari, 1938, Silén, 1941); California, Channel Islands, Mexico, Costa Rica (Osburn, 1950). DeptH: 2 to 49 fathoms (9 records). REMARKS: This appears to be the first fossil record of this species. Vou. XXIX ] SOULE AND DUFF: FOSSIL BRYOZOA 103 Genus Scrupocellaria van Beneden, 1845 Scrupocellaria regularis Osburn, 1940. (Non) Scrupocellaria cervicornis Busk, 1852, Cat. Mar. Polyzoa Brit. Mus., pt. 1, Cheilo., p. 24, pl. 52, figs. 1-4. Cellularia cervicornis SmitTt, 1872, K. Svensk. Vetens. Akad. Handl., vol. 10, pt. 11, p. 14, pl. 5, figs. 39-42. Scrupocellaria regularis Osspurn, 1940, N. Y. Acad. Sci., Sci. Sur. Porto Rico, Virgin Isl., vol. 16, pt. 3, pp. 384, 385. Scrupocellaria regularis, OsBuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, p. 144, pl. 18, figs. 3, 4, pl. 20, fig. 3. DescripTION: Zoaria erect, non-incrusting. Zooecia short, ovoid. Many frontal avicularia, elevated, small, found on both sides of the aperture. Lateral avicularia less numerous, small. Vibracula chambers large, later- ally placed. Fossil specimens lacking spines, vibracula and seuta. Ovi- cells, when present, globose, large, perforate. MEASUREMENTS: Aperture, 230/190 mp. Zooecia, 340/175 muy. LocaLitIES: Bath-house Beach, Santa Barbara, California, Pleistocene. OccURRENCE: Recent: Pacific: Gulf of California (Osburn, 1950). Af- lantic: Florida (Smitt, 1872, Osburn, 1914); Bermuda (Verrill, 1900) ; Puerto Rico (Osburn, 1940); Colombia (Osburn, 1947); Aruba Island, Caribbean Sea (Osburn, 1947). DeptH: Shore (intertidal) to 45 fathoms (5 records). Remarks: This species has not been previously recorded in the fossil records. Scrupocellaria varians Hincks, 1882. Scrupocellaria varians Hincks, 1882, Ann. Mag. Nat. Hist., ser. 5, vol. 10, pp. 461, 462, pl. 19, figs. 1-le. Scrupocellaria varians, Roperrson, 1905, Univ. Calif. Pub. Zool., vol. 2, no. 5, pp. 260, 261, pl. 8, figs. 38, 39, pl. 16, fig. 95. Scrupocellaria varians, O’DonoGHUE and O’DoNoGHUE, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 160. Scrupocellaria varians, O’DONOGHUE and O’DonocHuE, 1925, Pub. Puget Snd. Biol. Sta., vol. 5, p. 98. Scrupocellaria varians, O’DonocHuE and O’DonoGHuer, 1926, Contr. Canad. Biol. Nish: as) vole 3. NO. 3, D. 8. Scrupocellaria varians, OSBURN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 149, 150, pl. 19, fig. 5, pl. 20, fig. 6. 104 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Description: Zoaria erect, non-inerusting. Zooecia elongated, nar- rowed. Frontal avicularia oblique, triangular, elevated. Vibracular cham- ber shortened, dorsally placed. Ovicell, when present, ovoid, imperforate. MrasureMENts: Aperture 253/126 mp. Zooecia 470/190 mu. Locauities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. OccuRRENCE: Recent: Pacific: Queen Charlotte Islands (Hincks, 1884) ; Vancouver Island region (O'Donoghue and O’Donoghue, 1923, 1926) ; Puget Sound (O’Donoghue, 1925); California coast (Robertson, 1905, Osburn, 1950). DeptH: Intertidal to 138 fathoms. Remarks: Not previously found as a fossil. DIVISION VI, CRIBROMORPHA Harmer, 1926 Family CRIBRILINIDAE Hincks, 1880 Genus Reginella Jullien, 1886 Reginella mucronata (Canu and Bassler), 1923. Metracolposa mucronata CANu and Basster, 1928, U. S. Nat. Mus. Bull. 125, p. 92, pl. 35, fig. 4. Reginella mucronata, Ospurn, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 180, 181, pl. 28, fig. 4, pl. 29, fig. 3. DescriIPpTION: Zoaria incrusting. Zooecia distinct, elongate, convex, with 10 costules somewhat obscured by secondary calcification. Aperture wide, rounded above, and nearly straight on the proximal edge. On some zooecia the proximal rim of the aperture bears a central prominent blunt muera, which appears bifid in the well preserved specimens. Ovicells globose, large, with a well defined keel along the mid-longitudinal line. MeasurREMENTS: Aperture, 81/188 mp. Zooecia 552/356 mu. LocaLtities: Bath-house Beach, Santa Barbara, California, Pleistocene. OCCURRENCE: Pleistocene: Santa Barbara, California (Canu and Bass- ler, 1923). Recent: California, Channel Islands, Baja California, Gulf of California, Mexico (Osburn, 1950). DeptH: 10 to 150 fathoms (17 records). VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 105 Genus Colletosia Jullien, 1886 Colletosia radiata (Moll), 1803. Eschara radiata Mou, 1803, Seerinde, p. 63, pl. 4, fig. 7. (fide Hincks, Hist. Brit. Mar. Polyzoa, vol. 1, pp. 185-190, pl. 25, figs. 1-9). Cribrilina radiata, Hincxs, 1883, Ann. Mag. Nat. Hist., ser. 5, vol. 11, pp. 442, 443. Puellina radiata, CANU and BASSLER, 1920, U. S. Nat. Mus. Bull. 106, p. 294, fig. 84g—-h, pp. 295, 296, pl. 41, figs. 14-18. Puellina innominata CANu and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, pp. 90, 91, Die tbs hie Ws: Puellina radiata forma scripta CANU and BAsster, 1923, U. S. Nat. Mus. Bull. 125, pp. 89, 90, pl. 15, fig. 12, pl. 35, fig. 1. Puellina radiata forma rarecosta CANU and BAss LER, 1923, U. S. Nat. Mus. Bull. 1125s p. 90: Cribrilina radiata var. radiata O’DONOGHUE and O’DoNnoGHuRr, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 1, p. 172. Puellina radiata, O’DONOGHUE and O’DoNnoGHuk, 1923, Univ. Wash. Pub. Puget Snd. Biol. Sta. Trans., vol. 5, p. 101. Puellina radiata var. radiata, O’DonoGuurE and O’DoNnoGHuE, 1926, Contr. Canad. BOIL, JONG, Me Say WO, Bh Os 3 1s Yi Puellina radiata, CANU and BAsstrr, 1930, Proc. U. S. Nat. Mus., no. 2810, vol. 76, art. 13, pp. 3, 13. Puellina innominata, CANU and BaAss ter, 1930, Proc. U. S. Nat. Mus., no. 2810, vol. KOmantelo.spD: oF dice Colletosia radiata, OsBpuRN, 1950, Allan Hancock Pac. Expds., vol. 14, no. 1, pp. 187, 188, pl. 29, figs. 2, 2a. DescriPpTION: Zoaria inerusting. Zooecia distinct, frontal costate, con- vex, in our fossil material heavily coated with secondary calcification. Some of the zooecia showing one or two small round pores just proximal to the aperture on the midline. Aperture rounded distally, with a straight prox- imal border. Distal rim bearing the sears of five spines. Vicarious avicularia rare. Ovicell large, globose. MEAsSuREMENTS: Aperture, 46/69 mp. Zooecia, 357/219 muy. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, south location, Newport, California, Pleistocene. Bath-house Beach, Santa Barbara, Cali- fornia, Pleistocene. OCCURRENCE: Pleistocene: Santa Barbara, California (Canu and Bass- ler, 1923); Santa Monica, California (Canu and Bassler, 1923); Dizodo Beds, Boso Peninsula, Japan (Sakakura, 1935). Pliocene: Italy (Neviani, 1905); Panama (Canu and Bassler, 1928); France (Canu, 1925). IJiocene: Austria, Hungary (Canu, 1924); Austria (Gillard, 1938); Florida (Canu and Bassler, 1923); North Carolina (Canu and Bassler, 1923); Virginia (Canu and Bassler, 1923). Oligocene: Leeward Island (Canu and Bassler, 106 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 1923); Alabama (Canu and Bassler, 1923); Alaska (Howe, 1942); Africa (Roger, 1943). Eocene: Belgium (Canu, 1910); North Carolina, South Carolina, Georgia, Florida (Canu and Bassler, 1920); South Carolina, Ala- bama (Canu and Bassler, 1923). Recent: Pacific: Queen Charlotte Islands (Hineks, 1883); Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound (O’Donoghue and O’Donoghue, 1925); Hawaii (Canu and Bassler, 1927); South China Sea, Japan (Silén, 1942); Galapa- gos Islands (Canu and Bassler, 1930, Osburn, 1950); Oregon, south to Peru (Osburn, 1950); California, Mexico, Baja California, Gulf of California, Panama, Colombia (Hancock stations); Australia (Livingstone, 1927). At- lantic: Madeira ( Hincks, 1883, Busk, 1858, Norman, 1909) ; Britain (Hincks, 1880); Tunis (Canu and Bassler, 1930); Naples, Italy (Waters, 1885, Harmer, 1902); Azores, Cape Verde Islands, Canary Islands (Waters, 1918, Calvet, 1931); France (Calvet, 1907); Spain (Barroso, 1921); Flor- ida, Georgia, Cuba, Gulf of Mexico, Puerto Rico (Osburn, 1940, Canu and Bassler, 1928, Smitt, 1873); Tortuga and Aruba islands (Osburn, 1947) ; Brazil (Marcus, 1937, Kirkpatrick, 1888). DeptH: Shore (intertidal) to 268 fathoms (92 records). ASCOPHORA Levinsen, 1909 Family HiproTHorpaE Levinsen, 1909 Genus Hippothoa Lamouroux, 1821 Hippothoa hyalina (Linné), 1767. Cellepora hyalina LINNE, 1767, Systema Naturae, ed. 12, vol. 1, pt. 2, p. 1286. Schizoporella hyalina, Hincks, 1883, Ann. Mag. Nat. Hist., ser. 5, vol. 11, p. 445. Schizoporella hyalina, RoBertson, 1900, Proc. Wash. Acad. Sci., vol. 2, p. 326. Schizoporella hyalina, RosperTson, 1908, Univ. Calif. Publ. Zool., vol. 4, no. 5, pp. 289-290, pl. 19, figs. 43-45. Hippothoa hyalina, CANuU and Bass ter, 1920, U. S. Nat. Mus. Bull. 106, p. 326, figs. 93a-i. Hippothoa hyalina, CANU and BAssLeEr, 1923, U. S. Nat. Mus. Bull. 125, pp. 92-94, Diesh, fis. 4: Hippothoa hyalina var. rugosa CANU and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, pp. 94, pl. 35, fig. 9. Schizoporella hyalina, O’DoNOGHUE and O’DONOGHUE, 19238, Contr. Canad. Biol., n. si, vol. 15 no: 10, pp) 177, 178: Hippothoa hyalina, O’DoNOGHUE and O’DoNOGHUE, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta. Trans., vol. 5, p. 101. Hippothoa hyalina, O’DoNoGHUE and O’DoNnocHuRr, 1926, Contr. Canad. Biol. Fish., meess,, Vol. 3;.ps 100! Vou. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 107 Hippothoa hyalina, Hastines, 1930, Proc. Zool. Soc. Lond., 1929, p. 720. Hippothoa hyalina, OsBurN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, p. 277, pl. 30, figs. 1-5. DescrIPTION: Zoaria forming either single layered incrustations, or multilaminar formations. Zooecia either subeylindrieal, elongate distinct, disposed in radiating rows separated by large pores, or stubby, barrel- shaped, thick-walled, wrinkled. Orifice well rounded, wide, with a broad shallow sinus, with or without a pair of small denticles. Ovicells large, globose, with or without pores. MEASUREMENTS: Aperture, 67/70 mp. Zooecia, 330/207 muy. LocauitIEs: Bath-house Beach, Santa Barbara, California, Pleistocene. OCCURRENCE: Pleistocene: Santa Barbara, California, Dead Man’s Island, San Pedro, California, Los Angeles, California, Santa Monica, Cali- fornia (Canu and Bassler, 1923). Pliocene: England (Busk, 1859, Hincks, 1880). Recent: Pacific: Queen Charlotte Islands (Hincks, 1883); Alaska (Robertson, 1900); California (Robertson, 1908, Canu and Bassler, 1923) ; Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926) ; Puget Sound (O’Donoghue and O’Donoghue, 1925); Japan (Okada, 1929) ; Galapagos (Hastings, 1930); California Channel Islands, Baja California, Mexico, Peru, Ecuador, Galapagos Islands, Washington, British Colum- bia (Hancock stations). Atlantic: British Isles (Hassall, 1841, Johnston, 1847, Hincks, 1862, 1880, Mar. Biol. Ass’n, 1931); Norway, Sweden (Nor- man, 1894, Nordgaard, 1906, 1918, 1927, Broch, 1928, Borg, 1933); Adri- atic (Heller, 1867); Iceland, Greenland, Labrador (Packard, 1863, Verrill and Smith, 1873, Hincks, 1877, Osburn, 1912, 1919, 1936, Borg, 1933) ; Hudson Bay (Osburn, 1923, 1932, Borg, 1933); Chesapeake Bay, Mt. Des- ert Region, Maine, Long Island Sound, Woods Hole, Massachusetts (Osburn, 1912, 1932, 1944, Hutchins, 1945, Rogick, 1945, Verrill and Smith, 1873) ; Netherlands (Lacourt, 1949). DeprtH: Shore (intertidal) to 100 fathoms (94 records). Family PerrauimparE Levinsen, 1909 Genus Hippopodina Levinsen, 1909 Hippopodina feegeensis (Busk), 1884. Lepralia feegensis Busk, 1884, Rep. Voy. H.M.S. Challenger, vol. 10, pt. 3, Polyzoa, pp. 144, 145, pl. 22, fig. 9. Hippopodina feegeensis, CANu and BaAsstrr, 1920, U. S. Nat. Mus. Bull. 106, p. 532, figs. 156a—j. 108 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Hippopodina feegeensis, CANU and BAsster, 1923, U. S. Nat. Mus. Bull. 125, p. 168, fig. 29a. Hippopodina feegeensis, HAastines, 1930, Proc. Zool. Soc. Lond., 1929, p. 729. Hippopodina feegeensis, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 292, 293, pl. 31, figs. 6-8. DEscRIPTION: Zoaria incrusting. Zooecia distinct, alternate, large with a porous frontal. Aperture large, arched above and below, but slightly narrower below with a pair of lateral cardelles. Peristome low, well defined. Our specimens, although well preserved, lacking ovicells. Few avicularia present, placed along the aperture, pointing distally. MEASUREMENTS: Zooecia 815/580 mp. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OcCURRENCE: Recent: Pacific: Ceylon (Thornely, 1905) ; Colombia (Hast- ings, 1930); Philippines, Hong Kong (Busk, 1884); Singapore (Levinsen, 1909); Australia (MacGillivray, 1891). No Haneock stations. Atlantic: Curacao (Osburn, 1927); West Indies (Levinsen, 1909) ; specimens collected in Florida by R. C. Osburn. DeptH: Unknown. RemMaArKs: This species has not been recovered previously as a fossil. Family SCHIZOPORELLIDAE Bassler, 1935 Genus Arthropoma Levinsen, 1909 Arthropoma cecili (Audouin), 1826. Flustra cecili Aupoutn, 1826, in Savigny, J. C., Descr. de l’Egypti Hist. Nat., vol. 1, Dt 4,aps 8, fiz. 3. Schizoporella Cecilii, Hincks, 1883, Ann. Mag. Nat. Hist., ser. 5, vol. 11, p. 445. Schizoporella cecili, Ropertson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, p. 288, pl. 19, fig. 42. Arthropoma cecilii, CANU and BASsLER, 1920, U. S. Nat. Mus. Bull. 106, p. 351, fig. 105a-d. Arthropoma cornuta CANU and BASSsLER, 1923, U. S. Nat. Mus. Bull. 125, p. 97, pl. 116; fie. 3: Schizoporella cecilii, O’ DONOGHUE and O’DoNoGHUE, 1923, Contr. Canad. Biol., n. s., voli) mo: LO; pede Arthropoma cecilii, O’ DONOGHUE and O’DonoGHuE, 1926, Contr. Canad. Biol. Fish., nN: S., Vol. 3, no. 3, Dp. 104: Arthropoma cecili, CANU and BAsSster, 1930, Proc. U. S. Nat. Mus., no. 2810, vol. 76, art. 1135 p: 6: VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 109 Arthropoma cecili, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 333, 334, pl. 38, figs. 1-3. DEscriIPTION: Zoarium inecrusting. Zooecia obscure because of secondary calcification, alternate punctate. Aperture rounded distally, with a straight proximal margin containing a distinct sinus. No ovicells on our specimen. MEASUREMENTS: Aperture, 80/69 mp. Zooecia not possible to measure. Locauities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. OccuRRENCE: Pleistocene: Dizodo Beds, Japan (Sakakura, 1935). Plio- cene: no records. Miocene: Florida (Canu and Bassler, 1923). Recent: Pacific: Japan (Ortmann, 1890, Sakakura, 1935, Okada and Mawatari, 1935); Queen Charlotte Islands (Hineks, 1883); Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Galapagos Islands (Canu and Bassler, 1930); California (Robertson, 1908, Canu and Bassler, 1930); Australia (Livingstone, 1927); Baja California, Mexico (Hancock stations. ) Atlantic: England (Busk, 1857, Hincks, 1862, 1880); France (Calvet, 1902, 1907); Brazil, (Mareus, 1937); Africa (O’Donoghue, 1935). DeptH: 11 to 398 fathoms (12 records). Remarks: Not previously known as a fossil from the Pacifie coast. Genus Schizoporella Hincks, 1877 Schizoporella cornuta (Gabb and Horn), 1862. Reptescharellina cornuta Gass and Horn, 1862, Jour. Acad. Nat. Sci., Phila., ser. 2, vol. 5, pt. 2, pp. 147, 148, pl. 20, fig. 31. Schizoporella biaperta Hincks, 1883, Ann. Mag. Nat. Hist., ser. 5, vol. 11, pp. 445, 446. Schizoporella biaperta, Hincks, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 13, pp. 57, 58. Schizoporella biaperta, Hincxs, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 13, pp. 211, 212. (?) Schizoporella biaperta, Ropertson, 1900, Proc. Wash. Acad. Sci., vol. 2, p. 326. Schizoporella biaperta, RopertTson, 1908, (partim), Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 287, 288, pl. 19, fig. 41. Stephanosella biaperta, CANuU and BAssLer, 1923, (partim), U. S. Nat. Mus. Bull. 125, pp. 99-101, figs. 4-8. (?) Schizoporella biaperta O’DoNogHUE and O’DoNnoGHur, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10=p. 177. (?) Stephanosella biaperta, O’DoNoGHUE and O’DonoGHuE, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., vol. 5, p. 102. (?) Stephanosella biaperta, O’DoNoGHUE and O’DoNoGHur, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, p. 104. 110 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. (2) Stephanosella biaperta, Hastines, 1930, Proc. Zool. Soc. Lond., 1929, p. 721. Schizopodrella (Stephanosella) biaperta, CANU and BAssLeErR, 1930, Proc. U. S. Nat. Muss no 28t0-cvol. 16;,att. 3. ppalo;iais ple 2, fe. 192: Schizoporella cornuta, OsBurN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 320, 321, pl. 37, figs. 9-11. DescripTION: Zoaria incrusting. Zooecia ovate, slightly convex, frontal porous, though in the fossil specimens well obscured by secondary ealci- fication. Orifice subcireular, proximal border almost straight and having distinct sinus. No oral spines. At each side of the orifice is situated an elevated avicularium. Ooecia globular, front wall flattened and traversed by radiating lines. MEASUREMENTS: Aperture, 108/98 mp. Zooecia 465/345 muy. Loealities: Bath-house Beach, Santa Barbara, California, Pleistocene. OcCURRENCE: Pleistocene: Santa Barbara, California (Gabb and Horn, 1862; Dizodo Beds, Japan (Sakakura, 1935); Los Angeles, Santa Monica, Dead Man’s Island, San Pedro, California (Canu and Bassler, 1923). Pliocene: (?) New Zealand, Italy (Waters, 1887); (?) Sicily (Waters, 1878); (?) England (Hincks, 1880). Recent: Queen Charlotte Islands (Hinecks, 1883, 1884); California (Robertson, 1908); Galapagos Islands (Canu and Bassler, 1930). Hancock stations in California, Channel Islands, Baja California, Mexico, Panama, Ecuador, Colombia, and Galapagos Islands are known Pacific areas. It is certain that checking on specimens of S. biaperta from the Atlantic will reveal that some have a perforate frontal and hence should be reclassified as S. cornuta. Schizoporella linearis inarmata (Hincks), 1884 Schizoporella linearis, Hassatt, form inarmata Hincxks, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 13, p. 212. Schizoporella linearis HASSALL, subsp. inarmata, RORERTSON, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, p. 291, pl. 20, fig. 48. Schizoporella linearis HASSALL, var. inarmata, O’DONOGHUE and O’DONOGHUE, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 178. Schizopodrella linearis (HASSALL) var. inarmata, O’DONOGHUE and O’DONOGHUE, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., vol. 5, p. 102. Schizopodrella linearis (HASSALL), 1841, var. inarmata, O’DONOGHUE and O’DoN- OGHUE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, p. 104. Schizoporella linearis var. inarmata, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 319, 320, pl. 37, figs. 4, 5. DeEscRIPTION: Zoaria incrusting. Zooecia elongate, in a rough linear series, somewhat obscured by secondary calcification. Frontal porous, un- even. Aperture rounded above with a “V’-shaped sinus in the proximal rim. Avicularia wanting. No ovicells on the specimens. Vou. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 111 MEASUREMENTS: Aperture, 78/92 mp. Zooecia 414/320 mu. Locauities: West Second and Pacifie Avenue, San Pedro, California, Pleistocene. OccuRRENCE: Recent: Pacific: Queen Charlotte Islands (Hineks, 1884) ; California (Robertson, 1908); Puget Sound, Washington (O’Donoghue and O’Donoghue, 1925); Vancouver Island, British Columbia region (O’Don- oghue and O’Donoghue, 1923, 1926). Hancock stations, Mexico, Gulf of California, Baja California, Costa Riea. DeptH: 10 to 57 fathoms (8 records). ReMArRKS: This appears to be the first appearance of the species in the fossil record. Genus Schizomavella Canu and Bassler, 1917 Schizomavella auriculata cuspidata (Hincks), 1880. Schizoporella auriculata var. B. (cuspidata) HincKs, 1880, Hist. Brit. Mar. Polyzoa, vol. 1, pp. 261-263, pl. 29, fig. 8. DEScRIPTION: Zoaria inerusting. Zooecia distinet, frontal coneave, ar- ranged in linear series. Aperture rounded above, proximal margin with a distinct sinus. Avicularia suboral, raised. Ovicells large globular, pune- tate, distally bearing a sharp prong-like process on the mid-line. MEASUREMENTS: Aperture, 90/57 mp. Zooecia 435/218 muy. LocaLities: Bath-house Beach, Santa Barbara, California, Pleistocene. OCCURRENCE: Recent: Isle of Guernsey, Mediterranean (Hincks, 1880). DeptH: Unknown. Remarks: This variety, so far as we can determine, has not been pre- viously reported as fossil. Genus Hippodiplosia Canu, 1916 Hippodiplosia insculpta (Hincks), 1882. Schizoporella insculpta Hincxs, 1882, Ann. Mag. Nat. Hist., ser. 5, vol. 10, pp. 251-252. Schizoporella insculpta, HincKks, 1883, Ann. Mag. Nat. Hist., ser. 5, vol. 11, p. 447, figs. 5, 5a. Schizoporella insculpta, Rorertson, 1900, Proc. Wash. Acad. Sci., vol. 2, p. 326. 112 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Schizoporella insculpta, ROBERTSON, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 290, 291, pl. 20, figs. 46, 47. Schizoporella insculpta, O’DoNoGHUE and O’DonoGHur, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, pp. 178, 179: Schizoporella insculpta, O’DoNoGHUE and O’DoNoGHUE, 1925, Univ. Wash. Pub. Puget Snd. Biol. Sta. Trans., vol. 5, p. 102. Schizoporella insculpta, O’DoNOGHUE and O’DoNoGHUE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, p. 1038. Hippodiplosia insculpta, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 341, 342, pl. 40, figs. 1, 2. DeEscRIPTION: Zoaria incrusting. Zooecia large, alternate. Frontal wall with large pores. Aperture rounded above, lower margin occupied almost entirely by a wide shallow sinus. Ovicells large, abundant, sometimes cov- ering two-thirds of the zooecia above, and occasionally punctured about the base. Ooecial walls covered by granular ridges radiating from the oral arch. Avicularia lacking. MEASUREMENTS: Aperture, 69/103 mp. Zooecia 460/241 muy. LocaLities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. OcCURRENCE: Recent: Pacific: Alaska (Robertson, 1900); Queen Char- lotte Islands (Hincks, 1882, 1883); Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound (O’Donoghue, 1925); Cali- fornia (Robertson, 1908); Hancock stations, Oregon, California, Mexico, Gulf of California, Costa Riea. DeptH: Shore (intertidal) to 126 fathoms (19 records). Remarks: Not previously recorded as fossil. Family HippoporInipaE Osburn, 1952 Genus Hippoporella Canu, 1917 Hippoporella gorgonensis Hastings, 1930. Hippoporella gorgonensis HAstines, 1930, Proc. Zool. Soc. London, 1929, pp. 723, 724, pl. 12, figs. 62-72, pl. 17, figs. 119, 121. Hippoporella gorgonensis, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 348, 349, pl. 45, figs. 10-12. DeEscripTION: Zoaria incrusting. Zooecia with frontal wall finely gran- ular, with lateral areolar pores small. Ovicell nonporous, shallow rough- ened. Avicularia variable, may be found on both sides of the aperture, or on just one side, replacing lateral processes. There may be an avicularium on the frontal wall, directed away from the apertures. VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 113 MEASUREMENTS: Aperture, 115/149 mp. Zooecia 380/234 muy. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OccuRRENCE: Recent: Pacific: Panama, Colombia, Galapagos Islands (Hastings, 1930); Hancock stations, California, Mexico, Baja California, Panama, Costa Rica, Colombia, Ecuador, Galapagos. Atlantic: Brazil, St. Helena (Marcus, 1937, 1938). Depru: Shore (intertidal) to 150 fathoms (20 reeords). ReEMaRKS: This species is new to the fossil list. yenus Hippoporidra Canu and Bassler, 1927 Hippoporidra edax (Busk), 1859. Cellepora edax Busk, 1859, Monogr. Fossil Polyzoa Crag, London, p. 59, pl. 9, fig. 6. pl. 22, fig. 3. Cellepora minuta CANU and Basster, 1923, U. S. Nat. Mus. Bull. 125, p. 182, pl. 25, figs. 10-13. DeEscrRIPTION: Zoaria forming heavy incrustations. Zooecia irregularly placed, marginal area distinctly punctured. Aperture rounded anteriorly, somewhat contracted below the middle, forming a small distinct denticle on either side. Posterior edge arched proximally. A few small oral avicu- laria, situated close to the aperture, may be found. Ovicells, when pres- ent, rounded. MEASUREMENTS: Aperture, 920/690 muy. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. LACMIP loc. 136, Newport Bay road, Newport, Cali- fornia, Pleistocene. OccURRENCE: Pliocene: South Carolina (Canu and Bassler, 1923, 1928); England (Busk, 1859, Kendall, 1931). Miocene: North Carolina (Canu and Bassler, 1923, 1928). Oligocene: Australia (Waters, 1882, 1885). Recent: Atlantic: North Sea (Prenant, 1931); England (Busk, 1861, Hincks, 1862, 1880); Yucatan, Gulf of Mexico (Canu and Bassler, 1928); Tortugas Islands (Osburn, 1914); Florida (Smitt, 1873); New Jersey (Osburn, 1940). Depru: 15-79 fathoms, (4 records). Remarks: Not reported as living from the Pacifie coast. 114 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER Genus Stephanosella Canu and Bassler, 1917 Stephanosella biaperta (Michelin), 1842. Eschara biaperta MIcHELIN, 1842, Iconogr. zoophytol., p. 330, pl. 79, fig. 3. (2?) Schizoporella biaperta, Rosertson, 1900, Proc. Wash. Acad. Sci., vol. 2, p. 326. Schizoporella biaperta, ROBERTSON, 1908, (partim), Univ. Calif. Pub. Zool., vol. 4, NOW| DD 2st. 2os. Die 19: hie. 48 Stephanosella biaperta, CANU and BASSLER, 1923, (partim), U. S. Nat. Mus. Bull. 125, pp. 99-101, fig. 9. (?)Stephanosella biaperta, HAstines, 1930, Proc. Zool. Soc. London, 1929, p. 721. Stephanosella biaperta, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 368, 369, pl. 42, figs. 1, 2. DeEscrRIPTION: Zoaria incrusting, rough. Zoecia ovoid, somewhat raised, the frontal not perforate, thickened and some specimens obscured by sec- ondary calcification. Recent specimens having 4 or 5 areolar pores to each side that are obliterated frequently in the fossil material, limiting positive identification to only the extremely well preserved specimens. Aperture rounded, widened, with a sinus in the proximal border. No ovicells found on our fossil material. MEASUREMENTS: Aperture, 115/140 mp. Zooecia 640/375 muy. Locauities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. OccURRENCE: Pleistocene: Los Angeles, Dead Man’s Island San Pedro, Santa Monica, California (Canu and Bassler, 1923). Recent: California (Robertson, 1908); (?) Alaska (Robertson, 1900); Specimens in the Han- eock collection from Alaska, collected by G. E. MacGinitie. DeptH: 20 to 40 fathoms. Family MicroporELLIpAE Hineks, 1880 Genus Microporella Hincks, 1877 Microporella ciliata (Pallas), 1766. Eschara ciliata PALLAS, 1766, Elenchus Zoophytorum, p. 38. Microporella ciliata, Hincks, 1883, Ann. Mag. Nat. Hist., ser. 5, vol. 11, p. 443. Microporella ciliata, CANU and BAssuer, 1923, U. S. Nat. Mus. Bull. 125, pp. 119, 120, pl. 20, figs. 1-6, pl. 36, figs. 4, 5. Microporella ciliata, O’7 DONOGHUE and O’DONOGHUE, 1923, Contr. Canad. Biol., n. s., vol. 15 no, 410, ppadii3nine Vou. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 115 Microporella ciliata, O'DONOGHUE and O’DoNoGHUR, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta. Trans., vol. 5, p. 103. Microporella ciliata, O'DONOGHUE and O’DoNnoGHuR, 1926, Contr. Canad. Biol. Fish., NeSVOls oy Os oF DDS LlOe Tate Microporella ciliata, Hastines, 1930, Proc. Zool. Soc. London, 1929, p. 727. Microporella ciliata, OsBuRN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, p. 377, pl. 44, fig. 1. DescriIPTION: Zoaria incrusting. Zooecia distinet, elliptical. Frontal a porous tremocyst, either smooth or roughened. Aperture semicireular, with the proximal edge almost straight. A semilunar ascopore is located imme- diately proximal to the aperture. One or two avicularia located at the side of, or just proximal to, the aperture. The ovicell globular, large. MEASUREMENTS: Aperture, 92/103 mp. Zooecia 613/368 my. Locauities: Pleistocene: West Second and Pacific Avenue, San Pedro, Bath-house Beach, Santa Barbara, California, LACMIP loe. 66-2, New- port Harbor mesa, Newport, LACMIP loc. 136, Newport Bay Road, Newport, California. OCCURRENCE: Pleistocene: Japan (Sakakura, 1935); Italy (Neviani, 1891, 1896, ?1900); Santa Monica, Santa Barbara, Dead Man’s Island, San Pedro, California (Canu and Bassler, 1923). Pliocene: England, Aus- tria, Italy (Hineks, 1880); Italy (Waters, 1879). Miocene: Italy (Waters, 1879) ; New Zealand (Waters, 1887); Maryland (Ulrich and Bassler, 1904) ; Florida, Maryland, North Carolina (Canu and Bassler, 1923). Oligocene: Iran (Roger, 1943). Recent: Pacific: Queen Charlotte Islands (Hineks, 1883) ; California (Canu and Bassler, 1923); Vancouver, British Columbia region (O’Donoghue and O’ Donoghue, 1923, 1926); Puget Sound (O’Donoghue and O'Donoghue, 1925); Galapagos Islands, Panama, Colombia (Hastings, 1930); Japan (Okada, 1934, Sakakura, 19385, Okada and Mawatari, 1936) ; Australia, New Zealand (Hineks, 1880, 1883, Waters, 1879); Philippines (Canu and Bassler, 1929); Hancock stations: California, Channel Islands, Mexico, Baja California, Gulf of California, Costa Riea, Panama, Ecuador, Peru, Galapagos Islands. Atlantic: British Isles (Johnston, 1838, 1847, d’Orbigny, 1839, Hincks, 1862, 1880, 18838, Waters, 1879, Smith, 1932, O’Donoghue, 1935, Jones, 1939, Bassindale, 1943); Netherlands (Lacourt, 1949) ; Spitzbergen, Norway, Greenland (Hincks, 1880, 1883, Waters, 1879, Barroso, 1912, Nordgaard, 1918, 1927, Borg, 1933); Florida, Gulf of Mex- ico, Puerto Rico, Tortugas Island, Curacao, Panama (Hincks, 1880, 1883, Smitt, 1873, Waters, 1879, Osburn, 1914, 1927, 1940, 1947, Canu and Bass- ler, 1928); Mediterranean, Africa, Italy (Hincks, 1880, 1883, Waters, 1879, 116 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 1908, 1915, Harmer, 1902, Canu and Bassler, 1930); Brazil (Marcus, 1937) ; Adriatic (Heller, 1867); Cape Verde Islands (Waters, 1918); Labrador, Woods Hole, Chesapeake Bay, Long Island (Osburn, 1912, 1932, Hutchins, 1945); Hudson Bay region (Osburn, 1923, 1932). DeptH: Shore (intertidal) to 150 fathoms (137 records). Microporella californica (Busk), 1856. Lepralia californica Busk, 1856, Quart. Jour. Micro. Sci., vol. 4, p. 310, pl. 11, Hess Gere Microporella ciliata form californica Hincks, 1883, Ann. Mag. Nat. Hist., ser. 5, vols dh p 444 ople lie ties Se Microporella californica, Roperrson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 281, 282, pl. 18, figs. 32-34. Microporella californica, CANU and BAssLeER, 1923, U. S. Nat. Mus. Bull. 125, pp. 1238, 124, pl. 36, figs. 8-10. Microporella californica, O’DONOGHUE and O’DoNnoGHUE, 1923, Contr. Canad. Biol., mas vol, nox 10s ip) Ave: Microporella californica, O’DoNoGHUE and O’DoNnoanuk, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta. Trans., vol. 5, p. 103. Microporella californica, O’DoNoGHUE and O’DoNoGHUE, 1926, Contr. Canad. Biol. Mish= nas woleo.nO. oa, De wdil: Microporella californica, OsBpuRN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, DD. Osis S82. pl, 445 fis. 2: DESCRIPTION: Zoaria inerusting. Zooecia distinet, ovoid. Frontal wall punctate (tremoeyst). Aperture rounded above, with proximal margin straight. The distal rim of the aperture bears the scars of 4-6 spines. On each side and proximal to the aperture are avicularia, pointing obliquely forward. Ascopore lunate. Ovicells globose. MEASUREMENTS: Aperture, 103/120 mp. Zooecia, 759/400 muy. . Locauities: Bath-house Beach, Santa Barbara, California. West See- ond and Pacific Avenue, San Pedro, California, LACMIP loc. 136, New- port Bay road, Newport, California, Pleistocene. OccURRENCE: Pleistocene: Santa Monica [Long Wharf Canyon], Santa Barbara, San Pedro, California (Canu and Bassler, 1923). Recent: Cali- fornia (Busk, 1856), Robertson, 1908); Queen Charlotte Islands, British Columbia (Hineks, 1883); Vaneouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound (O’Donoghue, 1925); Hancock sta- tions, California, Mexico, Baja California, Galapagos Islands. DeptH: Shore (intertidal) to 81 fathoms (14 records). VoL. XXIX ] SOULE AND DUFF: FOSSIL BRYOZOA italy Genus Fenestrulina Jullien, 1888 Fenestrulina malusi (Audouin), 1826. Cellepora malusii AUDOUIN and Savieny, 1811, Desecr. de Egypt Hist. Nat., p. 239, pl. 8, fig. 8. Cellepora malusii AupOoUIN, 1826, Explic. de Savigny, p. 239. Cellepora californiensis Gass and Horn, 1862, Jour. Acad. Nat. Sci., Phila., ser. 2, Walls JA, 70k PA 1, TRIO, Tolle UI) ike, De Microporella Malusii, H1ncks, 1883, Ann. Mag. Nat. Hist., ser. 5, vol. 11, p. 444. Microporella Malusii, H1ncks, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 13, p. 57. Microporella malusi, Ropertson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 282, A333 joll, ISS ake By, Bio, Microporella malusi, CANU and BAsster, 1920, U. S. Nat. Mus. Bull. 106, pp. 418, 419, fig. 123 f-n. Fenestrulina malusi, CANU and Basser, 1923, U. S. Nat. Mus. Bull. 125, pp. 114- 117, pl. 36, figs. 2, 3, fig. 18 e-i, fig. 19 a-j. Microporella malusii, O'DoNoGHUE and O’DonoGHurE, 1923, Contr. Canad. Biol., n. s., Vole enOs Os ps daar Fenestrulina malusii var. umbonata O’DonoGHUE and O’DoNnoGHur, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta. Trans., vol. 5, p. 103. Fenestrulina malusii, O'DONOGHUE and O’DonoaHuE, 1926, Contr. Canad. Biol. Fish., TES VOL ono. om DDS LOO dalO: Fenestrulina malusi, Ospurn, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 387, 388, pl. 45, fig. 3. DEscRIPTION: Zoaria incrusting. Zooecia regular, well defined, large, the frontal surface of some of the better preserved specimens with stellate pores. Usually the pores are obscured by secondary calcification. About half-way down the frontal is a median areuate ascopore. The aperture is rounded above, with a nearly straight proximal edge. Ovicells globular, striated, large. MeASUREMENTS: Aperture, 86/149 mp Zooecia 667/452 muy. Locauities: Bath-house Beach, Santa Barbara, California, Pleistocene. OccURRENCE: Pleistocene: Santa Barbara, California (Gabb and Horn, 1862, Canu and Bassler, 1923). Dizodo Beds, Japan (Sakakura, 1935). Pliocene: England (Kendall, 1931). Miocene: Australia (MacGillivray, 1895). Recent: Pacific: California (Robertson, 1908); Queen Charlotte Islands, British Columbia (Hineks, 1883, 1884); Vancouver Island region (O'Donoghue and O'Donoghue, 1923, 1926); Puget Sound (O’Donoghue and O'Donoghue, 1925); Australia (Livingstone, 1926, 1927); Japan (Okada, 1929): India (Robertson, 1921); Hancock stations in California, Channel Islands, Gulf of California, Mexico, Baja California, Galapagos Islands. 118 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Atlantic: England (Johnston, 1847, Hincks 1862, 1880, Prenant, 1931, Jones, 1939); Norway, Greenland (Borg, 1933); St. Helena (Marcus, 1938); Puerto Rico, Gulf of Mexico (Osburn, 1940, Canu and Bassler, 1928) ; Adriatic (Heller, 1867); Spain (Norman, 1909, Barroso, 1912, 1921). DeptH: Shore (intertidal) to 150 fathoms (33 records). Family EURYSTOMELLIDAE Levinsen, 1909 Genus Eurystomella Levinsen, 1909 Eurystomella bilabiata (Hincks), 1882. Lepralia bilabiata Hincxs, 1882, Ann. Mag. Nat. Hist., ser. 5, vol. 10, pp. 253, 254. Lepralia bilabiata, Hincxs, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 13, p. 49, pl. 3, fig. 1. Lepralia bilabiata, Ropertson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 298-300, pl. 21, figs. 61-64. Eurystomella bilabiata, CANU and BAsster, 1928, U. S. Nat. Mus. Bull, 125, p. 142, pl. 37, fig. 6. Eurystomella bilabiata, O’DonoGHuUE and O’DoNnoGHUE, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 104. Eurystomella bilabiata, O’DoNoGHUE and O’DonoGHuE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, pp. 111, 112. Eurystomella bilabiata, OspurN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, p. 389, pl. 58, fig. 5. DescripTIOoN: Zoaria incrusting. Zooecia distinct (in spite of heavy secondary calcification), large, short and wide, alternate. Aperture large, bell-shaped. No avicularia. No ovicells found on our specimens. MEASUREMENTS: Aperture, 218/368 mp. Zooecia 578/465 muy. LocaLities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. OccURRENCE: Pleistocene: Dead Man’s Island, San Pedro, California (Canu and Bassler, 1923). Dizodo Beds, Japan (Sakakura, 1936). Recent: Queen Charlotte Islands, British Columbia (Hincks, 1882, 1884); Puget Sound region (Robertson, 1908, O’Donoghue and O’Donoghue, 1925); Van- couver Island (O’Donoghue and O'Donoghue, 1926); California (Robertson, 1908); Haneock stations, California, Channel Islands, Mexico. DepruH: Shore (intertidal) to 35 fathoms (4 records). VoL. XXIX } SOULE AND DUFF: FOSSIL BRYOZOA 119 Family SMITTiInmDaE Levinsen, 1909 Genus Porella Gray, 1848 Porella porifera (Hincks), 1884. Porella marsupium form porifera Hincks, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 13, pp. 50, 51, pl. 4, fig. 4. Smittina porifera, CANU and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, p. 147, pl. 38, fig. 9. Porella marsupium var. porifera, O'DONOGHUE and O’DoNOGHUE, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, pp. 182, 183. Cystisella aviculifera CANuU and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, p. 152, pl. 38, fig. 8. Smittina marsupium var. porifera, O’DoNoGHUE and O’DoNOGHUE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, p. 115. Porella porifera, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 395, 396, pl. 46, figs. 9-11. DESCRIPTION: Zoaria incrusting. Zooecia small, short, distinct, with a few large scattered frontal pores as well as marginal pores. Aperture with low lyrula, and surrounded by a very low peristome. The frontal wall contains small, raised, suboral avicularia. Ovicells globular, low, possibly perforate. MEASUREMENTS: Aperture, 81/103 mp. Zooecia, 402/218 mu. Locauities: LACMIP loc. 66-2 Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OCCURRENCE: Pleistocene: Santa Monica, Santa Barbara, California (Canu and Bassler, 1923). Dizodo Beds, Japan (Sakakura, 1935). Plio- cene: New Zealand (Waters, 1887). Recent: Pacific: Queen Charlotte Islands, British Columbia (Hincks, 1884); Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Hancock stations, California, Channel Islands, Mexico, Baja California, Colombia, Costa Rica. Atlantic: Greenland, Norway, Spitzbergen (Nordgaard, 1895, 1918, Norman, 1903, Bidenkap, 1905, Osburn, 1936); Canada, Labrador (Osburn, 1912, Borg, 1933, Osburn, 1932, Hinecks, 1888). DeptH: Shore (intertidal) to 130 fathoms (25 records). Genus Smittina Norman, 1903 Smittina landsborovi (Johnston), 1847. Lepralia Landsborovii JoHNston, 1847, Hist. Brit. Zoophytes, 2nd ed., p. 310, pl. 54, fig. 9. 120 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Smittia landsborovi, Ropertson, 1908, Univ. Calif. Publ. Zool., vol. 4, no. 5, p. 305, Dl 235 fic.) wae Smittia landsborovi, CANU and BASSLER, 1920, U. S. Nat. Mus. Bull. 106, p. 457. Smittia landsborovii, O’DoNoGHUE and O’DoNoGHUE, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, pp. 184, 185. Smittina landsborovii, O’DoNnoGHuE and O’DonoGHuEr, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 104. Smittina landsborovii, O'DONOGHUE and O’DonoGuuE, 1926, Contr. Canad. Biol. Fish., Te Son AVON Bi, 1aKo)s Sh D0, JI als} Smittina landsborovi, OsBpuRN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 400, 401, pl. 47, figs. 1, 2. DeEscRIPTION: Zoaria incrusting. Zooecia distinct, alternate, with mar- einal pores. Aperture with broad lyrula, surrounded by a peristome. Sub- oral avicularia small, pointing proximally. No ovicell on specimens. MEASUREMENTS: Aperture, 173/195 mp. Zooecia 862/437 muy. Locauities: Bath-house Beach, Santa Barbara, California, Pleistocene. OccuRRENCE: Pliocene: Australia, New Zealand (Waters, 1885, 1887). Recent: Pacific: California (Robertson, 1908); Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound (O’Donoghue and O'Donoghue, 1925); Japan (Okada and Mawatari, 1936); Indian Ocean (Thornely, 1912); Haneock stations, California, Channel Islands, Mexico, Baja California, Galapagos Islands. Atlantic: England (Johnston, 1847, Alder, 1857, 1864, Hineks, 1862, 1880), Arctic, Norway, Greenland, Spitz- bergen (Busk, 1861, Hincks, 1880, Nordgaard, 1895, Bidenkap, 1905). DeptH: 6 to 81 fathoms (25 records). Remarks: This appears to be the first fossil record of this species from the Pacifie coast of North America. Smittina maccullochae Osburn 1952. Smittina maccullochae OsspurN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 405, 406, pl. 48, figs. 5, 6. Porella collifera CAnu and Basster, 1923, U.S. Nat. Mus. Bull. 125, pp. 148, 149, pl. 38, figs. 10-15. DerscrIpTION: Zoaria inerusting. Zooecia large, with a perforate frontal. Aperture surrounded by a high tubular peristome with a proximal sub- oral avieularia. Lyrula large. Ovicell globose, porous. MEASUREMENTS: Aperture, 164/130 mp. Zooecia, 978/459 mu. Locauities: Bath-house Beach, Santa Barbara, California, Pleistocene. OcCURRENCE: Pleistocene: Santa Monica, Santa Barbara, California VoL. XXIX ] SOULE AND DUFF: FOSSIL BRYOZOA 121 (Canu and Bassler, 1923). Recent: Pacific: California, Channel Islands, Mexico (Osburn, 1952). Genus Parasmittina Osburn, 1952 Parasmittina trispinosa (Johnston), 1825. Discopora trispinosa JoHNSTON, 1825, Edinbh. Phil. Jour., vol. 13, p. 222. Lepralia trispinosa, Busk, 1856, Quart. Journ. Mic. Sci., vol. 4A ps Lis: Smittia trispinosa, HrncKs, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 18, p. 51. Smittia trispinosa, RoBerTsON, 1900, Proc. Wash. Acad. Sci., vol. 2, p. 327. Smittia trispinosa, Ropertson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 302, 303, pl. 22, figs. 68-70. Smittia trispinosa, Canu and Basster, 1923, U. S. Nat. Mus. Bull. 125, pp. 148, 144, pl. 22, figs. 7-14. Smittia trispinosa, O’DoNoGHUE and O’DonoGHUE, 1923, Contr. Canad. Biol:; n: s., Woll, al, sO, IMO, jos Ibs. Smittina trispinosa, O’Donocuur and O’DonocHuE, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 105. Smittina trispinosa, O’DoNoGHUE and O’DonoGHuur, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, pp. 113, 114. Smittina trispinosa, Hastines, 1930, Proc. Zool. Soc. London, 1929, pt. 4p. 126, iol; Ill, ies, Sey Smittina trispinosa, CANU and BAsster, 1930, Proc. U. S. Nat. Mus., no. 2810, vol. 76, art. 13, pp. 27—29, pl. 4, figs. 1-5. Parasmittina trispinosa, OsBpuRN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 412-414, pl. 49, figs. 7, 8. Description: Zoaria inerusting. Zooecia distinet on some specimens, obseured by secondary calcification on others, alternate. Aperture with well developed lyrula, surrounded by a low peristome. Avicularia scattered near the aperture, with no trace of sub-oral avicularia. Few olgantic interzooecial avicularia, which may be almost as large as the zooecia. The distal rim of the peristome bears the sears denoting the points where 2 or 3 spines were once attached. Ovicell large, globular, porous. Measurements: Aperture, 76/86 mp. Zooecia, 471/264 my. Locauities: Bath-house Beach, Santa Barbara, West Second and Pa- cific Avenue, San Pedro, LACMIP loc. 130-7, opposite Berth 79, Los Angeles Harbor, Timms’ Point formation, California, Pleistocene. OCCURRENCE: Pleistocene: South Carolina, Florida (Canu and Bass- ler, 1923); Sicily (Neviani, 1896). Plocene: Florida, South Carolina, (Canu and Bassler, 1923). Miocene: North Carolina (Canu and Bassler, 1923): Australia (Waters, 1882). Recent: Pacific: Mexico (Busk, 1856) ; 122 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Alaska (Robertson, 1900, 1908); Queen Charlotte Islands, British Colum- bia (Hineks, 1884); Vancouver Island (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound (O’Donoghue and O’Donoghue, 1925); Panama (Hastings, 1930); Galapagos Islands (Canu and Bassler, 1930); Aus- tralia (Livingstone, 1926, 1927; Japan (Sakakura, 1935); India (Robert- son, 1921); Haneock stations, California, Channel Islands, Gulf of Cali- fornia, Mexico, Baja California, Colombia, Galagapos Islands, Peru. Af- lantic: England (Johnston, 1838, 1847, Alder, 1857, Hincks, 1862, 1880, Smith, 1932); Denmark, Norway, Iceland, Arctic (Hincks, 1877, 1880, Levinsen, 1894, Nordgaard, 1895, 1907, 1912, 1918, 1927); Florida, Tortu- gas, Venezuela (Smitt, 1873, Osburn, 1914, 1947); Labrador, Woods Hole, Massachusetts, Newfoundland (Packard, 1863, Hincks, 1877, Osburn, 1912); Brazil (Marcus, 1937). DeptH: Shore (intertidal) to 150 fathoms (99 records). Parasmittina californica (Robertson), 1908. Mucronella californica Ropertson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 308, 309, pl. 23, fig. 80. (non) Smittina californiensis CANU and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, p. 146, pl. 37, figs. 7-9. Parasmittina californica, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 415, 416, pl. 51, figs. 8-11. DESCRIPTION: Zoaria incrusting. Zooecia distinct, alternate. Frontal porous, bears small raised sub-oral avicularia, occasional very large lateral spathulate avicularia, placed to one side of the aperture, may be found. Aperture surrounded by low peristome, with a lyrula. Ovicells globose, perforate. MEASUREMENTS: Aperture, 124/138 mp. Zooecia, 567/383 muy. LocaLities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. Bath-house Beach, Santa Barbara, California, Pleistocene. OCCURRENCE: Recent: California, Santa Catalina Island (Robertson, 1908); Haneock stations, Channel Islands, Mexico, Gulf of California, Galapagos Islands. DeptH: 23 to 150 fathoms (9 records). ReMARKS: Not previously recorded as fossil. Parasmittina collifera (Robertson), 1908. Smittia collifera Roperrson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 304, 305, pl. 23, figs. 72, 73. Vout. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 123 (non) Porella collifera CANU and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, pp. 148, 149, pl. 38, figs. 10-15. Smittia collifera, O'DONOGHUE and O’DonoGHusr, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 185. Smittina collifera, O’DONOGHUE and O’DoNnocHuR, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 104. Smittina collifera, O’DONOGHUE and O’DoNnoGHur, 1926, Contr. Canad. Biol. Fish., MEESeVOlaro. NOn oa. Parasmittina collifera, OspurRN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 416-418, pl. 49, figs. 9-11. DEscrIPTION: Zoaria incrusting. Zooecia distinct, alternate, with mar- ginal pores. Aperture with lyrula and surrounded by peristome. No sub- oral avieularia. Occasional lateral avicularia located at the side of, and pointing toward, the aperture. No ovicells present on our specimens. MEASUREMENTS: Aperture, 127/138 mp. Zooecia, 437/243 mu. Locauities: Bath-house Beach, Santa Barbara, California, Pleistocene. OccuRRENCE: Recent: Pacific: California (Robertson, 1908); Van- couver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound (O’Donoghue and O'Donoghue, 1925); Hancock stations, California, Channel Islands, Mexico. DeptH: Shore (intertidal) to 81 fathoms (7 records). RemMarKs: This is the first record of this species from fossil deposits. Genus Mucronella Hincks, 1880 Mucronella microstoma (Norman), 1864. Lepralia microstoma NorMaAn, 1864, Ann. Mag. Nat. Hist., ser. 3, vol. 13, p. 87, ple i, fic: 2: (non) Mucronella microstoma O’DoNOGHUE and O’DoNoGHUE, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 188. DeEscrIPTION: Zoaria incrusting, heavily coated by secondary calcifica- tion. Zooecia distinct, ovoid. Aperture small, surrounded by a thick mu- eronate peristome. No intact ovicells on our specimens. MEASUREMENTS: Aperture, 92/115 mp. Zooecia, 322/218 muy. Locauities: LACMIP loe. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OccURRENCE: Recent: Atlantic: British Isles (Norman, 1864, Hincks, 1880); France (Canu, 1924). 124 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. DeptH: 15 to 205 fathoms (4 records). Remarks: Not previously known as fossil. Family RETEPORIDAE Smitt, 1867 Genus Phidolopora Gabb and Horn, 1862 Phidolopora labiata Gabb and Horn, 1862. Phidolopora labiata Gass and Horn, 1862, Jour. Acad. Nat. Sci., Phila., ser. 2, vol. 5, pt. 2, pp. 138, 139, pl. 19, fig. 21. Retepora pacifica Roperrson, 1908, Univ. Calif. Publ. Zool., vol. 4, no. 5, pp. 310, 311, pl. 24, figs. 81-84. Phidolopora pacifica, CANU and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, p. 154, pl. 39, figs. 1-7. Phidolopora labiata, CANU and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, pp. 154, 155, pl. 39, figs. 13-17. Retepora pacifica, O’DoNoGHUE and O’DoNnoGHUE, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 189. Phidolopora pacifica, O’DONOGHUE and O’DONOGHUE, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 106. Phidolopora pacifica, O'DONOGHUE and O’DoNnoGHUE, 1926, Contr. Canad. Biol. Fish., n.s., vol. 3, no. 3, pp. 118, 119. Description: Zoaria in the fossil material consisting of fragments of what were in life large convoluted masses, erect. Fenestrae oval, twice as long as wide. On the dorsal side at the base of each fenestra is located an elevated avicularium, with the mandible portion directed downward. Zooecia small, frontal wall granular, sloping upward from the sides to the midline. On some older zooecia there is a high peristome, with a well-defined sinus below, which develops an elevated avicularium. Ovi- cells, when present, small and set far back, united secondarily to the zooecia by growth of the peristome. MEASUREMENTS: Peristome dia. 162 mp. Zooecia 667/360 my. LocaLities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. Bath-house Beach, Santa Barbara, California, Pleistocene. LACMIP loe. 64, “Hilltop Quarry,” Lomita Marl, San Pedro, California, Pleistocene. OccuRRENCE: Pleistocene: Santa Monica, California, Santa Barbara, California, Dead Man’s Island, San Pedro (harbor), California (Canu and Bassler, 1923); Santa Barbara, California (Gabb and Horn, 1862). Recent: Pacific: California (Robertson, 1908); Vancouver Island, British Columbia (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound region (Robert- Vou. XXIX] SOULE AND DUFF: FOSSIL BRYOZOA 125 son, 1908, O’Donoghue and O'Donoghue, 1925); Hancock stations, Oregon, California, Channel Islands, Mexico, Baja California, Gulf of California, Costa Rica, Galapagos Islands, Peru. DeptH: 8 to 131 fathoms (86 records). Genus Rhynchozoon Hincks, 1895 Rhynchozoon tumulosum (Hincks), 1882. Schizoporella tumulosa Hincks, 1882, Ann. Mag. Nat. Hist., ser. 5, vol. 10, p. 252. Schizoporella tumulosa, Htncks, 1883, Ann. Mag. Nat. Hist., ser. 5, vol. 11, pp. 447, 448, pl. 18, fig. 2. Schizoporella twmulosa, ROBERTSON, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 293, 294, pl. 20, fig. 53. Schizoporella tumulosa, O’DoNOGHUE and O’DoNoGHukr, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 179. Schizoporella tumulosa, O’DoNOGHUE and O’DonoGHUE, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 102. Schizoporella tumulosa, O’DONOoGHUE and O’DONOGHUR, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, p. 102. Rhynchozoon tumulosum, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 458, 459, pl. 54, figs. 4, 5, 12. DESCRIPTION: Zoaria incrusting, heavily calcified. Zooecia large, ovoid, frontal wall raised along center line, sloping to lateral margins. Aperture circular, containing a shallow sinus in the proximal rim. Below the aper- ture and to one side may be found evidence of an elevated avicularium. Avicularia more common in the older zoaria, also found frontally, but must be distinguished from the sub-oral type. Ovicells smooth, globular, immersed. MEASUREMENTS: Aperture, 57/80 mp. Zooecia, 402/276 mu. Locauities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. LACMIP loc. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OccuRRENCE: Recent: Pacific: Queen Charlotte Islands (Hincks, 1882, 1883); Vancouver Island Region (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound region (O’Donoghue and O’Donoghue, 1925); Cali- fornia (Robertson, 1908); Hancock stations, California, Channel Islands, Mexico, Baja California, Oregon. DeprtH: Shore (intertidal) to 235 fathoms (48 records). REMARKS: Not previously known as fossil. 126 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Family CHEILOPORINIDAE Bassler, 1936 Genus Hippopodinella Barroso, 1924 Hippopodinella adpressa (Busk), 1854. Lepralia adpressa Busk, 1854, Cat. Mar. Polyzoa Brit. Mus., pt. 2, p. 82, pl. 102, figs. 3, 4. Lepralia adpressa, BuskK, 1856, Quart. Jour. Mic. Sci., vol. 4, p. 178. Hippopodinella adpressa, HASTINGS, 1930, Proc. Zool. Soc. London, 1929, pt. 2, p. 279. Hippopodinella adpressa, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, De Bt ol NG wes OS DESCRIPTION: Zoaria incrusting. Zooecia ovoid, immersed, somewhat indistinct. Frontal wall pitted. Aperture rounded distally, contracted sharply just below the middle, with the proximal rim only slightly curved. Lateral walls of the aperture bearing two prominent cardelles. Ovicell endozooecial. MEASUREMENTS: Aperture, 162/138 mp. Zooecia 506/325 muy. I p. p LocaLities: West Second and Pacifie Avenue, San Pedro, California, Pleistocene. OCCURRENCE: Recent: Pacific: India (Robertson, 1921); Panama, Galapagos Islands (Hastings, 1930); Mexico (Busk, 1856); Hancock sta- tions, Mazatlan, Mexico, Baja California, Gulf of California, Costa Riea, Panama, Ecuador, Galapagos. Atlantic: England (Hincks, 1862); France (Calvet, 1902); Italy (Norman, 1909). DreptH: Shore (intertidal) to 92 fathoms (29 records). Remarks: Not previously found as a fossil. Family PHYLACTELLIDAE Canu and Bassler, 1917 Genus Lagenipora Hincks, 1877 Lagenipora punctulata (Gabb and Horn), 1862. Entalophora punctulata GABB and Horn, 1862, Jour. Acad. Nat. Sci., Phila., ser. 2, vol. 5, pt. 2, pl. 171, pl. 21, fig. 61. Lagenipora spinulosa Hincxks, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 13, pp. 57, 210, 211, pl. 3, fig. 4, pl. 9, fig. 4. Lagenipora spinulosa, RoBERTSON, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 283, 284, pl. 18, fig. 37. Tubucellaria punctulata, CANU and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, p. 170, pl. 40, figs. 1-4. Vou. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 127 Lagenipora erecta O’DoNoGHUE and O’DonocHur, 1928, Contr. Canad. Biol., n. s., vol. 1, no. 10, pp. 175, 176, pl. 3, fig. 22. Lagenipora erecta, O’DoNOGHUE and O’DonoGHuE, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 106. Lagenipora erecta, O'DONOGHUE and O’DonocHuE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, p. 120. Lagenipora punctulata, OspuRN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 485, 486, pl. 60, figs. 1, 2. DescripTION: Zoaria erect, consisting of tubular stems arising from a small base. Zooecia tubular or flask-shaped, the lower, somewhat swollen portion, forming a part of the stem, with the upper tubular neck stand- ing well out. The neck is wider at the oral extremity, with a tendency to flare. The top of the neck may or may not have traces of small avicularia borne on low processes. Aperture oval. Ovicells, when present, lie at the lower end of the tubular neck, between it and the stem. It has a large crescent-shaped flattened area in front, perforated by pores. MEASUREMENTS: Diameter peristome, 184 mp. Diameter aperture, 126 muy. Locauities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. Bath-house Beach, Santa Barbara, California, Pleistocene. LACMIP loc. 66-2, Newport Harbor mesa, Newport, California, Pleisto- eene. LACMIP loc. 64, “Hilltop Quarry,” Lomita Marl, San Pedro, Cali- fornia, Pleistocene. LACMIP loc. 130-7, opp. Berth 79, Los Angeles har- bor, Timm’s Point formation, Pleistocene. OccuURRENCE: Pleistocene: Santa Barbara, California (Gabb and Horn, 1862); Santa Barbara, Santa Monica, Dead Man’s Island, San Pedro [har- bor], California (Canu and Bassler, 1923). Recent: Pacific: Queen Char- lotte Islands, British Columbia (Hincks, 1884); Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound, Washington (O’Donoghue and O’Donoghue, 1925); California (Robertson, 1908); Han- cock stations, California, Channel Islands, Baja California, Mexico, Galapa- 20s Islands. DeptH: Shore (intertidal) to 150 fathoms (121 records). Family CELLEPORIDAE Busk, 1852 Genus Holoporella Waters, 1909 Holoporella brunnea (Hincks), 1884. Cellepora brunnea Hincxks, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 13, p. 56. Smittia californiensis RoperTsON, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, pp. 303, B04 sple22s figs tle 128 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Cellepora brunnea, O'DONOGHUE and O’DonoGHuE, 1926, Contr. Canad. Biol. Fish., NES VOLO ENON OD. lial Holoporella brunnea, HAstTines, 1930, Proc. Zool. Soc. London, 1929, p. 731, pl. 26, figs. 108-110. Holoporella brunnea, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 496, 497, pl. 62, figs. 10-12. DescripTION: Zoaria inerusting. Zooecia crowded, barrel-shaped, with some elevated, others immersed. Surface smooth, punctured along the margin. Aperture arched above, lower lip curved slightly outward, with a small notch rounded below, and contracted at the opening with two minute denticular projections. Large avicularia scattered among the zooecia. MEASUREMENTS: Aperture, 105/149 mp. Zooecia 546/348 mu. Locauitres: West Second and Pacific Avenue, San Pedro, California, LACMIP loc. 66-2, Newport Harbor mesa, Newport, California, LACMIP loc. 136, Newport Bay road, Newport, California, Pleistocene. OccURRENCE: Recent: Pacific: Queen Charlotte Islands, British Colum- bia (Hincks, 1884); Vancouver Island region, (O’Donoghue and O’Don- oghue, 1926); California (Robertson, 1908); Galapagos, Panama, Colombia (Hastings, 1930); Haneock stations, California, Channel Islands, Mexico, Baja California, Galapagos, Panama, Costa Rica. DeptH: Shore (intertidal) to 150 fathoms (118 records). RemMarKS: Not previously recorded as a fossil. Genus Costazia Neviani, 1895 Costazia costazi (Audouin), 1826. Cellepora costazii AuDOUIN, 1826, in Savigny, Descr. de l’Egypte Hist. Nat. vol. 1, pt. 4, p. 237, pl. 7, fig. 4. (non) Cellepora costazi, Ropertson, 1908, Univ. Calif. Pub. Zool., vol. 4, no. 5, p. 3138, pl. 24, fig. 89. Costazzia (Cellepora) costazzii, CANU and Basser, 1920, U. S. Nat. Mus. Bull. 106, p. 603, fig. 180a—c. Cellepora costazii, O’DONOGHUE and O’DONOGHUE, 1923, partim, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 190. (?) Costazzia costazgzii, O'DONOGHUE and O’DoNoGHUE, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 106. Costazzia costazii, O'DONOGHUE and O’DoNOGHUE, 1926, partim, Contr. Canad. Biol., Fish., n. s., vol. 3, no. 3, p. 121. Costazia costazi, OspuRN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, p. 506, pl. 62, figs. 3, 4. VoL. XXIX ] SOULE AND DUFF: FOSSIL BRYOZOA 129 DerscriPTION: Zoaria small, inerusting. Zooecia irregular, raised, with many marginal pores. Aperture subcireular, with a well defined rounded sinus. Peristome raised, bearing evidence of a small avicularium on each side of the aperture. Ovicells, when present, rounded, semi-lunar in ap- pearance. MEASUREMENTS: Aperture, 161/115 mp. Zooecia, 470/353 mup. Locauities: LACMIP loc. 66-2, Newport Harbor mesa, Newport, Cali- fornia, Pleistocene. OccURRENCE: Pliocene: Italy (Manzoni, 1870, Waters, 1879). Recent: Pacific: India (Robertson, 1921); Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound, Washington (O’Donoghue and O’Donoghue, 1925); Japan (Okada, 1929, 1933, 1934, Okada and Mawatari, 1935); Haneock stations, California, Channel Islands. Atlantic: British Isles (Hincks, 1880, Prenant, 1924, O’Donoghue and Watteville, 1935, Bassindale, 1941, Waters, 1879); Azores, Canary Islands (Calvet, 1907, 1931, Barroso, 1912, Jullien and Calvet, 1903); France (Calvet, 1902, 1907, 1927, 1931). Arctic, Norway, Sweden (Smitt, 1867, Nordgaard, 1896, 1906, Levinsen, 1909); Italy (Waters, 1879, Hincks, 1880, Marcus, 1920, Barroso, 1912); South Africa (O’Donoghue and Watteville, 1935, Stephenson, 1938, Eyre, 1939); North Africa (Canu and Bassler, 1930) ; Brazil, (Mareus, 1937); Puerto Rico (Osburn, 1940). DeptH: 6 to 1069 fathoms (25 records). Costazia robertsonae (Canu and Bassler), 1923. Costazzia robertsoniae CANU and BASsLeErR, 1923, U. S. Nat. Mus. Bull. 125, p. 181, pl. 39, figs. 10-12. Cellepora costazi, RoBERTSON, 1908, partim, Univ. Calif. Pub. Zool. vol. 4, no. 5, p. 313, pl. 24, fig. 190. Cellepora costazii, O’ DONOGHUE and O’DoNOGHUE, 1923, partim, Contr. Canad. Biol., Nese VOL es NOs LOS py 9 0: Costazzia costazii, O’DONOGHUE and O’DONOGHUE, 1926, partim, Contr. Canad. Biol. IMBIN Ws Sky Wolk Bh wos Se os dal Costazia robertsonae, OSBURN, 1952, Allan Hancock Pac. Expds., vol. 14, no. 2, pp. 507, 508, pl. 62, figs. 1, 2. Description: Zoaria small nodular. Zooecia rough, raised, with areolar pores partially obscured by secondary calcification. Aperture oval, rounded above, with a proximal sinus that is ‘“V” shaped in the well preserved specimens. Peristome with a pair of oral avicularia, with a sear of an ocea- sional third avicularia distally. Interzooecial avicularia wide, ovoid. Ovi- cells wanting on our specimen. 130 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. MEASUREMENTS: Aperture, 156/104 mp. Zooecia, 543/248 muy. Locauities: LACMIP loe. 76, “lumber yard,” San Pedro, California, Pleistocene. OccuRRENCE: Pleistocene: Santa Monica [Rustic Canyon], California, (Canu and Bassler, 1923). Recent: Pacific: California (Robertson, 1908) ; Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Hancock stations, California, Channel Islands, Mexico. DeptH: Shore (intertidal) to 50 fathoms (12 records). Sub-order CycLostomata Busk, 1852 Division |, TUBULIPORINA Hagenow, 1851 Family ONcousoEcnHDAE Canu, 1918 Genus Proboscina Audouin, 1826 Proboscina major (Johnston), 1847. Alecto major JOHNSTON, 1847, Hist. Brit. Zoophytes, 2nd Ed., p. 281, pl. 49, figs. 3, 4. Stomatopora major, Hincxs, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 13, p. 204. Stomatopora major, O'DONOGHUE and O’DoNOGHUE, 1923, Contr. Canad. Biol., nr. s., voll, no. 10; p. 153: Diaperoecia major, O'DONOGHUE and O’DonoGHuEr, 1926, Contr. Canad. Biol. Fish., Nn. S- Vol: 3, NO: 3; Ds 69: Oncouoecia (Proboscina) major, CANU and BASSLER, 1930, Proc. U. S. Nat. Mus., no. 2810, vol. 76, art. 13, pp. 46, 47. Proboscina major, OSBURN, 1953, Allan Hancock Pac. Expds., vol. 14, no. 3, pp. 621, 622, pl. 65, fig. 5. DESCRIPTION: Zoaria incrusting. Zooecia tubular, erect. Apertures free, arranged in rows of one or two zooecia in linear succession. Ovicell on our specimen broken, low, elongate, bulbous. MEASUREMENTS: Diameter peristome, 460 my. Diameter aperture 187 mu. LocaLities: West Second and Pacific Avenue, San Pedro, California, Pleistocene. OCCURRENCE: Pliocene: England, Italy (Waters, 1887). Recent: Pa- cific: Queen Charlotte Islands (Hincks, 1884); Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Hawaii (Canu and Bassler, 1927); Galapagos Islands (Canu and Bassler, 1930); Hancock stations, Mexico, Ecuador, Galagapos Islands. DeptH: Shore (intertidal) to 170 fathoms (14 records). Vou. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 131 Remarks: This is the first record of this species as fossil on the Pacific coast of North America. Genus Filisparsa d’Orbigny, 1853 Filisparsa clarki Canu and Bassler, 1923. Filisparsa clarki CANu and BASSLER, U. S. Nat. Mus. Bull. 125, pp. 195, 196, pl. 41, figs. 11-19. Description: Zoaria erect, branching, with transverse dorsal striations. Zooecia connate, except at distal ends, which rise as tubular extensions. MEASUREMENTS: Diameter peristome, 207 mp. Diameter aperture, 126 mp. Locauitires: Bath-house Beach, Santa Barbara, California, Pleistocene. OCCURRENCE: Pleistocene: Santa Barbara, California, Dead Man’s Island, San Pedro [harbor], California (Canu and Bassler, 1923). Depry: At this time there are no records of this species in living form. Family DIAPEROECHDAE Canu, 1918 Genus Diaperoecia Canu, 1918 Diaperoecia californica (d’Orbigny), 1852. Idmonea californica D’ORBIGNY, 1852, Paleon. Francaise, terr. cret., vol. 5, Bryo- zoaires, p. 732. Idmonea californica, Conrad, 1855, Proc. Acad. Nat. Sci., Phila., vol. 7, p. 441. Idmonea californica, GABB and Horn, 1862, Jour. Acad. Nat. Sci., Phila., ser. 2, vol. 5, pt. 2, pp. 168, 169, pl. 21, fig. 56. Idmonea californica, ROBERTSON, 1910, Univ. Calif. Pub. Zool., vol. 6, no. 12, pp. 253, 254, pl. 23, figs. 39-41. Idmonea californica, CANU and BASSLER, 1923, U. S. Nat. Mus. Bull. 125, p. 199, pl. 43, figs. 1-9. Idmonea californica, O’DoNoGHUE and O’DONOGHUE, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 154. Idmonea californica, O’DONoGHUE and O’DONOGHUE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3, no. 3, pp. 73, 74. Diaperoecia californica, OsBuRN, 1953, Allan Hancock Pac. Expd., vol. 14, no. 3, pp. 642-644, pl. 67, figs. 1, 2. DescripTION: Zoaria erect. Zooecia tubular, long, extending one behind the other. Zooecial openings may be found in transverse rows, with as many as five or six to a row. 132 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. MEASUREMENTS: Diameter aperture 207 mp. Diameter peristome 312 mu. LocaLities: Bath-house Beach, Santa Barbara, California, Pleistocene. West Second and Pacific Avenue, San Pedro, California, Pleistocene. LACMIP loe. 130-7, opposite Berth 79 Los Angeles harbor, Timm’s Point formation, Pleistocene. LACMIP loe. 76, “lumber yard,” San Pedro, Cali- fornia, A. T. Tieje, collector, Pleistocene. OccURRENCE: Pleistocene: Santa Barbara, California (Conrad, 1855, Gabb and Horn, 1862); San Pedro, California (Conrad, 1855); Santa Monica, Santa Barbara, California, Dead Man’s Island, San Pedro, Cali- fornia (Canu and Bassler, 1923). Recent: Pacific: California (d’Orbigny, 1852, Robertson, 1910); Vancouver Island region (O’Donoghue and O’Don- oghue, 1923, 1926); Hancock stations, California, Channel Islands, Mex- ico, Gulf of California, Costa Rica. DeptH: Shore (intertidal) to 150 fathoms (84 records). Family TuBuLiporiDAE Johnston, 1838 Genus Tubulipora Lamarck, 1816 Tubulipora tuba (Gabb and Horn), 1862. Semitubigera tuba Gass and Horn, 1862, Jour. Acad. Nat. Sci., Phila., ser. 2, vol. 5, Dt-2) Ds 169s pls 2i) fist ebi. Tubulipora occidentalis Rosertson, 1910, Univ. Calif. Pub. Zool., vol. 6, no. 12, p. 249, pl. 22, figs. 29-31. Tubulipora occidentalis, CANU and Basster, 1920, U. S. Nat. Mus. Bull. 106, p. 757, fig. 246 k. Tubulipora tuba, CANU and Basster, 1923, U. S. Nat. Mus. Bull. 125, p. 198, pl. 42, figs. 18-23. Tubulipora occidentalis, O’ DONOGHUE and O’DonoGuur, 1923, Contr. Canad. Biol., nm. s., vol. 1, no. 10; p. 150. Tubulipora tuba, O’DoNoGHUE and O’DoNnoGHuUE, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 95. Tubulipora tuba, O’DonoGHUE and O’DonocuvsE, 1926, Contr. Canad. Biol. Fish., n. s., vol. 3; no: 3, pp. “0; fae Tubulipora tuba, OspuRN, 1953, Allan Hancock Pac. Expd., vol. 14, no. 3, pp. 650, pl. 68, fig. 9. Description: Zoaria incrusting. Zooecia tubular, erect, arranged mono- serially or biserially, with many tubes. Ovicells globose, spreading among two or three series of tubes. Ooeciostome short, tubular, erect, located near first zooecial tube of series. VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 133 MEASUREMENTS: Diameter peristome, 126 my. Diameter aperture, 70 mp. LocauitIes: Bath-house Beach, Santa Barbara, California, Pleistocene. West Second and Pacific Avenue, San Pedro, California, Pleistocene. OccuRRENCE: Pleistocene: Santa Barbara, California (Gabb and Horn, 1862); Santa Monica, Santa Barbara, Dead Man’s Island, San Pedro [har- bor], California (Canu and Bassler, 1923). Recent: California (Robert- son, 1910); Vancouver Island region (O’Donoghue and O’Donoghue, 1923, 1926) ; Puget Sound (Robertson, 1910, O’Donoghue and O’Donoghue, 1925) ; Haneock stations, Channel Islands. DeptH: 4 to 81 fathoms (9 records). Tubulipora tuba var. fasciculifera (Hincks), 1884. Tubulipora fasciculifera Hincks, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 13, pl. 9, figs (6: Tubulipora occidentalis Ropertson, 1910, (pars), Univ. Calif. Pub. Zool., vol. 6, no. 12, p. 249, pl. 22, figs. 29-31. Tubulipora fasciculifera, CANU and BASsLER, 1923, U. S. Nat. Mus. Bull. 125, p. 197, pl. 42, figs. 9-17. Tubulipora fasciculifera, O’DONOGHUE and O’DonoGHur, 1923, Contr. Canad. Biol., n. s., vol. 1, no. 10, p. 150. Tubulipora fasciculifera, O’DONoGHUE and O’DoNoGHuUER, 1925, Univ. Wash., Pub. Puget Snd. Biol. Sta., Trans., vol. 5, p. 95. Tubulipora fasciculifera, O'DONOGHUE and O’DonoGHuE, 1926, Contr. Canad. Biol. INTs, We Shy AWOL, Bi, wo, 3 105 70s Tubulipora tuba var. fasciculifera, OSBURN, 1953, Allan Hancock Pac. Expd., vol. 14, no. 3, pp. 651, 652, pl. 68, fig. 10. DeEscrIPTION: Zoaria flat, incrusting, our specimens fragmentary. Zooecia tubular, radiating, erect, free, arranged monoserially or biserially with few tubes. Ovicells small, placed among the rows of zooecial tubes. Our specimens do not show intact ooeciostomes. MEASUREMENTS: Diameter peristome, 138 my. Locauitres: Bath-house Beach, Santa Barbara, California, Pleistocene. LACMIP loc. 66-2, Newport Harbor mesa, Newport, California, Pleisto- cene. OCCURRENCE: Pleistocene: Santa Barbara, California (Canu and Bass- ler, 1923). Recent: Pacific: Queen Charlotte Islands (Hincks, 1884); Van- ecouver Island region (O’Donoghue and O’Donoghue, 1923, 1926); Puget Sound (O’Donoghue and O’Donoghue, 1925); Hancock stations, California, Channel Islands. DeptH: 8 to 131 fathoms (16 records). 134 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Division II], ARTICULATA Busk, 1859 Family Crisupar Johnston, 1838 Genus Crisia Lamouroux, 1812 Crisia serrulata Osburn, 1953. Crisia serrulata OspurN, 1953, Allan Hancock Pac. Expd., vol. 14, no. 3, pp. 679, 680, pl. 72, fig. 2. Crisina serrata GAsBB and Horn, 1862, Jour. Acad. Nat. Sci., Phila., ser. 2, vol. 5, Pte) pp ita. a5, pl. 21h ne. 166: Crisia pacifica Ropertson, 1910, Univ. Calif. Pub. Zool., vol. 6, no. 12, pp. 242, 243, pl. 20, figs. 16, 17. Crisia serrata, CANU and BAssLer, 1923, U. S. Nat. Mus. Bull, 125, pp. 196, 197, pl. 42, figs 1-7. Crisia pacifica, O’DoNoGHUE and O’DonoGHuE, 1923, Contr. Canad. Biol., n. s., vol. 1, p. 149. Crisia serrata, O’DONOGHUE and O’DonoGHUE, 1926, Contr. Canad. Biol., Fish., n. s., vol. 3, no. 3, p. 64. Description: Zoaria caleareous, erect, with a low longitudinal keel along the mid-dorsal line. Zooecia tubular, connate. Ovicell prominent, adhering in its entire length to the ventral zoarial wall. MEASUREMENTS: Diameter peristome, 103 mp. Diameter aperture, 92 mp. Diameter zooecia, 138 muy. LocauitiEs: Bath-house Beach, Santa Barbara, California, Pleistocene. West Second and Pacific Avenue, San Pedro, California, Pleistocene. OccuRRENCE: Pleistocene: Santa Barbara, California (Gabb and Horn, 1862); Santa Barbara, Santa Monica, California (Canu and Bassler, 1923). Recent: California (Robertson, 1910); Vancouver Island region (O’Don- oghue and O’Donoghue, 1923, 1926); Hancock stations, Oregon, California, Channel Islands, Gulf of California, Mexico, Galapagos Islands. DeptH: 8 to 86 fathoms (26 records). SUMMARY In reviewing the material covered in this report, we find that of 56 species of Bryozoa noted, the following 19 have not previously been re- ported from the fossil record: Conopeum commensale Kirkpatrick and Metzelaar, 1922; Antropora tincta (Hastings), 1930; Microporina borealis (Busk), 1855; Labioporella sinuosa Osburn, 1940; Thalamoporella califor- nica (Levinsen), 1909; Amastigia rudis Busk, 1852; Scrupocellaria regularis Vou. XXIX ] SOULE AND DUFF: FOSSIL BRYOZOA 135 Osburn, 1940; Scrupocellaria varvans Hincks, 1882; Hippopodina feegeensis (Busk), 1884; Schizoporella linearis inarmata (Hineks), 1884; Schizo- mavella auriculata cuspidata (Hineks), 1880; Hippodiplosia insculpta (Hineks), 1882; Hippoporella gorgonensis Hastings, 1930; Parasmittina californica (Robertson), 1908; Parasmittina collifera (Robertson), 1908; Mucronella microstoma (Norman), 1864; Rhynchozoon tumulosum (Hinecks), 1882; Hippopodinella adpressa (Busk), 1854; Holoporella brun- nea (Hineks), 1884. In addition to those listed above, we have recorded another 10 species that have been reported from the fossil record prior to this time, but not from the Pacific coast of America. Finally, there are 4 species, namely, Schizomavella auriculata cuspidata (Hincks), 1880; Hippoporidra edax (Busk), 1859; Mucronella microstoma (Norman) 1864; and Filisparsa clarki Canu and Bassler, 1923, that are as yet unreported as living forms from the waters of the Pacific coast of North America. LITERATURE CITED ALDER, JOSHUA 1856. A catalogue of the zoophytes of Northumberland and Durham. T'rans- actions of the Tyneside Naturalist’s Field Club, vol. 3, pt. 2, pp. 93- 162, pls. 3-10. Barroso, M. G. 1912. Briozoos de la estacion de biologia maritima de Santander. Trabajos del Museo National de Ciencias Naturales, no. 5 (Zool. ser. no. 3), 1912, pp. 1-63. 1922. Notas sobre briozoos marinos espanoles, X. Especies de Mahon, Ballares. Boletin de la Real Sociedad Espanola de Historia Natural, Madrid, vol. 22, no. 1, pp. 88-101, 8 figs. BASSINDALE, R. 1941. The invertebrate fauna of the southern shores of the Bristol Channel and Severn Estuary. Proceedings of the Bristol Naturalist’s Society, 4th series, vol. 9, pt. 2, 1940, pp. 143-201. BIDENKAP, OLAF 1905. Fortegnelse over de arktiske bryozoer. Bergens Museums Aarbog, no. 9, pp. 1-79. Bore, FOLKE 1930. On the bryozoan fauna of Skelderviken. Arkiv for Zoologi, band 21A, no. 24, pp. 1-13. 1933. Die Bryozoen III Teil, Die marinen Bryozoen (Stenolaemata und Gymn- nolaemata) des arktischen Gebietes. Fauna Arctica, band 6, V Lief- erung, pp. 515-551. 136 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH SER. Brock, HJALMAR 1927. Untersuchung iiber die Marine Bodenfauna bei Lindeness im Juni 1926. Avhandlinger utgitt av Det Norske Videnskaps-Akademie i Oslo. I Matem.-Naturv. Klasse, no. 5, pp. 1-382. Busk, GEORGE 1852. An account of the Polyzoa, and sertularian zoophytes, collected in the voyage of the Rattlesnake, on the coasts of Australia and the Louisiade Archipelago. Appendix 4, in MacGillivray, John, Narrative of the Voyage of H.M.S. Rattlesnake, vol. 1, pp. 343-402, pl. 1. 1852. Catalogue of Marine Polyzoa in the British Museum, pt. 1, Cheilostomata, pp. I-VIII, 1-54, pls. 1-68. 1854. Catalogue of the Marine Polyzoa in the British Museum, pt. 2, Cheilo- stomata, pp. 55-120, pls. 69-124. 1855. Zoophytology. Quarterly Journal of Microscopical Science, vol. 3, pp. 253-256, pls. 1 & 2. 1856. Zoophytology. Quarterly Journal of Microscopical Science, vol. 4, pp. 93-96, pls. 5 & 6, pp. 176-180, pls. 7 & 8, pp. 308-312, pls. 9-12. 1857. Zoophytology. Quarterly Journal of Microscopical Science, vol. 5, pp. 172-174, pls. 15 & 16. 1858. On some Madeiran Polyzoa collected by J. Yates Johnson, Esq. Quarterly Journal of Microscopical Science, vol. 6, pp. 261-264, pls. 20 & 21. 1859. A monograph of the fossil Polyzoa of the Crag. The Paleontological Society of London, pp. 1-136, pls. 1-22. 1861. Description of new Polyzoa collected by J. Y. Johnson, Esq., at Madria in the years 1859 and 1860. Quarterly Journal of Microscopical Science, vol. 1, new series, pp. 77-80, pls. 32 & 33. 1861. Description of new or imperfectly known Polyzoa, no. 1. Quarterly Jour- nal of Microscopical Science, vol. 1, new series, pp. 153-156, pls. 34 & 35. 1884. Report on the Polyzoa collected by H.M.S. Challenger during 1873-1876. Report on the Scientific Results of the voyage of H.M.S. Challenger 1873-1876, Zoology, vol. 10, part 30, Polyzoa, pp. I-XXIV, pp. 1-216, pls. 1-36. CALVET, L. 1902. Materiaux pour servir a l’histoire de la faune des Bryozoaires des cotes francaises II, Bryozaires marine des cotes de Corse. Travauz de V'Insti- tut de Zoologie de VUniversite de Montpellier, 2nd ser., mem. 12, pp. 1-52, 2 pls. 1907. Bryozoaires, Expeditions scientifiques du “Traivailleur et du Talisman” pendant les anness 1880, 1881, 1882, 1883, vol. 8, pp. 355-495, pls. 26-30, Paris. 1931. Bryozoaires provenant des campagnes scientifiques du Prince Albert I de Monaco. Results Campagnes scientifique par Albert I, fase. 83, pp. 1-152, 2 pls. CANU, FERDINAND 1924. Bryozoaires in Dollfuss, R. P., Contribution a la faune des invertebres du bane de Rockall. Bulletin de l'Institut oceanographique de Monaco, no. 438, pp. 1-28 (Bryozoa pp. 13-17). VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 137 Canu, FERDINAND, and Ray S. BASSLER 1920. North American Early Tertiary Bryozoa. United States National Mu- seum Bulletin 106, two vols. text pp. 1-879, with 279 text figs., pls. 1-162. 1923. North American Later Tertiary and Quaternary Bryozoa. United States National Museum Bulletin 125, pp. 1-244, pls. 1-47. CLARK, ALEX 1931. The cool-water Timms Point Pleistocene horizon at San Pedro, Cali- fornia. Transactions of the San Diego Society of Natural History, vol. 7, No. 4, pp. 25-42. Desor, E. 1848. Ascidioidian polyps or Bryozoa (from Nantucket). Proceedings of the Boston Society of Natural History, vol. 3, 1848-1851, pp. 66 and 67. EYRE, JOYCE 1939. The South African intertidal zone and its relation to ocean currents. Annals of the Natal Museum, vol. 9, pt. 2, pp. 283-306, pls. 18-22. Gaps, Wn. M. and G. H. Horn 1862. Monograph of the fossil Polyzoa of the Secondary and Tertiary forma- tions of North America. Journal of the Academy of Natural Sciences of Philadelphia, new series, vol. 5, pt. 2, art. 3, pp. 111-179, pls. 19-21. HARMER, SIDNEY F. 1902. On the morphology of the Cheilostomata. Quarterly Journal of Micro- scopical Science, new series, vol. 46, no. 182, pp. 263-350, pls. 15-18. 1915. The Polyzoa of the Siboga Expedition. Siboga Expeditie, vol. 28 a, pt. 1, Entoprocta, Ctenostomata and Cyclostomata, pp. 1-180, pls. 1-12. 1926. The Polyzoa of the Siboga Expedition. Siboga Expeditie, vol. 28 b, pt. 2, Cheilostomata Anasca, pp. 181-501, pls. 13-34. 1934. The Polyzoa of the Siboga Expedition. Siboga Expeditie, vol, 28 ¢, pt. 3, Cheilostomata Ascophora. I, pp. 503-640, pls. 35-41. HASSALL, ARTHUR H. 1841. Supplement to a catalogue of Irish zoophytes. Annals and Magazine of Natural History, ser. 1, vol. 7, pp. 276-287, 363-373, pls. 6-10. Hastines, A. B. 1927. Zoological results of the Cambridge Expedition to the Suez Canal, 1924. Report on the Polyzoa. Transactions of the Zoological Society of London, vol. 22, pt. 3, pp. 3381-354, text figs. 82-88. 1930. Cheilostomatous Polyzoa from the vicinity of the Panama Canal, col- lected by Dr. C. Crossland on the cruise of the S.Y. “St. George.” Pro- ceedings of the Zoological Society of London, 1929, pt. 4, no. 53, pp. 697-740, pls. 1-17. HASWELL, WM. A. 1880. On some Polyzoa from the Queensland coast. Proceedings of the Lin- nean Society of New South Wales, vol. 5, pt. 1, pp. 33-44, pls. 1-3. 138 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. HELLER, C. 1867. Die Bryozoen des adriatischen Meeres. Verhandlungen Zoologisch- botanischen Gesellschaft in Wien, Jahrgang 1867, band 17, pp. 77-136, pls. 1-6. HINcKS, THOMAS 1862. A catalogue of the zoophytes of South Devon and South Cornwall. An- nals and Mazagine of Natural History, ser. 3, vol. 9, pp. 22-30, pl. 7, pp. 200-207, pl. 12, pp. 303-310, pl. 12, pp. 467-475, pl. 16. 1877. On British Polyzoa, Part I. Annals and Magazine of Natural History, ser. 4, vol. 20, pp. 212-218. 1877. On British Polyzoa, Part II Classification. Annals and Magazine of Nat- ural History, ser. 4, vol. 20, pp. 520-532. 1880. Contributions toward a general history of the marine Polyzoa. Annals and Magazine of Natural History, ser. 5, vol. 6, pp. 69-91, pls. 9-11, pp. 376-384, pls. 16 & 17. 1880. A history of British marine Polyzoa, vol. I text, pp. I-CXLI, 1-601, vol. II plates 1-83. 1881. Contributions toward a general history of the marine Polyzoa. Annals and Magazine of Natural History, ser. 5, vol. 7, pp. 147-161, pls. 8-10. 1881. Contributions toward a general history of the marine Polyzoa. Annals and Magazine of Natural History, ser. 5, vol. 8, pp. 1-14, pls. 1-14. 1882. Polyzoa of the Queen Charlotte Islands: Preliminary Notice of new species. Annals and Magazine of Natural History, ser. 5, vol. 10, pp. 248-256. 1882. Report on the Polyzoa of the Queen Charlotte Islands. Annals and Maga- zine of Natural History, ser. 5, vol. 10, pp. 459-471, pls. 19 & 20. 1883. Report on the Polyzoa of the Queen Charlotte Islands. Annals and Maga- zine of Natural History, ser. 5, vol. 11, pp. 442-451, pls. 17 & 18. 1884. Report on the Polyzoa of the Queen Charlotte Islands. Annals and Maga- zine of Natural History, ser. 5, vol. 13, pp. 49-58, pls. 3 & 4, pp. 203-215, pl. 9. 1884. Note on Professor G. Seguenza’s list of Tertiary Polyzoa from Reggio (Calabria). Annals and Magazine of Natural History, ser. 5, vol. 13, pp. 265-267. 1884. Contributions toward a general history of the marine Polyzoa. Annals and Magazine of Natural History, ser. 5, vol. 13, pp. 356-369, pls. 13 & 14. 1886. The Polyzoa of the Adriatic: a supplement to Prof. Heller’s ‘Die Bryo- zoen des adriatischen Meeres,” 1867. Annals and Magazine of Natural History, ser. 5, vol. 17, pp. 254-271, pls. 9 & 10. 1887. Critical notes on the Polyzoa. Annals and Magazine of Natural History, ser. 5, vol. 19, pp. 150-164. 1887. The Polyzoa of the Adriatic: a supplement to Prof. Heller’s “Die Bryo- zoen des adriatischen Meeres,” 1867. Annals and Magazine of Nat- ural History, ser. 5, vol. 19, pp. 302-316, pl. 9. 1888. The Polyzoa of the St. Lawrence: a study of arctic forms. Annals and Magazine of Natural History, ser. 6, vol. 1, pp. 214-227, pls. 14 & 15. VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 139 Hincks, THomas—Cont’d 1889. The Polyzoa of the St. Lawrence: a study of arctic forms. Annals and Magazine of Natural History, ser. 6, vol. 3, pp. 424-433, pl. 21 1891. Contributions toward a general history of the marine Polyzoa, 1880-91, Appendix. Annals and Magazine of Natural History, ser. 6, vol. 8 pp. 86-93, 169-176, 471-480. 1893. Contributions toward a general history of the marine Polyzoa, 1880-91. Appendix. Annals and Magazine of Natural History, ser. 6, vol. 12, pp. 140-147. , Howe, HENrRy VY. 1942. Fauna of the Gendon formation at its type locality. Journal of Paleon- tology, vol. 16, no. 2, pp. 264-271. HutTcHins, L. W. 1945. An annotated check-list of the saltwater Bryozoa of Long Island Sound. Transactions of the Connecticut Academy of Arts and Sciences, vol. 36, pp. 533-549. JOHNSTON, GEORGE 1825. Contributions to the British fauna. Edinburgh Philosophical Journal, XIII, 1825, pp. 218-222. 1838. A descriptive catalogue of the recent zoophytes found on the coast of North Durham. Transactions of the Natural History Society of North- umberland, Durham and Newcastle upon Tyne, vol. 2, pp. 239-272, pls. 7-12. 1838. A history of British Zoophytes, pp. 3-341, pls. 1-44. 1847. A history of British Zoophytes, 2nd ed., vol. 1, text pp. 1-488, vol. 2 plates 1-74. JONES, N. L. 1939. Some recent additions to the off-shore fauna of Port Erin. Report for 1939 (no. 52) of the Marine Biological Station at Port Erin, Isle of Man, pp. 18-32. JULLIEN, J, and Louris CALVET 1903. Bryozoaires provenant des compagnes de l’Hirondelle (1886-1888). Re- sultats des Campagnes Scientifique par Albert I, fase. 23, pp. 1-188, pls. 1-28. KENDALL, Percy F. 1931. The red crag of Walton-on-the-Naze. Geological Magazine, vol. 68, no. 807, pp. 405-420, 1 fig. KIRKPATRICK, R. 1888. Polyzoa of Mauritius. Annals and Magazine of Natural History, ser. 6, vol. 1, pp. 72-84, pls. 7-10. 1890. Polyzoa in Ridley, H. N., Notes on the zoology of Fernando Noronha. Journal of the Linnean Society of London, Zoology, vol. 20, nos. 124— 125, pp. 478-570 (Bryozoa pp. 504-506), figs. 1 & 2. 140 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. KIRKPATRICK, R., and J. MrETZELAAR 1922. On an instance of commensalism between a hermit crab and a polyzoon. Proceedings of the Zoological Society of London, 1922, pp. 983-990, pls. 1 & 2. Lacourt, A. W. 1949. Bryozoa of the Netherlands. Archives Neerlandaises de Zoologie, tome 8, livraison 3, pp. 1-33, pls. 13-17, text figs. 1-5. LEVINSEN, G. M. R. 1894. Mosdyr, in Zoologia Danica (Schiodte), vol. 4, afd. 1, pp. 1-105, pls. 1-9. 1909. Morphological and systematic studies on the cheilostomatous Bryozoa, Kjobenhavn, pp. I-VII, 1-431, pls. 1-27. LIVINGSTONE, A. A. 1926. Studies on Australian Bryezoa. Records of the Australian Museum, vol. 5, no. 1, pp. 79-99, pls. 5-8. 1927. Studies on Australian Bryozoa. Records of the Australian Museum, vol. 6, no. 1, pp. 50-69. MacGibuivray, P. H. 1859. On some new Australian Polyzoa. Transactions of the Philosophical Institute of Victoria, vol. 4, art. 10, pp. 97, 98, pl. 1, Melbourne. 1887. A catalogue of the marine Polyzoa of Victoria. Transactions and Pro- ceedings of the Royal Society of Victoria, vol. 23, art. 16, pp. 187-224, pls. 1 & 2. 1887. Polyzoa, in McCoy, Prodromus of the Zoology of Victoria, vol. 2, decade 14, pp. 187-150, pls. 136-138. 1891. Descriptions of new or little known Polyzoa. Proceedings of the Royal Society of Victoria, vol. 3, (new series), pp. 77-83, pls. 9 & 10. 1895. A monograph of the Tertiary Polyzoa of Victoria. Transactions of the Royal Society of Victoria, vol. 4, 1895, pp. 1-166, pls. 1-22. MANZONI, A. 1875. Le conchiglie morte e i Briozi del lido presso Venezia. Atti della So- cieta Toscana di Scienze Naturali, vol. 2, fasc. 1, pp. 65-67. Marcus, E. 1920. Mittelmeer-Bryozoen aus der Sammlung des Zoologischen Museums zu Berlin. Sitzwngsberichte der Gesellschaft Naturforschender Freunde zu Berlin, no. 2 und 3, pp. 65-108. 1937. Bryozoarios marinhos Brasileiros I. Boletins da Faculdade de Philosophia Sciencias e Letras, Zoologica no. 1, pp. 1-224, pls. 1-29. 1938. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16, LXIX Bryozoa from St. Helena. Videnskabelige Meddelelser fra Dansk Nat- urhistorisk Forening i Kobenhavn, bind 101, pp. 183-252, text figs. 1-32. 1939. Bryozoarios marinhos Brasileiros III. Boletins da Faculdade de Filo- sofia, Ciencias e Letras Universidade de Sdo Paulo, XIII, Zoologia no. 3, pp. 111-300, pls. 5-31. VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 141 Marcus, E.—Cont’d 1950. Systematical remarks on the bryozoan fauna of Denmark. Videnska- belige Meddelelser fra Dansk naturhistorisk Forening i Kobenhavn, bind 112, 1950, pp. 1-34, 5 text figs. 1953. Notas sobre briozoos marinhos Brasileiros. Arquivos do Museu Nacional, vol. XLII, pp. 273-342, pls. 1-8. NEVIANI, A. 1891. Contribuzione all conoscenza dei briozoi fossili Italiani—Briozoi post plio- cenici del sottosuolo di Livorno. Bolletino della Societa Geologica Italiana, vol. 10, pp. 99-148, 1 pl. 1896. Briozoi postplioceni di Spilinga (Calabria). Atti della Accademia Gioe- nia di Scienze Naturali in Catania, ser. 4, vol. 9, memoria 9, pp. 1-66, 382 text figs. 1900. Briozoi terziari e postterziari della Toscana. Bolletino della Societa Geologica Italiana, vol. 19, pp. 349-375. NORDGAARD, O. 1896. Systematisk fortegnelse over de i Norge hidtil observerde arter af marine polyzoa, I Cheilostomata. Bergens museum Aarbog for 1894-95, no. 2, pp. 1-33, pls. 1 & 2. 1906. Die Bryozoen des westlichen Norwegens. Bergens Museum, Meeresfauna von Bergen, hett 2, pp. 73-107, pls. 1 & 2. 1907. Bryozoen von dem norwegischen Fischereidampfer “Michael Sars” in den Jahren 1900-1904 gesammelt. Bergens Museum Aarbog, 1907, no. 2, pp. 1-21, 1 pl. 1912. Revision au universitetamuseets samling av Norske Bryozoer. Det Kongelige Norske Videnskabers Selskabs Skrifter, 1911, no. 3, pp. 1-27. 1918. Bryozoa from the arctic regions. Tromso Museums Aashefter, 40, 1917, no. 1, pp. 1-99. 1927. The Folden Fiord Bryozoa. Tromso Museums Skrifter, vol. 1, pt. 9, pp. 1-10. NORMAN, ALFRED M. 1864. On undescribed British Hydrozoa, Actinozoa and Polyzoa. Annals and Magazine of Natural History, ser. 3, vol. 13, pp. 82-90, pls. 9-11. 1894. A month on the Trondhjem Fiord. Annals and Magazine of Natural History, ser. 6, vol. 13, pp. 112-133, pls. 6 & 7. 1906. Greenlandic Polyzoa. Annals and Magazine of Natural History, ser. 7, vol. 17, pp. 90-93. 1909. The Polyzoa of Madeira and neighbouring islands. Journal of the Linnean Society of London, Zoology, vol. 30, pp. 275-312, pls. 33-42. O’DonocuurE, C. H., and EL_sir O’DONOGHUE 1923. A preliminary list of Bryozoa (Polyzoa) from the Vancouver Island region. Contributions to Canadian Biology, new series, vol. 1, no. 10, pp. 143-201, pls. 1-4. 1925. List of Bryozoa from the vicinity of Puget Sound. Puget Sound Biological Station, Transactions, vol. 5, pp. 91-108. 142 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. O’DonoGHUE, C. H., and Etstr O’DonogHuE—Cont’d 1926. A second list of Bryozoa (Polyzoa) from the Vancouver Island region. Contributions to Canadian Biology and Fisheries, new series, vol. 3, no. 3, pp. 49-131, pls. 1—5. O’DonoGHuE, C. H., and DorRA DEWATTEVILLE 1935. A collection of Bryozoa from South Africa. Journal of the Linnean Society of London, Zoology, vol. 39, no. 265, pp. 203-218, pls. 5 & 6. OKADA, Y. 1929. Report of the Biological Survey of Mutsu Bay, 12. Cheilostomatous Bryozoa of Mutsu Bay. Science Reports of the Tohoku Imperial Uni- versity, 4th ser. (Biology), vol. 4, no. 1, fase. 1, pp. 11-35, pls. 1-5. 1933. On a collection of Bryozoa from the Northern Kurile Expedition. Bul- letin of the Biogeographic Society, Tokyo, vol. 4, no. 3, pp. 213-216. 1934. Bryozoa fauna in the vicinity of the Shimoda Marine Biological Station. Science Reports of the Tokyo Bunrika Daigaku, sect. B, no. 26, vol. 2, pp. 1-20, pls. 1 & 2. OKADA, Y., and SuHizuo MAWATARI 1935. Bryozoa fauna collected by the ‘‘Misago” during the zoological survey around Izu Peninsula (I). Science Reports of the Tokyo Bunrika Daigaku, sect. B, no. 35, vol. 2, pp. 127-147, pls. 10 & 11. 1936. Bryozoa fauna collected by the ‘“Misaga” during the zoological survey around Izu Peninsula (II). Science Reports of the Tokyo Bunrika Daigaku, sect. B, no. 49, vol. 3, pp. 53-73, pls. 9 & 10. 1938. On the collection of Bryozoa along the coast of Wakayama-ken, the middle part of Honsyo, Japan. Annatationes Zoologicae Japonenses, vol. 17, nos. 3 & 4, pp. 445-462, 7 text figs. D’ORBIGNY, A. 1847. Voyage dans l’Amerique meridionale. Zoophytes, vol. 5, part 4, pp. 7-28. pls. 1-13. ORTMANN, A. 1890. Die Japanische Bryozoenfauna. Archiv fiir Naturgeschte, vol. 56, band 1, pp. 1-74, pls. 1-4. OSBURN, RAYMOND C. 1912. The Bryozoa of the Woods Hole region. Bulletin of the Bureau of Fish- eries, vol. 30, 1910, document no. 760, pp. 203-266, pls. 18-31. 1914. The Bryozoa of the Tortugas Islands, Florida. Papers from the Tortugas Laboratory of the Carnegie Institution of Washington, vol. 5, pub. 182, paper no. 11, pp. 181-222, 23 text figs. 1919. Bryozoa of the Crocker Land Expedition. Bulletin of the American Mu- seum of Natural History, vol. 41, art. 19, pp. 603-624. 1923. Bryozoa of the Canadian Arctic Expedition. Report of the Canadian Arc- tic Expedition 1913-18, vol. 8, pt. D, pp. 1-13. VoL. XXIX | SOULE AND DUFF: FOSSIL BRYOZOA 143 OspuRN, RAYMOND C.—Cont’d 1932. Biological and oceanographic conditions in Hudson Bay 7, Bryozoa from Hudson Bay and strait. Contributions to Canadian Biology and Fisheries, n. s. vol. 7, no. 29, (series A, general, no. 26), pp. 361-376, 1 fig. 1933. Bryozoa of the Mount Desert region. Biological Survey of the Mount Desert Region, Wistar Institute 1933, pp. 3-66, pls. 1-15. 1936. Bryozoa collected in the American arctic by Captain R. A. Bartlett. Journal of the Washington Academy of Sciences, vol. 26, no. 12, pp. 538-543, 1 text fig. 1940. Bryozoa of Porto Rico with a resume of the West Indian bryozoan fauna. New York Academy of Sciences, Scientific Survey of Porto Rico and the Virgin Islands, vol. 16, pt. 3, pp. 321-486, pls. 1-9. 1944. A survey of the Bryozoa of Chesapeake Bay. Chesapeake Biological Lab- oratory Publication no. 63, pp. 3-58, pls. 1-5, text figs. 1-28. 1947. Bryozoa of the Allan Hancock Atlantic Expedition, 1939. Allan Hancock Atlantic Expedition, Report no. 5, pp. 1-66, pls. 1-6. 1950. Bryozoa of the Pacific Coast of America, pt. 1, Cheilostomata-Anasca. Allan Hancock Pacific Expeditions, vol. 14, no. 1, pp. 1-269, pls. 1-29. 1952. Bryozoa of the Pacific Coast of America, pt. 2, Cheilostomata-Ascophora. Allan Hancock Pacific Expeditions, vol. 14, no. 2, pp. 271-611, pls. 30-64. 19538. Bryozoa of the Pacific Coast of America, pt. 3, Cyclostomata, Ctenosto- mata, Entoprocta and Addenda. Allan Hancock Pacific Expeditions, vol. 14, no. 3, pp. 613-841, pls. 65-82. PACKARD, A. S. JR. 1863. A list of animals dredged near Caribou Island, Southern Labrador, dur- ing July and August 1860. Canadian Naturalist and Geologist, vol. 8, no. 6, art. 28, pp. 401-429, pls. 1 & 2. PALLAS, P. S. 1766. Elenchus Zoophytorum, pp. 3-451. PRENANT, MARCEL 1931. Sur une collection de bryozoaires de la Mer du Nord appartenant au Musee royal d’Histoire Naturelle de Belgique. Bulletin du Musee royal d Histoire Naturelle de Belgique, tome 7, no. 17, pp. 1-6. PRENANT, MARCEL, and GEORGES TEISSIER 1924. Notes ethologiques sur la faune marine sessile des environs de Roscoff. Cirripedes, Bryozaires, Hydraires. Travaux de la Station Biologique de Roscoff, fase. 2, pp. 1-49. RIcHARDS, H. G., and ANNE HARBISON 1942. Miocene invertebrate fauna of New Jersey. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 94, pp. 167-250, pls. 7—22. ROBERTSON, ALICE 1900. Papers from the Harriman Alaska Expedition, VI The Bryozoa. Pro- ceedings of the Washington Academy of Sciences, vol. 2, pp. 315-340, pls. 19-21. 144 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. ROBERTSON, ALICE—Cont’d 1905. Non-incrusting chilostomatous Bryozoa of the West Coast of North America. University of California Publications in Zoology, vol. 2, no. 5, pp. 235-322, pls. 4-16. 1908. The incrusting chilostomatous Bryozoa of the West Coast of North America. University of California Publications in Zoology, vol. 4, no. 5, pp. 253-344, pls. 14-24. 1910. The cyclostomatous Bryozoa of the West Coast of North America. Uni- versity of California Publications in Zoology, vol. 6, no. 12, pp. 225-284, pls. 18-25. 1921. Report on a collection of Bryozoa from the Bay of Bengal and other eastern seas. Records of the Indian Museum, vol. 22, pt. 1, no. 8, pp. 33-65, text figs. 1-11. RoceEr, J., J. P. Nicotas, and E. Bucs 1943. La microfaune du Sahelien d’Oran. Bulletin du Museum National @d’ Histoire Naturelle, 2nd series, tome 15, pp. 248-255. Roeick, Mary D. and HANNAH CROASDALE 1949. Studies on marine Bryozoa III, Woods Hole region Bryozoa associated with algae. Biological Bulletin, vol. 96, no. 1, pp. 32-69, pls. 1-10. SAKAKURA, KATUHIKO 1935. Pliocene and Pleistocene Bryozoa from the Boso Peninsula, I Bryozoa of the Dizodo Beds. Journal of the Faculty of Science Imperial Uni- versity of Tokyo, sect. 2 (geology), vol. 4, pt. 1, pp. 1-48, pls. 1-7, 10 text figs. 1936. Studies on fossil Bryozoa from the Boso Peninsula (prelim. rept.). Journal of the Geological Society of Tokyo, vol. 43, pp. 422-423. Scorti, PIETRO 1936. Briozi fossili del miocene della Collina di Torina (Collezione Rova- senda). Atti della Reale Accademia della Scienze di Torino, vol. 71, tomo 1, pp. 402-431, pls. 1 & 2. SEGUENZA, G. 1880. Le formazioni terziarie nella provincia di Reggio. Atti della Reale Accademia del Linei, ser. 2, 1879-80, Memorie, pp. 1-143, pls. 1-17. SEN, LARS 1942. Cheilostomata Anasca (Bryozoa) collected by Prof. Sixten Bock’s ex- pedition to Japan and the Bonin Islands, 1914. Arkiv for Zoologi, band 33A, no. 12, pp. 1-130, pls. 1-9. SmitH, J. E. 1932. The shell gravel deposits and the infauna of the Eddystone Grounds. Journal of the Marine Biological Association, Plymouth, pp. 243-278, 4 text figs. VoL. XXIX ] SOULE AND DUFF: FOSSIL BRYOZOA 145 Smirt, F. A. 1872. Floridan Bryozoa collected by Count L. F. de Pourtales, Part I. Kong- liga Svenska Vetenskaps-Akademiens Handlingar, n. s., vol. 10, (1871), art. 11, pp. 1-20, pls. 1-5. 1873. Floridan Bryozoa collected by Count L. F. de Pourtales, Part II. Kong- liga Svenska Vetenskaps-Akademiens Handlingar, n.s., vol. 11, (1872), art. 4, pp. 1-84, pls. 1-13. STEPHENSON, T. A., ANNE STEPHENSON, and K. M. F. Bricur 1938. The South African intertidal zone and its relation to ocean currents, IV The Port Elizabeth district. Annals of the Natal Museum, vol. 9, pt. 1, no. 25, pp. 1-20, pls. 1-4, 1 text fig. THORNELY, LAURA R. 1912. The marine Polyzoa of the Indian Ocean from H.M.S. Sealark. Trans- actions of the Linnean Society of London, Zoology, second series, Vole 5. pts ievart: 105 pps 137-157, pl. 8: Unricn, EB. O., and Ray S. BASSLER 1904. Bryozoa, in Maryland Geological Survey, Miocene, 1904, pp. 40-429. VERRILL, A. E. 1873. Report upon the invertebrate animals of Vineyard Sound and the adja- cent waters, with an account of the physical characters of the region. United States Commission of Fish and Fisheries, Report on the con- dition of the sea fisheries of the south coast of New England in 1871 and 1872, part I, pp. 295-778, pls. 1-38. 1900. Additions to the Tunicata and Molluscoidea of the Bermudas. Transac- tions of the Connecticut Academy, vol. 10, pp. 588-594, pl. 70, 2 text figs. WatTerRS, ARTHUR W. 1878. Bryozoa (Polyzoa) from the pliocene of Bruccoli (Sicily). Transac- tions of the Manchester Geological Society, vol. 14, pp. 465-488. 1879. On the Bryozoa (Polyzoa) of the Bay of Naples. Annals and Magazine of Natural History, ser. 5, vol. 3, pp. 28-48, 114-126, 192-202, 267-— 281, pls. 8-15, 23 & 24. 1882. On fossil chilostomatous Bryozoa from Mount Gambier, South Australia. Quarterly Journal of the Geological Society of London, vol. 38, pp. 257-276, pls. 7-9. 1882. On chilostomatous Bryozoa from Bairnsdale (Gippsland). Quarterly Journal of the Geological Society of London, vol. 38, pp. 502-513, pl. 22. 1885. On the use of the avicularian mandible in the determination of chilo- stomatous Bryozoa. Journal of the Royal Microscopical Society, ser. 2, vol. 5, pp. 774-779, pl. 14. 1887. On the Tertiary chilostomatous Bryozoa from New Zealand. Quarterly Journal of the Geological Society of London, vol. 43, pp. 40-72, pls. 6-8, pp. 337-350, pl. 18. 1891. North Italian Bryozoa. Quarterly Journal of the Geological Society of London, vol. 47, pp. 1-34, pls. 1—4. 146 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. WatTeERS, ARTHUR W.—Cont’d 1892. 1908. 1918. 1921. North Italian Bryozoa, part II, Cyclostomata. Quarterly Journal ofthe Geological Society of London, vol. 48, pp. 153-1638, pl. 3. Reports on the marine biology of the Sudanese Red Sea, from collections made by Cyril Crossland; together with collections made by Dr. R. Hartmeyer—XII The Bryozoa, Part I Cheilostomata. Journal of the Linnean Society, Zoology, vol. 31, pp. 123-181, pls. 10-18. Some collections of the littoral marine fauna of the Cape Verde Islands, made by Cyril Crossland in the summer of 1904, Bryozoa. Journal of the Linnean Society, Zoology, vol. 34, pp. 1-45, pls. 1-4. Observations upon the relationship of the (Bryozoa) Selenariadae, Conescharellinidae, etc., fossil and recent. Journal of the Linnean Society, Zoology, vol. 34, pp. 399-427, pls. 29 & 30. YANAGI, NAOKATSU, and YAICHIRO OKADA 1918. On a collection of Japanese cheilostomatous Bryozoa, I. Annotationes Zoologicae Japonenses, vol. 9, pt. 4, pp. 407-429, pl. 6, text figs. 1-7. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. XXIX, No. 5, pp. 147-256, 10 maps, 6 plates November 5, 1957 A MONOGRAPH OF THE GENUS HYDRANGEA By ELIZABETH McCLINTOCK Assistant Curator of Botany, California Academy of Sciences PREFACE Hydrangea, a woody genus of the Saxifragaceae, has been a popular garden and greenhouse plant since the importation of Hydrangea macro- phylla into England from China by Sir Joseph Banks in 1789. The genus is of interest to botanists because of its disjunct distribution. It occurs in the temperate regions of eastern Asia and eastern North America and extends southward into the tropics of both hemispheres. A temperate group with a deciduous habit has had its greatest development in eastern Asia, while a tropical group with an evergreen habit has had its greatest development in Central and South America. The plants in cultivation have come ehiefly from the deciduous group. Throughout its wide range in both the Old and New worlds Hydrangea is fairly abundant and has been much collected over a period of years. Many species have been deseribed from these collections and also from cultivated plants. This has resulted in a multiplicity of names in the genus. No revision of the genus has been done for many years; therefore, for horticultural, as well as for botanical purposes, it seemed desirable to revise the genus thoroughly, to find out the morphological and geo- graphical limits of the species, and to find out the relationship which exists between the New World and the Old World groups. [147] 148 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H SEr. ACKNOWLEDGMENTS To the officers in charge of the various herbaria consulted, the au- thor wishes to express her gratitude for the privilege of studying their material, whether loaned or seen on visits. Specimens from the following herbaria are cited as follows (the abbreviations are according to Lanjouw and Staffleu, 1954) : WVA Arnold Arboretum of Harvard University, Cambridge, Massachusetts. Bailey Hortorium of Cornell University, Ithaca, New York. British Museum of Natural History, London, England. California Academy of Sciences, San Francisco, California. University of North Carolina, Chapel Hill, North Carolina. Duke University, Durham, North Carolina. The Royal Botanic Garden, Edinburgh, Scotland. Conservatoire et Jardin Botaniques, Geneva, Switzerland. University of Georgia, Athens, Georgia. Gray Herbarium of Harvard University, Cambridge, Massachusetts. The Royal Botanic Gardens, Kew, England. The Wallich Herbarium of the Royal Botanic Gardens, Kew. Universitatis Jagellonicae, Krakow, Poland. The University of Kentucky, Lexington, Kentucky. Herbario de la Facultad Nacional de Agronomia, Medellin, Colombia. The University of Michigan, Ann Arbor, Michigan. Missouri Botanical Garden, St. Louis, Missouri. New York Botanical Garden, New York, New York. Botanisk Museum, Oslo, Norway. Bebb Herbarium of the University of Oklahoma, Norman, Oklahoma. Muséum National d’Histoire Naturelle, Laboratoire de Phanérogamie, Paris, France. Naturhistoriska Riksmuseet, Botanical Department, Stockholm, Sweden. The University of Tennessee, Knoxville, Tennessee. Botanical Institute, University of Tokyo, Tokyo, Japan. Institute of Systematic Botany, Botanical Garden and Botanical Museum of the University of Uppsala, Uppsala, Sweden. The University of California, Berkeley, California. National Museum, Smithsonian Institution, Washington, D. C. Naturhistorisches Museum, Vienna, Austria. Botanisches Institut und Botanischer Garten der Universitat Wien, Vienna, Austria. West Virginia University, Morgantown, West Virginia. To Dr. Rogers MeVaugh, of the University of Michigan, under whose direction this work was done in partial fulfillment of the requirements for the degree of Doctor of Philosophy, the author wishes to express her gratitude for his advice and suggestions. To Mr. J. Robert Sealy of the Royal Botanic Gardens, Kew, the author expresses her thanks for his help in interpreting the specimens in the Wallich Herbarium, and to Dr. Hiroshi Hara of the Botanical Institute, Faculty of Science, University of Tokyo, Vou. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 149 for supplying photographs of type specimens from the herbarium of the University of Tokyo. To the Council and the Director of the California Academy of Sciences the author expresses her gratitude for a leave of absence during the summer of 1954 for the purpose of working on Hy- drangea in European herbaria. INTRODUCTION The name Hydrangea was first used by Gronovius (1739) in his Flora Virginica for a plant collected by John Clayton. Linnaeus (1753) adopted this name when he described Hydrangea arborescens in the Species Planta- rum. This, the first species to be described in the genus, was followed by H. radiata, which was named by Thomas Walter (1788) in the Flora Carolinaana, and H. quercifolia, named by William Bartram in 1790 in the account of his travels through the Carolinas, Georgia, and Florida. Ruiz and Pavon made the first collection of a species of the genus in South America, but they referred their plant to the genus Cornidia (1798). Cornidia umbellata Ruiz & Pavén is now referred to Hydrangea Presli Briquet. The first Asiatic species of the genus were placed in Viburnum. In 1781 the younger Linnaeus described one species, and in 1784 Carl Thunberg described five in his Flora Japonica. Because of the ornamental value of some species of Hydrangea, many of the early collections of the genus were actually made for the purpose of introducing the plants into gardens. Hydrangea arborescens, the first species to be cultivated in England, was said by William Curtis (1799) to have been introduced into England as early as 1736. Philip Miller (1768) stated that it had “been brought within a few years past to EKu- rope, and is now preserved in gardens for the sake of variety more than its beauty.” Apparently, this species was never as popular a garden plant as the next hydrangea to be introduced into England, the one we know today as Hydrangea macrophylla subsp. macrophylla. Through the agency of Sir Joseph Banks a living plant of this hydrangea was brought to the gardens at Kew from China in 1789. It was named H. hortensts by Sir James E. Smith (1792). Hydrangea hortensis, however, was the same as the plant described by Thunberg in 1784 as Viburnum macro- phyllum. Thunbere’s name was overlooked until Seringe (1830) trans- ferred it from Viburnum to Hydrangea. The plant, however, was not overlooked. As Hortensia opuloides it was described by Lamarck (1790). This name was based on a specimen cultivated at Mauritius and probably sent to France by Commerson. The Portuguese missionary, Loureiro, who spent many years in Indo-China, in 1790 described a plant as Primula mutabilis which was apparently a cultivated specimen of Hy- drangea macrophylla subsp. macrophylla (see Merrill, 1935). Other names were given to this plant by Siebold who during the several years which 150 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. he spent in Japan collected a number of horticultural forms of hydrangea. Siebold’s names were published and described (1829) in a synopsis of the Japanese species of Hydrangea. Later (1840) some of these were illus- trated in Siebold and Zucearini’s Flora Japonica. During the early part of the nineteenth century other Asiatic species were discovered and named. Hydrangea anomala, H. aspera, and H. het- eromalla were found in the eastern Himalayas by Francis Buchanan- Hamilton and Nathaniel Wallich. These same species were collected again by J. D. Hooker and Thomas Thomson during their explorations of the eastern Himalayas in 1848-1850. So far as known no material was intro- duced into cultivation from these collections. The first complete taxonomic treatment of the genus was that of Seringe (1830) in DeCandolle’s Prodromus, which brought together the species from both the Old and New worlds known to that time. In treating the South American species he apparently overlooked the genus Cornidia of Ruiz and Pavoén, but he did include Sarcostyles of Presl, in which was placed Sarcostyles peruviana Presl, now referred to Hydrangea Preslu. The other South American species which he included were H. peruviana Moricand and H. scandens Poeppig, now referred to H. serratifolia. He included among the Asiatic species not only the Japanese species de- seribed by Thunberg (1784) and Siebold (1829) but also H. oblongifolia Blume (now placed in synonymy under H. aspera) from Java, and the species from the eastern Himalayas deseribed by Wallich (1826) and David Don (1825). Maximowiez (1867) published an important revision of the eastern Asiatic species known at that time. These species he divided into two sections: Euhydrangea, for those which are erect in habit and have petals which fall separately and possess elliptic seeds, and Calyptranthe for those which are scandent in habit and have petals which are coherent and fall as a cap and possess circular seeds. Euhydrangea he divided into Series 1, Petalanthe, for those in which the petals are persistent and re- flexed during anthesis and the seeds are not caudate, and Series 2, Pipto- petalae, for those in which the petals are usually deciduous in early anthesis and the seeds are caudate. He divided Petalanthe into two groups, Americanae, for the North American species, and Japonico-sinensis, for Hydrangea hirta, H. virens, and H. chinensis, Koehne, treating only the cultivated species of the groups previously considered by Maximowicz, divided them into two subgenera, Hortensia for H. opuloides, and Hor- tensiella for the remainder of the species. This latter subgenus he divided into Euhydrangea and Calyptranthe. Engler’s summary of the genus (1891) recognized Euhydrangea and ‘alyptranthe as originally constituted by Maximowicz, and in addition established a third section, Cornidia, for the scandent evergreen species of Vout. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 151 Central and South America. Schneider’s treatment (1906) of the ecul- tivated species was based essentially on that of Engler, except that he called Engler’s sections subgenera. Rehder (1911), treating only the Chi- nese species, recognized Euhydrangea and Calyptranthe as constituted by Maximowicz. Euhydrangea he divided into three subsections. In Petalanthe he included only those species which Maximowiez had grouped together as Japonico-sinensis. Heteromallae was a new subsection for H. paniculata and H. heteromalla, and Asperae was new for H. aspera. Later (1927), in a treatment of the cultivated species, Rehder added a fourth group under Euhydrangea, Americanae, for the North American species. He refers to these subdivisions as “Groups” in 1927, but in 1940 and 1949 he refers to them as “Series.” Engler (1930), in a brief treatment of the genus, recognized Rehder’s subdivisions which he called subsections. Briquet (1919) revised the Central and South American species and fol- lowed Engler (1891) by placing them in the section Cornidia. Briquet di- vided this section into two subsections, Monosegia and Polysegia, based on the structure of their inflorescences. The present treatment of Hydrangea differs from previous ones in two respects. First, only two sections, Kuhydrangea, which must be ealled section Hydrangea according to a new rule adopted at the meeting of the International Botanical Congress held in Paris in 1954 (Stafleu, 1954), and Cornidia are recognized. The section Calyptranthe, established by Maximowiecz and recognized by everyone since, is made a subsection under the section Hydrangea. The characters on which Calyptranthe was based do not appear to be of the same importance as those on which the sections Hydrangea and Cornidia are constituted in the present treatment. Sec- ond, Hydrangea macrophylla (H. opuloides and H. serrata) is removed from subsection Petalanthe, in which it had been placed by Rehder and later by Engler, and put into a subsection by itself, designated in the present treatment as subsection Macrophyllae. MORPHOLOGY AND INFRAGENERIC DIVISIONS The characters used for dividing Hydrangea along sectional lines are the following: Hasir. Erect or scandent. When scandent the plants cling by means of aerial rootlets. Fouacr. Evergreen or deciduous. LEAF TEXTURE. Coriaceous or chartaceous. FLORAL BRACTS. Rounded or lanceolate. Rounded bracts, several in number, occur in a row at the base of the inflorescence. These envelop the unopened inflorescence, and leave noticeable scars on falling. Lanceo- late bracts, usually numerous, occur throughout the inflorescence. These 152 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. may cover, but do not envelop the unopened inflorescence and leave no sears on falling. The characters used to distinguish and delimit the two sections are the following: HYDRANGEA CORNIDIA Habit usually erect. Habit usually seandent. Fohage deciduous. Foliage evergreen. Leaf texture chartaceous. Leaf texture coriaceous. Floral bracts usually lanceolate. Floral bracts rounded. The characters found useful in delimiting the species in the present study are relatively clear cut and overlap very little from one species to another. The use by previous workers of extremely variable characters in describing new species, such as leaf pubescence and shape and size of the leaves, has given the genus an unwarranted reputation for being difficult. A discussion of the variation found in the principal morpho- logical characters follows: Leaves. In shape the leaves are usually ovate, varying from narrowly so to broadly ovate. Their margins are usually serrate or entire, although in two species they are pinnately lobed. In size, they vary in length from about two to thirty-five centimeters. In certain species shape and size of leaves are relatively constant, while in others both of these are extremely variable. A dimorphic condition of the leaves, which exists in the climbing species of the section Cornidia, is associated with a sterile, and perhaps juvenile, growth form. In this form the leaves are usually 1.5-3 em. long, dentate, chartaceous, and are borne on slender, epiphytic thread- like stems. According to Standley (1937), this growth form is very fre- quent in occurrence in the temperate region of Costa Rica, growing on tree trunks and fence posts. Because the sterile or juvenile leaves are very different from those on the flowering branches, the relationship be- tween the two kinds of leaves was long unrecognized. Specimens with the sterile leaves only have been seen from Panama, Guatemala, El Sal- vador, Nicaragua, Peru, and Chile. The specimens from El Salvador and Nicaragua are of particular interest because no flowering material has been collected from either of these countries. Hydrangea anomala, a climber from eastern Asia and a member of the section Hydrangea, also has sterile or juvenile leaves, as shown by two specimens in the herbarium at Kew, collected by Hooker and Thomson at Darjeeling. PuBESCENCE. The type of pubescence is usually uniform within each species, but its density may vary. Only in Hydrangea arborescens and H. quercifolia is there more than one kind of hair. In section Hydrangea the hairs are slender, about 0.5-1 mm. long, and white. Under a magnifi- VoL. XXIX | McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 153 eation of 35 or 45 diameters their surfaces appear rough, like frosted glass; under magnification of about 350 diameters their surfaces are seen to be covered with minute tubercules (see plate 3, figure 4). These hairs are sometimes stiff and appressed and at other times soft and curled. In H. arborescens subsp. radiata the hairs have no tubercules and are smooth. In H. querctfolia both the smooth and rough surfaced hairs are combined on the lower leaf surfaces. In section Cornidia the more common kind of hair is short (0.1-0.3 mm. long), stiff, light brown, and borne in stellate clusters. The less common type consists of single, longer hairs (0.4—0.6 mm. long), which are eurled, and light brown or ferruginous in color. INFLORESCENCE. The inflorescence in Hydrangea may be described in general terms as a dendritically branched, terminal cluster, usually many- flowered, and either rounded or pyramidal in shape. The rounded cluster is the more common, being found in all but two species, Hydrangea quer- ctfolia and H. paniculata, which have pyramidal clusters. The dendritie branching of the inflorescence is arranged in such a way that the flowers are held erect and more or less in the same plane. The ultimate members of the dendritically branched system, of which the rounded clusters are composed, consist of single flowers or small elus- ters of two to six flowers. These clusters correspond to modifications of a simple dichasium, which, as defined by Rickett (1943, p. 489), consists of a “short branch bearing a terminal flower and two prophylls which sub- tend as many lateral flowers.” Rickett (1948), in discussing the inflor- escence of Crataegus, interprets the terminal flower groups in the inflores- cence of C. Crus-galli as being dichasia. In some dichasia the terminal flower opens first, and in others the three flowers open approximately together. However, there are other flower groups with different numbers of flowers. Such groups represent condensations of dichasia, in which one or two flowers have failed to develop and which are aggregated in a rather irregular fashion upon a central axis. The inflorescence of Hy- drangea is made up of terminal units similar to those of Crataegus. These terminal units appear to represent modified dichasia with an irregular number of flowers which appear to open at approximately the same time. The entire inflorescence consists of an aggregation of irregular groups upon a central axis, several of which are joined together on a main rachis to form a rounded compound cluster. For such an inflorescence Rickett has proposed to use the term “cyme,” redefining it so that it does not refer to a sequence of flowering of the inflorescence. The pyramidal cluster found in Hydrangea quercifolia and H. pan- iculata has been called a panicle, or occasionally a thyrsoid ecyme. Ex- amination of the ultimate units of the inflorescences of these two species shows them to be single flowers or small clusters which appear to be modi- fications of simple dichasia and not different from those of the species 154 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. having the rounded clusters. These ultimate units are aggregated into branches which are scattered upon an elongate central rachis, and the whole inflorescence differs from the cyme, as redefined by Rickett, in its pyramidal form. For this inflorescence the term thyrse (Rickett, 1944), that is, a compound flower cluster of more or less pyramidal form whose ultimate units are simple dichasia, may be used. In this definition no reference is made to sequence of flowering of the inflorescence. However, in both species of Hydrangea having this kind of inflorescence there is a sequence of flowering from the bottom to the top of the inflorescence, the lowermost clusters opening before the uppermost. The main branches of the inflorescence in the section Cornidia all arise at one point, giving the inflorescence an “umbellate’ appearance. The arrangement of these umbel-like cymes is used as a basis for divid- ing the section into two subsections. The cymes occur singly in the sub- section Monosegia, while in the subsection Polysegia there are several which occur one above another along a main axis. In the section Hy- drangea the main branches do not arise at the same point, although sometimes their points of origin are close enough to present an ‘‘umbellate” appearance. The type of Hydrangea uwmbellata Rehder, now considered to belong to H. scandens subsp. chinensis, has such an inflorescence. In only one instance are a pair of closely related species separated by the dif- ferences in their inflorescences. The chief difference between H. heteromalla and H. panicuata is the rounded, eymose inflorescence of the former and the pyramidal one of the latter. FERTILE FLOWERS. Variations in floral structure involve in some de- gree most of the parts of the fertile flowers. Hypantuia. The ovary is fused wholly or in part with the receptacle and this fused portion is here called the hypanthium. When the fusion is complete, the top of the hypanthium, called the disk, is flat. When fusion is incomplete and only the lower part of the ovary is fused to the re- ceptacle, the disk is conical. SEPALS. The sepals are deltoid, relatively short (usually less than 1 mm. long) and inconspicuous. Their number is the same as that of the petals, that is, either four or five. Petats. The petals are always valvate and vary little in size (usually about 2 mm. long). In shape they are somewhat ovate or obovate. When ovate, they are truncate at the base; when obovate, they are narrowed at the base (see plate 6, figures 2B and 4B). In one species the petals cohere and fall as a cap or calyptra (see plate 5, figure 4B). STAMENS. The stamens are usually 8 or 10, that is, twice as many as the sepals or the petals. In only one taxon (Hydrangea anomala subsp. VoL. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 155 petiolaris) is their number three or four times that of the sepals or the petals. Pistits. Variation in the pistil is found in the number of its parts and in its relation to the hypanthium. The number of locules in the ovary and the number of styles vary from two to five, often with variation in the same species or even on the same plants. The relation of the ovary to the hypanthium has already been mentioned. The position of the ovary is a useful character for delimiting subsections and species in the see- tion Hydrangea. In the section Cornidia the ovary is always inferior. Placentation is always axile. In the inferior ovary the placentas are at- tached to the central axis which is at the same level as the attachment of the sepals to the hypanthium, while in the half superior ovary the cen- tral axis extends upward and under the conical disk. In the half superior ovary the styles arise from the top of the conical disk and their length in comparison with the height of the disk is used as a subsectional char- acter in two instances. The styles, normally separate, show a tendency to adhere during early anthesis in two species of the section Cornidia, Hydrangea diplostemona and H. tarapotensis, becoming separate by late anthesis (see plate 4, figures 7 and 8). The stigmatic surfaces develop from the apex of the styles downward along the inside surface in the form of noticeable papillae. The stigmatic surface may be present and receptive to pollen when the flower opens and the pollen is shed. However, from herbarium specimens it is not possible always to be certain when the stigmatic surface is receptive. In the two North American species the styles appear turgid when the flowers first open and the stigmatic surface is confined to a line extending from the top to a short distance down on the inside surface of the style. Later the stigmatic surface increases and the styles become less turgid. These two species which constitute the sub- section Americanae are most closely related to the Asiatic subsection Pipto- petalae on the basis of their both having an inferior ovary. In the sub- section Piptopetalae the styles from the beginning of anthesis appear slen- der and have a fairly well developed stigmatic surface. It would thus appear that in the individual flowers of the subsection Americanae the stigmas are not receptive to pollen until after the pollen has been shed, while in the flowers of subsection Piptopetalae the stigmas are receptive at the same time that the pollen is shed. Observations on living material are necessary to confirm this. In section Cornidia there are several species in which it is assumed that the stigmatic surfaces do not develop until after the pollen has been shed. In Hydrangea Oerstedu and H. serratifolva the styles remain smooth and do not elongate until some time after the flowers are open, and in H. diplostemona and H. tarapotensis the styles are adherent during early anthesis and the stigmatic surfaces are not exposed at this time. 156 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. CapsuLes. The capsules in all species open at the apex between the styles. The shape of the capsules depends on the position of the ovary. When the ovary is inferior, the capsule is truncate at the apex, and when the ovary is half superior, the capsule is conical. SEEDs. The seeds are numerous, elliptical to oval, and very small (about 1 mm. long). In some species the seeds are caudate, by virtue of the seed coat being attenuate into a short tail-lke extension at each end of the seed (see plate 5, figures le and 2c). In one species (Hydrangea anomala, see plate 5, figure 4D), the body of the seed is surrounded by a circular wing. The mature seeds in all species are variously striate or reticulate. STERILE FLOWERS. Sterile flowers, formed by the enlargement of the sepals into petal-like structures, are larger and more conspicuous than the fertile ones and occur around the periphery of the inflorescences. In most species, sterile flowers are present in addition to the fertile ones; however, occasionally (as in Hydrangea arborescens), they may be pres- ent in some individual plants and absent in others. In a few species only fertile flowers are present. The occurrence of inflorescences consisting entirely of sterile sepaloid flowers led to the early cultivation of Hydrangea macrophylla as an ornamental plant. There are similar forms, also in cultivation, of H. arborescens and H. paniculata. These inflorescences probably originated from a mutation in wild populations and the plants on which they oc- curred were brought into cultivation. I have seen no collections from the wild of either H. macrophylla or H. paniculata with this type of inflores- cence; however, collections of such have been made in the eastern United States for H. arborescens. In H. aspera, as judged from a number of her- barium specimens, this condition occurs in the wild. The condition also occurs in H. Oerstedi, as has been observed in a single specimen (Linden 1139). The development of sterile sepaloid flowers has taken place in Cardiandra, Platycrater, and Schizophragma, genera which are consid- ered to be related to Hydrangea. Superficially, the enlargement of the sepals in some species of Hydrangea gives these plants a resemblance to certain species of the unrelated genus, Viburnum. In Viburnum, however, it is the petals of the showy flowers which have become enlarged and sterile. The presence of sterile flowers has given rise to various speculations as to their function. Kerner von Marilaun (1902) suggested that in Hy- drangea quercifolia they are a means of protecting the pollen of the fertile flowers from rain. Their role in attracting insects to the inflores- cence has been considered. Plateau (1898) did not consider this to be of prime importance but felt that pollination of the fertile flowers would be accomplished regardless of the presence or absence of the sterile ones VoL. XXIX|] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 157 because, first, in the case of the inflorescences having only a few sterile flowers the insects proceed directly to the fertile flowers, and second, inflorescences having mostly sterile flowers are seldom visited by insects. The peripheral sterile flowers ordinarily consist of only the enlarged calyx. However, in some of the garden forms of Hydrangea macrophylla in which the inflorescences consist almost entirely of sterile flowers, petals, stamens, and pistils may be present also. Such flowers are not sterile although they probably do not often set seed. Perriraz (1911) reported unsuccessful attempts to obtain seeds from such flowers. He further reported that plants of the garden form known as H. macrophylla Otaksa having infloresences composed ot enlarged showy flowers, changed the composition of their inflorescences over a period of years so that fewer showy flowers and more fertile ones are produced. The floral structure in Hydrangea is essentially the same throughout the genus. The two sections are separated on the basis of vegetative characters and habit; not a single floral structure can be found to sep- arate them. The half superior ovary present in some species of Hydrangea may be considered more primitive than the inferior ovary. However, the development of the inferior ovary is not associated with a single other advanced structure in those species in which it occurs. In addition to the inferior ovary present in some species, the reduced number of stamens (which are double the number of the petals) is also an advanced char- acter. The separate actinomorphie petals, numerous seeds, and the large spreading cymose inflorescences with numerous small flowers are primitive. This combination of characters, together with the uniform floral pattern throughout the genus, becomes somewhat more significant when consid- ered in relation to the method of pollination in Hydrangea. The Saxifragaceae as a family is placed by Grant (1949) in a group, designated as “Promiscuous Plants,” pollinated by miscellaneous insects. Grant made a survey of the taxonomic characters used to distinguish species in various families of angiosperms and he found a correlation between the importance of floral structures used in classification and method of pollination. Species pollinated by miscellaneous insects are less frequently found to have differences and trends in floral structures than are those pollinated by specific insects. The only observations avail- able regarding pollination in Hydrangea are those made by Robertson (1892) on H. arborescens. These showed that certain unspecialized mem- bers of the Hymenoptera, Diptera, Coleoptera, and Lepidoptera visited this species. Although similar observations are not available for other species in the genus, it may be assumed that since the floral structure and the characters of the inflorescence in H. arborescens follow the pat- tern for that of the genus, the same type of insect visitors would be pres- ent on other species. 158 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. The presence of the sterile peripheral flowers and numerous small flowers arranged in cymose inflorescences gives Hydrangea a superficial resemblance to certain species of Viburnum (Caprifoliaceae). In Vibur- num, however, there are several advanced characters not present in Hydrangea: the corolla is united, the stamens are reduced to the same number as the parts of the corolla, and the ovules are reduced to one in each carpel. The presence of these characters places Viburnum in a family considered unrelated to the Saxifragaceae and shows a beginning in the direction of specialization in both floral structure and methods of pollina- tion found in the more highly developed genera of the family. Robertson’s (1898) observations on insect visitors to Viburnum pubescens Pursh and V. prunfolium L. showed much the same aggregation of miscellaneous insects as he had observed on Hydrangea arborescens. Therefore, in spite of the presence of the several advanced characters just mentioned for Viburnum, this genus would be classed with the group of Promiscuous Plants to which Hydrangea and other saxifragaceous plants belong. An- other group of plants which have numerous small flowers aggregated into a dense inflorescence is the Umbelliferae which has an inferior ovary and reduced number of stamens and seeds, but (as Hydrangea) separate actino- morphie petals. Undoubtedly, dense inflorescences made up of numerous small flowers serve as a landing place for various unspecialized insects which effect pollination through their movements. This is a device found to be successful in several unrelated families of angiosperms and one not associated with diversified or specialized floral structures. Natural hybrids in Hydrangea are rare. The only report known to the writer of the occurrence of such a hybrid is that made by Makino (1932). He described a plant, growing on Izu Peninsula, Honshu Island, Japan, which he considered to represent a hybrid between H. hirta and H. scan- dens. These two very closely related species differ chiefly in the shape and margin of their leaves. An illustration of the reputed hybrid showed leaves more or less intermediate between the two putative parent species (Hiyama, 1934). Hybrids have been reported in cultivation but crosses must be difficult to effect since the reports have been few. Haworth-Booth (1950) lists the following: 1. A hybrid between Hydrangea macrophylla var. rosea and H. pan- iculata, reported by Monsieur L. Foueard, horticulturist in Orleans, France, in 1912. The plant was pictured in Revue Horticole (1912, p. 324) and showed corymbose (not paniculate) inflorescences consisting entirely of sterile flowers. 2. A hybrid between Hydrangea macrophylla var. rosea and H. petio- laris, called H. & hortentiolaris, was made by Monsieur Henri Cayeux of Le Havre, France. Vout. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 159 According to Haworth-Booth, both of these plants were lost before they became established in the horticultural trade. Sax (1931) reported chromosome counts for several species in the see- tion Hydrangea. Hydrangea arborescens, H. radiata, H. cinerea, H. petio- laris, H. paniculata, and H. xanthoneura all had a diploid number of 36, with the exception of 72 in a horticultural form of H. paniculata. Sugiura (1931) reported a haploid count of 18 for H. virens, and Haworth-Booth (1950) reported counts made of various garden forms of H. macrophylla by Dr. Janaki Ammal which also showed a diploid count of 36. So far as known, no counts have been made of species in the section Cornidia. GEOGRAPHIC DISTRIBUTION The occurrence of Hydrangea in eastern Asia and eastern North Amer- ica places it in that group of genera which indicate a relationship between the floras of these two regions. The problem of this disjunct distribution has recently been discussed and reviewed by Li (1952). Li lists two groups of genera which are especially confined to eastern Asia and eastern North America, those which are distributed in temperate regions in both areas, and those distributed in the temperate regions and also in the tropical (or subtropical) regions in one or both areas. Hydrangea falls into the latter group, since both in Asia and America there are species found in temperate and subtropical regions. Map 1 shows the distribution of the genus. The section Hydrangea occurs in the temperate regions of eastern Asia and eastern North America. Its greatest development is in eastern Asia where nine of its eleven species are found. In this area it extends from the Himalayas eastward across central and southern China to Sak- halin, Japan, the Ryukyu Islands, the Philippine Islands, Formosa, Su- matra, and Java. The center of this large area (with reference to Hydrangea) is western and central China, where six of the nine species occur. This region includes Kweichow, Yunnan, Szechuan, and parts of Kwangsi, Hunan, and Hupeh, and within it are the Yanetze Valley and the high mountains of southwestern China. The vegetation characteristic of the middle elevations of this region is a deciduous mesophytie forest known to be the richest of all present day temperate floras. The richness of this flora may be attributed to the topography and mild climatic condi- tions of the region and also to the lack of extensive Pleistocene glaciation. The occurrence of Hydrangea within this area follows the distributional pattern of many of the genera which are disjunct between eastern Asia and eastern North America. Li (1952) pointed out that in eastern Asia the distribution of species in these disjunct genera is often widespread. Such genera occur in the same regions where the widespread species of Hydrangea are found. The 160 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. extension of some genera into the warmer subtropical areas such as the mountains of the Philippines, Sumatra, and Java is not uncommon, but these islands may be considered to represent peripheral areas in the dis- tribution of these genera. In discussing the relationship between the floras of Formosa and southwestern and central China, Wilson (1920) stated that the climate of Formosa is warm temperate, but that in the mountains such cool temperate genera as Fagus, Ulmus, Carpinus, Alnus, and Malus reach their southern limits. Merrill (1934) pointed out that certain southern Asiatic species make a jump to the higher altitudes of Sumatra, Java, Borneo, and the Philippines but do not occur in the Malay Peninsula. Hydrangea, as many genera of the eastern Asiatic—eastern North Amer- ican relationship is found in Japan. Some species of Hydrangea found in Japan are widespread in China, while others are confined to Japan or to Japan and such adjacent areas as Formosa, Korea, and the Ryukyu Islands. The relationship of the flora of Japan to that of central and western China has been discussed by Hu (1940) who attributes this relationship in part to the fact that Japan was connected to the Asiatic mainland up to the middle of the Tertiary period. The two species of Hydrangea found in North America occur in the deciduous mesophytie forest of the eastern United States. Both appear to be centered in the southern Appalachian Mountains. The Appalachian area, being an upland plateau and not glaciated, is known to have been continuously available for plant life since long before the beginning of the Tertiary period. It represents an area of refuge for the eastern Asiatic—eastern North American disjunct genera which were more wide- spread during the Tertiary period than they are now. Cain (1943) has pointed out the Tertiary affinity of many of the genera of the cove hard- wood forests of the southern Appalachians. In contrast to the temperate occurrence of the species belonging to the section Hydrangea, those of the section Cornidia are found in the subtropical regions of Central and South America, where twelve species extend from Guatemala through Costa Rica and Panama and into the Andes of South America. A single restricted species occurs in the Sierra Madre in Durango, Mexico. All of the species of Cornidia in this region occur in the mountainous areas where temperatures are moderate and where there are mists or frequent rains. This subtropical element of Hydrangea differs from the temperate element in its thick coriaceaus evergreen leaves and generally climbing habit. The species of Cornidia are not confined to the New World; a single species, Hydrangea integrifolia, occurs in the mountains of the Philippines and Formosa. The present day disjunct distribution of the temperate mesophytic forests of eastern Asia and eastern North America can be better under- 161 VoL. XXIX|] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA ‘sotoeds [ISsSOy O ‘soloeds SUIATIT @ ‘DabuDIph JO sotoods [Issoy pue SUIATT JO UOTINGLUSIG “T av Se Bae es Hees oe ete Oe baptestes.° S ie x BS 7 os ———— > : \, : A ay iE ws . | NSE 162 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. stood by a knowledge of the past distribution of a similar forest which was composed of many of the same woody genera. Fossil records indi- eate that a deciduous mesophytie forest extending as far north as Man- ehuria, Greenland, and Alaska existed during the early and middle parts of the Tertiary period over a considerable portion of the northern hemi- sphere. This forest belongs to the so-called Arcto-Tertiary flora. The northwestern part of North America has provided a fairly com- plete chronological record of the history of the Areto-Tertiary flora in this region. The changes in the composition of this flora have been out- lined by Chaney (1940) on the basis of a series of fossil floras. The vari- ous fossil floras from Eocene through Miocene times in western North America show that there were for any period of time latitudinal and alti- tudinal variations in composition of the vegetation, but that as the climate changed from warm and wet in the earlier part of the Tertiary to cool and dry in the later part, the vegetation changed also. The climatic trend toward lower temperature and precipitation resulted in the elimi- nation of an evergreen neotropical flora, which existed as far north as Oregon, by the end of the Oligocene, and in the southward migration of a temperate deciduous flora of higher northern latitudes to replace it. By early Miocene times the redwood and broad-leaved, deciduous trees were dominant in Oregon. The Bridge Creek flora of the John Day Basin, late Oligocene or early Miocene, contains (Chaney, 1944) redwoods and broad-leaved deciduous trees characteristic of a wet summer climate. Other Miocene floras of the same age were of similar composition. How- ever, during the Miocene, the Cascades were uplifted and with the changes in topography and climate, the redwoods and broad-leaved trees began to disappear. The Maseal flora of later Miocene showed that trees such as black and live oaks, box elders, madrono, syeamores, and poplars began to replace the trees requiring more moisture. By the Pliocene, redwoods and other moisture requiring trees had nearly disappeared. Hydrangea, which belonged to this mesophytie element, is not reported after the Miocene in western North America. LaMotte (1952) lists six fossil species of Hydrangea from Eocene, Oligocene, and Miocene deposits in southern Alaska, Washington, Oregon, California, Idaho, and Colorado. These species and their places of col- lection are as follows: H. alaskana Hollick. Paleocene or Eocene. Jaw Mountain, Alaska Pen- insula, Alaska. H. californica MacGinitie. Eocene. Ione Gravels: Chalk Bluffs, Nevada. County, California. H. russellii Chaney & Sanborn. Eocene-Oligocene. Fisher: Goshen, Lane County, Oregon. H. fraxinifolia (Lesquereux) Brown. Oligocene. Florissant, Colorado. VoL. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 163 H. reticulata MacGinitie. Oligocene. Weaverville, Hay Fork, Trinity County, California. H. bendiret (Ward) Knowlton. Miocene. Maseall: Grant County, Ore- gon. Latah: Spokane, Washington. Whitebird, Idaho. Trout Creek, Harney County, Oregon. Hydrangea is thus seen to be a member of several fossil floras in west- ern North America from early to middle Tertiary (see Map 1). Its typi- cal occurrence was in the temperate mesophytic forest characteristic of the Arcto-Tertiary flora; however, it is known also from one of the warm humid neotropical floras present in the Pacific Northwest during early Tertiary times. Its presence in the Goshen flora indicates that the sub- tropical element of the genus, occurring now in Central and South Amer- ica, may have had a northward extension in the Pacific Northwest. As this neotropical flora died out and was replaced by temperate floras, the subtropical element of Hydrangea was also lost in the area. These records of Hydrangea in the Tertiary floras of the west consti- tute the only fossil records of this genus in North America. In eastern North America, Tertiary fossil plants are known only from the south- eastern Gulf Coast. Although Hydrangea occurs today in the eastern United States, it is not found in any of these deposits. In China, Hydrangea is reported from the Miocene Shanwang flora in Shantung Province by Hu and Chaney (1940), who show that the ma- jority of Shanwang genera are wide ranging in China today and that this fossil flora includes elements which now characterize the forests of eentral and south China and the Yangtze Valley. Many of these are temperate northern genera, and although they are found today in middle and upper levels in the mountains, they are rare or absent from lower altitudes in these areas. The Shanwang flora is the only record of Mio- cene vegetation in China. As in western North America, Hydrangea is not found in China in deposits later than the Miocene. Pliocene floras of the northern interior of China lack the mesophytic elements present in the Shanwang flora. Hu and Chaney also showed that the modern forests of Korea and Japan were very similar to the Miocene vegetation of northern China, but that Pliocene floras from Japan are not different from central Japanese forests of today. THE SPECIES IN CULTIVATION Hydrangeas were cultivated in China and Japan long before their introduction into Europe and America, and many of our garden forms eame from the Orient. There is no way of knowing how long these forms had been grown by the Chinese and Japanese; we merely know that many Europeans, including Thunberg, Siebold, Fortune, and Maximowicz, found 164 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. these plants as well established horticultural subjects in China and Japan before they sent them to Europe. As with other horticultural plants hav- ing forms which are the result of many generations of selection, their origin is lost in obscurity. A discussion of the origin of the cultivated forms of Hydrangea macrophylla is given under that species. Haworth-Booth (1950) has discussed the horticultural aspects of the genus in his recent book, The Hydrangeas. Since Haworth-Booth has covered the subject adequately, the present discussion of Hydrangea as a garden plant will only list the species cultivated, together with some of the synonyms by which they are often known. Complete synonymy is given under each of the species mentioned. 1. Hydrangea macrophylla. Of the species listed, this one has always been the most popular because of its versatility, both as a garden plant and as a florists’ plant. Hydrangea macrophylla subsp. macrophylla var. macrophylla (H. Hortensia, H. hortensis, H. opuloides, Hortensia opu- loides). This is the form to which belong the many globose-headed plants with all sterile flowers called hortensias. Hydrangea macrophylla subsp. macrophylla var. normalis is the form to which belong the plants with a few sterile flowers surrounding a cluster of fertile flowers. Haworth-Booth calls these lacecaps. Hydrangea macrophylla subsp. serrata (H. serrata) is said to be in cultivation, but it is so similar to subsp. macrophylla that it is very difficult to recognize garden forms as actually belonging to this subspecies. 2. Hydrangea arborescens subsp. arborescens. A form of this sub- species with all sterile flowers, called H. arborescens grandiflora, is often cultivated. Subspp. discolor (H. cinerea) and radiata (H. radiata) are oe- easionally cultivated. 3. Hydrangea quercifolia. 4. Hydrangea paniculata. A form of this species with all sterile flowers, known as H. paniculata grandiflora, is called the ‘“‘peegee” hydrangea. 5. Hydrangea heteromalla (H. xanthoneura, H. Bretschneideri, H. vestita). 6. Hydrangea anomala subsp. petiolaris (H. petiolaris, H. scandens). This plant is often erroneously called Schizophragma hydrangeoides, a quite different Asiatic climber. 7. Hydrangea involucrata. A form of this species with inflorescences consisting of all sterile flowers has been cultivated in Japan. It appar- ently has not been introduced into the United States. 8. Hydrangea aspera subsp. aspera (H. villosa). H. aspera subsp. strigosa (H. strigosa, H. aspera var. macrophylla). H. aspera subsp. ro- busta (H. longipes). H. aspera subsp. Sargentiana (H. Sargentiana). 9. Hydrangea serratifolia (H. integerrima). The only species of the section Cornidia to be cultivated. VoL. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 165 SYSTEMATIC TREATMENT Hydrangea Linnaeus, Genera Plantarum 189. 1754. Hortensia Commerson ex Jussieu, Genera Plantarum, 214. 1789. Cornidia Ruiz & Pavon, Florae Peruvianae et Chilensis Prodromus, 53, t. 35. 1794. Sarcostyles Presl ex Seringe in DeCandolle, Prodromus, 4:15. 1830. Much branched shrubs, erect, or scandent and clinging by means of aerial rootlets; deciduous or evergreen; inflorescences, young branchlets, and leaves variously pubescent; leaves opposite, simple, chartaceous or leathery, penninerved, margins entire, denticulate, or serrate, rarely pin- nately lobed; inflorescences usually terminal, occasionally axillary, on current year’s wood, consisting of compound, much branched clusters, rounded or pyramidal in shape, made up of complicated, dendritically branched systems, the ultimate members of which bear a single flower or small clusters of two to seven flowers; flowers fertile or sterile; fertile flowers small, perfect, usually very numerous, comprising the main body of the inflorescence; calyx tubes adnate to ovary, forming the hypan- thium, apices flat or conical; sepals 4 or 5, inconspicuous, deltoid; petals 4 or 5, ovate or spatulate, valvate, deciduous during anthesis, falling separately or rarely together as a cap; stamens 8 or 10, rarely as many as 20, filaments usually as long as petals or longer, filiform, anthers oblong to somewhat rounded; ovary inferior or half superior, 2 to 4, rarely 5 locular, styles 2 to 4, rarely 5, separate when ovary is inferior, united at their bases on top of conical apical disk of the hypanthium when ovary is half superior, stigmas terminal to more or less decurrent on inner surface of styles; ovules numerous, borne on axile placentas; cap- sules hemispherical or turbinate, dehiscent at apex between the styles; seeds numerous, small, linear or elliptic, sometimes caudate, rarely cir- cular with a wing surrounding body of seed; sterile flowers consisting of 3 to 5 enlarged petaloid sepals, usually lacking other flower parts, conspicuous, few in number, arranged on the periphery of the inflores- cences, or in some cultivated forms numerous and comprising the entire inflorescence. The generic name, Hydrangea, is derived from two Greek roots, hydor, meaning water, and angeion, diminutive of angos, meaning vessel or con- tainer, in allusion to the shape of the mature capsules. GEOGRAPHICAL DISTRIBUTION: eastern Asia, from the Himalayas east- ward to central and southern China, Sakhalin, Japan, the Ryukyu Islands, Formosa, the Philippine Islands, Sumatra, and Java; eastern North America; mountains of Central America southward into the Andes of South America. TYPE SPECIES: Hydrangea arborescens Linnaeus. 166 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. KEY TO THE SECTIONS, SUBSECTIONS, AND SPECIES la. Plants deciduous, erect or scandent, unopened inflorescence not enveloped by broadly ovate floral bracts (except in H. involucrata). Section Hydrangea 2a. Ovary inferior in both anthesis and fruit, capsules truncate at apex, styles separate to their bases. 3a. Seeds with rounded wing surrounding body of seed, petals falling to- gether as a cap or calyptra; plants scandent, clinging by means of aerial rootlets. Subsection Calyptranthe 6. H. anomala 3b. Seeds without rounded wing, petals falling separately, plants erect. 4a. Seeds caudate. (Plants of China and Japan.) Subsection Asperae 5a. Leaves pinnately lobed. 3. H. sikokiana 5b. Leaves not lobed. 6a. Bracts broadly ovate to rounded, few, enveloping the un- opened inflorescence, leaving noticeable scars on falling; capsules pubescent. 4. H. involucrata 6b. Bracts lanceolate, numerous, scattered through the inflo- rescence, covering but not enveloping the unopened in- florescence; capsules glabrous. 5. H. aspera 4b. Seeds not caudate. (Plants of the southeastern United States.) Subsection Americanae 7a. Inflorescence a flat-topped cluster; leaves not lobed. 1. H. arborescens 7b. Inflorescence a pyramidal cluster; leaves pinnately lobed. 2. H. quercifolia 2b. Ovary half superior in both anthesis and fruit; capsules conical at their apices. 8a. Seeds caudate; petals truncate at their bases; styles in fruit shorter than conical disk of capsule. Subsection Heteromallae 9a. Inflorescence a pyramidal cluster, calyx lobes 0.5-1 mm. long. 9. H. paniculata 9b. Inflorescence a flat-topped cluster, calyx lobes 0.8-3 mm. long. 10. H. heteromalla 8b. Seeds not caudate; styles in fruit longer than conical disk of capsule; petals clawed at their bases. Subsection Petalanthe 10a. Inflorescences consisting entirely of fertile flowers, stamens 2 to 3 times as long as styles, leaves with serrations 3-8 mm. deep. 7. H. hirta 10b. Inflorescences consisting of both fertile and sterile flowers; sta- mens same length as styles to as much as two times their length, leaves entire or with serrations less than 3 mm. deep. 8. H. scandens 2c. Ovary inferior in anthesis, becoming half superior in fruit; capsules conical at their apices. Subsection Macrophyllae 11. H. macrophylla 1b. Plants evergreen, shrubs or lianes clinging by aerial rootlets; leaves leathery; floral bracts broadly ovate, few, enveloping the unopened inflorescence, leaving Vou. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 167 noticeable scars on falling. (Plants of Mexico, Central and South America, and one species in the Philippine Islands and Formosa. ) Section Cornidia lla. Inflorescence consisting of a single, terminal, flat-topped or rounded cluster; flowers pink or white; steriie flowers present or absent. Subsection Monosegia 12a. Calyx lobes and petals 4 or 5. 13a. Leaves oval, broadest near middle; sterile flowers present; calyx lobes broadly deltoid, less than 0.5 mm. long. 14a. Branchlets of inflorescence glabrous. (Plants of the Sierra Madre Occidental in the state of Durango, Mexico.) 12. H. Seemannii 14b. Branchlets of inflorescence pubescent with stellate hairs borne on short epidermal papillae. (Plants of Philippine Islands and Formosa.) 14. H. integrifolia 13b. Leaves obovate, broadest above middle; sterile flowers not present; calyx lobes deltoid to semi-orbicular, 1-2 mm. long. (Plants of Guatemala. ) 19. H. Steyermarkii 12b. Calyx lobes and petals 4. (Plants of Central and South America, from Costa Rica southward into the Andes.) 15a. Flowers white; leaves generally 5-14 cm. long, 3—5.5 em. wide; branchlets and inflorescence and lower leaf surface ferruginous tomentose with stellate hairs, which may be partly deciduous with age. 13. H. asterolasia 15b. Flowers pink; leaves generally 10-25 em. long, 4-15 em. wide; branchlets and inflorescence pubescent with single or stellate hairs, hairs 0.1—0.5 mm. long. 16a. Styles usually 3; sterile flowers absent. 18. H. Preslii e®16b. Styles usually 2; sterile flowers usually present. 17a. Styles during anthesis 0.3-1 mm. long and without evident stigmatic papillae; stamens 2—4.5 mm. long. 15. H. Oerstedii 17b. Styles during anthesis 1-2 mm. long and with evi- dent stigmatic papillae; stamens 0.4-3 mm. long. 18a. Stamens 0.4-1.6 mm. long, shorter than or rarely as long as styles; styles always separate. 16. H. peruviana 18b. Stamens 1.5-3 mm. long, longer than styles; styles usually coherent in bud and early anthesis, sepa- rating later. 17. H. diplostemona 11b. Inflorescences consisting of a series of rounded clusters, one above another; flowers white; sterile flowers always absent. (Plants of the Andes of South America. ) Subsection Polysegia 19a. Styles coherent in bud and occasionally during anthesis. 21. H. tarapotensis 19b. Styles always separate. 20a. Stamens 1-1.5 mm. long, not more than one and one-half times as long as styles. 22. H. Jelskii 20b. Stamens 3-7 mm. long, three to six times as long as styles. 21a. Inflorescence and lower leaf surface densely ferruginous pubescent with stellate hairs. (Plants of northern Peru.) 23. H. Mathewsii 168 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 21b. Inflorescence and lower leaf surface glabrous or lightly pubescent with stellate or simple hairs. (Plants of central Chile and adjacent Argentina. ) 20. H. serratifolia Section I. HYDRANGEA Hydrangea, sect. Huhydrangea Maximowicz, Mémoires, Académie imperiale des Sciences de St. Pétersbourg, ser. 7, 10(16):6 (Revisio Hydrangearum Asiae Orientalis), 1867. Hydrangea, subgen. Hortensiella Koehne, Deutsche Dendrologie, 187, 1893. In part. Hydrangea, subgen. Euhydrangea (Maximowicz) Schneider, Handbuch der Laub- holzkunde, 1:384, 1905. Deciduous shrubs, usually erect, rarely scandent and clinging by aerial rootlets; leaves membranaceous or chartaceous; bracts lanceolate, scattered throughout inflorescence (except in Hydrangea involucrata), bracts ocea- sionally absent; stamens usually 10; ovary inferior or half superior; capsule truncate or conical at apex; seeds elliptic to oval, sometimes caudate. TYPE SPECIES: Hydrangea arborescens Linnaeus. The repetition of the generic name for the section containing the type species is in accordance with a rule adopted by the Nomenclature See- tion of the International Botanical Congress held in Paris in 1954 (Stafleu, 1954). Subsection 1. AMERICANAE Hydrangea, sect. Huhydrangea, subsect. Americanae (Maximowicz) Engler, in Engler & Prantl, Natiirlichen Pflanzenfamilien, 3, 2a:204, 1930. As Group, Rehder, Manual Cultivated Trees and Shrubs, p. 291, 1927; as Series, Rehder, Manual Cultivated Trees and Shrubs, ed. 2, p. 286, 1940. Hydrangea, sect. 1, Euhydrangea, ser. 1. Petalanthe, Americanae Maximowicz, Mémoires Académie imperiale des Sciences de St. Pétersbourg, ser. 7, 10(16): 6 (Revisio Hydrangearum Asiae Orientalis), 1867. Flowers white; ovary inferior; capsules truncate at apex; seeds not caudate. North American species. TYPE SPECIES: Hydrangea arborescens Linnaeus. 1. Hydrangea arborescens Linnaeus, Species Plantarum, 937. 1753. Much-branched shrub, 1 to 3 m. tall; deciduous; young branchlets and inflorescence strigose pubescent, or puberulent; leaves ovate, elliptic- ovate, or broadly ovate, acute or acuminate at apex, rounded, subcordate, or cuneate at base, broadest near center or toward base, margins serrate, 6-16 em. long, 2.5-12 em. wide, their length usually 1.5 to 2.5 times their width, lower surface glabrous or nearly so, or pilose or tomentose, upper surface glabrous, or with occasional scattered hairs; inflorescence a rounded, compound, much-branched cluster, 5-10 em. wide; fertile flowers white, very numerous; hypanthium 0.8-1 mm. long; calyx lobes 5, deltoid, VoL. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 169 0.5 mm. long; petals 5, oblong, 2 mm. long, falling separately; stamens 10, 1.5-4 mm. long, the 5 longer ones about twice as long as the shorter ones; ovary inferior; styles 2, or rarely 3, 1 mm. long; capsule 2—-2.5 mm. long, 2-3 mm. wide at apex, somewhat conical at apex, styles 1.2-1.5 mm. long, ribs of capsule 8 or 9, conspicuous; seeds elliptical, blunt at ends, 0.5—0.8 mm. long, 0.4-0.6 mm. wide, prominently striate; sterile flowers white, sometimes absent, usually 4-lobed, occasionally 3-lobed, lobes ovate or orbicular, 8-15 mm. in diameter. (See plate 4, figure 1.) DIsTRIBUTION: Eastern United States from southern New York south- ward to the southern Appalachian Mountains, westward to the Ozarkian Plateaus. (See map 2.) Key TO THE SUBSPECIES OF HyDRANGEA ARBORESCENS 1. Lower leaf surface green, glabrous or nearly so, with hairs, when present, only along the midrib and secondary veins; sterile flowers usually not present. a. subsp. arborescens 1. Lower leaf surface white or gray, variously pubescent; sterile flowers present or absent. 2. Lower leaf surface gray, velutinous, pilose, or tomentose, hairs usually not dense enough to be matted; hairs when examined under magnification of about 50 diameters are seen to be conspicuously roughened by numerous tubercles. b. subsp. discolor 2. Lower leaf surface usually white or silvery, rarely gray, with felt-like indumen- tum, hairs densely matted; hairs when examined under magnification of about 50 diameters are seen to be smooth or with only a few scattered tubercles. c. subsp. radiata la. Hydrangea arborescens Linnaeus subsp. arborescens. Hydrangea arborescens Linnaeus, Species Plantarum, 937. 1753. Hydrangea vulgaris Michaux, Flora Boreali-Americana, 1:268. 1803. A specimen under this name, without locality or other data, in the Michaux Herbarium of the Natural History Museum, Paris, may be the type. Hydrangea arborescens var. vulgaris (Michaux) Seringe in DeCandolle, Prodromus, 4:14. 1830. Hydrangea arborescens var. kanawhana Millspaugh, Preliminary Catalogue of the Flora of West Virginia, 363. 1892. Type from West Virginia, Millspaugh 291 (WVA). Hydrangea arborescens var. australis Harbison, American Midland Naturalist, 11:255. 1928. Type from Union Springs, Alabama, Harbison June 18, 1902. (Biltmore Herb. No. 1339i, US.) (Note: A number of additional synonyms, listed by Rehder in the Bibliography of Cultivated Trees and Shrubs, pp. 199-200 (1949) under Hydrangea arborescens, are not included here. These could not be verified; in some cases, the names were based on cultivated plants and the type specimens were not designated; in other cases, the type specimens could not be located.) Lower leaf surface glabrous or nearly so, with hairs, when present, along midrib; individual hairs conspicuously tubercular, 0.3-1 mm. long, 10-25 microns wide; sterile flowers often not present. 170 CALIFORNIA ACADEMY OF SCIENCES | Proc. 4TH SER. TYPE LOCALITY: Virginia. Type collection: ‘““Dr. Clayton, ex Ameriea, An. 1734 num. 79” (BM); ILLUSTRATIONS : The Botanical Magazine, vol. 13, table 437. 1799. Gleason, H. A. Illustrated Flora of the Northeastern United States and Adjacent Canada, 2: 275. 1952. New Flora and Silva, 4: figure 79, 1932 (habit); 9: figure 34, 1937 (habit), both as Hydrangea arborescens grandiflora. DISTRIBUTION: Southern New York southward through the Appa- lachian Mountains to the coastal plain of central Alabama, westward to the Ozarkian Plateaus. ALABAMA: Bullock, Cullman, DeKalb, Etowah, Hale, Henry, Lee, Madison, Montgomery, St. Clair, Tuscaloosa, and Win- ston counties. ARKANSAS: Baxter, Benton, Clay, Carroll, Craighead, Craw- ford, Faulkner, Independence, Logan, Marion, Perry, Pike, Polk, Pope, Saline, Washington, and Yell counties. Disrricr of COLUMBIA. GEORGIA: Bartow, Catoosa, Clarke, Clay, Cobb, DeKalb, Fannin, Floyd, Fulton, Gilmer, Gordon, Gwinnet, Habersham, Hall, Hancock, Lumpkin, Madison, Pickens, Rabun, Rockdale, Stephens, Towns, and White counties. ILLINOIS: Champaign, Jackson, Johnson, Madison, Randolph, Richland, Shelby, and Union counties. InpIana: Cass, Clinton, Crawford, Floyd, Harrison, Howard, Huntington, Jefferson, Knox, Lawrence, Marion, Martin, Mon- roe, Morgan, Ohio, Orange, Owen, Parke, Perry, Putnam, Ripley, Scott, Vermillion, and Warren counties. Kentucky: Allen, Anderson, Bath, Bell, Boyd, Boyle, Breathitt, Carter, Casey, Christian, Daviess, Edmon- son, Estill, Fayette, Greenup, Grayson, Hancock, Harlan, Jefferson, Jessamine, Johnson, Letcher, Lyon, Madison, McCreary, Ohio, Pike, Union, Warren, and Wolfe counties. LourstANa: West Feliciana Parish. Maryann: Allegany, Cecil, Frederick, Harford, Howard, Montgomery, and Prince Georges counties. Mississtppt: Tishomingo and Wayne counties. Missouri: Barry, Butler, Callaway, Christian, Crawford, Dent, Dunklin, Franklin, Gasconade, Greene, Hickory, Howell, Iron, Jefferson, MeDon- ald, Miller, Newton, Oregon, Ozark, Pulaski, Ripley, St. Charles, St. Genevieve, Shannon, Stoddard, Stone, Taney, Wayne, Webster, and Wright counties. New Jersey: Burlington, Mercer, and Warren counties. New York: Chemung, Steuben, and Tioga counties. NORTH CAROLINA: Alexander, Alleghany, Ashe, Avery, Buncombe, Cherokee, Craven, Cum- berland, Durham, Forsyth, Graham, Haywood, Jackson, Iredell, Macon, Madison, Montgomery, Orange, Polk, Rockingham, Stanley, Stokes, Surry, Swain, Transylvania, Wake, Watauga, Wilkes, Yadkin, and Yancey coun- ties. Onto: Adams, Belmont, Butler, Columbiana, Fairfield, Green, Ham- ilton, Jefferson, Lawrence, Meigs, Pike, Ross, and Scioto counties. OKLA- HoMA: Cherokee, Delaware, Johnston, Leflore, and McCurtain counties. Vou. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA i q(al PENNSYLVANIA: Allegheny, Blair, Bradford, Bucks, Butler, Cameron, Centre, Chester, Clearfield, Elk, Fayette, Franklin, Fulton, Laneaster, Lehigh, Luzerne, Lycoming, Mifflin, Monroe, Montgomery, Northampton, Perry, Philadelphia, Schuylkill, Snyder, Sullivan, and Westmorland coun- ties. SoUTH CAROLINA: Oconee County. TENNESSEE: Bedford, Blount, Campbell, Carter, Cheatham, Cocke, Coffee, Cumberland, Davidson, Fen- tress, Franklin, Grainger, Hamilton, Knox, Pickett, Rhea, Sevier, Shelby, Sullivan, Unicoi, and Washington counties. Vircinta: Arlington, Au- gusta, Bedford, Botetourt, Carroll, Fairfax, Fauquier, Giles, Green, Greens- ville, Isle of Wight, James City, Loudoun, Louisa, Madison, Mecklenburg, Montgomery, Middlesex, Page, Patrick, Rappahannock, Roanoke, Rock- bridge, Rockingham, Scott, Shenandoah, Smyth, Southampton, Surry and York counties. West VirGINIA: Barbour, Boone, Braxton, Cabell, Cal- houn, Doddridge, Fayette, Grant, Greenbrier, Hampshire, Harrison, Jef- ferson, Kanawha, Marion, Marshall, Mercer, Mineral, Monongalia, Mon- roe, Morgan, Nicholas, Ohio, Pendleton, Preston, Raleigh, Randolph, Ritchie, Summers, Taylor, Tucker, Tyler, Upshur, Wayne, Webster, Wetzel, and Wood counties. lb. Hydrangea arborescens subsp. discolor (Seringe) MeClintock, Journal Arnold Arboretum, 37: 373. 1956}. Hydrangea arborescens var. discolor Seringe in DeCandolle, Prodromus, 4:14. 1830. Hydrangea arborescens var. Deamii St. John, Rhodora, 23:208. 1921. Type from Jackson County, Indiana, Deam 28122 (isotype A). Hydrangea arborescens var. Deamii f. acarpa St. John, 1. c. Type from Montier, Shannon County, Missouri. Bush 6116 (holotype GH, isotypes MO, US). Hydrangea Ashei Harbison, American Midland Naturalist, 11:256. 1928. Type ma- terial said to be from Montgomery County, Arkansas, collected by Ashe, but no actual specimen was cited. Placed here because the leaves were described as being pubescent. Hydrangea cinerea Small, Bulletin Torrey Botanical Club, 25:148. 1898. In part. Small cited five syntypes. Of these Lamson-Scribner, June 29, 1890 (NY, US), and Kearney, July 16, 1894 (US, NY, MO), both collected at White Cliff Springs, Tennessee, and Small, August 7-12, 1895 (A, BH), Ringgold, Catoosa County, Georgia, are subsp. discolor. Lower leaf surface velutinous, pilose, or tomentose, usually grayish; individual hairs conspicuously tubercular, 0.83-1 mm. long, 10-25 microns wide; sterile ray flowers present or absent. Type collection of Hydrangea arborescens var. discolor: A cultivated specimen collected by Seringe in 1824 (G). 1 Herbarium specimens belonging to this taxon were annotated before the writer had seen the specimen typifying Seringe’s varietal epithet. Because there was some doubt as to the proper appli- cation of this name previous to this time a combination based on St. John’s var. Deamii was used to annotate the specimens studied earlier. 172 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. DISTRIBUTION: Occurring with subsp. arborescens except in the north- eastern part of the range of that subspecies. ALABAMA: Bibb, Calhoun, Clay, De Kalb, Jackson, Jefferson, Lauderdale, Madison, Marshall, and Tus- ealoosa counties. ARKANSAS: Baxter, Benton, Carroll, Cleburne, Crawford, Cross, Garland, Hot Springs, Newton, Pulaski, Phillips, Poinsett, Pope, St. Francis, Stone, Van Buren and White counties. GEORGIA: Catoosa, Cobb, Dade, Walker and Whitfield counties. ILLINoIs: Calhoun, Cham- paign, Jackson, Jersey, Johnson, Peoria, Pope, St. Clair, Vermillion, and Williamson counties. INpIANA: Bartholomew, Brown, Gibson, Jackson, Knox, Morgan, Monroe, Perry, Posey, and Tippecanoe counties. KEN- TucKY: Livingston and Union counties. Missourr: Cape Girardeau, Car- ter, Dallas, Jefferson, Madison, Newton, Oregon, Ozark, Phelps, Pike, Reyn- olds, Ripley, St. Genevieve, St. Louis, Scott, Shannon, Stoddard and Wash- ington counties. OKLAHOMA: Leflore County. TENNESSEE: Blount, Carroll, Davidson, Decatur, Franklin, Grundy, Hamilton, Marion, Monroe, Obion, Polk, Roane and Van Buren counties. le. Hydrangea arborescens subsp. radiata (Walter) McClintock, Jour- nal Arnold Arboretum, 37: 373. 1956. Hydrangea radiata Walter, Flora Caroliniana, 251. 1788. Hydrangea nivea Michaux, Flora Boreali-Americana, 1:268. 1803. A specimen under this name, without locality or other data, in the Michaux Herbarium, Natural History Museum, Paris, is doubtless the type. Hydrangea nivea Michaux var. conformis Seringe in DeCandolle, Prodromus, 4:14. 1830. Hydrangea cinerea Small, Bulletin Torrey Botanical Club, 25:148. 1898. In part. Small cited five syntypes. Of these Curtiss 833 (June, 1882) (MO, US, GH, UC, BM, W, G, K), Chilhowee Mountains, Blount County, Tennessee, and Kearney, June 24, 1893 (NY), Chilhowee Gap, Blount County, Tennessee, are H. arbo- rescens subsp. radiata. Lower leaf surface covered with a conspicuous white felt-like indu- mentum, occasionally a grayish tomentum; individual hairs either smooth (without any tubercules), 1-3 mm. long, 5-10 microns wide, or with a few scattered tubercules, 1-3 mm. long, 5-20 microns wide; sterile flowers usually present. Type Locauity of Hydrangea radiata: “Carolina.” Type collection: Walter s.n. (BM). DISTRIBUTION: Occurring with the other two subspecies in the southern section of the Blue Ridge Mountains, extending to the piedmont area in the vicinity of the Seneea River in the northwestern corner of South Carolina. Plants with indumentum composed of smooth hairs: GEorGIA: Rabun County. NortH Carotina: Buncombe, Graham, Henderson, Jackson, VoL. XXIX|] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 173 Macon, Madison, Polk, Swain, and Transylvania counties. SoutH Caro- LINA: Anderson, Greenville, Oconee, and Pickens counties. TENNESSEE: Blount County. Plants with indumentum composed of inconspicuously tubercular hairs: NortH CarouiIna: Buncombe, Henderson, Macon, Polk, Swain, and Tran- sylvania counties. TENNESSEE: Blount County. VARIATION IN Hydrangea arborescens Variation in Hydrangea arborescens involves the kind and amount of vesture on the lower leaf surface, the presence or absence of sterile per- ipheral flowers, and the size and shape of the leaves. The flower parts show no variation. The vesture characters of the lower leaf surface have been used to divide the species into subspecies as follows: 1. Lower leaf surface glabrous or nearly so, with only a few scattered hairs along the midrib (plate 1, figure 1). Subsp. arborescens. 2. Lower leaf surface pubescent with erect or spreading, straight, rather firm hairs which vary in amount from sparsely seattered to dense enough to cover the surface and form a gray tomentum (plate 1, figures 2 and 3; plate 2, figure 3). The hairs are covered with numerous tuber- cules, giving them a roughened appearance (plate 3, figure 4), and are 0.5-1 mm. long and 10-25 microns wide. These hairs are found in subsp. discolor. The few hairs along the midveins of subsp. arborescens have also these same dimensions and markings. 3. Lower leaf surface with intertwined fine woolly hairs, usually very dense and forming a white, or rarely gray, felt-like indumentum (plate 1, figure 4). The hairs have a smooth surface (plate 3, figure 1) and are 1-3 mm. long and 5-10 microns wide. Subsp. radiata. 4. Lower leaf surface with indumentum resembling that of 3 above, except for its gray color (plate 2, figure 2) and with hairs which differ in having a few scattered tubercules (plate 3, figures 2 and 3). The hairs are 1-3 mm. long and 5—20 microns wide. These hairs appear more like the hairs of subsp. radiata and therefore the individuals having them are included in this subspecies. The geographical distribution of the three subspecies is given on Map 2. It is seen from this map that subsp. arborescens is the most widespread of the three; that subsp. discolor occurs in the western and southeastern, but not in the northeastern part of the range of subsp. arborescens; and that subsp. radiata occurs only in the southeastern part of ranges of the other subspecies, which is the southern Appalachian Mountains. The dis- tribution of the three subspecies in the southern Appalachians is shown on Map 3. In this area occur the two forms of subsp. radiata, the one having smooth hairs and the other having hairs with seattered tubercules. 174 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Here also occur individuals of subsp. discolor having both “sparse” and “dense” pubescence, as well as subsp. arborescens. Sterile flowers are present in the inflorescences of some individuals and absent in others. There is evidence to show from herbarium specimens that their presence is associated in part with type of pubescence and in part with geographic range. Of 231 individuals of subsp. arborescens in the eastern part of its range, from Virginia to South Carolina, 65, or 27 per cent, had sterile flowers, while in the western part of its range, from Tennessee and Kentucky to Missouri and Arkansas, of 243 indi- viduals, 167, or 69 per cent, had sterile flowers. In the western part of the range of subsp. discolor (Missouri, Arkansas, Indiana, Illinois, and Ohio), of 87 individuals, 14, or 17 per cent had sterile flowers, but in the eastern part of its range, Georgia, Alabama, and Tennessee, where the pubescence is “sparse” to “dense,” of 39 individuals, 36, or 92 per cent, had sterile flowers. In subsp. radiata, 50 individuals of the form having smooth hairs were examined, and 49, nearly 100 per cent, had sterile flowers, and of the form having a few tubereules on its hairs, 9 out of 19, or 47 per cent, had sterile flowers. Since all of the subspecies are found in the southern Appalachian Mountains, and often more than one have been found by different collectors at, or near, the same locality, it seemed of interest to determine whether more than one subspecies occur together, and if so, whether the form of subsp. radiata with seattered tubercules on its hairs would be found. In the summer of 1948, the writer visited this region and collected Hydrangea material in about 50 localities. It was found that for the most part the plants were uniform at a given locality. In only two localities were there more than one subspecies. In Madison County, North Carolina, about 414 miles west of Walnut Gap, there were typical plants of both subsp. arborescens and subsp. radiata. The other locality was in the Pisgah Na- tional Forest, near White Pine Annex Campground, Transylvania County, North Carolina. The three subspecies were in this locality along with several atypical individuals. No. 48-59B (see plate 2, figure 2) was like the one previously referred to subsp. radiata. There were several other individuals (Nos. 48-61B, 48-59A, see plate 2, figure 1, and No. 48-60, see plate 1, figure 5) in which the hairs were short (0.5-1 mm.) and had Map 2. Distribution in the eastern United States of oO Hydrangea quercifolia Hydrangea arborescens @ subsp. arborescens A subsp. discolor W@ subsp. radiata Vou. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 175 176 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. numerous tubercules as in subsp. discolor, but their width was less (5-20 microns) than is typical for this subspecies. Another individual, No. 48-61C, placed in subsp. arborescens because of being nearly glabrous, had a few scattered hairs along the veins which were like those of the individuals of subsp. discolor just described. A search for similar atypical individuals in other localities in the Pisgah Forest area was not successful. It would appear that such indi- viduals are rare. Dunean (Castanea 15: 153, 1950) reported finding subspp. arborescens and radiata growing together in Rabun County, Geor- gia, but he found no “intermediates” in a large number of plants in- spected. Perhaps if the localities where those individuals of subsp. radiata having ineonspicuously tubereulate hairs have been found were searched, other intermediates would turn up. However, it would appear that intermediates are not common and those few which have appeared have not caused the character of the populations to change. 2. Hydrangea quercifolia Bartram, Travels, xviii, nom., 382, t. 6, deser., 408, nom. 1791; ed. 2, xii, 380, t. 6, 406, 1792.2 Species Plantarum, ed. 4, C. L. Willdenow, 2: 634. 17993. Hydrangea radiata Smith, Icones Pictae Plantarum rariorum, t. 12. 1791. Non Walter, 1788. Stout erect shrub, much branched from base, 1-2 m. tall; young branchlets and petioles densely tomentose with long, fine, reddish hairs; branches of inflorescence pilose with short, coarse hairs; leaves suborbicular or oval in outline, 8-25 em. long, mostly 5-lobed, the lobes coarsely serrate, petioles one-eighth to one-third as long as blades; upper leaf surface gla- 2 According to Merrill (Bartonia, 23:10-35, 1945), the citation of H. quercifolia given in Index Kewensis isnot correct. The citation here is Merrill’s correction. 3 Because of the doubt which is attached to Bartram’s Travels as a legitimate place to begin nomen- clature, Willdenow’s publication may be considered to be the earliest valid one for this name. See Merrill, 7. c. pp. 17 and 27. Map 3. Distribution in the southern Appalachian Mountains of Hydrangea arborescens @ subsp. arborescens subsp. discolor A Sparsely pubescent @ tomentose subsp. radiata H@ indumentum white and hairs smooth * indumentum gray and hairs inconspicuously tubercular McOCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA Vou. XXIX] | “Tallule Y uospt, % ow, 9A0D sapen Nese ulmunoW i 178 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. brous; lower leaf surface pilose or tomentose with two kinds of hairs— short, coarse, tubercular ones, and longer, fine, smooth ones which form an intertwining network; inflorescence a compound, much branched cluster, pyramidal in shape, 15-25 em. long; fertile flowers white, very numer- ous; hypanthium 1-1.5 mm. long; calyx lobes 5, deltoid, 0.5-0.8 mm. long; petals oblong, acute, concave, 2—-2.5 mm. long; stamens 10, 3-6 mm. long; styles 2, 3, or 4, 1-1.4 mm. long, somewhat thickened toward apex; cap- sules 1.5-2.5 mm. long, 2-2.5 mm. wide at apex, apex truncate, ribs 7 or 8, inconspicuous; styles 1.2-1.5 mm. long; seeds elliptical, 0.7-0.8 mm. long, 0.3-0.4 mm. wide, prominently striate, blunt at each end; sterile flowers present, usually a single one terminating each lateral branch of the inflorescence, four-lobed, lobes orbicular, 10-15 mm. in diameter. (See plate 4, figure 2.) TYPE LOCALITY: GrorGIA: Crawford County, Sweet Water Brook —= Knoxville Branch (see Merrill, Bartonia 23:27, 1945). TyPE COLLECTION: W. Bartram s. n. (holotype BM). ILLUSTRATIONS : Curtis’s Botanical Magazine, table 975, 1806. New Flora & Silva 4: fig. 80. 1932 (habit). Gleason, H. A. 1952. Illustrated Flora of the Northeastern United States and Adjacent Canada, 2: 275. DISTRIBUTION: Southeastern United States, on the piedmont and coastal plain of western Georgia, Alabama, and Mississippi. (See Map 2.) AvAaBAMA: Blount, Calhoun, Clarke, Coneeuh, Cullman, Dallas, DeKalb, Etowah, Franklin, Jackson, Jefferson, Marshall, Shelby, St. Clair, Tallapoosa, Tuscaloosa, and Winston counties. FLoripA: Calhoun, Gadsden, Jackson PLATE 1. Vesture types in Hydrangea arborescens. Figure 1. Subsp. arborescens. Lower leaf surface glabrous except for few scat- tered hairs along veins. (from McClintock 48-44) Figure 2. Subsp. discolor. Lower leaf surface sparsely pubescent. (from Pal- mer 12640) Figure 3. Subsp. discolor. Lower leaf surface with slightly more hairs than in figure 2. (from Deam 2248) Figure 4. Subsp. radiata. Lower leaf surface covered with white felt-like indu- mentum. (from McClintock 48-53) Figure 5. Subsp. discolor. Lower leaf surface moderately pubescent with hairs finer than those in figure 3. (from McClintock 48-60) The photographs in this plate and in Plate 2 were taken by Richard Johnson. Magnification in all figures on both plates is 19x. Vou. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 179 180 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. and Liberty counties. Grorcta: Bibb, Chattahooche, Clarke, Clay, Craw- ford, Dade, Decatur, Early, Floyd, Harris, Heard, Merriwether, Polk, Randolph, Stewart, Talbot, Taylor, and Upson counties. Louistana: East Feliciana and West Feliciana parishes. Mississippi: Adams, Attala, Copia, Grenada, Hinds, Holmes, Itawamba, Tallahatchie, Tishomingo, and War- ren counties. TENNESSEE: Lincoln, Marion, and Shelby counties. Discussion: Hydrangea quercifolia is remarkably uniform throughout its range. It is characterized by its pyramidal inflorescence and its lobed leaves, by which characters it is distinguished from H. arborescens. In geographic distribution these two species differ, although they overlap in part in the southern part of the range of H. arborescens. Hydrangea quercifolia is found chiefly in the area of the piedmont and coastal plain of western Georgia, Alabama, and Mississippi. Hydrangea arborescens subsp. arborescens and subsp. discolor occur infrequently in this area. Subsection 2. ASPERAE Hydrangea, sect. Euhydrangea Maximowicz, subsect. Asperae Rehder, Plantae Wil- sonianae, 1:39. 1911. Hydrangea, sect. Euhydrangea Maximowicz, subsect. Piptopetalae Maximowicz, Mémoires Académie imperiale des Sciences de St. Pétersbourg, ser. 7, 10(16):8 (Revisio Hydrangearum Asiae Orientalis), 1867. In part. Hydrangea, subgenus Euhydrangea, sect. Japonico-sinensis, subsect. Piptopetalae Schneider, Handbuch der Laubholzkunde, 388, 1905. In part. Flowers white or blue; ovary inferior; capsule truncate at apex; styles free to their bases; seeds caudate. Chinese and Japanese species. Type species: Hydrangea aspera Don. PLATE 2. Vesture types in Hydrangea arborescens and Hydrangea aspera. Figure 1. Hydrangea arborescens subsp. discolor. Lower leaf surface moderately pubescent with hairs somewhat finer, longer and more dense than those on Plate 1, figure 5. (from McClintock 48-59A) Figure 2. Hydrangea arborescens subsp. radiata. Lower leaf surface covered with gray indumentum, hairs not as dense as in Plate 1, figure 4, and having scat- tered tubercules. (from McClintock 48—-59B) Figure 3. Hydrangea arborescens subsp. discolor. Lower leaf surface gray to- mentose. (from McClintock 48-82.) Figure 4. Hydrangea aspera subsp. strigosa. Hairs straight, appressed along raised veins. (from Fang 15568) Figure 5. Hydrangea aspera subsp. aspera. Hairs tending to be erect and spreading. (from Fang 19721) 181 MONOGRAPH OF GENUS HYDRANGEA McCLINTOCK: Vou. XXIX] 182 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 3. Hydrangea sikokiana Maximowicz, Bulletin scientifique Académie im- périale des Sciences de St. Pétersbourg, 31:42. 1887. Shrub, 1-2 m. tall; branchlets, inflorescence and pedicels pubescent with upwardly appressed hairs 0.5-1 mm. long; leaves pinnately lobed with 4 to 6 lobes, 8-21 em. long, 8-20 em. wide, lower surface pubescent along veins and veinlets with hairs 1-1.5 mm. long, upper surface with seat- tered hairs along veins; petioles 2-18 em. long, pubescent; inflorescence a flat-topped, compound, several-branched cluster, 12-30 em. across; bracts few, scattered through inflorescence, lanceolate to ovate, 10-30 mm. long, pubescent as leaves; fertile flowers purplish; hypanthium 1—1.6 mm. long; ealyx lobes 5, broadly deltoid, 0.5 mm. long; petals 5, 2-4 mm. long, 1.5— 2 mm. wide, truncate at base; stamens 10, rarely 8 or 9, 3.5-5.5 mm. long; styles 2, 1-1.5 mm. long; ovary inferior; capsule 2-3 mm. long, styles 1.5-2 mm. long; seeds elliptical, 1 mm. long, tapering at each end into minute tail-like extensions; sterile flowers always present, purplish, 4-lobed, lobes rounded, 1-3 em. in diameter. (See plate 5, figure 1.) TYPE LOCALITY: Japan, Honshu Island, Kii Province (now Wakayama and Mie prefectures). Type collection: Tanaka. Specimen not seen by the writer. DISTRIBUTION: Woods and mixed forests on Honshu and Kyushu islands in Japan, from 100 to 1,000 meters. (See Map 4.) JAPAN. HonsHu IsuaNp: Nara Prefecture: Tonomine, Mt. Odaigahara. Wakayama Pre- fecture: Mt. Koya. Kyusuu Isuanp: Oita Prefecture: Bungo. Kagoshima Prefecture: Mt. Hirishima. Kumamoto Prefecture: Hitoyoschichi. Discussion: Hydrangea sikokiana is distinguished from H. mvo- lucrata, the only other closely related Japanese species, by its pinnately lobed leaves. The American species, H. quercifolia, and H. sikokvana are the only species in the genus which have lobed leaves. PLATE 3. Surface appearance of hairs from lower leat surfaces of Hydrangea arborescens. Figure 1. Smooth hairs of subsp. radiata. (from McClintock 48-27) Figure 2. Hairs with very few tubercules from subsp. radiata. (from Cor- rell 3717) Figure 3. Hairs with few tubercules from subsp. radiata. (from Davis 7683) Figure 4. Hairs with numerous, conspicuous tubercules from subsp. discolor. (from Palmer 35469) Drawings made with a camera lucida. Magnification 350. neg Vou. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA FIG. 1 oo % ‘. 2 ; ft z # \ aN } > & 183 184 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 4. Hydrangea involucrata Siebold, Nova Acta: Academia (caesarea) Leopoldino-Carolina, 14(2):691. 1829. Hydrangea longifolia Hayata, Journal, College of Science, Imperial University, Tokyo, 25(19):91. 1908. Type from Formosa, Kawakami & Nakahara 690, not seen, photo CAS. Shrub, 1-2 m. tall; branchlets, inflorescence and pedicels pubescent with upwardly appressed hairs 0.5 mm. long; leaves ovate, 10-26 em. long, 5-17 em. wide, their length 1.5 to 2.5 times their width; both surfaces pubescent with appressed hairs; petiole 1.5-8 mm. long; inflorescence rounded, compound, many-branched; involucral bracts subtending the inflorescence, 1.5—2.5 em. long, broadly obovate, pubescent with upwardly appressed white hairs on outside, glabrous on inside, caducous, leaving a row of scars at base of inflorescence when falling; in addition, a few bracts, ovate or lanceolate in shape, are scattered through the inflorescence; fertile flowers lavender, hypanthium 1.2-1.5 mm. long; calyx lobes 5, deltoid, 0.2-0.6 mm. long; petals 5, 2-3 mm. long, truncate at base; stamens 10, rarely 11 or 12, 3.5-6 mm. long; ovary inferior; styles 2 or 3, 1-2 mm. long; capsules 3-4 mm. long, pubescent, styles 2-2.5 mm. long; seeds elliptical, 0.8-1 mm. long, tapering at each end into minute tail-like extensions; sterile flowers always present, lavender, 4-lobed, lobes rounded, 1-3 em. in diameter. (See plate 5, figure 5.) PLATE 4. Flowers and capsules of some American species of Hydrangea. Figure 1. Hydrangea arborescens. a. flower (without petals and some of sta- mens. Biltmore herb. 1339c). b. capsule (Biltmore herb. 8212d). Figure 2. Hydrangea quercifolia. a. flower. b. capsule. Figure 3. Hydrangea asterolasia. a. flower (Skutch 3342). b. capsule (A. Smith H470). Figure 4. Hydrangea peruviana. a. flower (Allen 534). b. capsule (Standley 39080). Figure 5. Hydrangea Mathewsii. Flower (Mathews 3051). Figure 6. Hydrangea Preslii. a. flower (Weberbauer 6776). b. flower (Skutch 2630). Figure 7. Hydrangea diplostemona. a. flower (Spruce 4328). b. young cap- sule (Spruce 4328). Figure 8. Hydrangea tarapotensis. a. flower (Smith 2627). b. young capsule (Lawrence 775). Figure 9. Hydrangea integrifolia. a. flower (Clemens June 1924). b. capsule (Wilson 9691). Figure 10. Hydrangea serratifolia. a. flower (Morton Dec. 3, 1906). b. capsule (Sargent Jan. 12, 1906). VoL. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 185 186 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TYPE LocALITty: Japan. Type collection: Collected in Japan by von Siebold. The specimens collected in Japan by von Siebold are in the Rijksherbarium in Leiden. Presumably the type of H. involucrata is there but the writer has not seen it. ILLUSTRATION: Siebold and Zucearini. Flora Japonica, plates 63 and 64. 1839 or 1840. DistRIBUTION: Woods and mixed forests on Honshu Island in Japan, from near sea level to 1,500 meters, and on Formosa. JAPAN. HoNsHuU IsLAND: Chiba Prefecture: Owari, Kiyozumi-yama. Fukushima Prefee- ture: Mt. Haguro. Gifu Prefecture: Norikura, Washiga-take. Gumma Pre- fecture: Ikaho. Kanagawa Prefecture: Yokohama, Miyanoshita, Hakone Mts., Kamakura, Mt. Takao. Nagano Prefecture: Usui-toge, Mt. Izuna, Mt. Tsubakura, Kuramoto, Kiso near On-take-san, Asamayama. Shiga Prefecture: Shinano. Tochigi Prefecture: Nikko. Tokyo Prefecture: Tokyo, Mt. Takao, Hachijo. Yamanashi Prefecture: Motsuko. (See Map 4.) Discussion: Hydrangea involucrata bears a close resemblance to H. aspera in flower and fruit structures, and in the shape of the leaves. The two species may be distinguished by the tomentose capsules and the in- volueral bracts of H. involucrata. These bracts which envelop the un- opened inflorescence resemble those found in the section Cornidia. The distribution of Hydrangea involucrata as listed above includes localities from Japan only. The species, however, occurs in Formosa as indicated by a specimen collected by Kawakami and Nakahara at Taito on the southeast coast in January 1906. This specimen, the type of Hydrangea longifolia Hayata, was not seen by the writer. A photograph of the specimen shows the sears left by deciduous involucral bracts, the presence of which identifies it with H. amvolucrata. PLATE 5. Flowers and capsules of some Asiatic species of Hydrangea. Figure 1. Hydrangea sikokiana. a. flower (Sakurai August 16, 1909). b. cap- sule (Komeda E53). c. seed (Komeda E53). Figure 2. Hydrangea heteromalla. a. flower (Wilson 2184). b. capsule (Li 23). c. seed (Li 23). Figure 3. Hydrangea paniculata. a. flower (Chung & Sun 469). b. single petal. c. capsule (Canton Christian College herb. no. 8738). d. seed (as preceding). Figure 4. Hydrangea anomala. a. flower (Wilson 4901). b. united petals or calyptra (as preceding). c. capsule (Steward et al. 502). d. seed (as preceding). Figure 5. Hydrangea involucrata. a. flower (Dorsett & Morse 980). b. capsule (Suzuki December 31, 1948). c. seed (Suzuki December 31, 1948). VoL. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 187 y Vent ve ttn we tty Ret wea) OH 188 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 5. Hydrangea aspera D. Don, Prodromus Florae Nepalensis, 211. 1825. Shrub or small tree, 1-4 m. tall; branchlets and inflorescence pubescent with stiff and upwardly appressed or curling and erect hairs; leaves variable in shape, lanceolate to ovate, broadly ovate or deltoid, the lanceo- late or ovate ones cuneate or rounded at base, 5-35 em. long, 1.5-15 em. wide, their length 2.5 to 7 times their width, serrate or serrulate along margin, petioles 0.5-6.5 em. long, the broadly ovate or deltoid ones rounded or cordate at base, 10-25 em. long, 7-17 em. wide, serrate or doubly ser- rate along margin, petioles 4-21 em. long; vesture on lower leaf surface variable and similar to that of stem; inflorescence a flat-topped, many- branched cluster, 10-30 em. broad; bracts lanceolate, densely hairy on outer surface, glabrous on inner, caducous or remaining; fertile flowers white or blue, numerous, hypanthium 1—1.5 mm. long; sepals 5, 0.4-0.5 mm. long; petals 5, 2-2.5 mm. long, truncate at base, deciduous before stamens; stamens 10, 2-7 mm. long, usually alternately long and short; ovary com- pletely inferior; styles 2, 3 or 4, 0.5-1.5 mm. long; capsules 3-5 mm. long; seeds elliptical, 0.7-0.8 mm. long, 0.3-0.4 mm. wide, tapering at each end into minute tail-like extensions; sterile flowers always present, white, 4-lobed, lobes 1-3.5 em. long, usually slightly longer than wide, with entire or fimbriate margins. (See plate 6, figure 6.) DistriIBuTION: In woods or thickets, eastern Himalaya Mountains from 1,500 to 2,300 meters; western China from 700 to 4,000 meters; central and eastern China from 250 to 2,100 meters; Formosa, Sumatra and Java from 1,000 to 2,500 meters. (See Map 4.) KEY TO THE SUBSPECIES OF HYDRANGEA ASPERA 1. Stems, lower leaf surface, and petioles strigose or velutinous. 2. Leaves lanceolate or narrowly ovate, their length usually two to five times as great as their width, rounded at base; petioles wsually 1-5 cm. long. 3. Lower leaf surface strigose, hairs appressed, straight. b. subsp. strigosa Map 4. Distribution in eastern Asia of Hydrangea aspera HM subsp. strigosa A subsp. aspera (inset Sumatra and Java) Vv subsp. robusta * subsp. Sargentiana @ subspecies not determined A Hydrangea sikokiana o Hydrangea involucrata MONOGRAPH OF GENUS HYDRANGEA 189 McCLINTOCK: Vou. XXIX] 190 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH SER. 3. Lower leaf surface velutinous, hairs erect, somewhat curled, occasionally long coarse pili along midvein, petiole and stem. a. subsp. aspera 2. Leaves ovate or broadly ovate, usually less than two times as long as wide, base truncate or cordate; petioles wsually 5-17 cm. long. c. subsp. robusta 1. Stem, lower leaf surface along midveins, and petioles velutinous, and in addi- tion, interspersed with conspicuous, fleshy trichomes, 2—5 mm. long, split at apex; leaves large, 15 cm. or more long, lower surface velutinous. d. subsp. Sargentiana 5a. Hydrangea aspera D. Don subsp. aspera. Hydrangea aspera D. Don, Prodromus Florae Nepalensis, 211. 1825. Hydrangea oblongifolia Blume, Bijdragen tot de Flora van Nederlandsch Indié, 920. 1826. Type from Java, Blume, s. n. (isotypes W-U, NY). Hydrangea vestita var. fimbriata Wallich, A Numerical List of Dried Specimens of Plants in the East India Company’s Museum, published in 1828. Name only. Type from Nepal, Wallich in 1821 (K—W, listed in Wallich’s Catalogue under No. 440). Hydrangea longipes Hemsley var. lanceolata Hemsley, Journal Linnaean Society Botany, 23:274. 1887. Type from Patung, Hupeh, Henry 1786. (K, according to Rehder in Plantae Wilsonianae 1:40, this is the type.) Hydrangea aspera var. strigosior Diels, Botanische Jahrbiticher, 29:375. 1900. Type from Szechuan, Bock & von Rosthorn 2546 (holotype O). Hydrangea Kawakamii Hayata, Journal, College of Science, Imperial University, Tokyo, 25(19):90. 1908. Type from Formosa, Kawakami & Mori 1875 (type not seen, photo CAS). Hydrangea aspera var. scabra Rehder, Plantae Wilsonianae, 1:31. 1911. Type from Kangting (Tatsienlu), Sikang, Wilson 4485 (holotype A). Hydrangea aspera var. velutina Rehder, 1. c. 30. Type from Wa-Shan, Szechuan, Wilson 2405 (holotype A, isotypes K, E). Hydrangea fulvescens Rehder, 1. c. 39. Type from Wushan, Szechuan, Wilson 1373 (holotype A, isotypes EK, K, W). PLATE 6. Flowers and capsules of some Asiatic species of Hydrangea. Figure 1. Hydrangea hirta. a. flower (Wilson 6958). b. petal (as preceding). c. capsule (Wilson 7680). d. seed (as preceding). Figure 2. Hydrangea macrophylla subsp. serrata. a. flower (Henry M83). b. petal (as preceding). c. immature capsule (as preceding). d. mature capsule (Wilson 7621). e. seed (as preceding). Figure 3. Hydrangea macrophylla subsp. macrophylla, immature capsule (Su- zuki July 1917). Figure 4. Hydrangea scandens subsp. chinensis. a. flower (Tso 21020). b. petal (as preceding). c. capsule (Steward & Cheo 1003). Figure 5. Hydrangea scandens subsp. liukiuensis. a. flower (Beattie & Kuri- hara 10646). b. capsule (Wilson 7797). c. seed (as preceding). Figure 6. Hydrangea aspera. a. flower (Wilson 2401). b. capsule (Wilson 4337). Vou. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 1s 192 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Hydrangea glabripes Rehder, I. c. 30. Type from Fang Hsien, Hupeh, Wilson 2391 (holotype A, isotypes E, K, W). Hydrangea villosa Rehder, 1. c. 29. Type from Wenchuan Hsien, Szechuan, Wilson 1227 (holotype A, isotypes E, K). Hydrangea Rehderiana C. K. Schneider, Handbuch der Laubholzkunde, 2:940. 1912. Type from Nanto, Hupeh, Wilson 1533 (holotype W, isotypes NY, K). Premna Merinoi Léveillé, Sertum Yunnanense, 3. 1916. Type from “brousses des montagnes a Siai-Ho, 2700 m.”, Yunnan, Maire May-July, 1912 (fragment of holotype A). Leaves lanceolate or narrowly ovate, their length usually two to five times as great as their width, rounded at base; petioles usually 1-5 em. long; lower leaf surface velutinous, hairs erect, or somewhat curled, oec- easionally long coarse pili along midvein, petiole and stem. (See plate 2, figure 5.) Type LocaLity of Hydrangea aspera: Narainhetty, Nepal. Type col- lection: Buchanan September 26, 1802 (holotype BM). ILLUSTRATIONS : H. H. Hu & W. Y. Chun, 1927. Icones Plantarum Sinicarum, fase. I, plate 28. New Flora and Silva 7: plate opposite p. 177. 1935, as H. villosa. B. Hayata, 1908. Flora Montana Formosae in Journal, College of Sei- ence, Imperial University, Tokyo 25(19), plate 8, as H. Kawakami Hayata. DISTRIBUTION: CHINA. ANHweI: Kimen, Wang Shan. CHEKIANG: T’ien-mu Shan, King Yuan, Suian, Yentang Shan, Taishun. Hunan: Anfu, Sinning, Changning, Sikwangshan district. HupeH: Changyang, Ch’ing young Hsien, Enshih Hsien, Fang Hsien, Hsing Shan Hsien, He-tan-kon, Ichang, Liang sung you, Peh Yang Tsai, Mo Yu Ping, Nanto, Paokang, Patung, Sui Yeh See, Siao-Ya-tze, Wushan, Wuchang. Krtanost: Kiu Chi Hsien, Yungshiu. Kwanest: Pai-yun-an, Ling-chwan district. KWANGTUNG: Lokchong district. KwrtcHow: Kweiting, Pi-tsieh-Hsien, Fan Ching Shan, Tsunyi Hsien, Tungtze. Sr1kaNnG: Lushan via Paohing (Mouping), Paohing, Yaan. SZECHUAN: Chung Hsien, Hsi-Chang between Huangnipu and Yaan, Huangnipu, Kwanhsien, Lungan, Nanchuan, Omei Shan, Opien, Wanhsien, Yun-ching Hsien. YUNNAN: Salwin Valley near Champutong, Io chan, Kiao me ti, Ku-long-tschang, Lanping, Lungling, Lu-se, Lichiang, Mengtsz, Po- shang, Pingpien, Shuang-chiang, west flank Shweli-Salwin divide, banks of Shweli River, Shunning, Shang pa, between Tengyueh and Lungling, Weisi, Wenshan. BURMA. Bhamo district near Sinlun Kaba, Kachin Hills. BHU- TAN. Tunbje Punakha. INDIA. West BENGAL: Darjeeling. SIKKIM: Rumbhi Jhora. TIBET. Tsarong, Ngawchang Valley. SUMATRA. Habin- saran, Seriboe, road from Siantar to Brastagi. FORMOSA. Arisan, Kiraiki. Vout. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 193 5b. Hydrangea aspera D. Don subsp. strigosa (Rehder) McClintock, Journal Arnold Arboretum, 37: 373. 1956. Hydrangea strigosa Rehder, Plantae Wilsonianae, 1:31. 1911. Hydrangea aspera D. Don var. macrophylla Hemsley, Journal Linnaean Society, Botany, 23:273. 1887. Type from ‘“Nanto and mountains to the northward,” Hupeh, Henry s. n. (K). Hydrangea aspera D. Don var. angustifolia Hemsley, 1. c. 273. Type from Ichang, Hupeh, Henry s. n. (K). Hydrangea aspera D. Don var. typica Diels, Botanische Jahrbiicher, 29:375. 1900. Type from Nanchuan, Szechuan, Bock & von Rosthorn 299 (holotype O). Hydrangea aspera D. Don var. sinica Diels, l. c. 375. Type from Nanchuan, Szechuan, Bock & Rosthorn 629 (holotype O). Hydrangea Hemsleyana Diels var. lanceolata (Hemsley) Diels, J. c. 376. (Based on H. longipes Hemsley var. lanceolata Hemsley. ) Hydrangea aspera D. Don var. strigosior Diels, l. c. Type from Wei kuan, Szechuan, Bock & von Rosthorn 2546 (holotype O). Hydrangea villosa Rehder var. Mairei Léveillé, Catalogue des Plantes du Yunnan, 254. 1917. Type from Kiao me ti, Yunnan, Maire July—October 1912 (isotype A). Hydrangea villosa Rehder forma sterilis Rehder, Journal Arnold Arborteum, 11:161. 1930. Type from Changyang, Hupeh, Wilson 1473a (holotype A, isotypes K, EH, Wale Leaves lanceolate or narrowly ovate, their length usually two to five times as great as their width, rounded at base; petioles usually 1—5 em. long; lower leaf surface strigose, hairs appressed, straight. (See plate 2, figure 4.) Type Locatity of Hydrangea strigosa: Techang, Hupeh, China. Type collection: Wilson 765 (holotype A, isotypes K, US, E, W). ILLUSTRATIONS : New Flora and Silva, 9: figure 33, 1937 (habit), as Hydrangea strigosa var. macrophylla. Curtis’s Botanical Magazine, table 9324, 1933, as H. strigosa Rehder. DistripuTION: CHINA. Huprn: Changyang, Chienshih, Ichang, On tan tsian. Kwanost: Tzu Yuen district. KwricHow: Kweiting, Lao Shan, Luneli, Pichieh, Pin fa, Pi-tsieh-Hsien, Tating, Touchan. Srkana: Kang- ting, Lushan via Paohing (Mouping), Ta Kwan. SzecHuan: Chia-ting, Kwanhsien, Omei Shan, Opien, Tien-Tsian, Wa-Shan, Wenchwan Hsien, Weiknan. YuNNAN: Champutong, Chih-tse-lo, Chungtien on bank of Yangtze, Ku-long-tschang, Liang-shan, Lichiang, Long ky, Lon pon, Lou Lan, Ma chou, Menetsz, Paoshan, Pingpien, Sichour-hsien, Shang pa, Tao Shan between Chao and Hungai, Tengyueh, Tchen fong chan, Tsekou, Tsan Shan Range near head of Yangpi Valley, Tunghai, Weisi, between Yangpi and Yungchang-fu, mountains of Yangpi River drainage basin, Yaoan. 194 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. BURMA. Hpimaw. NEPAL. Barpak Village, Narainhetty, Thanjey. JAVA. West side of Jang Plateau at Besoeki, Dieng Plateau, Mt. Kendeng, Mt. Malabar, Preanger, Mt. Soembing, Tjibodas on Mt. Gedah. FOR- MOSA. Mt. Taihei, Mt. Nanko, Nai-taroko. 5e. Hydrangea aspera Don subsp. robusta (Hooker f. & Thomson) Me- Clintock, Journal Arnold Arboretum, 37: 373. 1956. Hydrangea robusta Hooker f. & Thomson, Journal Linnaean Society of London, Botany, 2:76. 1858. Hydrangea robusta var. Griffithii C. B. Clarke in J. D. Hooker, Flora of British India, 2:404. 1878. Type from “Eastern Bhotan,” Griffith 2495 (holotype K, isotype P). Hydrangea longipes Franchet, Nouvelles Archives, Muséum d’histoire naturelle, Paris, ser. 2, 8:228 (Plantae Davidianae, 2:45). 1885. Type from Paohing (Moupin), Sikang, David in 1869 (holotype P, isotypes P). Hydrangea longipes, Hemsley, Journal Linnaean Society of London, Botany, 23:273. 1887. Type from Hupeh, Henry s. n. (K). Hydrangea Rosthornii Diels in Engler’s Botanische Jahrbiicher ftir Systematik, 29:375. 1901. Type from Nanchuan, Szechuan, Bock & von Rosthorn 471 (isotype W). Hydrangea Hemsleyana Diels, 1. c. 376, based on Hydrangea longipes Hemsley, not Franchet. Hydrangea aspera var. longipes (Franchet) Diels, 1. c. 374, based on Hydrangea longipes Franchet. Hydrangea Mavrimowiczii Léveillé, Bulletin Académie international de Géographie Botanique, 12:114. 1903. Syntypes from Kweichow, Martin & Bodinier 1654 (H, P), and Cavalerie 22 (FE). Leaves ovate or broadly ovate, usually less than two times as long as wide, base truncate or cordate; petioles usually 5-17 em. long; lower leaf surface strigose, hairs appressed, straight. Type LocaLity of Hydrangea robusta: Sikkim. Type collection: J. D. Hooker s.n. (holotype K, isotypes K, NY, BM, C, G, P, W, 8S). ILLUSTRATIONS : H. H. Hu and W. Y. Chun, Icones Plantarum Sinicarum, fase. III, plate 140, as Hydrangea longipes Rehder, plate 141, as Hydrangea Ros- thorn Diels. DistripuTION: CHINA. Honan: Lushih, Sunghsien. Hunan: Anfu. Hureu: Changlo Hsien, Changyang, Chienshih, Hsing Shan Hsien, Hiang sung you, Ichang, Nanto, Paokang, Patung, Wan Tsao Chan. Kansv: Lower Tebbu country between Hera Sambaku & Dayaya gorge, Wantsang. Kwanest: Chuen Yuen, Kweilin district, Sankiang Hsien. KWEICHOWw: Lao Shan, Tunyun. SuHanst: Locality not stated. SHenst: Mt. Kin-quan- san, Tai-pei-shan. Skane: Kangting, Paohing, Wu Ya Ling. SzecHUAN: VoL. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 195 Hungya, Nanchuan, Omei Shan, Opien, Wa-shan, Wenchwan Hsien, Weik- nan. YUNNAN: Kiukiang Valley (Taron), Long ky, Shang pa, Tchen fong ehan. BURMA. Bhamo. BHUTAN. Mirichoma Timpu, Trashiyangsi. INDIA. Urtar PrapescH: Garhwal. West BenGAauL: Darjeeling. SIKKIM: Khursiang, Rhikisun, Sureil. TIBET. Yigrong Valley. FORMOSA. Ari- san, Ariko-Banti. 5d. Hydrangea aspera Don subsp. Sargentiana (Rehder) McClintock, Journal Arnold Arboretum, 37: 373. 1956. Hydrangea Sargentiana Rehder, Plantae Wilsonianae, 1:29. 1911. Stem, lower leaf surface along midveins, and petioles velutinous, and in addition, interspersed with conspicuous, fleshy trichomes, 2-5 mm. long, split at apex; leaves broadly ovate, 15 em. or more long. Type Locauity: Hsing Shan Hsien, Hupeh Province, China. Type dpa?) collection: Wilson 772 (holotype A). ILLUSTRATIONS: H. H. Hu & W. Y. Chun, 1933. Icones Plantarum Sinicarum, fase. ITI, plate 138. New Flora and Silva, 12: figure 39, 1940 (habit), and 9: figure 32, 1937 (habit). Curtis’s Botanical Magazine, table 8447. 1912. DistRIBUTION: CHINA. Hupen: Hsing Shan Hsien, Wan Tsao Chan. Discussion oF Hydrangea aspera Hydrangea aspera is a wide-ranging species, occurring from the eastern Himalayas across western and south-central China and on the islands of Formosa, Sumatra, and Java. It varies throughout this range in size and shape of the leaves and in the pubescence of the lower leaf surface. Most of the names listed in the synonymy were based on such variants. The leaves vary in shape from lanceolate to ovate, often broadly ovate; their bases are cuneate, rounded or truneate, or occasionally cordate. In actual size they vary in length from 8 em. to as much as 30 em. The petiole leneth varies also, and is often associated with leaf shape. Thus a broadly ovate leaf usually has a longer petiole than one which is lanceolate. The pubescence may be strigose with straight hairs which are appressed and arranged along the veins and veinlets more or less at right angles to them in a regular fashion, or arranged along the veins somewhat irregularly, or the pubescence may be velutinous with curled erect hairs. Although various combinations of leaf shape and vesture occur, most individuals fall into the following categories: 1, leaves lanceolate or ovate, and 2, 196 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. leaves ovate; often broadly so. In the first of these categories, the pubes- cence may be either strigose, referred here to subsp. strigosa, or velutinous, referred here to subsp. aspera. In the second category, the pubescence is mostly strigose and appressed, referred here to subsp. robusta. Most of the herbarium material examined can be referred to these three taxa, but there are some specimens which clearly are intermediate. A fourth group, subsp. Sargentiana, has large, broadly ovate leaves, but its chief distinguishing character is the presence of conspicuous, more or less turgid trichomes on the stem, petioles, and lower leaf surface along the midvein. Don stated in his deseription of Hydrangea aspera that this species was based on a collection made by “Hamilton” at Narainhetty, Nepal. In the British Museum (Natural History) there is a specimen from the Lam- bert Herbarium collected by “Dr. Buchanan” at Narainhetty, September 26, 1802. It may be considered to be the type of H. aspera. Stapf in discussing the variation of Hydrangea strigosa Rehder (Cur- tis’s Botanical Magazine, 156: tab. 9324, 1933) referred to the relation of this taxon to H. aspera Don, stating that the two may be given specific rank or be treated as minor groups within H. aspera sensu lato (1. H. aspera protonyma and H. aspera strigosa). Stapf clearly stated that he recognized the first alternative. Since Stapf did not accept the cate- gory referred to by the name H. aspera strigosa, his publication of this combination is not valid according to Article 43 of the International Rules which states that a name is not valid which is not accepted by the author who published it. Subsection 3. CALYPTRANTHAE Hydrangea, sect. Hydrangea, subsect. Calyptranthe (Maximowicz) McClintock, Journal Arnold Arborteum, 37:373. 1956. Hydrangea, sect. Calyptranthe Maximowicz, Mémoires Académie imperiale des Sciences de St. Pétersbourg, ser. 7, 10(16):6 (Revisio Hydrangearum Asiae Orientalis), 1867. Schneider, Handbuch der Laubholzkunde, 1:384, 1905, as subgenus. Flowers white; petals cohering at apex and falling together as a cap or calyptra; stamens 10, 15 or 20; ovary inferior; capsule truncate at apex; styles free to their bases; seeds with a rounded wing surrounding body of seed. Only one species, Hydrangea anomala D. Don, widespread in eastern Asia. 6. Hydrangea anomala D. Don, Prodromus Florae Nepalensis, 211. 1825. Woody climber; branchlets and inflorescence sparsely pubescent with upwardly spreading hairs, 0.5 mm. long; leaves ovate, cuneate, truncate, or cordate at base, acute or acuminate at apex, 6-15 em. long, 3.5-10.5 em. wide, their length 1.5 to 2 times their width, glabrous except for scattered Vou. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 197 hairs or clumps of hairs along midvein on lower surface; inflorescence a flat-topped, several-branched cluster; involucral bracts lanceolate or ovate- lanceolate, membranous; fertile flowers white; hypanthium 1.5-2 mm. long; calyx lobes 5, 0.5-0.9 mm. long; petals 5, 2-3 mm. long, remaining at- tached and falling as a cap before the stamens elongate, cap apiculate or rounded at apex; stamens 9-20, 3-5 mm. long, 3 to 4 times as long as styles; ovary inferior; styles 2 or 3, 1-1.3 mm. long; capsule glabrous, 2.5-4.5 mm. long, styles 1-3 mm. long; seeds 1.5-2 mm. long, 1-1.5 mm. wide, nearly round, winged all around; sterile flowers usually present, white, sepals rounded, 1-2 em. in diameter. (See plate 5, figure 4.) DISTRIBUTION: Woods and mixed forests in Japan from sea level to 2,000 meters, on Honshu, Kyushu and Hokkaido islands; in Korea on Dagelet and Quelpaert Islands; in the eastern Himalaya Mountains of northern India, from 1,200 to 2,500 meters, and in west central to east. central China, from 600 to 3,800 meters. (See Map 5.) Kry TO THE SUBSPECIES IPS EA TIVE TSS OY UO milli. eet ree a oe a on ee ee eee cee subsp. anomala lee Stamens a5) to 20) 2 ee SF dre Oa ee a subsp. petiolaris 6a. Hydrangea anomala D. Don subsp. anomala. Hydrangea anomala D. Don, Prodromus Florae Nepalensis, 211. 1825. Hydrangea altissima Wallich, Tentamen Florae Napalensis, 2: t. 50. 1826. Type from Nepal, Wallich Catalogue No. 439/1. Stamens 9 to 15. ILLUSTRATIONS: N. Wallich, Tentamen Florae Napalensis, 2: t. 50, 1826. As Hydrangea altissima. H. H. Hu & W. Y. Chun, Icones Plantarum Sinicarum, fase. III, plate 142. 1933. DISTRIBUTION: Eastern Himalayas and western and central China. CHINA. ANuHwer: Hwang Shan. CuHEKkiane: Tihtai Shan; Kingyuan. Hunan: Yun Shan near Wukang. Hupen: Hsin Tien-tsze; Paokang; Iehang; Hingshan; Patung. Kwanest: Ling Wan district; Lucheng; Shan Chuen to Tzeyuan district. KwericHow: Fan Ching Shan; Shihtsien district. SzecHUAN: Omei Shan; Nanechwan; Kwanhsien; Wushan; O’Pien Hsien; Ma-pien Hsien; Wa-shan; Tchen-keou-tin. YUNNAN: Tali Range; Mt. Gitsa west of Mekong and north of Weisi; Yungshan; Weisi; Shang-pa; Kungshan (Chamutang, Champutong); Niangonang; Mingkwong Valley; Kani Pass, N. Maikha-Salween divide; Shweli-Salween divide; Ma Nung Mt. near Tali. INDIA. Assam: Talung Dzong, Balipara frontier. Urrar [ Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES *(SpUBIS] OUTAAI[IYd JOSUL) sysuauryo ‘dsqns suapunos nabuniphiy Me pue (SedAv[eullzZ Ut9}SeM JOST) DIJDWOUD DabUDAPAH @® JO vIsy Udosvo UL UOTINGIAISIG ‘“G dVIN Vou. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 199 PrapvesH: Dehra Dunn Dist., Deohan; Chakrata; Siwalik and Jaunsar divi- sions; Chekerli, Siwahk and Jaunsar divisions; Landour; Dhanolti; Garh- wal Dist., Tehri Dist. Sikkim: without definite locality. HmacHaL Pra- DESH: Chamba Dist.; Simla district. Punsasp: Chenab Valley on ascent to Pass Punjab; Kangra Dist.; Nagkanda; Dharmsala; Kulu; Jibhi; Bushire; Manali in Lahul subdivision; Kanugadh; Kumaun dist. Wrest BENGAL: Darjeeling. NEPAL. Doti; Montes Shandaghery, Sherpore; Sermore; Thare. TIBET-BURMA FRONTIER. Seinghku Wang; Adung Valley; Kangfang route. BHUTAN. Tilageng Punakha; Kandam Punakha; Shamb- lng, Kuru chu valley; Transmiyangse Dzone. 6b. Hydrangea anomala D. Don subsp. petiolaris (Siebold & Zucearini) datas L956; ( Hydrangea petiolaris Siebold & Zuccarini, Flora Japonica, 1:108, t. 54. 1839 or 1840. McClintock, Journal Arnold Arboretum, : Hydrangea ? bracteata Siebold & Zuccarini, 1. c. 172, t. 92. 1841. No type specimen was designated for this nor the following name. The plates may be considered to represent the types in each case. Hydrangea cordifolia Siebold & Zuccarini, J. c. 113, t. 59, fig. 2. 1839 or 1840. Hydrangea scandens Maximowicz, Mémoires Académie imperiale des Sciences de St. Pétersbourg, ser. 7, 10(16):16. 1867. Non Seringe 1830, nec Poeppig 1830. Based on the above two taxa of Siebold and Zucecarini. Hydrangea glabra Hayata, Journal College of Science, Imperial University, Tokyo, 25 (art. 19):89, plate 6. 1908. Type from Formosa, Kawakami & Mori 1787. (Type not seen, photo CAS.) Hydrangea tilaefolia Léveillé, in Fedde’s Repertorium Novarum Specierum 8:282. 1910. Three syntypes, Taquet 809 (K, E, G), Fawrie 1654 (H, G, BM), 358 (B, BM), all from Dagelet Island, off the south coast of Korea. Stamens 15 to 20. TYPE LOCALITY: Japan. Type specimen: No specimen is mentioned. The plate (Flora Japonica 1: t. 54) may, therefore, be considered to rep- resent the type. ILLUSTRATIONS : B. Hayata. Flora Montana Formosae in Journal, College of Science, Im- perial University, Tokyo, 25 (art. 19): plate 6. 1908. As Hydrangea glabra Hayata. Curtis’s Botanical Magazine, plate 6788. 1884. As H. petiolaris. Siebold and Zucearini. 1839 or 1840. Flora Japonica, 1 : t. 59, figure IT, as H. cordifolia, and t. 54, as H. petiolaris. New Flora and Silva, 1: figure 43, 1929 (habit). New Flora and Silva, 9: figure 31, 1937 (habit). This, and the above illustration, as H. petiolaris. 200 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH SER. DISTRIBUTION: SOVIET FEDERATED SOCIALIST REPUBLIC. Sakhalin (Karafuto) : Oko. JAPAN. Honsuu Isuanp: Aomori Prefecture: Tanabu; Hakkoda; Mts. of Shichinoki. Osaka Prefecture: Mt. Kongosan. Akita Prefecture: Akita. Gifu Prefecture: Mt. Kasagi; Mitaka. Hyogo Prefecture: Tii-mura; Ashiu. Nagano Prefecture: Otake-gawa; Mt. Izuna; Shinano. Wakayama Prefecture: Sitigawa-mura. Yamagata Prefecture. Nara Prefecture: Tonamine; Mt. Misen. Kanagawa Prefecture: Mt. Ha- kone. Tochigi Prefecture: Chuzenji; Shiobara. Shizuoka Prefecture: Sho- jiko; Mt. Amagi on Idzu Peninsula. Iwate & Miyagi border, Mt. Kurikoma. Yamanashi Prefecture: between Fujiyoshida and Lake Shoji. Gumma Prefecture: Mt. Akagisan; Aze. HoxkKatpo Isuanp. Ishikari Subprefecture: Sapporo; Maru-yama. Abashiri Subprefecture : Kitami Fuji. Oshima Sub- prefecture: Hakodate. Hidacha Subprefecture: Erimo Cape. Iburi Sub- prefecture: Noboribetsu; Muroran. Shiribeshi Subprefecture: Moribetsu. Soya Subprefecture: Soya Cape (Cape Romanzoff). KyusHu ISLaANnp. Kago- shima Prefecture: Yaku-shima. Fukuoka Prefecture: Hikosan. SHIKOKU Istanp. Kochi Prefecture. Formosa. Arisan. Korea. Dagelet and Quel- paert islands. Discussion oF Hydrangea anomala Throughout its wide geographical range, Hydrangea anomala varies but little. The most obvious variation is in the number of stamens; the Himalayan and Chinese plants have 9 to 15, and the Japanese 15 to 20. The Japanese plants have been considered to be a separate species from the Himalayan and Chinese plants. However, the differences between the two groups do not seem to warrant specific status for them and they are here treated as subspecies. Don stated in the Prodromus that he based Hydrangea anomala on a collection made by Wallich in Nepal. Since there was no such specimen in the Lambert Herbarium in the British Museum it was assumed that it must be in the Wallich Herbarium at Kew. In the Wallich Herbarium there are three collections of H. anomala written up as follows under Wal- lich’s name for the species in his Numerical List: “439 Hydrangea altissima Wall. in Herb. 1824. 1. Napalia 1821 2. Kumaon, R. B. 3. Sermore, Lieut. Gerard.” Although most of the material in the Wallich Herbarium was brought by Wallich to England on his return from India in 1828 some had been sent ahead previous to this datet. Material under No. 439 was received 4 Wallich so stated this in his explanation of the purpose of his Numerical List; see Kew Bulletin for 1913, p. 256. Vout. XXIX] McCLINTOCK: MONOGRAPH OF GENUS HYDRANGEA 201 in England in 1824 and was available to Don before work on the Prodromus was completed. A significant point in Don’s description of Hydrangea anomala is the statement that the flowers are small and uniform and that there are no sterile flowers. It may have been for this reason that Hooker and Thom- son in 1857° placed H. anomala in synonymy under Adamia cyanea, a species considered today to belong in the genus Dichroa. Don’s description of H. anomala further states that it has two short, thick, recurved styles. While it is true that Dichroa has no sterile flowers, it has more than two styles, and these are not short, thick, or recurved. Of the three specimens of ‘Hydrangea altissima” in the Wallich Herbari- um under No. 439, only one, No. 439/1, was collected by Wallich. This speci- men consists of a branch bearing two young inflorescences on which sterile flowers are not evident, and a detached portion of an inflorescence which is in anthesis and has sterile flowers. The label reads: ‘‘Sheopore. Montes Chandaghery. May 1821.” Since the specimen was collected by Wallich in Nepal, and consists of a branch without evident sterile flowers, it presumably was the specimen on which Don based his H. anomala. In the material recognized in this paper as H. anomala there were several specimens which lacked sterile flowers. The detached inflorescence men- tioned above may not have been part of the specimen which Don saw, but may have been added later when the specimen was mounted. Wallich published Hydrangea altissima in the Tentamen the year following the publication of Don’s Prodromus. H. altissima was illustrated on plate 50, but no description accompanies this plate and there is no indication of the specimen on which Wallich based his name. The plate of H. altissima appears