7 wai) +N ay ane i : ay is he V 7 AK ; if i ‘2 fone WA, whe me ica | mY ‘y if i ji a) | in nek Ba ae : | te irs PROCEEDINGS OF THE California Academy of Sciences FOURTH SERIES Vol. XXXI SAN FRANCISCO PUBLISHED BY THE ACADEMY 1960-1965 COMMITTEE ON PUBLICATION Dr. GEorGE E. Linpsay, Chairman Dr. Epwarp L. KEssE, Editor Dr. LEo G. HERTLEIN No. No. ; No. a0: CONTENTS OF VOLUME XXXI WitiiaMs, Francis X. The Wasps of the Genus Pleno- culus (Hymenoptera: Sphecidae, Larrinae) (90 figures). Pu blshedeaiulyer Se lOO Oi ae. sae we a MALNATE, EpMoND V., and HARoLp E. MUNSTERMAN. In- terpopulation Variation in the Colubrid Snake WNatrix pryeri from the Riukiu Islands, with Description of a New Subspecies (2 figures). Published July 8, 1960 PuiLip, CORNELIUS B. Further Records of Neotropical Tabanidae (Diptera) mostly from Peru (1 figure). Pub- lnseclaniulyemos S09 OOr .sthet Be ek aE Es be: ee Ae YonceE, C. M. Mantle Cavity, Habits, and Habitat in the Blind Limpet, Lepeta concentrica Middendorff (2 figures). ublshedeseptemberson 1960.2 ee YonceE, C. M. Further Observation on Hipponix antiqua- tus with Notes on North Pacific Pulmonate Limpets (4 figures). Published September 6, 1960 FREUDE, HEINZ. Revision Der Chilenischen Misolampini: Gattungen Heliofugus Guerin und Myrmecodema Gebien ( = Myrmecosoma Germain) (Coleoptera: Tenebrionidae) (17 figures). Published September 6, 1960 _..._ FOLLETT, W. I., AND LILLIAN J. DEMpsSTER. First Records of the Echeneidid Fish Remilegia australis (Bennett) from California, with Meristic Data (1 plate). Published Sep- tember 3.0): mr OGO 82 ar oe eS re Gates, G. E. On Some Earthworms of Eisen’s Collection. Eublished Warchie/e 902s ee ee eee Kevan, D. Kerra McE. Pyrgomorphidae (Orthoptera: Acridoidae) Collected in Africa by E. S. Ross and R. E. Leech, 1957-1958, with Descriptions of New Species (6 iieures)). -seublished: March 7, 1902) 2.2 SACHET, MARIE HELENE. Flora and Vegetation of Clipper- ton Island (12 figures, 1 map). Published March 7, 1962 BANTA, BENJAMIN H., and ALAN E. Leviton. Remarks on the Colubrid Genus Chilomeniscus (Serpentes: Colu- bridae) (10 figures). Published February 28, 1963 Leviton, ALAN E., and STEVEN C. ANDERSON. Third Con- tribution to the Herpetology of Afghanistan (5 figures). Bublishedahebruaty: 20-91903. == ee. ee a PAGES 51-67 69-102 103-110 111-119 121-168 169-184 185-225 227-248 249-307 309-327 No. No. ula: wale : LG: mil Ti 18. El 9: 20: ale iy igh enon 24. Appicott, WARREN QO. Interpretation of the Invertebrate Fauna from the Upper Pleistocene Battery Formation near Crescent City, California. Published May 20, 1963 __ SoveREIGN, H. E. New and Rare Diatoms from Oregon and Washington (2 plates). Published May 20, 1963 Leviton, ALAN E. Remarks on the Zoogeography of Philippine Terrestrial Snakes (1 figure). Published Sep- tember 10} (1963: (=. EE ee ANDERSON, STEVEN C. Amphibians and Reptiles from Iran (15 figures; 14 tables). Published September 10, 1963 __ Gorn, CoLEMAN J., and Doris M. CocHran. Two New Genera of Leptodactylid Frogs from Colombia (2 figures). Published December 920, 1903) === as eee GuRNEY, ASHLEY B., and Davin C. RENTz. A New Spine- Throated Grasshopper from the White Mountains of Cali- fornia (Orthoptera: Acrididae: Cyrtacanthacridinae) (19 figures). (Published june 19) 31904 EE CorréA, Divia Diniz. Nemerteans from California and Oregon. Published’ June@eo: 1904-0 =. eee SANDERSON, Mitton W. Phyllophaga saylori, n. sp., from Nuevo Leon, Mexico (Coleoptera: Scarabaediae) (7 fig- utes) seublished: January 15>) 1965. 2.) ene ee Quast, JAY C. Osteological Characteristics and Affinities of the Hexagrammid Fishes, with a Synopes (3 figures). Published@january #15, 1965 ___.. = a RADFORD, KEITH W., RoBert T. Orr, and Cart L. Husss. Reestablishment of the Northern Elephant Seal (Mi- rounga angustirostris) off Central California (6 figures). Published’ January, 15, 1965 <--Se e Gans, Cart. On Amphisbaena heathi Schmidt and A. carvalhoi, new species, small forms from the Northeast of Brazil (Amphisbaenia: Reptilia) (12 figures). Published January 15, 1965 LOUKASHKIN, ANATOLE S., and NorMAN GRANT. Be- havior and natural reactions of the Northern Anchovy, Engraulis mordax Girard, under the Influence of Light of Different Wave Lengths and Intensities and Total Dark- ness (11 figures). Published January 15, 1965 Index to Volume XXXI Errata 341-347 349-368 369-416 417-498 499-502 503-513 * 515-558 559-562 563-600 601-612 613-630 631-692 693-732 US PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. XXXI, No. 1, pp. 1-49, 90 figs. July 8, 1960 THE WASPS OF THE GENUS PLENOCULUS (HYMENOPTERA: SPHECIDAE, LARRINAE) BY FRANCIS X. WILLIAMS Research Associate California Academy of Sciences My study of these small wasps was begun many years ago and has extended, with interruptions, up to 1959. About 850 specimens of Pleno- culus were assembled for study. IT wish to extend my thanks to the many entomologists and to the insti- tutions with which they may he connected, for the very generous assistance given me. Mr. P. H. Timberlake, Associate Entomologist, Emeritus, Uni- versity of California, Citrus Experiment Station, Department of Biological Control, Riverside, California, has made available the large collection of Plenoculus, which he obtained chiefly in southern California. A great deal of material, also from desert regions and elsewhere, has come from the collee- tions of the California Insect Survey, Division of Entomology and Para- sitology, University of California, at Berkeley, and the University of California at Davis, with Drs. P. D. Hurd and R. M. Bohart and their associates doing the collecting. Dr. George D. Butler, Jr., of the Depart- ment of Entomology, University of Arizona; Dr. Howard E. Evans, of the Department of Entomology, Cornell University; and Dr. R. R. Dreisbach, of Midland, Michigan, have likewise loaned considerable and interesting material. Dr. Herbert F. Schwarz, of the American Museum of Natural History, has donated several specimens of Plenoculus. Finally, Dr. Karl V. [1] 2 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. Krombein, of the Division of Inseet Identification and Parasite Introduc- tion Laboratories, U. S. Department of Agriculture, Washington, D. C., has been unfailingly helpful in the loan of specimens, including the entire material of Plenoculus davist davist Fox from his own collection and also that from the collection of the U. S. National Museum. He has also com- pared specimens, and in his useful correspondence has made helpful sug- gestions and given references to pertinent literature. The excellent drawing of the head of Plenoculus gillaspyi Krombein (fig. 33) is the work of Mr. Arthur D. Cushman, of the U. S. National Museum. Through the courtesy of Dr. Krombein, I was able to study an important collection of Plenoculus and Solierella made by Dr. W. R. M. Mason in the Colorado and Mojave deserts. The author has also made collections of Plenoculus, chiefly in the Colorado desert. The collection of the California Academy of Sciences was also very helpful. The facilities made available by both the California Academy of Sciences, San Francisco, and the San Diego Society of Natural History Museum are here gratefully acknowledged. In the Synoptic Catalog of Hymenoptera of America North of Mexico (Muesebeck, Krombein, Townes, and others, 1951, U.S.D.A. Monograph No. 2), and the first supplement to this Catalog for 1958, prepared under the direction of Dr. Krombein, a total of seven species and one subspecies are listed. In the Old World, two species have been described, one from Transeaspia, U.S.S.R. (as Pavlouskia tadzhika by Gussakovskij), the other from Portugal and Spain, by Andrade. In the present paper ten species and four subspecies of Plenoculus are deseribed as new, while two forms of Plenoculus previously listed as valid are here regarded as mere color forms. This brings the total, as here accepted, to 17 species and 5 subspecies, as follows: 1, Plenoculus sinuwatus,; new species . . 3. =. | Page 7 21 Plenoculus qillaspy: Krombein 2) 9 ssene eee ee 2. Plenoculus hurdi, new species . |. aleceeso eee toe} 4, Plenoculus timberlakei, new species... © . .| . Sy total ®. Plenoculus cuneatus, mew species’ . . . . « % veer 1 ©: , Llenaculus cockerelli Nox ... i | +..5. 4 ae ee Aaele 1. Plenoculus mexcanus, new species . = = 4:05. pene aged | 8. Plenoculus boregensis, new species : “aneO 8a. Plenoculus boregensis perniger, new subspecies . Saul Us. Plenoculus porvus-Kox:. .) 4.55 Anneoneeeee ee 10. Plenoculus palmarum, new species . . . . .. Cae Se Ii. Pienoculus propmquis Fox... 2 sy eee 25 12. Plenoculus deserti, new species . . : . . . | Saakz eo—aN Scutellum UMetancrum y » _—Propodeum Scutum. p ~ 2 4 ees) y ihe: 3 Pronotum. ; \ NOONE Cx Lat. lob a at ~n \ A ¢, Lobe Sx NY ST \or? = \ SS \_¢ f \ ao —— = > \ ~Mesopleuron es 7 7 \ x Metapleuron Proepisternum : Prepecrus(-Epicnemium) Figure 1. Plenoculus propinquus. Female. Length 6.2 mm. From Riverside, Cali- fornia. At A, thoracic sternites 2 and 3 of a male from Nevada. Figure 2. Plenoculus timberlakei. Female. Thorax, from side. From five miles south of Palm Springs, California. 10 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. to the distance between the inner margin of the compound eye and the outer margin of the antennal sockets. Discussion. Obviously, this species is rather closely related to Pl. hurdi, new species. Plenoculus hurdi Williams, new species. (Figure 34). FreMaLe, holotype. Length, 4.75 mm. Head and thorax subshining, abdo- men shining, sculpture very fine. Black; there are no pale markings, the mandibles are nearly uniformly black, the tibial spurs pale, the tarsi nearly black, the venation blackish brown, and the apex of tergites 1-3 very nar- rowly testaceous. The head is wide, chiefly finely reticulate or coriaceous and the preoccipital carina rather narrowly obsolete dorsad. The elypeus is short, and the anterior margin sinuate and is strongly tumid, rounded and polished along the shallow median emargination, and has an inconspicuous tooth over the base of the mandibles. The mandibles are quite slender, somewhat erooked before their middle length, broadly excavate beneath and with no inner tooth; the antennae are moderately stout, the flagellum subfusiform, articles 3 and 4 subequal; the ocelli in somewhat less than a right-angle triangle. The dise of the propodeum rounded, very finely reticulate, its median trough shallow, commencing well beyond the base, not infringed upon by silvery pile, its pleurae coriaceous, the posterior face with some transverse striae and a narrow median eleft expanding toward the dorsum. Tarsal comb bristles shorter than the diameter of the segments from which they arise. Abdomen generally very finely reticulate. The dise of the pyei- dium is entirely black, polished and with moderately fine punctures, it is rather narrow with the sides nearly straight. Vestiture: The moderate sil- very pile has evidently been obscured by the effects of moisture. HloLotypr, female. In unworn condition, Teotihuacan Pyramid, Mexico, July 6, 1951 (P. D. Hurd). Pararypr, 1 female, also in fine condition (ecol- lection, University of California at Davis), Teotihuaean, Mexico, July 21, 1956 (Rand and K. Dreisbach). Discussion. This species is most nearly related to Pl. gillaspyi Krombein described from Round River, Williamson County, Texas. It differs from Pl. gulaspy? in its wider elypeal emargination, and in the fact that the distance between the inner margin of the antennal sockets is much ereater than the distance between the outer margin of the antennal sockets and the inner margin of the compound eye, the ratio being approximately 3.5 to 2.2, or a coefficient of 6.29. Named for Dr. Paul D. Hurd of the Department of Entomology, Uni- VoL. XXXT] WILLIAMS: THE GENUS PLENOCULUS 11 versity of California, Berkeley, who with his fellow entomologists has col- lected so much material in this genus of wasps. Unfortunately, the labrum of the holotype was damaged through my dis- section, but in the paratype, the labrum shows as a dark brownish, almost opaque hair-fringed strip that is closely appressed to or fused with, the elypeus and stretches inconspicuously across its median emareination. Plenoculus timberlakei Williams, new species. (Figures 2, 11, 19-22, 31, 38, 47, 49, 52, 77.) FEMALE, holotype. Length, 6 mm. Shining except dise of propodeum. Black, mandibles reddish beyond base to near apex, antennae from apex of scape onwards, brownish, all tibiae reddish apically with yellow base, tegulae transparent testaceous, bases of principal veins more or less creamy yellow, spines honey yellow, abdomen red. Frons coriaceous, vertex finely pune- tate; elypeus evenly rounded out mesad; mandibles evenly curved, with a notch or step-down beneath near base; antennae short, subclavate, article 3 longer than 4; ocelli in slightly more than a right-angle triangle; preoecipital carina interrupted dorsad; secutum and seutellum with rather fine separate punctures, postscutellum more finely and closely punctured, mesopleurae with some fine obscure wrinkles posteriorly. Dise of propodeum coriaceous, with fanning basal carinulae and a shallow, more or less cross-earinulate median groove, the pleurae generally polished and with some carinulae and punctures, posterior face polished, with a narrow obcuneate groove and a few transverse carinulae. Tibiae and tarsi, except for the more lightly armed fore tibiae, with rather numerous short stout bristles. Tergite 5 with strong well-spaced punctures. Pygidium well maregined, the dise sparsely punctured and somewhat constricted subspatulate apically. Vestiture: rather sparse sil- very pile, the specimen being rather worn. MALE, allotype. Length, 3.8 mm. Head and thorax rather densely clothed with silvery pile; abdomen banded with silvery pile. Black; elypeus except base of produced portion and most of the sides, mandibles except apex, fore femora beneath except at base, all tibiae except in part beneath, pale yellow to brownish yellow; tegulae translucent, base of wings more or less creamy; antennae yellowish brown, excepting the first 6-7 segments above; abdomen reddish. Clypeus rather narrowly produced mesad, apically cuneate, the area at the base of this production is well depressed and gives rise on each side to a short outeurved process upon which the mandibles when folded seem to rest. Mandibles without inner tooth in line with the blade. Antennae stout, fusiform-clavate, the pedicel constricted basally, article 7 lobed apically on outer side above and provided with one longer and several shorter hairs, shorter hairs being present also on 2-6; preoecipital carina interrupted dor- 12 CALIFORNIA ACADEMY OF SCIENCES [Proc. 41TH SER. sad. Dise of propodeum opaque, with fanning basal and some transverse sarinulae, no obvious median eroove, but there is a slightly depressed area at apex of the dise; pleurae opaque, with fine carinulae; posterior face with transverse carinulae and a median incision that widens dorsad. Second sub- marginal cell with a short petiole. Abdomen finely punctate, not tuberculate beneath. Aedeagus as in PI. cockerellu. Ho.otyre and ALLOTYPE (Citrus Experiment Station, Riverside) ; holo- type, 5 miles south of Palm Springs, California, June 28, 1941, on Dalea spinosa; allotype, Beaver Dam, Arizona, June 20, 1939, on Eriogonum tri- chopodum; PARATYPES, 2 females, Beaver Dam, Arizona, June 20, 1939, on Eriogonum trichopodum; 2 males, 6 miles south of Palm Springs, Colorado Desert, California, June 8, 1930, on Hriogonum trichopodum; 2 females, 4 miles south of Palm Springs, June 25, 1941, on Dalea spinosa. Not consid- ered a paratype is a female dissected for study and bearing the label: 6 miles south of Palm Springs, Colorado Desert, California, June 8, 1930. All these specimens were collected by P. H. Timberlake. Not considered paratypes are 2 females, Palm Springs, June 25 and 28, 1941 (E. C. Van Dyke). Discussion. In the female the evenly curved mandibles, the pygidium somewhat spatulate at tip and the lack of pale femoral markines will serve to distinguish Pl. timberlakei from Pl. cockerellii, while the curiously de- veloped elypeus and the somewhat modified antennae will serve that purpose in the male. Plenoculus cuneatus Wiliams, new species. (Figures 10, 17, 18, 24, 25.) FEMALE, holotype. Length, 4.5 mm. Shining, except dise of propodeum. Black; rim of median portion of elypeus, mandibles except at darker base, and apex of scape narrowly beneath, dull reddish brown; tibiae and tarsi in part reddish; antennae dull brownish beneath, the apical segment a little paler; tegulae and basal portion of wings, particularly alone the costa, pro- thoracic lobes apically, fore femora for apical half beneath, a small spot api- cally on intermediate femora, the very tip of hind femora, and all tibiae above, largely creamy yellow; abdomen all red. Sculpture of head and thorax largely obscured by the generous silvery pile. The frons evidently coriaceous Figure 3. Plenoculus cuneatus. Male. To show part of the venation of the fore- wing. From Glendale, Nevada. Figure 4. Plenoculus cockerellii ?. Male. From Tucson, Arizona. Figure 5. Plenoculus stygius. Female, holotype. From Palm Springs, California. VoL. XXXI] WILLIAMS: THE GENUS PLENOCULUS 13 Marginal V. _3°¢ Tr Cubital y. Transverse Median V. 274 Ty Cubital V. Sy 1§* Tr. Cubital V. ae Vg 7: HEE Median V. | Submedian Vv. | — 18¥ Recurrent V. - 270 Recerrent V. Se mecha, | rates oe Se Discoidat y 10 12 : Figure 6. Plenoculus davisi (=P. apicalis). Male, type series. From Northwest- ern Kansas. Figure 7. Plenoculus davisi. Male. From Gavilan, California. Figure 8. Plenoculus davis transversus. Female. Holotype. From Tulare County, California. Figure 9. Plenoculus palmarum. Male. From Palm Springs, California. Figure 10. Plenoculus cuneatus. Holotype. Female. From Imperial County, Calif. Figure 11. Plenoculus timberlakei. Female. From near Palm Springs, California. Figure 12. Plenoculus davisi. Probably a female. From Morongo Valley, Calif. 14 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. or granular. Clypeus with the well-bossed median portion chiefly shining, curving into its cuneate and well-rimmed extension; mandibles almost evenly curved, rather broadly notched beneath; antennae short, subclavate, article 3 longer than 4, article 12 subeonie; ocelli forming a triangle intermediate between an equilateral and a right-angle triangle; preocciptal carina com- plete dorsad. Seutum and seutellum strongly and closely punctate. Dise of propodeum rather closely hedged by dense silvery pile, the median line deli- eate and interrupted, the pleurae reticulate-striate, the posterior face sub- opaque and with a shallow median area. Comb of the basal article of the foretarsus consisting five bristles which are nearly twice as long as the diameter of this basal article. Other leg bristles moderate. Marginal cell of forewings somewhat shorter than usual in the female, and its apex is broadly and obliquely subtruneate and extending apically fully as far as does the third submarginal cell. The third transverse-cubital vein is outbowed and not sinuate. Abdomen broad, the tergites shining, the tergites 1-4 very fine, chiefly hair punctured, the usual large scattered punctures on tergite 5, the 6th tergite (pygidium) widely triangular, slightly constricted apically and provided with large scattered punctures, the bounding carinae low, hardly raised. The apices of tergites narrowly and obscurely pale yellowish, that of tergite 1 rather strongly banded by silvery pile. Mater, allotype. Leneth, 4 mm. somewhat less shining than the female. Markings about as in the female, except that the dise of the elypeus almost entirely, the mandibles basally, the scape beneath and the legs to a greater degree, are creamy yellow. The antennae have the first three articles dark brown above, becoming much lighter for the last ten articles, and beneath articles 2 and 3 are dark, anc all the remainder lighter than above. The vena- tion generally, testaceous and the abdomen all red. The elypeus is not eari- nate, its median portion is modeiately bossed, drawn out mesad, and with a tooth before its rather narrowly truncate apex. The clypeal margin is wide and rather bristly for its lower sides. Mandibles well notched beneath, al- most evenly curved, not toothed within. Antennae stout and subclavate, viewed dorsally article 3 is very little longer than 4 and narrower than the subquadrate pedicel (article 2), the pedicel however, as viewed from the side is subtriangular, but constricted basally; from the side the basal articles trend obliquely, and all the flagellum except articles 12 and 13 are rounded out below with a total crenulate effect. Article 13 is tapering and nearly as long as 11 plus 12. Ocelli in very slightly more than an equilateral triangle. Preoccipital carina complete dorsad. Dise of propodeum subopaque, finely reticulate, a short median carina and an apical depression, the dise hedged by silvery pile. Marginal cell short, the third submarginal cell extending about equally toward apex of wing. Tergites with fine dense punctures, with silvery pile bands on apices of 1-4; tevgites 6 and 7 with quite sparse larger VoL. XXXII] WILLIAMS: THE GENUS PLENOCULUS 15 punctures. Terminalia of the Pl. cockerellu type. No examination was made of the terminalia of the allotype of Pl. cuneatus. Hototyprr, female, and ALLOTYPE, male, from Pinto Wash, Imperial County, California, May 5, 1958 (F. X. Williams), on Euphorbia polycarpa var. hirtella. PARATYPE: 2 females and 3 males, same data as type material. Specimens not considered paratypes: 1 male, Thousand Palms, Riverside County, California, April 23, 1955 (W. R. M. Mason); 1 male, Borego, San Diego County, April 25, 1954 (P. D. Hurd), on Croton californicus, and 1 male in the U. S. National Museum (No. 1119, B 29), from Glendale, Nevada, October 3, 1929, on Chrysothamnus paniculatus, and collected by David E. Fox. Discussion. This distinct species is readily separated from Pl. timber- lakei, its nearest relative, in the female by its cuneate clypeal margin, the narrower ocellar triangle, the generally shorter marginal cell and the pres- ence of femoral and tibial markings; the male of Pl. cuneatus is differen- tiated by the form of the elypeus and the unmodified antennae. Plenoculus cockerellii I'ox. (Figures 4, 14, 15,16, 42, 48, 50, 51, 70, 80.) Plenoculus cockerellii Fox, 1893, Proc. Acad. Nat. Sci. Phila., 45:538. Female. Las Cruces, New Mexico (T. D. A. Cockerell). Fema.e. Plenoculus cockerellii, as represented by a specimen in the U.S. National Museum, and taken in the type locality in 1928, by Cockerell, is about 6 mm. long. It is black with the elypeus and mandibles largely reddish, the antennae brownish with the apex of the last article paler, the first and second femora with pale creamy white apically beneath, the legs are gener- ally darker and lighter reddish, and the abdomen red. The clypeus is evenly rounded out, the antennae rather short and subclavate, article 3 plainly longer than 4; the ocelli form about a right-angle triangle, and the preoc- cipital carina is entire above; the tibial spines are moderate; tergite 5 is without large deep punctations; the pygidium is very little constricted be- fore the apex, the dise of which is evenly flattened. The punctation is fine, or the sculpture largely coriaceous. Mate (hitherto undescribed). Subopaque. It is black with the clypeus and mandibles, except their apex and lower tooth, brownish yellow, the antennae with the scape beneath lemon yellow, articles 3-6 dull yellowish beneath, the last articles being honey yellow; the margin of the prothoracic lobes and the base of the wings are creamy yellow, the tegulae are in part translucent, the rest creamy yellow; most of the fore femora beneath and the apex of all the femora, tibiae, except a stripe beneath, and the tarsi, 16 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. creamy yellow; the abdomen is reddish, obscurely black above and beneath at middle length. The elypeus is rounded out, slightly wedge-like mesad, with a slightly prominent translucent rim (the elypeus and mandibles are somewhat worn down); mandibles slightly elbowed, not dentate within; an- tennae rather slender for the basal part of the flagellum, articles 3 and 4 subsequal from above; ocelli forming very slightly more than a right-angle triangle; vertex finely granulate, preoecipital carina entire dorsad; thoracic notum finely and closely punctate; dise of propodeum finely granulate and with a shallow trough that has some transverse striae, the pleurae in part smooth and in part finely striate, the posterior face shining and with a tri- angular depression above. Vestiture is of silvery pile forming bands on the abdomen. Terminalia with the lobes of the aedeagus, as seen from above, extending cone-like laterally. This deseription of the male is based on somewhat worn specimens from: Four miles east of Eden, Riverside, California (P. H. Timberlake). Males from La Paz, Baja California, Mexico, are much darker. And intermediates occur elsewhere. SPECIMENS STUDIED: ARIZONA—CocuHisk County: 17 miles east of Douglas, 2 females, August 8, 1958 (R. M. Bohart); Willcox, 2 males, 1 female, August 14, 1958 (D. D. Linsdale); Madera Canyon, Santa Rita Mts., 1 female, July 31, 1958 (R. M. Bohart). GranHam County: San Carlos, 1 female, July 15, 1955 (N. J. Nerney), swept/] alfalfa. Maricopa County: Wickenbureg, 1 female, August 25, 1927 (Cornell Univ., Lot 542, sub. 334). Pima County: Sabino Canyon, 1 female, July 17, 1954 (R. M. Bohart). Tueson, | ntalles May 30, 1920 (F. X. Williams, doubtfully this species); 10 females and 1 male, July 17, 1955 (G. D. Butler), Wislezenia. Pinan County: Eloy, 10 miles south, 1 female, July 3, 1953 (T. R. Haig). CALI- FORNIA—RiversipE County: Cathedral City, 1 female, October 8, 1946 (P. H. Timberlake), on Euphorbia polycarpa. Four miles east of Eden, 2 males, April 17, 1987 (P. H. Timberlake), on Hriogonum thomas. Palm Springs, 3 females, June 24, 29, 1954 (P. H. Timberlake), on Dalea spinosa. Twenty-nine Palms, 1 female, August 9, 1946 (P. H. Timberlake), on Wis- lezenia refracta; one female (earrying a pyralid caterpillar), September 5, 1946 (P. H. Timberlake). Thousand Palms, 2 males, April 16 and 23, 1955 (W. R. M. eee San Dieco County: Borego Desert, 1 female, April 27, 1954 (P. D. Hurd); 1 female, April 27, 1954 (M. Wasbauer), on Croton ene 1 female, May 26, 1954 (F. X. Williams), 1 female, April 27, 1955 (R. O. Schuster). NEW MEXICO—Dona Ana nee Pee Cruces: 3 females, August, 1928 (Cockerell) (Ckll. 4870 and CkIl. , Collee- tion U. S. National Museum). Albuquerque (desert), 5 males, i 5 1956 (R. and K. Dreisbach). Ilipanco Country: Rodeo, 2 females, August 21, 26, 1958 (R. M. Bohart); 18 miles north of Rodeo, 7 females, August 19, 25, VoL. XXXI] WILLIAMS: THE GENUS PLENOCULUS ibe and 26, 1958 (R. M. Bohart). TE XAS—E. Paso County: El Paso, 1 fe- male, July 24, 1914 (J. C. Bradley), Cornell U. Lot 684, sub. 35 (labeled, Plenoculus cockerell Fox 2, J. C. Bradley, 1923. Comp. with type). Luano County: Not far from Austin, 1 female, June 12, 1941 (J. E. Gillaspy) Col. Univ. Calif., Berkeley). MEXICO—Basa Catirornia: La Paz, 1 fe- male, October 12, 1954 (F. X. Williams); 33 females and 4 males, October 8, 1955 (F. X. Williams). GuERRERO: Mexeala, 1 female, June 29, 1951 tee Eurd)). Discussion. Easily separated from Pl. timberlakei by its elbowed man- dibles, this character being more obvious in the female, and by the unmodi- fied clypeus and antennae in the male. However, Pl. cockerellii shows con- siderable variation over its wide range. A female from Las Cruces, New Mexico, has a good deal of reddish about the mandibles. elypeus and femora; those from La Paz, Baja California, are considerably darker, with some of the California examples intermediate between the two. The pale femoral markings seem always present. The female from Llano County, Texas, and several from Arizona (Cochise, Graham, and Pima counties) have the ely- peus drawn out wedge shaped (figs. 15, 16). Males may be quite dark, and in material from the same locality and collected on the same date one speci- men has the clypeus quite black while another may have the clypeus of an old ivory color. Sometimes the clypeus shows a weak median ridge, as in the five males from Albuquerque, while a male from Tueson has the mandibles narrowly cleft within. Plenoculus mexicanus Williams, new species. (Figures 13, 23.) Frema.e, holotype. Length, 5.5 mm. Black; vertex and dorsulum shining, dise of propodeum subopaque. Clypeus narrowly reddish margined, man- dibles red, darker apically, seape black and obseure reddish, pronotal lobes dull reddish, the first and second pair of femora with a creamy yellow stripe beneath, tibiae and tarsi mainly reddish brown; abdomen red, with much blackish apically up to the second tergite; pygidium dark red. Head wider than high; elypeus with the median portion rather gibbous and drawn out subeuneate, with scattered punctures except rather narrowly mesad, thus giving it a subearinate aspect; mandibles very slender, distinctly elbowed at the basal third, feebly and widely excavate beneath, the bordering noteh low, no inner tooth, the blade being quite low and forming in profile only a gentle convexity; antennae stout, article 3 longer than 4; ocelli arranged in about a right-angle triangle; preoceipital carina complete dorsad but not reaching the gular suture below. Dorsulum very closely punctate; dise of propodeum finely and transversely aciculate, narrowly depressed apically, « 18 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. posterior face with a narrow wedge-like depression; marginal and submar- ginal cells extending equally apieally. Fifth tergite with the usual large punctures; pygidium slightly compressed subapically, with a few strong punctures. Pile silvery. Matz, allotype. Length, 4 mm. Head, thorax, except the largely creamy yellow pronotal lobes, and the abdomen, except a little reddish laterally at the base, black. Clypeus red apically, mandibles creamy yellow basad, red- dish apically; seape beneath and the last five or six articles increasingly brownish; tegulae and axillary selerites pale yellowish brown; first and sec- ond pairs of femora beneath from apex, all tibiae widely above, and the tarsi, ereamy yellow. Clypeus rounded subecuneate mesad, with a fine low, more or less divided carina; mandibles slender and elbowed, though less so than in the female, well excavated beneath; antennae fusiform, slender at the base, articles 3 and 4 subsequal, article 13 tapering; ocelli forming less than a right-angle triangle. Dorsulum and dise of the propodeum nearly opaque. Terminalia of the Pl. cockerellu type. Vestiture of silvery pile, forming bands on the abdomen. Hotoryper, female, and ALLOTYPE male, from Acapulco, State of Guerrero, Mexico, July 1, 1951 (H. E. Evans). Paratypss, all from Mexico, as follows: 5 females from Acapulco, date as above; 1 female, Tecolutla, State of Vera- eruz, June 19, 1951 (H. E. Evans); 4 females and 1 male from Veracruz, June 20, 1951 (H. KE. Evans); 1 male and 16 females, Acapuleo, July 1, 1951 (P. D. Hurd); 16 females, Veracruz, June 20, 1951 (P. D. Hurd); 3 females, Tecolutla, June 19, 1951 (P. D. Hurd), and 1 male, Alpuyeea, Morelos, Mexico, July 3, 1951 (P. D. Hurd). Discussion. Plenoculus mexicanus differs consistently from Pl. cockerellu in its more slender and more crooked mandibles, particularly in the female. In the male of Pl. mexicanus, of which sex only four specimens have been collected, the abdomen may be almost entirely black. Figure 13. Plenoculus mexicanus. Female. Holotype. Jaws and clypeus. From Acapulco, Mexico. Figure 14. Plenoculus cockerellii. Female. From La Paz, Baja California, Mexico. At A, clypeus of male. Same locality. Figure 15. Plenoculus cockerellii. Female. From Llano County, Texas. Figure 16. Plenoculus cockerellii. Female. From Tueson, Arizona. Figure 17. Plenoculus cuneatus. Female. Holotype. From Imperial County, Calif. Figure 18. Plenoculus cuneatus. Female. Holotype. Ocellar triangle. Figure 19. Plenoculus timberlakei. Female. Holotype. Five miles south of Palm Springs, Riverside County, California. Figure 20. Plenoculus timberlakei. Male. Beaver Dam, Arizona. Figure 21. Plenoculus timberlakei. Male. Antenna from side. VoL. XXXII] WILLIAMS: THE GENUS PLENOCULUS 19 pe Figure 22. Plenoculus timberlakei. Female. From near Palm Springs. Figure 23. Plenoculus mexicanus. Male. Allotype. From Acapulco, Mexico. Figure 24. Plenoculus cuneatus. Male. Clypeus. From Glendale, Nevada. Figure 25. Plenoculus cuneatus. Male. From Borego, San Diego County, Calif. 20 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Plenoculus boregensis Williams, new species. (Figures 26, 27, 28, 29.) FEMALE, holotype. Length, 6 mm. Black; elypeus diluted white, its mar- ein narrowly, except wedge-like mesad, reddish; mandibles reddish, the apices darker; scape of antennae pale yellow beneath; above, the flagellum is reddish brown to dusky brown; base of pronotum, outer portion of its lobes, tegulae, largely so; axillary selerites and basal wing veins, creamy white; fore femora postero-ventrad and mid femora ventrad, creamy white from apex to near base; posterior trochanters and femora reddish, all tibiae red- dish with yellowish white dorsad; tarsi reddish brown; abdomen reddish, shining, the apical margins of tergites 1-4 whitish, that on tergite 1 the most distinet. Vestiture: dense appressed silvery pile on head and thorax, conspicuous also on femora, and in certain lights it shows as three stripes on the mesonotum, and as narrowing bands at the apex of tergites 1-4. Cly- peus gently rounded outwardly, this lobe very slightly roughened and mod- erately gibbous; mandibles very slender, stepped down at basal third beneath rather than emarginate, and with no inner tooth; antennae subclavate, the flagellum rather fusiform, article 3 distinctly longer than 4, 12 about half again as long as thick and tapering rather sharply (collapsed ?); ocelli in slightly more than a right-angle triangle. Exposed dise of propodeum much narrowed by the invading pile, opaque, and with a line-lke groove that widens and flattens apically. Foretarsal rake particularly well developed for Plenoculus, the pale bristles extending to nearly twice the width of their respective segments; mid-tarsi with a rather dense row of long bristles; the bristles on the posterior tarsi moderate. Wing venation mostly pale testa- ceous, the long third submarginal cell extending apically fully as far as the marginal cell. Abdomen relatively broad; pygidium broadly triangular, the sides well carinate, the apex narrow, and the dise with a few large punctures. Mats, allotype. Length, 4 mm. Black; elypeus, mandibles at base, scape beneath, large spot on forecoxae, femora 1 and 2 beneath and at apex, apex of femora 3, tibiae and tarsi, base of wings to well along the costal margin, creamy white; scape of antennae above and pedicel in part and base of article 3 nearly black; otherwise the flagellum is orange yellow; last tergite reddish. Clypeus gently rounded outward, no hair tuft on either side; man- dibles slender, well excavate beneath, but with no inner tooth; antennae slender, subclavate, the pedicel thicker than, and fully as long as, the third antennal article which is about as wide as long and very little shorter than the fourth; the clavate portion of the antennae with no article, except the thirteenth, longer than wide, article 13 one and one-half times longer than 12 and tapering on one side from half its length; ocelli forming less than a right-angle triangle. Pronotum distir etly depressed mesad. Dise of propo- VoL. XXXTI] WILLIAMS: THE GENUS PLENOCULUS 21 deum narrowly exposed, depressed mesad, and with some transverse regulae; the pleurae tessellate, shining; the posterior face with a subfusiform depres- sion. Marginal cell rather narrowly truneate, slightly exceeding the third submarginal. Abdomen smooth ventrad. Terminalia: uneal lobes sharply pointed dorsad (as facing the last visible ventral segment) ; a row of spines, short, heavy and spike-like, on each volsella. Vestiture: abundant silvery pile, as in the female. Hototyre, female, Borego, San Diego County, California, April 30, 1954 (M. Wasbauer), on Croton californicus. ALLOTYPE, male, topotypical, April 25, 1954 (P. H. Timberlake), on Croton californicus. PARATYPE, 1 male, topotypical, April 25, 1954 (M. Wasbauer), on Croton californicus. All three specimens are in fine condition. Discussion. Plenoculus boregensis belongs nearest the Pl. cockerellu group, although the terminalia of the male of Pl. boregensis are quite differ- ent. I consider this species our finest example of Plenoculus. The yellow teg- umentary band on the pronotum of the female seems distinctive among wasps of this genus. Plenoculus boregensis perniger Williams, new subspecies. Mats, holotype. About 3 mm. long. Head, thorax, and abdomen shining black. Mandibles blackish at base, thenee reddish; antennae black, femora black, tibiae with some creamy white, all tarsi, except apex of last article, pale; venation of forewings generally dark testaceous, paler at base; the very tip of abdomen slightly reddish. Clypeus rounded out mesad, and except for its thick shining rim it is roughened. Mandibles not slender, slightly crooked, with a very slight notch within. Antennae thick, subclavate, article 3 as long as the pedicel and narrowed basad, article 4 very slightly shorter than 3, article 13 tapering and a little longer than 114 the length of 12. Ocelli forming a little less than a right-angle triangle. Vertex coria- ceous; dorsulum finely and closely punctate; dise of propodeum widely ex- posed, finely granulate. Marginal cell of forewings rather narrowly truncate and exceeding the third submarginal cell. Some patches of silvery pile. Ter- minalia as in Pl. boregensis boregensis; the parameres with the usual fring- ing bristles, a row of stronger erect bristles from the concave surface, and a row of shorter volsellar bristles, lobes of aedeagus acuminate on their ventral side (facing the last visible ventral segment). Hotoryeg, male, Thousand Palms, Riverside County, California, April 7, 1955 (W. R. M. Mason). Pararypes, 4 males. Two other males not con- sidered paratypes, topotypical, early to late April, 1955 (W. R. M. Mason). FEMALE, unknown. 22 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Discussion. This species differs from the nominate species chiefly in its black elypeus and abdomen. The largest of the series is 3.40 mm. long. Plenoculus parvus Fox. Plenoculus parvus Fox, 1897, Ent. News, 8:71—-72. Type, female, Las Cruces, New Mexico (Cockerell, 5173). FEMALE. The original description is as follows: Anterior margin of clypeus subtruncate in the middle, not incised or dentate; the flagellum strongly clavate; the first joint longer than the second, dorsum and scutellum finely and closely punctured; middle segments microscopically striated, not impressed above, posterior face more distinctly transversely striated; with a deep longitudinal central furrow; legs tolerably spinose, tarsal comb feebly developed; pygidial area with large sparse punctures, not margined or carinated laterally. Black; mandibles except apex, legs except coxae and extreme tip of abdomen, red; scape beneath and tegulae yellowish; flagellum beneath testaceous; entire insect clothed more or Jess with silvery pubescence, especially the head in front and the thorax on sides and beneath; apical margins of abdominal segments narrowly testa- ceous; wings hyaline, strongly iridescent, nervures testaceous. Length 3 mm. This is the smallest species of Plenoculus, and is not closely related to any of the others. Discussion. I am obliged to Dr. Karl V. Krombein for his additional findings from an examination of this unique female type, principally that there are two teeth at each lateral angle of the elypeus and that the pygi- dium is delimited, though rather weakly, by a carina. Plenoculus palmarum Williams, new species. (Higures' 9536, 37, 715 73; 78>) Mate, holotype. Leneth, 3.8 mm. Black; antennae brownish yellow, duskier toward base; forepart of elypeus reddish brown; mandibles, except teeth and apex, yellow; fore and mid-femora at apex beneath, all tarsi and Figure 26. Plenoculus boregensis. Female. Holotype. At A, mandible, from outer side. Borego. Figure 27. Plenoculus boregensis. Female. Mid tarsus. Borego. Figure 28. Plenoculus boregensis. Male. Clypeal outline. Borego. Figure 29. Plenoculus boregensis. A., pygidium of female; B., male, extremity of aedeagal lobes, ventral and lateral views. Borego. Figure 30. Plenoculus sinuatus. Male. Allotype. Mandible somewhat inclined from side. Borego. Figure 31. Plenoculus timberlakei. Male. Allotype. Mandible. To show low tooth, T, behind blade, B. Beaver Dam, Arizona. VoL. XXXII] WILLIAMS: THE GENUS PLENOCULUS bo a) Figure 32. Plenoculus sinuatus. Male. Allotype. Terminalia in part. From Borego. Figure 33. Plenoculus gillaspyi. Female. Type. From Williamson County, Texas. Drawn by Arthur D. Cushman. Figure 34. Plenoculus hurdi. Female. Holotype. From Teotihuacan, Mexico. Figure 35. Plenoculus sinuatus. Female. Holotype. From Borego. 24 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. tibiae, pale yellowish to yellowish brown; tegulae and basal venation nearly white; first three abdominal segments orange. Produced portion of elypeus subtruneate, no elypeal side whiskers; mandibles with a fang-like tooth within; antennae stout, the segments short, the thirteenth tapering; ocelli in slightly less than a right-angle triangle. Dorsulum shining, finely pune- tate; metapleurae above with some well-marked carinulae extending to near spiracle. Exposed dise of propodeum narrow, tongue-like, there being an imperfect bounding carina hedged in by silvery pile; there are also a few transverse and an imperfect longitudinal carina; posterior face with some fine carinulae and a narrow wedge-like cleft expanding to apex of the disc. Marginal cell very short. Abdomen smooth beneath. Lateral lobes of the aedeagus somewhat cuneate beneath, and apically above are provided with a tuft of bristles. The pile is silvery and abundant. FEMALE, allotype. Length, 4 mm. Body shining. Black; clypeus for apical portion, mandibles except apex, scape of antennae beneath at base, and flagellum beneath, fore and middle femora beneath beyond base, creamy white; posterior femora reddish brown apieally; tibiae and tarsi pale yellow- ish brown, but tibiae in part whitish dorsad; pronotal lobes apically, tegulae, axillary sclerites, and base of wings, creamy white; venation testaceous, abdomen entirely red; legs with spines on the palest areas whitish, otherwise spines and spurs pale brown. Head rather wide; produced portion of ely- peus gently lobed, narrowly rimmed and with three teeth on each side, the innermost tooth quite small; mandibles with a fang-like tooth within at base of blade; antennae moderately stout, article 3 slightly longer than 4, article 12 gently tapering; ocelli in about a right-angle triangle. Dorsulum with close punctures. Exposed dise of propodeum narrowing from base, and with two longitudinal carinulae and some shorter basal ones; pleurae finely reti- culate; posterior face with its depression in its upper portion. Marginal cell rather short, exceeding the third submarginal. Dise of pygidium broadly triangular, tapering slightly at apex, its bounding ecarinae not strong, the dise with a few punctures. Vestiture silvery pile in the usual places. Ho.otyrr, male (Citrus Experiment Station, Riverside), Palm Springs, Riverside County, California, April 24, 1988 (P. H. Timberlake), flying over ground. ALLoTYPE, female, Borego, San Diego County, April 29, 1955 (F. X. Williams). Paratypes, 2 males, Borego, April 29, 1954 (P. H. Tim- berlake), on Huphorbia polycarpa; 1 female. Borego, April 27, 1954 (P. D. Hurd). Other specimens not considered paratypes: 1 female, Blythe, River- side County, April 24, 1955 (W. R. M. Mason); 3 males, Pinto Wash, Impe- rial County, California, April 28, 1958, and 12 males and 4 females, topo- typical, May 5, 1958 (F. X. Williams), all at plants of Huphorbia polycarpa hirtella. VoL. XXXI] WILLIAMS: THE GENUS PLENOCULUS 25 Discussion. This small, stoutly-built species has the labrum gently bi- lobed in both sexes. It is evidently related to Pl. parvus Fox, of which only the female is known. The abdomen of Pl. palmarwm is entirely red, and there are three teeth on either side of the low elypeal lobe, while in Pl. parvus there are but two such on either side. Plenoculus propinquus I'ox. ChicunesMes4 Gah 5s 56, ole 2s) Plenoculus propinquus Fox, 1893, Proc. Acad. Nat. Sci. Phila., 45:537-538. Female, Colorado. Ashmead, 1899, Psyche, 8:337-339. Male and key to spp. Plenoculus propinquus var. rufescens COCKERELL, 1898, Proc. Davenport Acad. Sci., 7:144. Reported from Arizona, California, Colorado, New Mexico, Idaho, and now from Nevada, Oregon, Washington, and Utah. SPECIMENS EXAMINED: CALIFORNIA—Kern County: Randsburg, 1 male, May 1, 1952 (R. M. Bohart). Pumpkin Center, 1 male, July 30, 1956 (E. I. Schlinger). LAssEN County: Halleluja Junction, 1 male, July 4, 1951 (R. C. Bechtel); 3 females and 1 male, July 11, 1957 (R. M. Bohart). Mon- TEREY County: San Lucas, 1 male, August 20, 1933, on flowers of Hriogo- num gracile (P. H. Timberlake). Los ANGELES County: Clairmont, 3 fe- males ‘“PomC” (Baker). Glendale, 1 male, June 8, 1952 (E. I. Schlinger) ; 2 miles east of Lancaster, September 14, 1956 (KE. I. Schlinger). ORANGE County: Yorba Linda, 1 female, August 15, 1920, on flowers of anise (P. H. Timberlake). RrversiIpE County: Banning, 1 male, June 26, 1952 (J. W. MacSwain). Four miles east of Eden, 1 male, April 17, 1937, on Larrea diva- ricata (P. H. Timberlake) ; Riverside, 8 females and 3 males, distributed in April, June, July, September, and October, and the years 1927, 1928, and 1929, on Euphorbia marginata; 4 males and 3 females, comprising 6 speci- mens collected in June and 1 in September, during the years 1926, 1927, 1932, and 1941, on Eriogonum gracile; 1 female and 1 male, September, 1924, on annual Eriogonum; 1 female digging in loose soil. All collected by P. H. Timberlake; 3 females, August 7 and 10, 1956 (J. C. Hall and E. I. Schlinger). San BerNARDINO County: One mile east of Cajon Junction, 8 males, August, 1956 (E. I. Schlinger). VenrurA County: Santa Paula, 3 males, June 5, 1927, on or near ground (P. H. Timberlake). COLORADO— ALAMosa County: Great Sand Dunes, 6 females, July 20-21, 1954 (TH. E. and M. A. Evans). One female, No. 2276 Colorado; Boulder, 1 female, August 5, 1908; 1 male, August 4, 1908 (S. A. Rohwer, Det. S. A. Rowher), Collections U. S. National Museum. Limen, 1 female, August 25, 1951 (R. R. Dreisbach). NEVADA—CuurcuHILL County: Fallon, altitude 4,000 feet, 4 males, May 29, 1930 (EH. L. Bell), Acc. 30540, American Museum of Natural History, Minden, 2 females, August 24, 1952 (R. M. Bohart). WASHOE County: Sparks, 1 female, July 18, 1953. OREGON—DEscuHutTEs County : 26 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Tumalo Reservation, near Bend, 2 males, June 22-23, 1954 (K. I. Sehlin- over). UTAH—North fork Duchesne River, 1 female, July 13-14, 1927 (Cor- nell University Lot 542, sub. 305). Morcan County: Pine View, Ogden Can- yon, 1 female, July 21, 1922 (E. P. Van Duzee). WASHINGTON—Grant County: Columbia River, near Vantage, 1 male, August 27, 1954 (H. HE. and M. A. Evans). Discussion. The female of Pl. propinquus averages about 6 mm. long. It is black or with the basal part of the abdomen reddish, and the anterior tibiae in front yellowish brown. The elypeus is deeply cleft mesad, almost with a stoved-in effect. The dise of the propodeum is finely granulate, with a few earinulae and a shallow median suleus. The male has a rather gibbous elypeus, often shallowly cleft mesad or bluntly produced there, and its color is more or less pale yellowish or old ivory and sometimes sharply dark at the extreme base. With rare exceptions the seape is dark beneath. The 2-3 stout volsellar thorns appear to be its best character. Plenoculus deserti Williams, new species. (Figures 85, 87.) Mate, holotype. Leneth, 4 mm. Black; dise of clypeus anteriorly vellow- ish red; mandibles same color except base and apex; no pale ventral stripe on seape; a stripe on fore tibiae, on fore tarsi generally, a rather obscure stripe behind on the posterior tibiae, and an obscure basal and apical mark on middle tibiae, all pale dull yellowish; middle and posterior tarsi brown- ish, tegulae yellowish; venation testaceous, the costa and other veins forming the marginal cell, darker; abdomen orange red. Produced portion of elypeus with acute lateral angles, the low wide median lobe polished and somewhat downcurved apieally; a pale elypeal tuft far to either side; mandibles exea- vate beneath and with a strong tooth within; antennae rather slender, articles 3 and 4 subsequal, 13 curved on one side; frons and vertex sub- opaque a line forward from anterior ocellus, the ocelli forming barely less than a right-anele triangle, the space between the fore ocellus and each pos- terior one tumid. Dorsulum closely punctate, the meso- and metapleurae shining, the mesopleurae beneath with a small tubercle; dise of propodeum subopaque, broad and subtruneate, bounded apically by a curved carina, the dise is reticulate and with fanning and transverse carinulae, the median trough widens apically and is polished there, propodeal pleurae shining and very finely striate, posterior face shining, transversely striate and with a broad subeordate depression. Abdomen with very fine reticulations. Aedea- ous of the Pl. davisi type, but with about 8 strong volsellar spines. Vestiture moderate silvery pile. VoL. XXXII] WILLIAMS: THE GENUS PLENOCULUS bo -~] FEMALE, allotype. Length, 5 mm. Head and thorax generally subshining, the punctation finer than in the male. Black, the elypeus reddish except at base, only the posterior tibiae with a pale dirty yellow stripe posteriorly, tarsi largely brownish; venation as in holotype; abdomen reddish with a darker ill-defined median blotch on tergite 2. Produced portion of elypeus subtruneate, mostly bare and shining with a very few large punctures, strongly emarginate and rather depressed mesad, the margin sloping down from this emargination and provided with an angle and two teeth at each side; mandibles stout, emarginate beneath and toothed within; antennae rather slender, segments 3 and 4 subequal, the shghtly reddish segment 12 slightly longer than segment 11; ocelli forming very little less than a right- angle triangle, the line forward from the anterior ocellus about as long as the diameter of that ocellus, the space between the fore ocellus and each posterior ocellus rather tumid; head texture subcoriaceous. Length of spines of the fore tarsal comb up to about 1.3 the thickness of the segment from which they originate. Dise of propodeum generally opaque except the apical part of the very shallow median trough, and with very fine recurved striae; propodeal pleurae shining with exceedingly fine striate-reticulate surface; posterior face shining, with a V-shaped depression. Abdomen with tergites showing very fine reticulations and the usual strong punctures on the apical ones. Dise of pygidium subtriangular, not pinched apically, the sides being nearly straight, the dises with rather sparse though strong punctures having the effect of roughening its surtace. Houoryrer, male, in good condition, Borego, San Diego County, April 20, 1955 (F. X. Williams). ALLoTypPE, female, topotypical, April 2, 1953 (P. D. Hurd). Paratypes likewise from Borego are, 2 males, April 25, 1954 (P. D. Hurd), on Croton californicus; 1 female, April 9, 1955 (EF. X. Williams) ; 3 females, March 15, 1957 (F’. X. Williams). Specimens not regarded as para- types are, 1 male (with elypeus and mandibles black), Picacho Pass, Ari- zona, September 13, 1954 (P. H. Timberlake), on Huphorbia albomarginata; 1 male and 1 female, Thousand Palms, Riverside County, Mareh 28 and April 25, 1955 (W. R. M. Mason); 1 male, Fish Creek mountains, 300 feet, Imperial County, California, April 20, 1955 (W. R. M. Mason). Discussion. Plenoculus deserti seems to be the Colorado Desert represen- tative of Pl. propinquus Fox. It differs from the latter in its generally smaller size; in the male in the slightly different elypeus and in the more numerous and more spinelike volsellar armature; in the female in the more roughened pygidial dise. And admittedly, this rather weak species may be difficult to separate from some specimens of the desert Pl. davisi Fox. 28 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Plenoculus boharti Williams, new species. (Figures 40, 61, 83, 84.) FEMALE, holotype. Length, 4.5 mm. Frons subopaque, thoracic dorsum somewhat shining. Black; mandibles widely orange mesad, legs nearly black, fore femora at extreme base in part reddish, hind tibiae suffused with orange red, tarsi obscurely brownish, tegulae testaceous apically; abdomen red with a large central black spot on tergite 1, a still larger one on tergite 2, with tergite 3 almost entirely black, and the remaining tergites blackish. Frons densely and minutely punctate, less densely so on vertex which is shining; elypeus with the dise gently convex, the anterior part shining and with some large deep punctures, the margin shghtly areuate and with four strong teeth on each side of the rather narrow median emargination; antennae sub- clavate, rather stout, article 3 appearing slightly longer than 4; ocelli form- ing very slightly more than a right-angle triangle. Dorsulum very finely punctate. Dise of propodeum short, almost coriaceous, shghtly depressed and very minutely eross-wrinkled for its apical half, the pleurae with very fine longitudinal striae, shining; posterior face chiefly smooth, with a shal- low subtriangular depression from the upper part of which a pair of divere- ing earinulae arise. Legs moderately spinose; venation rather heavy, the marginal cell ending very slightly beyond the third submarginal eell, the second transverse-cubital and the second recurrent vein interstitial in one wing, nearly so in the other wing. Abdomen broad, the tergites generally reticulate, the fifth however with the usual large punctures; pygidium of the Figure 36. Plenoculus palmarum. Female. Paratype. From Eorego. Figure 37. Plenoculus palmarum. Male. From Palm Springs. Figure 38. Plenoculus timberlakei. Female. Head. to show dorsally interrupted preoccipital ridge. Figure 39. Plenoculus davisi. Male, with red abdomen. Ocellar triangle. From Plumas County, California. Figure 40. Plenoculus boharti. Female. Holotype. From Campo, California (U.S. N.M.). Figure 41. Plenoculus davisi. Female. From Midland County, Michigan. Figure 42. Plenoculus cockerellii. Female. From Douglas, Arizona. Figure 43. Plenoculus davisi, male, with red abdomen. Abdomen, from side, to show ventral ridges or undulations. From Big Bear Valley, San Bernardino Moun- tains, California. Figure 44. Plenoculus davisi. Male (of Fig. 43). Abdomen, from beneath, to show ridges. Figure 45. Plenoculus davisi. Female. Pygidium. From Olmsted County, Minn. Figure 46. Plenoculus propinquus. Female. To show bilobed labrum. From Riverside, California. Figure 47. Plenoculus timberlakei. Female. To show labrum very slightly emar- ginate mesad. VoL. XXXT] WILLIAMS: THE GENUS PLENOCULUS 29 Figure 48. Plenoculus cockerellii (?) male. From Tucson, Arizona. Figure 49. Plenoculus timberlakei. Female. Pygidium. From Beaver Dam, Ariz. Figure 50. Plenoculus cockerellii. Female. Pygidium. From Douglas, Arizona. Figure 51. Plenoculus cockerellii. Female. From Eloy, Arizona. Figure 52. Plenoculus timberlakei. Female. To show ocellar-occipital ratio. Figure 53. Plenoculus davisi. Female, with red abdomen. To show ocellar- occipital ratio. From Mill Creek, San Bernardino County, California. Figure 54. Plenoculus davisi gracilis. Female, holotype. The pale clypeus has unusual dentition. Six miles west of Indio, California. 30 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. usual triangular form, not constricted subapically, shining, with rather Sparse strong punctures except mesad. Mats, allotype. Leneth, 5 mm. Black; mandibles distad of broad part, reddish; extreme base of fore tibiae and a stripe posteriorly on the hind tibiae and their spurs, reddish brown; tarsi dark brown; abdomen red, with a transverse blackish spot on tergites 2 and 4, beyond it is blackish; a pair of small dark spots on sternite 2. Head with exceedingly fine close punctures, more scattered behind the posterior ocelli where it is shining; elypeus with the lateral hair tufts pale brown, clypeal dise moderately convex, the anterior part roughly sculptured though shining, the produced margin only gently arcuate and armed with four strong teeth; antennae rather stout, with the seape all dark, articles 3 and 4 subequal, 13 not bent; ocelli in slightly less than a right-angle triangle; a shining, slim fusiform line from the fore ocellus to between the bases of the antennae. Dorsulum shining, very finely and closely punctate, the pleurae striato-punctate and with a polished bossed area. Dise of propodeum widely exposed, shining, longitudinally and divere- ingly striato-punctate, the apex obeuneately depressed mesad, the pleurae striato-punctate, the posterior face largely shining, transversely striate and with a V-shaped depression mesad. Venation heavy, testaceous. Abdomen gently undulate ventrad except at base, above mostly reticulate. Terminalia as in Pl. davisi davisi (compared with a Pl. davisi from Midland County, Michigan, see fig. 79); laterodistal bristles of the parameres strong, other bristles weak or wanting. Vestiture: Sparse, some of it may have been rubbed off. Hotorypr, female (U. S. National Museum Collection, Type No. 58464, from kK. V. Krombein Collection), from Campo, San Diego County, Califor- nia, April 27 1939 (R. M. Bohart). Au1orypr, male, Glencliff Camp, near Buckman Springs, San Diego County, May 7, 1953 (F. X. Williams). Thus, the holotype and allotype were taken from localities seareely 15 miles apart. PaRatypEs, California, as follows: Los ANGELES County : Pasa- dena, 3 males, May 14, 1944 (Kk. W. Cooper), in the Karl V. Krombein Col- lection (U. S. National Museum Collection), 44e14. Tanbark Flat, San Gabriel Mts., 3 males, July 11, 1952; July 5 and 12, 1956 (R. M. Bohart); 1 male, July 18, 1952 (J. W. MacSwain); 1 male and 1 female, June 17, 1 female, June 20, 1956 (R. C. Bechtel). MonrErEy County: Arroyo Seco Camp, 3 females, June 6, 1956 (R. M. Bohart), 2 females, June 5, 1957 (R. M. Bohart). Other specimens not considered paratypes are: 1 male, Sequoia National Park, Tulare County, California (R. C. Bechtel), and 1 male, Samuel Spr., Napa County, California (R. M. Bohart). Discussion. The male of Pl. boharti is identified by its blackish elypeal dise whose edge is multidentate, more rarely, crenulate, and emarginate VoL. XXXI] WILLIAMS: THE GENUS PLENOCULUS 31 mesad much as in the female, by the entirely dark antennal scape and the red and black abdomen. The male from Sequoia National Park and the one from Samuel Springs have the hind tibiae and tarsi pale reddish. The female, on the other hand, except for the not-altogether-constant red- and-black color pattern of the abdomen and the frequently stronger pro- tuberant clypeal teeth, seems inseparable from females with the abdomen entirely black from the same locality, that I regard as a rather dark form of Pl. davisi. Associated with these “davist”-type females are typical pale- marked males of Pl. davisi. Plenoculus stygius Williams, new species. (Figures 5, 62, 90.) FEMALE, holotype. Length, 3.5 mm. Shining, the sculpture very fine. Black; mandibles except base and apex, margin of produced portion of ely- peus, the legs beyond femora, yellowish brown; the tarsi in part duskier; antennae brownish beneath; tegulae brownish except at apex; apex of pygi- dium reddish. Head coriaceous; elypeus short, smooth and thick alone middle part of margin, the dentition rather minute and consisting of three teeth on each side of the rather wide and shallow emargination; antennae subclavate; ocelli forming very slightly more than a right-angle triangle. Dorsulum finely and closely punctate; dise of propodeum coriaceous, under certain hehts with minute transverse wrinkles and a shallow median trough, the pleurae polished, exceedingly fine reticulate-punctate, posterior face of pro- podeum nearly smooth, with an inverted tear-shaped depression. Foretarsal comb not strong, leg bristles pale. Truncation of marginal cell very narrow. Tergites under lower magnification appearing smooth, but under 80 mag- nification with very fine transverse striae, and with a few strong punctures on tergite 5. Pygidium triangular, very little constricted apically, the dise with a few strong punctures. The silvery pile while not generally outstand- ing, is conspicuous on the pronotum, mesopleurae, and above on either side of the dise of the propodeum. Mats, allotype. Leneth, 3.2 mm. Much like the female but the sculpture is not quite so fine, although the propodeal pleurae appear smooth and pol- ished under moderate magnification. In addition to the pale yellowish brown tibiae and tarsi, the clypeus is dull yellow, as is also the scape beneath, the antennae being generally dull brownish above, paler beneath. There is the usual hair tuft at each side of the elypeus that is angled laterad and with a low median lobe. Antennae subclavate; ocelli forming a little less than a right-angle triangle. Abdomen with sternal cross ridges; each volsella with five stout bristles. Hotoryrr, female (Citrus Experiment Station, Riverside), from Palm 32 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. Springs, Riverside County, California, May 11, 1935 (P. H. Timberlake), on Euphorbia polycarpa; ALLOTYPE, male, Borego, San Diego County, May 12, 1955 (EF. X. Williams), on a mat Huphorbia. Pararypes: 1 female, Palm Springs, Riverside County, May 11, 1935 (P. H. Timberlake), on Euphorbia polycarpa; 1 female, 6 miles south of Palm Springs, Colorado Desert, Cali- fornia, June 8, 1930, on Eriogonum trichopodum, and 1 male near Palm Springs, June 8, 1930, on Croton californicus, both specimens collected by P. H. Timberlake; 4 males, Borego, San Diego County, April 29 and May 12, 1955 (F.. X. Williams); 1 male, Borego Valley, San Diego County, dunes, April 18, 1957 (R. M. Bohart). Other specimens not considered paratypes: 4 males, Borego, May 12, 1955 (EF. X. Williams); 6 females and 1 male, Thousand Palms, Riverside County, California, one in March, the rest in April, 1955 (W.R.M. Mason); 1 female, Blythe, Riverside County (W.R.M. Mason); 1 female, Cronise Valley, San Bernardino County, California, April 29, 1956 (M. Wasbauer). One female, Picacho Pass, Pinal County, Arizona, September 13, 1954 (P. H. Timberlake), “flying over the ground.” On the same mount as one of the California females taken by Timberlake is its prey, a small mirid bue. Discussion. Obviously this little desert dweller is closely related to Pl. davist Fox, but a combination of characters, at times somewhat variable, separates it from Pl. davist. The more obvious distinctions in Pl. stygius are its highly polished condition, the lack of any pale clear yellow markings, the more delicate clypeus dentition, the rather sac-like marginal cell, its narrow truneation that extends well beyond the third submarginal cell, and addi- tionally in the male, the generally stouter volsellar spines. Oceasionally Pl. stygius has the abdomen orange red, as in the above listed female from Blythe, and the one from Cronise Valley. Plenoculus davisi I'ox. (Figures 6, 7, 8, 12, 39, 41, 48, 44, 45, 53, 54, 57-60, 63-69, 74, 75, 76, 79, 86, 88.) Plenoculus davisi Fox, 1893, Psyche, 6:554-555; 18938, Proc. Acad. Nat. Sci. Phila., 45:537. Male and female. Michigan, Montana. Plenoculus abdominalis ASHMEAD, 1899, Psyche, 8:339. Male. Arizona (Prescott). Plenoculus apicalis WiLLIAMs, 1914 (1913), Kansas Univ. Sci. Bul., 8:175. Male and female. (Biol., 1. ¢., 207-208, pl. 30, fig. 120.) Distribution: Transcontinental in the United States. Canada, British Columbia (Vernon). Mexico, Baja Califor- nia Norte; Sinaloa (Mazatlan); Durango (Nombre de Dios), Nayarit (San Blas). The original description of this species is as follows: Plenoculus davisi 9 —Anterior margin of clypeus deeply incised, armed with five teeth on each side, the outer and inner tooth is generally the largest, the others being in some specimens indistinct; front very finely granulated, with a long, dis- VoL. XXXT] WILLIAMS: THE GENUS PLENOCULUS (vt) (St) tinct medial impressed line, which originates in a strong fovea, before the anterior ocellus and extends down between the base of the antennae; ocelli forming a tri- angle; on each side of anterior portion of vertex there is a curved furrow, running from each hind ocellus to the inner eye-margin; first joint of flagellum possibly a little shorter than either second or third, which are a little longer than any of the following joints, except the last; dorsulum and scutellum with very fine, close punc- tures, both strongly convex; suture between dorsulum and scutellum strong; meta- thorax above very finely granulated, rather strongly furrowed down the middle, the extreme base generally with a transverse series of small fossae, sides and posterior face very finely striated; abdomen finely and closely punctured, the pygidial area triangular, with large, sparse punctures, its lateral ridges not well defined; black; clypeus, except apex, tegulae, four anterior tibiae on outer side, except apical por- tion, yellowish; tarsi and apical margins of the abdominal segments testaceous; wings hyaline, iridescent; nervures testaceous, apical abdominal segment more or less rufous; head, thorax and abdomen more or less covered with silvery pile, which is most dense on the face, clypeus, mesopleurae and metathorax. Length 5-6 mm. Var. legs, except anterior tibiae entirely black. @. Anterior margin of clypeus slightly produced medially; antennae shorter than in the © and subclavate; clypeus entirely, scape beneath, tegulae, tubercles, apex of femora, the tibiae, except inner side of the two anterior pair, and the tarsi, bright yellow. Length 4144-5 mm. Agricultural College, Michigan (June and July). Collected by Mr. Gager C. Davis, to whom it is dedicated; Montana (Morrison). Coll. Amer. Entom. Society. DISTRIBUTION. Over five hundred specimens of what I consider to be Pl. davisi have been examined. The great majority of these are listed here, with their distribution as follows: ARIZON A—CocuiskE County: Douglas, 1 female, August 8, 1955 (R. R. Dreisbach). Maricopa County: Tempe, 2 males, August 3, 1917 (Cornell University, Lot 542). Pima County: Tucson, 1 male, May 30, 1920 (F. X. Williams). Dateland, 1 female, April 12, 1955 (Butler and Werner). CALIFORNIA—ALPINE County: Hope Valley, 4 females and 1 male, July 9 and 18, 1948 (J. W. MacSwain and P. D. Hurd). Contra Costa County: Antioch, 1 male, June 4, 1949 (F. X. Williams); 1 male and 1 female, July 8, 1954 (P. D. Hurd); Danville, 2 males, June 21 and 23, 1949; 2 females, August 6 and 10, 1949 (F. X. Williams). Ex Dorapo County: Echo Lake, 1 male, July 23, 1955 (E. I. Schlinger); Lake Fontanillis, 8500 feet, 1 female, July 21, 1955 (KE. I. Schlinger) ; two miles south of Meyers, 2 females, July 24, 1955 (KE. I. Schlinger). Imprr1aL County: Gordon’s Well, 3 miles west, 1 male, July 14, 1956 (KE. I. Schlinger) ; Pinto Wash, 3 females and 13 males, May 5, 1958 (F. X. Williams), either on Huphorbia polycarpa hirtella or Eriogonum inflatum; Fish Creek Mts., 300 feet, 2 males, March, 1 female, April, 1955 (W. R. M. Mason). Inyo County: Westgard Pass Plateau, 1 female, May 27, 1987 (C. A. Hamsher). Kern County: Randsburg, 1 male, May 1, 1921 (R. M. Bohart). Lake County: Midlake, 1 female, May 30, 1955 (KE. I. Schlinger). Lassen County: Bridge Creek Camp, 6 females, July 9, 1949 (J. W. MacSwain); Summit Camp, I female, 34 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH Ser. July 28, 1949 (P. D. Hurd) ; Westwood, 2 males, July 9, 1949 (P. D. Hurd); McCoy Flat, 3 females, July 8, 1949 (J. W. MaeSwain and P. D. Hurd). Los ANGELES County: Mt. Wilson Trail, Branigan, 1 female, August 10, 1915 (P. H. Timberlake); Claremont (Baker), 1 female (PemC); Chrystal Lake, 1 male, June 29, 1950 (F. X. Williams) ; 1 female, July 9, 1952 (R. M. Bohart); eight miles east of Lancaster, 1 male, September 1, 1956 (KH. I. Schlinger); Pasadena, 1 male, May 14, 1944 (44F14) (USNM) (K. W. Cooper, Coll.); Tanbark Flat, 1 male, June 17, 1956 (R. M. Bohart). Mon- TEREY County: Arroyo Seco Camp, 38 males and 6 females, August 6, 1956, and June 5, 1957 (R. M. Bohart) ; San Lucas, 1 male, August 20, 1935 (P. H. Timberlake), on Hriogonum gracile. Napa County : Samuel Springs, 1 female and 2 males, May 9, 1953 (E. I. Schlinger, R. M. Bohart); 1 female, May 28, 1953 (R. C. Bechtel); 1 female, May 29, 1953, and 1 female, May 24, 1956 (KE. I. Schlinger). NEvapa County: Sagehen Creek, near Hobart Mills, 1 female, May 29, 1 male, June 25, 1954 (R. M. Bohart); 3 females, July 2 and 9, 1954 (E. I. Sehlinger, R. M. Bohart); 1 male, July 25, 1956 (R. M. Bohart). PLAcER County: Lake Tahoe (6225 feet), 1 female, July 15, 1949 (EK. G. Linsley) ; Carnelian Bay, Lake Tahoe, 2 males and 1 female, July 22, 1957 (R. M. Bohart). PLumas County: Bucks (5070 feet), 1 male, July 23, 1937 (EF. X. Williams); Joinsville, 1 female, August 26, 1956 (R. M. Bohart). RIvERSIDE County: Andreas Canyon, Palm Springs, 1 female, April 24, 1932 (P. H. Timberlake), on Friogonum polifolium; Palm Springs Station, 1 female, June 26, 1952 (J. W. MacSwain); Perris, 3 miles west of, May 14, 1956 (U.S.N.M.) (U. N. Lanham, Coll.); Mt. Wilson Trail, Branigan, 1 Figure 55. Plenoculus propinquus. Female. From Riverside, California. Figure 56. Plenoculus propinquus. Male. From Riverside. Clypeal dentition of A, male from Santa Paula, California; B, male from Eden, California. Figure 57. Plenoculus davisi transversus. Female. Holotype. From Three Rivers, Tulare County, California. Figure 58. Plenoculus davisi. Female. Andreas Canyon, Palm Springs. Figure 59. Plenoculus davisi transversus. Female. Holotype. Mandible. From outer side. Figure 60. Plenoculus davisi. Female. The mandible is worn down. Figure 61. Plenoculus boharti. Female. Holotype. From Campo, San Diego County, California (U.S.N.M. collection). At A, clypeal dentition of Pl. boharti, male, allotype, from near Buckman’s Springs, San Diego County, California; at B, is the clypeal dentition of a male paratype from Tanbark Flat, San Gabriel Mts., Calif. Figure 62. Plenoculus stygius. Female. Holotype. The head has been rotated so that the lower side of the face is more forward than the vertex. From Palm Springs, California. Figure 63. Plenoculus davisi. Female, with red abdomen. From Riverside, Calif. Figure 64. Plenoculus davisi. Female. From Sergeant’s Bluff, Iowa. Figure 65. Plenoculus davisi. Male. From Riverside. VoL. XXXII] WILLIAMS: THE GENUS PLENOCULUS 35 Figure 66. Plenoculus davisi. Male, with red abdomen. From Plumas County, California. Figure 67. Plenoculus davisi. Female, with red abdomen. Clypeus probably worn down. From Imperial County, California. Figure 68. Plenoculus davisi. Female. From northwestern Kansas. Figure 69. Plenoculus davisi. Female. From Olmsted County, Minnesota. 36 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. female, August 10,1915 (P. H. Timberlake) ; One Thousand Palms, 1 female, . April 10, 1957 (P. H. Timberlake), on Hriogonum reniforme; Blythe, 1 female, April 24, 1955 (W. R. M. Mason) ; Thousand Palms, 13 males, April, 1955 (W. R. M. Mason); Riverside, 1 female, September 26, 1924, on annual Eriogonum, 1 female, October 15, 1926 on Eriogonum reniforme, 1 male, August 7, 1926, on Helianthus annuus, 1 female, May 14, 1926, and 1 female, September 8, 1927, on Euphorbia albomarginata, and 1 male, May 12, 1950, on EHriogonum fasciculatum, all of these collected by P. H. Timberlake; The Gavilan, 1 male and 1 female, May 31, 1937, 1 female, June 2, 1938 (P. H. Timberlake), all on Hriogonum fasciculatum. SAN BERNARDINO County: Big Bear Valley, 6700-6800 feet, 2 males, August 11, 1933, on Eriogonum wrighti, and September 14, 1934, on Eriogonum; Camp Baldy, 2 males, June 26, 1956 (R. M. Bohart); Dollar Line Trail, San Bernardino Mts., 1 male, July 11, 1956 (R. M. Bohart); Cronise Valley, 2 females, April 29, 1956 (P. D. Hurd, M. Wasbauer), Helotropus curassavicus, Prosopis; twelve miles east of Mentone, San Bernardino Mts., 3 males and 3 females, July 11, 1956 (R. C. Bechtel) ; Mill Creek, 6000 feet, 1 female, August 28, 1936 (P. H. Timberlake); Morongo Valley, 1 male, May 7, 1939 (P. H. Timberlake), on Eriogonum fasciculatum; Valley of the Falls, 1 female, August 18, 1945 (P. H. Timberlake), on flowers of Chrysothamnus; Verdemont, 1 male, May 17, 1946 (P. H. Timberlake), on Chorizanthe parryi; Helendale, 1 female, May 16, 1955 (W. R. M. Mason); Redlands, 1 male (R. F. Cole). San Dirco County: Borrego (or Borego), 10 males and 4 females, April 1 and 2, 19538 (P. D. Hurd); 9 males, 3 females, most with label “Croton californicus,” last part April, 1954 (P. D. Hurd); 2 males, April 25, 1954 (M. Wasbauer), on Croton californicus; 4 males and 2 females, April, May, 1955, and 1957 (F. X. Williams); Borrego Dunes, 1 male, April 18, 1957; Borrego, Palm Canyon, 1 male and 1 female, April 19, 1957 (R. M. Bohart) ; two miles north of Warner Springs, 1 female, July 8, 1956 (R. M. Bohart), on Croton californicus; Green Valley, 5 miles south of Cuyamaea, 1 female, August 8, 1951 (P. H. Timberlake), on Hugelia virgata. SAN Luts OBISPO County: Black Lake, 1 female, July 29, 1955, and 2 females, August 29, 1955 (R. M. Bohart). San Joaquin County: Tracy, 1 female, June 7, 1949 (J. W. MacSwain). Sanra Cruz County: Felton, 1 male, May 15-19, 1907, 300-500 feet (Bradley), Cornell U. lot 684, sub. 39, det. J. C. Bradley, 1928; 2 females, same locality, May 20-25, 1907 (J. C. Bradley) ; Glenwood, 1 female, May 27, 1908 (J. Chester Bradley). SHasta County: Hat Creek, 1 female, June 23, 1955 (J. W. MacSwain), 1 female, May 28, 1956 (J. W. MaeSwain) ; Moose Camp, 1 female, July 14, 1957 (E. I. Schlinger); Snow Mt. Road, 1 female, June 23, 1955 (J. W. MacSwain). Srerra County: Independence Lake, 2 females, July 17 and 20 (R. M. Bohart); 1 female, July 27, 1956 (R. M. Bohart). Sonoma County: Cloverdale, 1 male, August 26, 1953 (H.I. Schlinger). TeHama County: Four miles north of Paynes Creek, 1 female, VoL. XXXT] WILLIAMS: THE GENUS PLENOCULUS 37 May 21, 1955 (R. M. Bohart). TULARE County: Three Rivers, 3 miles west of, 1 female, May 10, 1938 (P.H. Timberlake), flying over ground (= Pl. davisi transversus). TUOLUMNE County: Dardanelles, 1 female, June 26, 1951 (C. A. Downing). VENTURA County: Santa Paula, 2 males and 2 females, June 5, 1927 (P. H. Timberlake). Yoto County: Davis, 32 males and 50 females, collected from June 7 to September, 1953, 1956, and 1957 (Collee- tors: R. M. Bohart, R. C. Bechtel, J. C. Downey, A. T. MeClay, and E. I. Schlinger), a female of one of these specimens is mounted with its prey, the nymph of a heteropterous bug (R. C. Bechtel, collector); Elkhorn Ferry, 4 females and 60 males, August 1, 1956 (R. M. Bohart). COLORADO—A.aAmosa County: Great Sand Dunes, 1 female, July 20- 21, 1954 (H. E. and M. A. Evans). CONNECTICUT—East Hartford, 1 male, September 4, 1947 (H. E. Evans) ; Poquonock, 1 male, June 27, 1905 (H. L. Viereck) (= Pl. davisi atlanticus Viereck) (Col. U.S.N.M.). FLORIDA—Welaka, 1 male, May 1-4, 1955 (H. E. and M. A. Evans) (= Pl. davisi atlanticus). IDAHO—Hollister, 1 female, No. 6, July 27, 1932, S. pestifer (David E. Fox) (Col. U.S.N.M.); Jerome, 1 female, June 27, 1932 (“Pole 20, trap 2, Wind Van Trap’’). INDIAN A—1I female, No. 2177 (1933). TOW A—Sergeant Bluff, 3 females and 3 males, August 3, 1933 (C. N. Ainslie); Sioux City, 1 male, May 25, 1931 (C. N. Ainslie). KANSAS—Criay County: 4 males, August 9, 1952 (Lin). GraHnam County: 1 male, August 16, 1912 (F. X. Williams); Grant County: 1 female, August 18, 1952 (Acc. No. 8021, H. E. Evans, Coll.). Paiuiies and Morton Counties: 22 males and 15 females, August, 1912 (F. X. Williams). PoTTAWATOMIE County: | female and 1 male, June, 1950 (H. E. Evans). MICHIGAN—Crawrorp County: 1 male, June 24, 1953. GuapwIN County: 1 male, June 14, 1953. MipLaNp County: 1 male, June 27, 1953. Otsreco County: 1 male, June 24, 1955. All R. R. Dreisbach, Collection. WASHTENAW County: Ann Arbor, 1 male, July 28, 1950 (U. N. Lanham) ; GLADSTONE County and Newaco County: 10 females, 1951, 1953 (R. R. Dreisbach) ; Mipuanp County: 1 female, 1 male, June 9, 1936 (R. R. Dreis- bach) (Det. K. V. Krombein, U.S.N.M.). MINNESOTA—OImstead, 2 females (C. N. Ainslie) (Det. Stevens, U.S. N.M.). MONTANA—Plenoculus davisi davisi Fox “type” (Det. K. V. Krom- bein, U.S.N.M.). NEVADA—Fernley, 1 male, August 5, 1953 (R. M. Bohart); Mt. Rose, 6,500 feet, 1 male, June 14, 1957 (R. M. Bohart). 38 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. NEW JERSEY—Clementon, 1 male, May 24, 1902 (J. C. Bradley). NEW MEXICO—Hmatco County: 2 males, Rodeo, August 26, 1958 (R. M. Bohart). NEW YORK—Ithaea, Kite Hill, 14 males, July 1 and 2 (P. P. Babiy); Kite Hill Campus, Ithaca, and Ithaca, 7 males and 1 female, August 19, 1929, June 15, 19387 (P. P. Babiy); June 26, June 29, 19385 (K. V. Krom- bein); 1 female, 6 miles from Ithaca, August 11, 1955 (with biological note) (H. E. Evans); Minetto, 3 females, June 27, 28, 1955 (H. EH. Evans). NORTH CAROLINA—Kill Devil Hills, 1 female, July 1, 1954 (K. V. Krombein) (U.S.N.M.). NORTH DAKOTA—Beach, 2 females, July 19, 1923 (C. N. Ainslie) (U.S.N.M.) (Det. Stevens). OREGON—Bend, 1 female, August 19, 1953 (R. M. Bohart). TENNESSEE—Memphis, 1 male, July 16, 1914 (J. C. Bradley); 1 female, June 4, 1918 (Cornell U., Lot 684, Sub. 36 and 37, Det. J. C. Brad- ley, 1923, Plenoculus davist 6 Fox). TEX AS—MeDade, 1 male, June 11, 1935 (J. E. Gillaspy) (Krombein Collection, and compared by Dr. Krombein with the type of Pl. abdomi- nalis Ashmead). UTAH—North Fork Duchesne River, 1 male, July 138-14, 1927 (Cornell U., Lot 542, Sub. 305); Logan, 1 female, July 1, 1955 (R. M. Bohart) ; Roose- velt, 5,000 feet, 1 female, July 27, 1953 (R. Dreisbach). WYOMING—Powder River, 1 female, August 1, 1950 (R. R. Dreisbach and K. K. Schwab). CANADA—RBnritisH CoLUMBIA: Vernon, 1 female, July 14, 1947 (H. B. Leech). Tunneling in sand. The burrow stored with immature Aphidae. MEXICO—Duvranco: Nombre de Dios, 1 male and 1 female, August 1, 1951 (P. D. Hurd). Baza Cauirornia: La Paz, 1 female, October 7, 1955 (EF. X. Williams). Nayarit: San Blas, 2 males, July 20, 1951 (P. D. Hurd). SinALoa: Mazatlan, 1 female, October 7, 1955 (F. X. Williams). DIAGNOSES OF THE SUBSPECIES Plenoculus davisi Abdomen red and black; clypeus black in both sexes, almost squarely truncate in the male, usually very low subcuneate in the female, and with a median cleft and poorly developed teeth or none; in the male the aedeagus the volsellar ridge is weakly armed............ Plenoculus davisi mojavensis Williams, new subspecies (86, 88). Abdomen chiefly reddish; head relatively wide; produced portion of clypeus truncate, its median emargination narrow and with two teeth far to each side (57), email eats ) NB: CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER. Kopf und Halsschildseiten mittelstark punktiert, + glanzend. Punktreihen der Hiyinenskrataes aber eleichmaBiger. ( Pattial ) <.<.<4:..2-2-8E4-6 LilsteGaSk oto Sisal Oba 9L+-Z-98T +2 CT/ST 8+6 CileGal €S-¥Z CZ cz 92 LOT T@l-Zo OT-OT AG eer 9L+3-61 +2 OE sietee Ee atno 8E-+Z-LE +2 8I-+3-26+2 GL oaielilste Cstbaitetar cI-+Z-9E-+T PL/GT 8+6 ¢‘I-¢'I 6-1 &@ GS GS “**dBuUrlUe] JO I1ed 4se] UO sapoIyUeq 6001006 "Trrsessess sregaysoryouRig "7" 1OLLIISOg soo TOLl8IUW [YyoIe YF 6 Suey Gino Oia Gr a.0id LOT19}S0g LOLLoJUW :Yole pag Ponce S cree ae porauc rondo 101.10}80q Biota tg. 0 Or0N om co aU otic LOloqUy :Yore pug Seb toparsyel cae tresses ess 1OTIOISOg IOLLaJUY :YoIe 4ST z+ SloYBA-[[ID {JUILINIOIg [edputigd SABI Tepneg Pra toro-a40,Grololg obim 0.6 G.0,0 csc SAB OTATOd soba istiomanenle dsmel lars Aackalewene sABl [R10100q stlensob ential evstouel cueyeUer rele 1810} ‘sea peuy asteveue mee ates sia uere sieves SABI-JOS [Bs.10q] Mo Suc O,OLGeUD GOL CLOT ‘ @BUTWIE] [ROSIC] syunog HO Didict.6 Go Clad Galo uc uu ‘YySuUe] oSIG 668 08g 162 98Z Z9% OSZaek ee ae sponses WU “YSU, prepuris ee eS i En OS ee ee repels) OSE IMA ee ‘UsWILDeds JO apIS JYSLt JO yUNOD JUaSa.de.l uaydaAy Jo YBa Ye soinsy Ye J Soinsty “Aydeisouessg Jo UOI{N{IWSUy SddLIogs—oys £9996 SVO £9996 SVO £9996 SVO £9992 SVD £9996 SVO LOJODADY OD ee -UWdWLVedS JO apis JJa] JO JUNOD JUaSe.IdaI uesyday Jo ‘TY “M APQ—UMC ‘seuUsIDg Jo AWapRoy BIULOJI[VO—=SVO penurju0opO—] WIAVL ST[RAISNV BLoo[LUtay fo suamwads gy fo spunoa Paj}oajas pup “YpHua) osip “y)hua) pavpunypy 176 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. DorsaL sort-RAys. Modally 25, ranging from 23 to 26. The published counts range from 20 (Smith, 1958:319) to 22 (Gunther, 1860a :401) ; possibly they do not include some of the anterior rays, which— because of the thick integument of this fish—mavy have been overlooked. ANAL RAYS. Modally 25, ranging from 23 to 26. Pending further inves- tigation, we regard the two anterior elements, which are embedded in thick integument, as unbranched rays rather than as spines.” The published counts range from 21 (Giinther, 1860a:401) to 24 (Ben- nett, 1840 :273). PECTORAL RAYS. Modallv 23, ranging from 21 to 24. The uppermost ray is invariably unbranched. We have not expressed the pectoral formulae in terms of branched and unbranched rays because, in the absence of alizarin preparations, the branching of the lowest rays in the smaller specimens does not appear satisfactorily determinable. The published counts range from 1/20 (Cadenat, 1953:681) to 24 (Waite, 1915 :340). PELVIC BAYS (1,5: When Bennett (1840:273) wrote ‘Ventral 5,” he had surely overlooked the concealed spine. When Giinther (1860b:382) said of Echeneis scutata, “the ventrals are ..., aS in all the species of the genus [cheneis|, composed of one spine... and four soft rays,” he apparently used the word “four” by inadvertence, since in his diagnoses of two other species which he re- ferred to the genus Echeneis, he noted that the ventrals were composed of one spine and five soft-rays (Gunther, op. cit. :377, 383). CAUDAL RAYS. Principal: 9+8. Procurrent: modally 13 in the upper series and 14 in the lower series, ranging from 12 in the upper series and 13 in the lower series to 15 in the upper and 15 in the lower. Since all our specimens have 17 principal caudal rays, it seems probable that the count of 20 by Bennett (1840:273) included the longest procurrent ray in one moiety of the caudal fin and the two longest procurrent rays in the other moiety. Counts of 13-++6 and 19-20 were noted by Waite (1915: 340; 1923 :185). GILL-RAKERS. First arch (anterior aspect) : modally 2+-17, ranging from 1 to 3 on the upper limb and from 14 to 19 on the lower limb. The gill-rakers of the upper limb are small, round structures, which are difficult to discern because they are masked by the long gill-rakers at and immediately below the angle of the arch. Frequencies of counts of the gill-rakers on the anterior and posterior aspects of all arches are presented in table IT. 2. In the order Echeneiformes, according to Berg (1940:495), there are “‘no spines in second dorsal and anal.” VoL. XXXT] FOLLETT AND DEMPSTER: ECHENEIDID FISH 177 TABLE II Frequencies of gill-raker counts in 18 specimens of Remilegia australis! (Both sides counted) Upper limb Lower limb i 2 3 ji HA ss iA ahs KS ey IY 2 ik BZ First arch: /NTMIGICON!P 5 ooacdoc 2 BA 3 6 15 1 2 IPOSteGlor a. 422) Path i) 1 als 6 Second arch: AMIECTION fee ac oe Zot: 3 ae) y eee eee Be 1 (ale 5 4 7 Aland al RoOsternionyeoea 5: (en Pe) eee ee te eee es ( alal tl 0 , a eee il Third arch: INMIOON? 5 ono > Golo se a 4 tab 9 3 2 IROStERIOR) 5 = «cis. 32 4 4 2" 4 6 Fourth arch: IANECTION fick. es APS Dik tee ees See ee 1 6 18 ii 2 1 POSterion jeao..- DOM ey tek 1 Te ales 8 1 1 1. All rudimentary gill-rakers are counted, and the gill-raker in the angle of the arch is included in the count of the lower limb. The published counts are ++ 14 (Waite, 1915 :341), 1 at angle and 14 on lower limb (Cadenat, 1953:681), 2+1+14 (Matsubara, 1955:1212), and 0+15 (Smith, 1958 :319). BRANCHIOSTEGAL RAYS. Modally 10-10; 9-9 in one specimen, 10—9 in two specimens. In the original deseription of Echeneis australis, Bennett (1840 :273) noted, ‘““Branchiostegous rays 10.” The original diagnosis of the genus Echeneis included the character ““Membr. branch. radiis X” (Linnaeus, 1758 :260). Giinther (1860b :376), however, regarded “branchiostegals seven” as a character of the genus Echeneis (to which he referred the present spe- cles; op. cit.:381, as Echeneis scutata). This statement of Giinther’s may have been accepted by those authors who noted seven branchiostegals as a character of the family Echeneididae (e.g., Gill, 1883 :565; Jordan and Gil- bert, 1883:416; Jordan and Evermann, 1898 :2266; 1905 :494; Evermann and Marsh, 1900:301; Meek and Hildebrand, 1928:895; Fowler, 1959 :498). BRANCHIOSTEGAL MEMBRANES. In all our specimens, the left branchioste- gal membrane overlaps the right. This asymmetry is in accordance with the general rule for all fishes in which the branchiostegal membranes are not united with each other or with the isthmus (Hubbs and Hubbs, 1945 :279; Crossman, 1960 :368). Size. Our specimens range from 105 to 399 mm. in standard length. The smallest example that we have found mentioned in the literature is 178 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. the 46-mm. specimen recorded by Liitken (1875:42). The largest of which we have found a published record is the 23-inch specimen mentioned by Giinther (1860b:382); possibly this specimen provided the basis for the statements that this species attains a length of 2 feet and a weight of 8 pounds (Giinther, 1880:461; 1886 :326) and that it attains a length of 600 mm. (Kamohara, 1950:237; Smith, 1958 :320). Disc LENGTH. Averaging 0.50 standard leneth; ranging from 0.48 stand- ard length to 0.52 standard length. A dise length of 0.5 standard length was recorded by Woods (1942 :192) and a dise leneth of 0.46 standard length, by Smith (1958:319). The dise leneth has been recorded as 21% in total length (Ginther, 1860a :401; Pe- rugia, 1881:17) and as 274 in total length (Lititken, 1875 :42). Cotor. In life, one specimen—the only one that we obtained alive—was blue with a narrow white margin on each fin. This specimen, which is shown in plate 1, upper figure, was photographed immediately after it had been preserved in formalin and before any observable change had occurred in the life colors. After months in alcohol, this specimen and 15 others are dark slaty blue and one specimen is brown. In life, the specimen from a whitebelly dolphin (see Hosts, infra) was reported to have been light slate gray with some hint of bluish background; after preservation in aleohol it has darkened to a bluish brown. The life colors of the whalesucker have been recorded as brown (Giin- ther, 1860a :401; 1860b :381; Liitken, 1875 :43) ; dark brown with white edge on dorsal and anal fins (Matsubara, 1955:1212); brownish (Kamohara, 1950 :237); uniform violet (Perugia, 1881:17); uniform dark slaty blue, edges of fins lighter, dise brown (Waite, 1915 :341); dark slaty-blue (Whit- ley, 1949:23); deep marine blue (Cadenat, 1953 :682); bluish white (Cade- nat, 1953 :682); grayish white (Ui, 1932 :253); uniform slate gray, margins of dorsal and anal and tips of tail white (Breder, 1936:43); uniform gray (Mahnken and Gilmore, 1960:134); almost black, margins of dorsal, anal, and pectoral and upper and lower margins of caudal white (Smith, 1958: 320). Hosts Our 17 specimens were all taken from blue whales, Sibbaldus musculus. The specimen (Seripps Institution of Oceanography no. 59-74) from off San Roque Rock, Baja California, was taken from a whitebelly dolphin, Delphinus bairdu (John E. Fitch, personal communication). Published records mention the following cetaceans as hosts of the whale- sucker: dolphin (Liitken, 1875:42); sulphur bottom whale [blue whale} (Carl and Wilby, 1945:29); Delphinus delphis (?) (Cadenat, 1953 :680) ; Vout. XXXII] FOLLETT AND DEMPSTER: ECHENEIDID FISH 179 sperm whale, Physeter (Krefft, 1953:278); porpoise (Smith, 1958 :320): spotted porpoise, Stenella plagiodon (Mahnken and Gilmore, 1960:134). One individual was attached to an oar (Perugia, 1881:17). DISTRIBUTION The present specimens were collected off California and Baja California. Remilegra australis appears to be of world-wide distribution in temper- ate and tropical seas. It is known from as far north as Vaneouver Island, British Columbia (Lat. 49° N.), and as far south as the Cape of Good Hope (at. 34°°S.)). The localities from which we have found this species recorded may be listed as follows: Pacific Ocean: Society Islands (Bennett, 1840, vol. 1:165). New Guinea (Myers, 1960:78). Japan (Ui, 1932:253; Kamohara, 1950:237; 1958:61; Matsubara, 1955:1211). British Columbia (Halkett, 1913:96). Between Lower California and Panama (Breder, 1936 :42). Peru (Krefft, 1953 :278). Atlantic Ocean: Gulf of Mexico (Woods, 1942:192: Mahnken and Gilmore, 1960:154). Chesapeake Bay (Massmann, 1957:157). Mid-Atlantic (Lititken, 1875:42). Adriatic Sea (Perugia, 1881:17). French West Africa (Cadenat, 1953 :680). Indian Ocean: South Africa (Whitley, 1949:22; Smith, 1958 :320). India (Gitnther, 1860b:381). Ceylon (Giinther, 1860a :401). Western Aus- tralia (Whitley, 1947:149). South Australia (Waite, 1915 :340). ACKNOWLEDGMENTS We wish to express our gratitude to Toshio Asaeda of the California Academy of Sciences, for translations of the Japanese literature; to Seth B. Benson of the University of California, for advice; to Charles Caito, Gaeton Caito, and John Caito, of the Del Monte Fishing Company and Western California Fish Company, for enabling us to obtain the first speci- mens of Remilegia australis known from California; to David K. Caldwell and Dale W. Rice, of the United States Fish and Wildlife Service, for the loan of their specimens and for permission to publish their data; to John E. Fitch of the California Department of Fish and Game, for the loan of the specimen from Baja California; to G Dallas Hanna and C. Edward Cromp- ton, of the California Academy of Sciences, for the colored plate; to Carl L. Hubbs of the University of California, for assistance and advice. 180 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. LITERATURE CITED AMERICAN FISHERIES SOCIETY COMMITTEE ON NAMES OF FISHES 1960. A list of common and scientific names of fishes from the United States and Canada. Special Publication no. 2. 2nd ed., 102 pp. American Fish- eries Society, Ann Arbor, Michigan. BENNETT, FREDERICK DEBELL 1840. Narrative of a whaling voyage round the globe, from the year 1833 to 1836. Comprising sketches of Polynesia, California, the Indian Archi- pelago, etc. With an account of southern whales, the sperm whale fish- ery, and the natural history of the climates visited. Vol. 1, xv + 402 pp.; Vol. 2, vii + 395 pp. Richard Bentley, London. Bere, Leo S. 1940. Classification of fishes, both Recent and fossil. Travaux de Institut Zoologique de Vv Académie des Sciences de VURSS, 5(2):87-517. (Litho- printed, J. W. Edwards, Ann Arbor, Michigan, 1947.) BREDER, C. M., JR. 1936. Scientific results of the second oceanographic expedition of the ““Pawnee,’ 1926. Heterosomata to Pediculati from Panama to Lower California. Bulletin of the Bingham Oceanographic Collection. Peabody Museum of Natural History, Yale University, 2(3):1—-56. CADENAT, J. 1953. Notes d’ichthyologie ouest africaine. IV—Les rémoras des cotes du Sénégal. Bulletin de VInstitut Francais d’Afrique Noire, 15(2) :672-688. Cart, G. CLirrorD, and G. V. WILBY 1945. Some marine fish records for British Columbia. Canadian Field-Natu- ralist, 59 (1) : 28-30. Carus, JULIUS VICTOR 1893. Prodromus faunae mediterraneae; sive, descriptio animalium maris mediterranei incolarum quam comparata silva rerum quatenus inno- tuit adiectis locis et nominibus vulgaribus eorumque auctoribus in commodum zoologorum. Brachiostomata. Mollusca. Tunicata. Vertebrata. Vol. 2, IX + 854 pp. Stuttgart. CLEMENS, W. A., and G. V. WILBY 1949. Fishes of the Pacific coast of Canada. Rev. ed. Bulletin of the Fisheries Research Board of Canada, 68:1-368. CROSSMAN, E. J. 1960. Variation in number and asymmetry in branchiostegal rays in the fam- ily Esocidae. Canadian Journal of Zoology, 38:363-—375. DE BUEN, FERNANDO 1934. Discocephali de Espana (Peces). Boletin de la Sociedad Espanola de Historia Natural, 34(7) :397—412. Vou. XXXT] FOLLETT AND DEMPSTER: ECHENEIDID FISH 181 EVERMANN, BARTON WARREN, and MILLARD CALEB MARSH 1900. The fishes of Porto Rico. Bulletin of the United States Fish Commission, for 1900, 20(1) : 49-350. FABER, G. L. 1883. The fisheries of the Adriatic and the fish thereof. A report of the Austro- Hungarian sea-fisheries, with a detailed description of the marine fauna of the Adriatic Gulf. xxvi + 292 pp. Bernard Quaritch, London. Fow Ler, HENRY W. 1959. Fishes of Fiji. vii + 670 pp. Government of Fiji, Suva. GILL, THEODORE 1862. Remarks on the relations of the genera and other groups of Cuban fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 1862: 235-242. 1864. Note on the nomenclature of genera and species of the family Echenei- doidae. Proceedings of the Academy of Natural Sciences of Philadel- phia, 1864:59-61. 1883. Note on the relationships of the echeneidids. Proceedings of the United States National Museum, 1882, 5:561—566. GRAHAM, Davip H. 1953. A treasury of New Zealand fishes. 404 pp. A. H. and A. W. Reed, Wel- lington. GRINNELL, JOSEPH 1933. Review of the Recent mammal fauna of California. University of Califor- nia Publications in Zoology. 40(2):71-234. GuDGER, E. W. 1926. A study of the smallest shark-suckers (Echeneididae) on record, with special reference to metamorphosis. American Museum Novitates, no. 234:1-26. GUNTHER, ALBERT 1860a. On the history of Echeneis. Annals and Magazine of Natural History, ser. 3, 5:386—-402. 1860b. Catalogue of the acanthopterygian fishes in the collection of the British Museum. Squamipinnes, Cirrhitidae, Triglidae, Trachinidae, Sciaeni- dae, Polynemidae, Sphyraenidae, Trichiuridae, Scombridae, Carangidae, Xiphiidae. Vol. 2, xxi + 548 pp. London. 1880. An introduction to the study of fishes. xvi + 720 pp. Adam and Charles Black, Edinburgh. 1886. 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Description of the fishes from the provinces of Tosa and Kishu, Japan. 3+ 288 + 5 + 48 + 27 pp. Bunkyo Kyokai, Kochi. (In Japanese. ) 1958. A catalogue of fishes of Kochi Prefecture (Province Tosa), Japan. Re- ports of the Usa Marine Biological Station, 5(1):1-76. KREFFT, GERHARD 1953. Ichthyologische Mitteilungen aus dem Institut fiir Seefischerei der Bun- desanstalt fiir Fischerei. I. Zoologischer Anzeiger, 150(11-12) : 275-282. LINNAEUS, CAROLUS 1758. Systema naturae per regna tria naturae, secundum classes, ordines, gen- era, species, cum characteribus, differentiis, synonymis, locis. 10th ed., vol. 1, 824 pp. Laurentii Salvii, Holmiae. VoL. XXXII] FOLLETT AND DEMPSTER: ECHENEIDID FISH 183 LUTKEN, CHR. 1875. Ichthyographiske Bidrag. V. Museets Sugefiske (Echeneidae). Videnska- belige Meddelelser fra den naturhistoriske Forening i Kjdbenhavn, 1875 (1-4) : 26-43; French summary, 4—5. MAHNKEN, THOMAS, and RAyMOND M. GILMORE 1960. Suckerfish on porpoise. 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The marine and fresh water fishes of Ceylon. xvi + 351 pp. Department of External Affairs, Canberra. Myers, G. S. 1960. Phylax telescopus. Copeia, 1960(1):75-78. NicHons: J. I. 1930. The fishes of Porto Rico and the Virgin Islands. Pomacentridae to Ogco- cephalidae. Scientific Survey of Porto Rico and the Virgin Islands. Vol. 10, no. 3, pp. 297-400. New York Academy of Sciences, New York. NINNI, EMILI0o 1912. Catalogo dei pesci del Mare Adriatico. vii + 271 pp. Venezia. PERUGIA, ALBERTO 1881. Elenco dei pesci dell’Adriatico. viii + 60 pp. Milano. SCHULTZ, LEONARD P., and ALLAN C. DELACY 1936. Fishes of the American Northwest. A catalogue of the fishes of Washing- ton and Oregon, with distributional records and a bibliography. Mid- Pacific Magazine, 49 (2) :127-142. 184 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH SER. Smirn, J. L. B. 1958. Rare fishes from South Africa. South African Journal of Science, 54(12): 319-323. TORTONESE, ENRICO 1958. Elenco dei leptocardi, ciclostomi, pesci cartilaginei ed ossei del Mare Mediterraneo. Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale in Milano, 97(4) :307—-345. Ur, Hozo 1932. Monograph of fishes of Kishu, Wakayama Prefecture. 3rd ed., 45 + 284 + 10 pp. Kindaibungeisha Book Co., Osaka. (In Japanese. ) WAITE, EpGAr R. 1915. A supposed incidental occurrence of a sucker fish (Hcheneis australis Bennett) in Australian waters. Transactions of the Royal Society of South Australia, 39:340-343. 1921. Catalogue of the fishes of South Australia. Records of the South Aus- tralian Museum, 2(1):1-208. 1923. The fishes of South Australia. 243 pp. British Science Guild, Adelaide. WHITLEY, GILBERT P. 1947. New sharks and fishes from Western Australia. Part 3. Australian Zool- ogist, 11(2) :129-150. 1948. A list of the fishes of Western Australia. Western Australia Fisheries Department, Fisheries Bulletin no. 2:1—35. 1949. Sucking fishes. Australian Museum Magazine, 10(1) :17-23. Woops, LOREN P. 1942. Rare fishes from the coast of Texas. Copeia, 1942(3):191-192. Pe ACE Upper figure. Remilegia australis (Bennett), whalesucker (CAS 26664), stand- ard length 262 mm., from a blue whale captured October 16, 1958, off San Mateo County, California. Kodachrome by W. I. Follett. Lower figure. Dise of a 380-mm. specimen of Remilegia australis (CAS 26663) with 28 pairs of laminae—one more pair than previously recorded for this species. Kodachrome by W. I. Follett. VoL. XXXT] FOLLETT AND DEMPSTER: ECHENEIDID FISH Ctrrceec Pea Oe WM ee ~— Nh i Ss Rates PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. XXXI, No. 8, pp. 185-225 March 7, 1962 ON SOME EARTHWORMS OF EISEN’S COLLECTION’ BY G. E. GATES University of Maine, Orono Gustav Hisen, pioneer of oligochaetology in North America, and for whom three genera and several species were named, collected in California and Mexico, and also during 1880-1903, while he was on expeditions to Central American republics. Most of his material was lost at the time of the San Francisco earthquake in 1906. Fortunately, a few lots survived the disaster. The present contribution reports on those belonging to the California Acad- emy of Sciences. This material was received by the author more than ten years ago. All of it is more or less softened. Some specimens were so macer- ated that even identification to family was impossible. Others were identi- fiable, but disintegrated more or less completely during dissection. The study of worms in such delicate condition is tedious and the results are not always commensurate with the time expended. However, some of the material has provided information of considerable interest. The author’s thanks are extended to Dr. Robert C. Miller for loan of the material. Especially appreciated is his understanding of the fact that a study of some of these specimens would result in their destruction. On CERTAIN BLoop VESSELS OF KARTHWORMS Vascular organs were not mentioned in many of the specifie descriptions, and in others only the location of the last hearts and the condition of the dor- 1. From research financed by the National Science Foundation. [185 ] 186 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. sal trunk (whether double or single) were stated. Such inadequate treatment of essential structures is doubtless responsible for part of the artificiality of the classical system. Much more information must be obtained before taxo- nomie and phylogenetic importance can be evaluated, but data even now available suggest that certain vessels can provide characters for definition of more natural taxa, and of course for a better understanding of evolution. Dorsal and ventral trunks are more likely than not to be complete, but abortions of short portions are known to be noteworthy in the oetochaetid Eutyphoeus. Extra-esophageal trunks, free in coelomic cavities of a pre- intestinal region of the body, may prove to be uniformly median to or lateral to the segmental commissures and the hearts in some genera and this is even true in some families. Posterior portions of these trunks, by their associa- tions with the esophagus and other major vessels, are expected to provide a ereater diversity of taxonomic characters. Latero-parietal trunks (cf. Gates, 1939), in contrast to the extra-esophageals, are on the body wall throughout nearly all of their lengths, rising into the coelom only in some anterior seg- ment to pass up toward the esophagus. Again, association with the gut or other major vessels may provide important characters. These trunks, like the subneural, often are empty or nearly so throughout most of their lengths, and then may be almost if not quite unrecognizable in field-preserved mate- rial. Posterior latero-parietals in Hutyphoeus, as well as in the oetochaetid Eudichogaster, do pass back to the hind end of the body and, according to evi- dence now available, are present only when a subneural is absent. Anterior latero-parietals, possibly even two pairs, have been detected only in Spar- ganophilus in which a subneural appears to be lacking. The subneural has been thought to end in the vicinity of xili—xiv, but probably is complete and continued to a bifureation at the anterior end of the body. Almost a century ago the segmental commissures or hearts that connect dorsal and ventral trunks were designated as laterals by Perrier. Hearts that connect supra-intestinal and ventral trunks he called intestinals. The term “‘latero-intestinal” was later coined by Bourne for hearts that open above into dorsal and supra-intestinal trunks. The supra-intestinal trunk of early authors long has been more properly called the supra-esophageal, and substitution of esophageal for intestinal in the Perrier-Bourne terminology (Gates, 1939, pp. 153-154) provided a more accurate characterization that was not so different as to require a glossary. Latero-esophageal hearts have been identified by various authors as dorso- esophageals, true intestinals, intestinals, dorso-supra-intestino-ventrals, and esophageals. Lateral hearts have been designated as dorso-ventrals and eso- phageals. Esophageal hearts have been called intestinals, supra-intestino- ventrals. The terms lateral and esophageal have also been used for any heart, regardless of its dorsal connections. Unfortunately, the meaning of a par- VoL. XXXT] GATES: EARTHWORMS OF EISEN’S COLLECTION 187 ticular characterization is not always determinable from the text or figures. Doubtless some of the confusion is attributable to the fact that one of the two dorsal bifureations of a latero-esophageal heart is often empty. When strongly contracted, or in poor preservation, an empty bifurcation may be very difficult to identify. Possibly only in Sparganophilus are hearts present in an intestinal region of the body, but in that genus the vessels are lateral. Family ACANTHODRILIDAE This family now comprises the Acanthodrilinae of the classical system (cf. Gates, 1959), the holonephrie genera of the Diplocardiinae (cf. Pickford, 1937), and the holonephrie genera with tubular prostates of ectodermal ori- gin that previously were in the Megascolecinae. The classical Acanthodrilinae probably did not constitute a natural group as Stephenson seems to have suggested when he wrote (1930:819) that “eon- venience is best served” by placing therein all genera ineligible for admission to other megascolecid subfamilies. Recent additions (Gates, 1959) to the family admittedly were for convenience only. Not until much more informa- tion about previously neglected somatie anatomy is available will a more natural arrangement of genera into subfamilies or families be possible. Genus Plutellus Perrier, 1873. This classical genus includes species with one gizzard, in v, vi, or vii, that may be vestigial or well developed, sometimes even in two segments, v—vi or vi-vii and then really two. Species without caleiferous glands or with glands of unknown structure which may be 2 pairs in xiv—xv or xv—xvi, 3 pairs in X—Xil, or X1-xili, 4 pairs in x—xiii or xii-xv or xiii—xvi, 5 pairs in ix—xiii. Spe- cies with an intestinal origin in xiv, xv, xvi or xvii, with hearts in x—xi, x—xii, x-xiil, the last pairs latero-esophageal, lateral, or even esophageal. By defi- nition, the two prostates open to the exterior in xviii but in some species the prostatic pores are in xix or xx. Male gonoducts open to the exterior in xviii near the prostatic pores or pass into prostatie duets within the body wall, just above the parietes, at various more ental levels to the end of the duct or even into the gland itself. Plutellus, accordingly, is a congeries which has in common only the two characters of its family and in addition the lumbricin arrangement of the setae. The distribution, as incongruous as the morphology, comprises Cey- lon, India, Burma, Australia, Tasmania, New Caledonia, New Zealand, Auck- land Islands, Queen Charlotte Island, a Pacifie coastal strip of the United States, Guatemala, and a northern portion of South America. The type spe- cies, supposedly from Pennsylvania, may not even be American. The andry is unknown and the holotype, though still in existence, seems never to have 188 CALIFORNIA ACADEMY OF SCIENCES [Proc. 41711 SEr. been re-examined nor has the species been recognized elsewhere. Until P. he- teroporus is more adequately characterized and some at least of its relation- ships are determined, perhaps more especially to North American species, the status of Eisen’s Argilophilus is likely to remain uncertain. Accordingly, the commonly accepted classical generic name is retained for the two species considered below. Plutellus papillifer (Hisen, 1893). Plutellus papillifer, GatTrs, 1941, Proc. California Acad. Sci. 23, p. 443. Colony Mill, Tulare County, California, 5415 feet, May 18, 1904, 0-1-0. Chas. Fuchs. Tamalpais, Marin County, California, 0—-2-10. Tamalpais Station, Marin County, California, 12-18. Eisen collection, no. 595, 0-0-2 (anterior fragments of 18 and 24 see- ments). EXTERNAL CHARACTERISTICS. Diameter 6-7 mm. (fragments), 4-5 mm. (other specimens). Segments, 139-147, 161 (Colony Mill). Prostomial tongue open posteriorly but with a transverse furrow (several specimens) slightly behind the anterior margin of segment i. Ventral setae of li are lacking (fragments) and follicle apertures either are closed or are now unrecogniz- able. Setae in preclitellar segments are ornamented near the tip with fine, transverse serrations. Clitellum in xili—xviil, anterior border indistinet, often apparently in xii, definitely at eq/xii (1 specimen) or much nearer 11/12 (1 specimen), in a third worm possibly at 11/12. The elitellum is protuberant (relaxed worms) or coneave like a waist (some relaxed and contracted specimens). Spermatheeal pores at B (see notes to key below), except as indicated to the contrary in the section on polymorphism, at.7/8-8/9. Female pores in a transversely elliptical area of epidermal thickening in BB which is opaque and protuberant except in one fragment where it is wholly translucent and depressed. Male fields more or less distinctly demareated, reach into xvii, as well as xix and are centered in AB. Each is longitudinally placed, widest at eq/xvill where a slight protuberance bears the male pore and follicle aper- tures. The epidermis in an anterior portion (mostly in xvii) and in an equi- sized posterior portion (mostly in xix) appears to be thicker but without translucence. Genital markings are located as follows: 8/9 (14 specimens), 9/10 (33), 10/11. (84), 11/12 (83), 12/13) (382), 18/14 (4), 14/15 (34), 15/16 arr 16/17. (24), 17/18. (3), 18/19°(1),.19720: (83), 20721 (28), 21722 (8) a 22/725 (1), 23/24 (1). A central portion of each marking is translucent except on VoL. XXXI] GATES: EARTHWORMS OF EISEN’S COLLECTION 189 specimen no. 17 which has two diserete translucent areas in each of the mark- ings from 12/13 posteriorly. INTERNAL ANATOMY. Septa 5/6 and those following are funnel-shaped and so large that the gizzard is back at the level of the eighth segment, 6/7— 10/11 increasingly thickened posteriorly. 11/12-12/13 decreasingly so, sub- sequent septa also slightly strengthened as is the horizontal subesophageal mesentery in x—xili. A special longitudinal muscle band at mD is unrecog- nizable, but behind the elitellum in these relaxed worms a spindle-shaped gap in the longitudinal musculature extends a quarter of the way through each segment from the intersegmental levels between. Gizzard large, strong, with thickened cuticular lining, in v (30 speci- mens). The ventral tvphlosole always is obvious in x-xiv even in the frag- ments in which the mucosa has disintegrated. Low longitudinal ridges on inner wall of gut in x—xiv are irregularly interrupted. Esophageal valve short, at region of insertion of 16/17. Intestinal origin in xvii (20 speei- mens) and recognizably so even when the valve is relaxed. Typhlosole rudi- mentary but very gradually increasing in height through xxi to region of XXVI-Xxvili, ending as follows: in 101st of 139 segments, 107th of 145 seg- ments, 108th of 143 (abnormal worm), 144 and 146 segments, 110th of 147 segments, leaving 35-388 metameres atyphlosolate. The dorsal trunk is complete, traceable in five worms to the brain under- neath whieh it bifureates, the branches passing ventrally along the cireum- pharyngeal nervous connectives. A supra-esophageal trunk is present in x— xili. Extra-esophageal trunks always are median to the hearts and segmental commissures. Posterior lateroparietal vessels small, recognizable only in Xvili-xiv of three specimens, pass up to the gut just in front of 14/15 but are not traceable to any of the major trunks. Segmental commissures of y—1x lateral, those of v in front of the gizzard, traceable to the ventral trunk only in vill-ix except in one worm and then in vil-ix. Last hearts in xii (30 specimens). Nephridia avesiculate, present from ii, in preclitellar segments of con- tracted specimens coiled in long and tight spirals. Holandrie and metagynous (except as noted below), male funnels rather small, plicate. Male gonoduets without epididymis, apparently united just in front of 12/13, passing into the prostate gland or somewhat more ectally and then at the junction of the duet and gland, sometimes with variation from one side to the other of the same worm. Seminal vesicles medium-sized or smaller, vertically placed on the posterior faces of the septa, acinous, not filling the coelomic cavities of xi-xiil. Lumen of prostate glands slit-like in eross section, in the ducts much smaller and cireular in section. Ducts 2 + mm. long, a slenderer ental portion with one or two u-shaped loops. 190 CALIFORNIA ACADEMY OF SCIENCES [Proc. 471m SER. Penisetal follicles conspicuously protuberant into coelomic cavities and reaching to the middle of the prostatic duct or (an early aclitellate worm) to a level three-fourths of the way up the duct. Spermatheeae large, in contact with the ventral parietes back to the sep- tum and then up on its anterior face, the appearance of a slight curvature presumably due to a greater elongation of the anterior wall of the duct. Ova- ries fairly large, fan-shaped, with numerous short eg@ strings. Mature ova distinguished, in present condition of these specimens, by an obvious opacity that is lacking elsewhere in the gonad. GENITAL POLYMORPHISM. Three specimens are of athecal morphs, even rudiments of spermathecae lacking as was demonstrated by removing the longitudinal musculature from the interior of the body wall. Each specimen is of a different morph. One worm (no. 33) has a small ovary but no female funnel on the right side of xiv and a testis but no male funnel on right side of xii. Differences between the other specimens are in genital markings and penial setae. The markings of no. 31 are at 9/10-10/11 and 14/15-20/21. Ventral follicles of xviii are only slightly larger than in adjacent segments. Shape and ornamentation of the setae are unknown as only middle frag- ments of setal shafts were found. (Hach fragment was enclosed in a sleeve of euticle thit extended for some distance beyond the jagged edges, the setae possibly broken as the follicles were pulled out of the parietes.) Genital markings of no. 32 are at 9/1U-12/13, 14/15-16/17, 19/20-20/21. Penial setae are aS usual in sexual specimens. First order intermediate morphs (cf. Gates, 1956, for terminology) are represented in the present lots by tive individuals. One of these (no. 34) has four very small spermathecae that protrude into the coelomic cavities just enough to permit distinguishing the duet and ampulla. The spermathecal duct is slender and shorter than the ampullary rudiment. Penial setae of this worm are as usual. Ovaries and prostates are mature though a clitellum is as yet unrecognizable. A more advaueed stage of evolution was shown by six of the seven worms previously examined (Gates, 1941, p. 449) in which ental ends of rudiments were just recognizable at the parietes. No indications of the differentiation of the duct and ampullary portions were recognized. In such evolutionary lines all spermathecae appear to have been simultaneously and equally affected. Whether further growth had been inhibited at an early stage of development or whether initial invagination had been too long de- layed remains to be learned. Two specimens lacked one of the posterior spermathecae, that on the left side. No rudiments were present at the sites of the missing organs. A quad- ritheeal individual, in which the right posterior spermatheca is shorter as well as slenderer than the others and without differentiation into duct and ampulla, may provide a somewhat earl’er evolutionary stage. VoL. XXXI1] GATES: EARTHWORMS OF EISEN’S COLLECTION 191] Three worms are bitheeal. One of the antero-bitheeal (spermathecal pores at 7/8 only) individuals is juvenile. Spermatheeae (of no. 36) are rudimentary and only slightly protuberant into the coelom. The prostates are fairly large but obviously are juvenile as are the testes, ovaries, and gono- dueal funnels. Genital markings are not distinguishable (anlage possibly not visible because of poor condition ?). The penisetal follicles, however, have attained adult size and the penial setae have the usual shape, size and orna- mentation. The other antero-bitheeal worm (no. 29) is adult but follicles of the ventral setae in xviii protrude only slightly into the coelom. The setae are penial but shorter and slenderer than usual and with somewhat coarser ornamentation. The postero-bitheeal worm (no. 30) appears to be normal except for the absence of the anterior spermathecae. Four worms are pseudo-intermediate morphs, so-called because they are not evolving in the direction of any of the major standard morphs such as A, R, and AR. Three worms (nos. 14, 24, 28) have an extra spermatheca on the right side of vii. The other (no. 25) has the extra organ on the left side. Each extra-spermatheca is normal (as are the other structures) except in no. 14 where the ampulla is only about one-quarter the usual size. Genital markings are located as follows: at 8/9-11/12, 14/15-15/16, 19/20-20/21 (no. 14), 9/10-12/13, 14/15-15/16, 19/20-20/21 (no. 24), 8/9-12/13, 14/15- 16/17, 19/20-21/22 (no. 25), 9/10-12/138, 14/15-16/17, 19/20 (no. 28). Addition of extra organs to a normal complement is much less common than deletion in genital polymorphism. The number of genital markings in these morphs of P. papillifer averages somewhat higher than in normally quadrithecal worms. Certainly there is little indication of deletion of mark- ings, and in some lines markings may have been added. No indications of any trend toward elimination of prostates or of preco- cious abortion of testes were detected. Aside then from reduction and elimi- nation of spermatheeae, reduction (and possibly elimination) of penial setae, the evolutionary trends that were noted are toward inerease in number of spermathecae and of gonads. The extra gonads represent a return to an ancient ancestral condition, but addition to the spermatheeal battery ap- pears to be a change never before made in the ancestry of P. papillifer. The ovaries seem to have matured precociously in an aclitellate specimen and may have done so in several clitellate individuals. The testes in some eli- tellate worms are very large, bushy, each of the numerous long digitiform lobes showing only the same early stage of spermatogenesis. Perhaps some modification in a system of endocrine control had delayed the initiation of male gametogenesis until it was too late for the sperm to be matured prior to breeding. ABNORMALITY. All organs of the left side from the first spermatheea back to the prostates are two segments behind their normal locations. Displace- 192 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ments are due to interealations of two metameres by halving of two meso- blastie somites at levels in front of the eighth (cf. Gates, 1960). Metamerism is abnormal in the region in front of what normally would have been 9/10. Genital markings are at 9/10-12/138, 15/16-16/17, 20/21-21/22. REPRODUCTION. Iridescence on the male funnels of the Tulare specimen proves that sperm had been matured. Ividescence within the seminal echam- bers in the spermatheecal ducts shows that copulation had been completed even though a elitellum is lacking. Reproduction in this specimen presum- ably could have been biparental. Maturation of sperm, even profusely, does not, however, guarantee that reproduction is biparental (cf. Gates, 1957). Even after copulation reproduction may be parthenogenetic. Spermatozoal iridescence was lacking on the male funnels and in the sper- mathecae of 29 specimens regardless of the condition of clitellum and ovaries. The breeding season for P. papillifer is unknown, and even if it were, collec- tion dates are lacking for most of these worms. Mere absence of spermatozoa, in these circumstances, searcely provides trustworthy evidence as to method of reproduction in most quadritheeal individuals. Evolution of genital polymorphism obviously is well advaneed in P. papil- lifer. This sort of polymorphism in earthworms appears, so far as now known, only when reproduction has become parthenogenetic. Individuals of athecal and of some of the intermediate morphs must reproduce partheno- genetically. Male sterility is anticipated in some of the quadrithecal worms with normal anatomy. Remarks. Most of the specimens probably had been scoured, then anes- thetized and preserved in a relaxed state. All now are more or less softened. The worms from which the anterior ends had broken off probably had been suddenly killed and preserved in such strong alcohol as to make them brittle. Floeculent masses of corpuscles filled the coelomic cavities of most seg- ments in the anesthetized worms. These masses may mask structures such as nephridia so that preservation is not as good as is needed for some studies. Presence of quite some amounts of the flocculent material in the coelom of v— vi was, in the present case, advantageous as it enabled easy recognition of the delicate septum 5/6 and its insertion on the gut just behind the gizzard. The septum is, however, attached at several points to the anterior margin of the gizzard which might, in less fortunate conditions, have been thought to be in vi (cf. notes on several species of Ramiellona below). Except as indicated to the contrary above, the external characteristics and internal anatomy of each worm are as in previous specimens (Gates, 1941). VoL. XXXII] GATES: EARTHWORMS OF EISEN’S COLLECTION 193 Hearts, of course, never were found in xiv. Possibly much distended an- terior portions of the posterior latero-parietal trunks rising from the body wall of xiv were mistaken by Eisen for hearts. Relationships with P. marmoratus remain to be worked out. Plutellus marmoratus (MWisen, 1893). Hisen Collection, 0-0-2. (No further data. ) EXTERNAL CHARACTERISTICS. Length, 80-83 mm. Diameter, 4 mm. Seg- ments, 140, 167. Setae, AA = or < BC, DD ea. = 14C anterior to clitellum, posteriorly AA > BC or CD, DD <14C. Dorsal pores unrecognizable. Ch- tellum on xili-xix/2 or xix. Females pores at A, midway between 13/14 and eq/Xiv, within a transversely elliptical presetal tumescence in BB. Male fields distinetly delimited, reaching to presetal secondary furrow of xix and the postsetal of xvil, median margins especially tumescent. Genital markings small and not quite as wide as AB, cireular, each with a single greyish translucent central area, centered about at A, paired, at 9/10, 15/16 (1 specimen only), 16/17, 19/20. INTERNAL ANATOMY. Gizzard probably in v (2 specimens) but 5/6 very delicate, possibly incomplete and apparently attached (but not inserted) or perhaps only adherent at several points to anterior marein of the eizzard. Gut valvular in xvi-xvii, and there scarcely if at all thicker than the dis- tended dorsal blood vessel, gradually widening in xvilii-xx, normal intesti- nal width attained only in xxi. Typhlosole rudimentary in xix—xxiii, fairly high and simply lamelliform from region of xxiv—xxvi posteriorly, ending abruptly in 110th of 140 and 127th of 167 segments. Last hearts in xiii (2). Nephridia avesiculate, ducts pass into parietes at C, D or, oceasionally, dorsal to D by a distance about equal to CD. Spermatheeae fairly large, erect in coelomic cavities and reaching up to dorsal parietes. Duet much shorter than ampulla, wider entally and there with a considerable protuberance (almost reniform) from anterior and me- dian face or from anterior half and in which a number of small seminal chambers are visible. REPRODUCTION. Iridescence on the male funnels of each specimen shows that sperm had been matured. Seminal chambers of two spermathecae of one specimen and all chambers of the other specimen contain sperm. Sperma- thecal ampullae are filled with a loose coagulum. As sperm are matured and then exchanged during copulation reproduction is assumed to be biparental. The worms may have been collected toward the end of breeding period. REMARKS. The specimens were macerated and fell apart during the dis- section. The coelomic cavities of x—xi were filled with a compact coagulum 194 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SEk. that came away with difficulty from the male funnels. The tips of penial setae are lacking or softened and deformed. Exeept as indicated to the contrary above these worms are like P. papil- lifer. Family OCTOCHAETIDAE This family now comprises the Octochaetinae of the classical system (cf. Gates, 1959), the meronephrie genera of the Diplocardiinae, and the mero- nephrie genera having tubular prostates of ectodermal origin that were pre- viously in the Megascolecinae. The octochaetids of Mexico and Central America have been referred, in the past, to seven classical genera: Hodrilus, Acanthodrilus, Trigaster, Di- chogaster, Howascolex, Ranuellona, and Ramiella. Consideration of Trigaster and Dichogaster is deferred to a subsequent oceasion. All of the other octo- chaetids belong in Ramiella. The necessity for placing a Central American endemie (Gates, 1957) in an Indian genus provided yet another demonstra- tion of the artificiality inherent in a system based primarily on esoteric phylogenies. Hisen’s collection now provides some reason for believing that further information with regard to taxonomically important somatie characters of American, as well as of Indian species, will enable at least two American genera to be morphologically distinguished from Ramiella. Although the American taxa cannot be so defined at present, resurrection of the first avail- able name may obviate some nomenclatural changes in the future. Howascolex by definition is “purely meganephridial” in an anterior por- tion of the body. Meganephridial, in the classical system, implied that exere- tory organs are two per segment only and also that each has a preseptal fun- nel as well as a coiled postseptal body with an epidermal nephropore in the same metamere as the loops. This is the condition that characterized the Eudrilidae, Glossoseolecidae, Lumbricidae, Ocnerodrilinae, Acanthodrilinae, and some of the Megascolecinae which were all defined (Stephenson, 1930) as meganephridial. Other excretory systems have several pairs to several hundred micronephridia per segment. In more highly evolved meronephrie systems, some micronephridia were found to be as large as, or even larger, than meeganephridia. To indicate whether a large excretory tubule was or was not one of several meronephridia in a segment, some qualification was needed, and the primitive condition (one pair per segment) became known as purely meganephridial. American species, so far as can be determined, are meronephrie and without holonephridia (as they should be called) in any region of the body. Accordingly, these species are exeluded by definition from /Towascolex in whieh there are “true meganephridia” (Stephenson, 1930 :843). VoL. XXXT] GATES: EARTHWORMS OF EISEN’S COLLECTION 195 Genus Ramiellona Michaelsen, 1935 The word Ramiellona, a modification of Ramiella which is based on a Hindu patronymie (Ram), by itself hints at the difficulties encountered while attempting to define, in the classical manner, an American taxon so as to distinguish it from an unrelated group of species endemic in the Indian peninsula. Similarities, expressed in the usual classical manner, actually are as follows: lumbricin setae, acanthodrilin male terminala, one esophageal gizzard in a single segment, micronephridial. Ramiella lacks caleiferous elands which means only that no pouches are constricted off from the gut. Ramiellona accordingly was distinguished by the presence of calciferous folds within esophageal widenings in xii and some of the preceding segments. Mention of esophageal widenings or of foldings of the inner wall cannot be expected in older descriptions as even highly evolved calciferous glands were derogated taxonomically. However, one Indian species, Ramiella nainana Gates, 1945, does have in vili-xii (cf. descriptions below) folds between which ealeareous granules were found. For the present. then, the two genera are distinguishable only by their geographical distributions. The Guatemala specimens probably were quickly killed in the field and preserved in a strongly contracted state. Most are folded, twisted, or coiled. During their stay in alcohol their tissues were browned with the resultant loss of optical differentiation. Soaking in a solution of picrie acid did im- prove matters somewhat, but not enough in several instances to permit char- acterization of the genital markings. The softening, usually associated with long stay of earthworms in any of the standard preservatives, obviates cer- tainty as to the number of intestinal caeca and sometimes even as to presence or absence of these structures. The nephridia, of course, are in poor condi- tion and microscopic structure was not determinable. Portions of the esopha- eus almost certainly are deformed in some individuals as a result of strong contraction. Extremes of deformation probably are recognizable, but states that seem more normal may not be recognizable after slight relaxation. Even before deterioration has set in, field preservation may not be good enough to permit detection of taxonomically important characters in the excretory system. Ramiellona guatemalana Gates, new species. Guatemala, vicinity of Totenicapan, highlands of Huehuetenango, May- Nov. 1902. 15-19-38. G. Eisen. EXTERNAL CHARACTERISTICS. Length 140-190 mm. Diameter 5 mm. Seg- ments, 287, 294, 320. Prostomium, prolobous (30 specimens), recognizable from dorsal side (3), visible in buceal cavity from anterior end (27), so deeply retracted as to be invisible from exterior (7). Peristomium soft, with 196 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. many longitudinal creases, some part (except in 3 specimens) withdrawn into the anterior end. Two furrows on the peristomium, next to mD, some- times appear to be deeper than the others and then may look like margins of a tanvlobous tongue. A deep transverse furrow, however, always demareates the prostomium from the real anterior margin of i. Pigmentation unrecog- nizable (aleoholie preservation). A postsetal secondary furrow is present from iv posteriorly, a presetal secondary from vill. These secondaries in the preclitellar region, may be as deep as the interseemental furrows so that ree- ognition of the latter becomes difficult. Tertiary furrows behind the clitel- lum, sheht, often apparently incomplete but searcely distinguishable from the secondaries. Toward the hind end the segments are very short and with- out secondary annulation. Nephropores unrecognizable and doubtless micro- scopic. First dorsal pore\at 12/138 (22), 212/138 (2), 218/14 (5), unrecog- nizable until behind elitellum (2 specimens). Setae lacking in 1-111 (20 specimens), also absent in iv (7), a and b alone present on right side of iv (3), ¢ and d often lacking on v or v—vi, at first very small and very closely paired. In front of the clitellum, AB= CD, AA < BC, DD ca.= AC, near the 37th segment CD becomes wider than AB, the lateral ranks very gradually becoming more irregular but only in region of the 150th segment beginning to get well into the dorsum. The size of the setae as well as the protuberance from the epidermis inereases pos- teriorly and as the quincuncial arrangement is attained, but decreases again in the last 4-6 setigerous segments. The enlarged posterior setae are rather spindle-shaped, their tips ornamented with about 15 rather widely and ir- regularly interrupted cireles of very fine serrations. Ventral setae of xvil and xix penial, usually markedly protuberant. Ventral setae of xviii present (37 specimens), seemingly displaced slightly toward mV, ornamented dis- tally with short transverse rows of fine serrations at 15-20 circumferential levels, a shorter terminal portion curved slightly to one side. Clitellum saddle-shaped, reaching down nearly to B, on xiv—xix. Spermathecal pores transversely slit-like when closed, nearly circular when open and then filled with a plug of hard coagulum or revealing inter- nally smooth and elistening anterior as well as posterior lips in contaet with each other, in AB at 7/8-8/9. Female pores minute, in setal annulus of xiv, just in front of or slightly anteromedian or anterolateral to a. Male pores somewhat larger, transverse slits, in seminal grooves and just at 17/18. Prostatic pores immediately lateral to apertures of penisetal follicles, usually unrecognizable except when traction on adjacent epidermis separates the margins, about at equators of xvii and xix. The penial setae at first appear to emerge from a single aperture on each side of xvii and xix, but after they have been pulled out, a dividing membrane can be seen to reach down almost to the level of the external surface of the epidermis. Seminal grooves with VoL. XXX] GATES: EARTHWORMS OF EISEN’S COLLECTION 197 tumescent margins, well lateral to B, curve mesially in xvii and xix to the prostatic pores. An equatorial portion of xvii and xix, about at AB on each side, seems to be somewhat tumescent and is protuberant in a rather conical manner, follicle apertures at the apex. Seminal grooves and genital mark- ings provide boundaries for a median male field that usually is slightly depressed. Genital markings unpaired and median, transversely placed, the anterior- most often about reaching mBC, size decreasing posteriorly, along interseg- mental furrows as follows: 16/17 (15 specimens), 19/20 (11), 20/21 (12), 21/22 (4). Posterior markings perhaps develop later than the one at 16/17 as they are lacking on the younger specimens. Hach marking probably has one transverse row of circular areas. The equatorial annulus of xx and xxi (4 specimens), between the genital markings, is conspicuously protuberant, apparently somewhat tumescent and with a distinetive white opacity that is especially obvious because of the aleoholic browning in adjacent areas. INTERNAL ANATOMY. Septa 5/6-9/10 large, funnel-shaped, posteriorly directed, 5/6 muscular, 6/7-9/10 thickly museular, 10/11 shehtly museular, 11/12 obviously thicker, 10/11-11/12 united peripherally except ventrally in a middle portion of CC and organs of xi unrecognizable in a dissection from the dorsal side until after the apparently very thick septum 10/11 has been separated into its constituent parts. A special longitudinal muscle band at mD is present from 11/12 or 12/13 and is very distinet. Large and strong muscle bands from the posterior margin of the gizzard are inserted in the parietes of xi or xii. Diagonal muscle bands, with median parietal insertions near nerve cord, are present in xvi-xix; two in xvi on each side that are wide provide a useful marker for the segment, sometimes only one is present on each side but then it is much wider, covering the parietes from 15/16 to 16/17. A transverse muscle band from the gut in xiii passes, on each side, straight laterally to a parietal insertion. Gizzard large and strong, with a thickened cuticular lining, in v (8 speci- mens), but because of the size of the septal funnels about at the level of vii— ix as indicated by intersegmental furrows externally. Postgizzard portion of esophagus narrow, high up in the coelomic eavities, little if any longer than the gizzard, or at most shorter than the section in i-v. Esophagus deeply constricted by 7/8-12/13, apparently with five pairs of sacs, the first two or three pairs gorged with blood so as to appear black, the last two pairs always white, the first two pairs apparently opening widely into the esophagus, the next three pairs more nearly constricted off, but without stalks and opening through smaller circular apertures. Caleiferous lamellae numerous, thin, high, with free margins centrally, in vili-xii and possibly reaching very slightly into vii. Lower and rounded longitudinal ridges are present on the inner wall of the esophagus in xili-xiv. Intestinal origin in region of inser- 198 CALIFORNIA ACADEMY OF SCIENCES [Proc. 471 Ser. tion of septum 14/15, apparently just in front of the insertion (4 specimens) or just behind (4)—septa 14/15-15/16 often in contact mesially and ad- herent to each other, an anterior portion of the intestine with insertions of one or both septa sometimes drawn back into the interior of the gut, the adja- eent portion of the intestine then bulged forward. Intestinal eaeca small, paired, arising just lateral to the level of the secondary typhlosole (which is indicated externally by a deep red band) and immediately behind the septa, in Rxxili—xxvi (1 specimen), xxiv—xxvi (1), xxiv—xxvii (1), xxv—xxvi (1), XXV—Xxvill (1), (xxiv—xxv?) xxvi-xxx (1) xxv—xxx (1), unrecognizable (1). Typhlosole rudimentary from xvi to region of xxli—xxiv, thence posteriorly fairly large, thicker distally and with three longitudinal ridges on its ventral face, ending abruptly in 162d of 287, 176th of 294, 168th of 320 seements. Lateral typhlosoles low but simply lamelliform, beginning and ending abruptly, uninterrupted, just median to the caeeal apertures, from the re- ei0n of Xxli-xxiy to Xxx—xxxii. Supra-intestinal glands lacking. Dorsal blood vessel single throughout, complete, bifureating just be- hind or under the brain, the branches passing ventrally alone the cireum- pharyngeal nervous connectives to unite and become the ventral trunk (2 specimens). Ventral trunk complete, in front of 4/5 joined by three pairs of blood-filled vessels and then bifureating over the subpharyngeal ganglia (3 specimens, in one of which the branches are traceable only a short way up the nervous connectives). Extra-esophageal trunks recognizable from 11, large and filled with blood anteriorly, median to segmental commissures, empty and not traceable posteriorly. Supra-esophageal trunk bound closely to gut In vili—xil, with a large branch to each ealeiferous gland of vili—xi, bifureating anteriorly in xii. Commissures of vi-x lateral, traced to the ven- tral trunk except in vi, each joined just before opening into the ventral trunk by one or two vessels as large as itself. Commissures of v lacking or unrecognized. Hearts of xi—xii latero-esophageal, anterior bifurcation of a heart of xii continuous with posterior bifurcation of the supra-esophageal. Subneural trunk not found (7 specimens) and probably lacking. Latero- parietal trunks not found. Exeretory system meronephric. A longitudinal band of closely crowded micronephridia is somewhat lateral to mV on each side in i1i—iv. Two duets emerge together from the anterior margin of the cluster, but separate almost at once and disappear from view on parietes of ii and ii. Smaller clusters of nephridia are present on the anterior faces of 4/5-11/12 or 12/13. One elus- ter usually is recognizable way up on the septum alongside the gut. Another cluster joins the duct which runs downward on the septum, near the ventral parietes. Nephridia are at the parietes from one of segments xii-xv but at first the loops are vertical against anterior faces of the septa, from xx pos- teriorly the tubules are arranged in a transverse row at the middle of each VoL. XXX] GATES: EARTHWORMS OF EHISEN’S COLLECTION 199 segment. At least five longitudinal ranks appear to be present on each side of the body (crowding and poor condition of the organs obviating more pre- cise characterization). The median nephridium on each side in a posttyphlo- solar region of the body is no larger than some others in the same seement, but it does have a preseptal funnel. Metandric (7 specimens), testes of xi bushy, male funnels large and pli- cate, the testicular chamber (formed by peripheral union of 10/11—11/12) containing more or less coagulum but no seminal vesicles. Male funnels of x rather small but plicate (no testes in x). Seminal vesicles finely acinous, one pair, filling the coelomic cavity of xii, at maximal development bulging pockets of 12/13 back into xiii or xiv or (1 specimen) rupturing 12/13 and penetrating into xiii where a number of the lobes are separated off (owing to maceration?). Male gonoducts without epididymis but sometimes shortly looped in xili-xvi, with thick wall (presumably muscular but without sheen), slightly narrower in xvii and there lateral to prostatic ducts, passing down into the parietes in front of a@ of xviii. Slender ducts from male funnels of x are traceable (1 specimen) back to junctions with other ducts in xii. Pros- tates tubular, coiled in xvii and xix, distending septa or reaching into xix and xx—xxul, ducts with muscular sheen and 2-3 mm. long. A thick muscular sae containing follicles of functional penial setae projects conspicuously into the coelomic cavities just median to each prostatic duet. Reserve penial setae often occur in a membranous sae that passes back from the apex of the mus- cular sac. Penial setae red or with some greenish color in an ectal portion of the largest shafts, thick, 2-4 mm. long. Shaft nearly straight or in a slight are like a parenthesis. Distally the shaft narrows abruptly or gradually, the tip band-like, curved over to one side and at low magnification often seem- ingly hook-shaped. The margins of the band usually are slightly curved toward each other, but when the narrowing is abrupt the short tip may have a shape more or less like that of the bowl of a spoon. Ornamentation is of 30 or more circles of fine serrations, irregularly and frequently interrupted. The teeth, on the tip, are fewer, coarser, and with pointed distal ends well away from the shaft. Spermathecae fairly large, to 5 mm. long, reaching up to the level of gut on the anterior faces of the septa, adiverticulate and without external demar- cation into duct and ampulla. The duct is the anterior vertical portion pass- ing straight down through the body wall and is thicker than the ampulla. The narrow lumen of the coelomie portion of the duct curves over posteri- orly. The long ampulla gradually narrows entally. The rather thickish wall is provided with numerous closely crowded annular ridges. The seminal chambers, 6-8, are in the thicker anterior wall of the duet. Ovaries fan- shaped, with several short egg strings, usually hidden against ventral pari- etes by muscle bands. Ovidueal funnels slightly folded, readily recognizable. 200 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rir Ser. ReEpRoDuCTION. Iridescence on the male funnels of three aclitellate and one clitellate specimens is brilliant. Iridescence also is recognizable in the spermathecal seminal chambers of the aclitellate worms, but was not distin- oulshable in two spermatheeae of a clitellate specimen. Reproduction, inas- much as sperm are matured and then exchanged in copulation, is assumed to be biparental. GROWTH STAGES. Juvenile 2 mm. thick. The epidermis in AB at equators of xvii and xix appears to be shghtly tumescent. The tips of the red setae are just distinguishable in a central depression of the tumescenece. The testes (x1 only, none in x) are bushy but seminal vesicles had not as yet appeared. The spermathecae just protuberant into coelom, slender but already bent over toward the posterior septa. The ventral follicles of xvii and xix are protuberant into coelom just enough so that they can be grasped with the fine forceps. Each follicle contained three red setae, one a mere tip, a second about half the leneth of the longest which has a ribbon-like and flat tip. The male funnels are recognizable in x as well as in x1, but are much smaller in x. Larger juveniles possess more prominent tumescences in xvii and xix, but still lack seminal @rooves. More mature worms, with distinet seminal grooves, were listed as aclitel- late. One such has no genital markings and may not have been mature enough for them to be recognizable. Testes, with early stages of spermato- genesis, are unusually large in one early aclitellate with quite juvenile pros- tates, male funnels and seminal vesicles. Aclhitellate worms with sperm in the spermathecae showed no signs of a elitellum having regressed, but the seminal vesicles are dark. Epidermal evi- dences of elitellar regression could have been unrecognizable because ot aleo- holic browning, but contrariwise the dark appearance of the vesicles could be due to the browning by aleohol and so not evidence for postsexual re@ression. Autotomy. Three worms obviously are very recent posterior amputees. Two to seven circumferential constrictions, always at intersegmental levels, are recognizable, on several other specimens, in the terminal forty milli- meters of the body. At some of those constrictions the body wall already had been ruptured on one side or the other. The species obviously autotomizes readily, perhaps rapidly and sometimes extensively. The stimulus to the rup- turing, in the present instances, may have been provided by grasping the worm posteriorly to drop it into the preservative. Individuals of species that do autotomize readily never have done so, in the author’s experience, in killing aleohol regardless of the strength employed. Autotomy then can be avoided by seizing the worm near the anterior end. An anterior rupture through which spermatheeae protrude (aclitellate specimen) is likely to have been produced by the instrument used for digging VoL. XXXI] GATES: EARTHWORMS OF EISEN’S COLLECTION 201 (or perhaps ploughing) up the worms. Eight anterior fragments probably were broken off in that way. Incesta. Earth, reddish. No bits of plant matter were recognized under the binocular. ABNORMALITY. Spiral metamerism involving segments xi-xvi (1 speei- men). Parasites. Large, blood filled blisters of the intestinal wall in region of XViI-xxx contain two parasites each for which it is hoped identification can be obtained. Remarks. Parietal insertions of some of the anterior septa almost cer- tainly are displaced from their proper intersegmental levels. Two adjacent membranous septa sometimes adhere to each other so completely as to result in erroneous enumerations. Painstaking manipulation and repeated checks against fixed-position organs and against external annulation often were necessary. Even so, certainty as to the segment of intestinal origin was not achieved. Some coelomic cavities were completely filled by a ecoagulum. No elands were recognized in the body wall above the genital markings. Several characters, including retraction of the prostomium, location of the first dorsal pore, quineuncial arrangement of the setae, genital markings —identieally located and each with a transverse row of circular areas, the trifid typhlosole, the ten ranks of nephridia in a postelitellar portion of the body, suggest relationships with the Salvadorean R. lasiwrus. Differences, however, are numerous and as follows: prostomium prolobous (epilohous), elitellum saddle-shaped and on xiv—xix (annular on xiii—xx), muscularity of 5/6 (septum not mentioned), gizzard in v (vi), ealeiferous saes in vili—xil (ix—xiii), segmental commissures and hearts in vi-xil (vii—x1), presence of a testicular chamber (not mentioned), metandrie (holandrie ? with seminal vesicles in xi?), spermathecae adiverticulate (with a sessile, rosette-shaped diverticulum). Guatemalan worms also appear to be slightly larger and to have more segments. Some of these supposed differences may be of little or no taxonomic significance (cf. R. lasiurus below) and the more important ones now appear to be those relating to the clitellum and to the spermatheeal diverticulum. To facilitate further discussion of relationships in this difficult group there are subjoined certain notes on those species of which no material has been available for study. Each of these forms, with one exception, is known only from the original specimens secured at a single locality. 202 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH Ser. Ramiellona balantina Gates, new species. Guatemala, vicinity of Totenicapan, highlands of Huehuetenango, May- Nov. 1902, 0-3-0. G. Eisen. EXTERNAL CHARACTERISTICS. Length, 116 mm. Diameter, 6 mm. (largest specimen). Segments 111 (old posterior amputee), 197, 203. Pigmentation not determinable (alcoholic preservation and considerable browning). Pro- stomium prolobous (2 specimens), withdrawn but visible from the anterior end or unrecognizable (third specimen). A postsetal secondary furrow is present from vi or vil posteriorly, a presetal secondary from vill or ix. See- ondary furrows behind the clitellum, slight and searcely distinguishable from the tertiaries. Setae possibly lacking in 11 as well as ili (part only ?), a and b of xvii-xvili present, @ and b of xix penial, very small and very closely paired in preclitellar region, AB ca.—=CD, AA < BC, DD ca.=Y,C throughout much of the body, toward the posterior end AB and CD some- what wider but only slightly smaller than AA, the d rank becoming some- what above mL, all more protuberant and probably somewhat larger. Setae in the region of exx ornamented with irregular interrupted circles of fine ser- rations, a short portion of the tip curved to one side so as to have a hook- shape. Nephropores unrecognizable and apparently microscopic. First dor- sal pore, at 12/13 (2), 2138/14 (1). Spermatheeal pores, fairly large, centered about at A, one pair, at 7/8 (3 specimens). Female pores minute, just in front of a and hence in equatorial annulus of xiv (1). Male pores, somewhat larger, in seminal grooves, lateral to B at 18/19 (3). Prostatie pores at ends of the seminal grooves, just lateral to apertures of penisetal follicles, at eq/xix. The penial setae at first appear to project from a common aperture but after they have been drawn out a vertical membrane can be seen to reach down almost to the level of the external surface of the epidermis. Seminal grooves, lacking in the smallest specimen, pass from the prostatic pore anterolaterally to 18/19 (larger worm) or to 17/18 (largest specimen). Ventrum of xix to mBC markedly tumeseent (3), reaching greatest elevation at A where there is a half-collar of special protuberance that is open laterally just beyond the penial setae. Genital markings unpaired, transversely placed, recognizable only on the largest specimen and there at 15/16-17/18, 19/20-21/22. Each marking probably has one transverse row of circular areas. INTERNAL ANATOMY. Septa 5/6—-9/10 large, funnel-shaped, posteriorly directed, 5/6 muscular, 6/7—9/10 thickly muscular, 10/11 museular, 11/12 thickly muscular, 10/11 and 11/12 united peripherally except ventrally in region of BB so that organs of xi are unrecognizable in a dissection from the dorsal side until after an apparently very thick septum 10/11 has been sepa- rated into its constituent parts. Spee al longitudinal muscle band at mD VoL. XXX] GATES: HARTHWORMS OF EISEN’S COLLECTION 203 quite distinet. Large muscle bands from the posterior margin of the gizzard pass back to parietes in region of x—xii. A strong band from the lateral face of gut, on each side of xiii, has a parietal insertion laterally in xiv. Diagonal parietal bands, strong and wide, have median insertions near the nerve cord in xviii, but similar bands are lacking in xvi-xvil. Gizzard large and strone, with thickened cuticular lining, in v (3 speei- mens) but about at level of vili-ix as indicated by intersegmental furrows externally, so elongated in one worm that its anterior opening is dorsal and its posterior aperture is ventral. Posteizzard portion of esophagus narrow, high up in eoelomie cavities, little if any longer than the gizzard, deeply con- stricted at insertions of 7/8-12/13, the portions in vili-ix thin dises of nearly circular outline between compressing septa, a somewhat larger but equally thin dise in x obviously bilobed. Caleiferous lamellae with free margins cen- trally, in vili—xii, of xi—xii within nearly spheroidal unstalked saes all four of whieh at first appeared to be in xii as separating tissues are very thin. Intestinal origin in the region of insertion of 14/15, seemingly (3 specimens) just in front of 14/15. Intestinal caeca small, recognizable only with diffi- eulty, in xxiv—-xxy (3), possibly a smaller caecum in xxvii (1). Tyvphlosole rudimentary but lamelliform in xvi to region of xxii, thence posteriorly with widened ventral portion bearing three longitudinally lamelliform ridges, ending abruptly in 123d of 197 segments but in the amputee becoming rudi- mentary in the 94th segment and unrecognizable behind the 108th. Lateral typhlosoles low but lamelliform, uninterrupted, in xxili-xxvill. Supra-intes- tinal glands lacking. Dorsal blood vessel single throughout. Extra-esophageal trunks empty and unrecognizable except anteriorly. Supra-esophageal trunk distinguish- able only in xi-xii. Subneural trunk not found (2 specimens) and probably lacking. Lateroparietal trunks unrecognizable. Segmental commissures of vi-x lateral, traceable to ventral trunk only in x. Hearts in xi-xii, latero- esophageal. Exeretory system meronephrie. A longitudinal band of closely crowded micronephridia parallels the nerve cord on each side of the body im ili—iv. Smaller clusters of micronephridia are present on the anterior faces of septa 4/5-11/12 or 12/13. Nephridia from xiii posteriorly, on the parietes, behind xx in at least four (possibly 5 or 6?) longitudinal ranks, the tubules trans- versely placed at the equator of each segment. The medianmost nephridium on each side, toward the hind end, has a preseptal funnel. Stomate nephridia of one specimen obviously are thicker than the astomate tubules of the same segment. Metandric (3 specimens), testes (present only in xi) bushy, the testicular chamber with little or no coagulum but reaching forward (because of a ven- tral pocket of 10/11) on floor of x nearly to 9/10 (largest specimen). Male funnels all plicate, those of x smaller, those of xi (the largest specimen ) 204 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. within the anterior pocket of the testicular chamber. Seminal vesicles finely acinous, one pair in xii. Male gonoduets without epididymis, united on each side in front of 12/13, seemingly with a muscular sheath though the sheen is unrecognizable, passing down into the parietes in the region of 18/19. Pros- tates in younger specimens obviously juvenile and coiled in xix, probably adult in the largest worm where they reach into xx. Duets about 2 mm. long, museular, looped. Penisetal follicles within two thick, muscular saes econ- spicuously protuberant into coelomic cavity of xix. Membranous saes pass- ing back from the apex into xxiii, each containing a large and a small reserve seta as well, sometimes as a mere tip of a third. Penial setae (functional), 2 + mm. long, thick, sometimes widest at ectal end which sometimes appears to be rather bulbous and abruptly narrowed to a terminal spine. The latter, usually softened, reveals under high magnification the shape of a spoon bowl. Shafts of reserve setae lack the bulbous widening at the tip and may narrow eradually to a thin band. Ornamentation is of frequently and irregularly interrupted cireles of fine serrations, the teeth slightly larger and rising slightly away from the shaft on the narrowed tip. Spermatheeal duct vertical, thick, wider entally, adherent to the posterior face of 7/8, with a rather flat dorsal surface. Ampulla of about the same size as the duet, on the ventral parietes, ovoidal, narrowing to a very short but still fairly thick neck that passes to the posterior face of the duct. The wall of the ampulla is opaque (not membranous) and without obvious ridges. The lumen is wide distally and filled with ecoagulum, but in the neck it is very small as in the duct. Six sight white rounded protuberances on the dorsal face of the duct anteriorly mark the sites of distended seminal chambers. One or two additional chambers may be present down deeper so as to be un- recognizable externally. Ovaries fan-shaped. REPRODUCTION. Iridescence on the male funnels of the largest worm is brilliant. The seminal chambers, in one spermatheea, are filled with a mate- rial in which iridescence is readily recognizable. Although there is no clitel- lum, the worm must have copulated. Reproduction, as sperm are matured and exchanged in copulation, is assumed to be biparental. InGestTA. Earth, fine textured, dark, of a reddish brown color, without macroscopically recognizable plant fragments. PARASITES. Six nematodes, red except at each end, were found in the coelomic eavity of xii (1 worm). Several small white cysts are present on an anterior portion of the intestine. REMARKS. Protuberances from xix presumably ean be inserted into the large spermathecal pores to function as temporary intromittent organs. The short seminal grooves of the less mature worm are slight and may not have attained full development in depth as well as in length. VoL. XXXT] GATES: EARTHWORMS OF EISEN’S COLLECTION 205 An esophageal typhlosole at mV, as in the previous and the next species, was not recognized. Somatie anatomy indicates close relationships to species that are desig- nated hereinafter as the “quatemalana” group. The major difference from others of the group (and which probably would have been considered very important in the classical system) is that the genitalia have undergone the metandrie and the balantin reductions. The present species, accordingly provides one more demonstration that genital anatomy is liable to more rapid evolutionary modification than the somatic. The persistent male funnels of segment x show that advent of metandry was recent. Holandrie individuals or even a holandrie subspecies may be found. The balantin reduction of the ancestral acanthodrilin male terminalia is much rarer than the microseoleein and is as yet little known. The peristomium, with numerous longitudinal wrinkles, is flaccid, a con- dition that appears not to be wholly attributable to postmortem softening, especially in view of the firmness of the prostomium. Apparently under way in the “quatemalana” group is an evolutionary derogation of the first seg- ment which ean be expected to result, perhaps in some hitherto uncollected species, in disappearance of intersegmental furrow 14. With loss of the setae of the next metamere, 1 and 11 would be indistinguishable and all fixed- position organs would appear to be one level in front of their usual locations. Just such a change appears to have been completed in two megascolecid In- dian genera, Tonoscolexr Gates, 1933 and Nelloscolex Gates, 1939. A similar process is well under way in at least one species of the American @lossoscole- eid genus Pontoscoles Schmarda, 1861, in which 1/2 has disappeared but setae of 11 still are present. The prostomium in P. corethrurus (Miiller, 1856) seemingly has disappeared, but in Tonoscolex and Nelloscolex was retained. Retention of the prostomium now seems more likely to characterize more advanced stages of peristomial derogation in Central America. Male apertures in the supposedly ancestral, acanthodrilin genitalia of the classical system are at eq/xvill, midway between the anterior and posterior prostatic pores which are at eq/xvil and eq/xix. Although all four elands appear to be equally developed (and presumably synchronously) in the “quatemalana” group, the male apertures are further anteriorly, at 17/18 and hence closer to the openings of the first pair of glands. Disappearance of the anterior prostates supposedly involves union of male and posterior pros- tatic pores. That has not happened in R. balantina where the male openings do not even get into xix, but are at 18/19 and so are at the same distance from the remaining prostatic pores as they previously were from the an- terior pair. 206 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4ru SER. Ramiellona strigosa Gates, new species. Guatemala, vicinity of Totenicapan, highlands of Huehuetenango, May- Nov. 1902, 0-5-1. G. Eisen. EXTERNAL CHARACTERISTICS. Length, ca. 97 mm. Diameter 3-315 mm. Seements, 181, 203, 208. Prostomium prolobous, just recognizable from dor- sal side (2 specimens), visible only from anterior end (4). Peristomium soft, with numerous longitudinal creases, some part withdrawn into anterior end (6). A postsetal secondary furrow is present from v, a presetal secondary from vill or ix. Tertiary furrows behind the elitellum, sheht, often appar- ently incomplete and seareely distinguishable from the secondaries. Setae present from 11, small, closely paired, DD ea. = 14C throuzhout, 4B = CD, AA < BC anterior to elitellum, behind the elitellum AA > BC, toward pos- terior end AA and BC only a little larger than AB and CD, ventral setae of Xvi and xix penial, ventral setae of xvili present (6 specimens) and median to seminal grooves. Nephropores unrecognizable and doubtless microscopie. First dorsal pore at 12/13 (6), in the elitellate worm a functional pore pres- ent at 13/14. Chtellum dark red, intersegmental furrows obliterated, dorsal pores oe- cluded, setae unrecognizable, probably at maximum tumescence, saddle- shaped, reaching down to B except in xvii-xix and there only to mBC, on Xiv—x1x, extending halfway through xx and slightly into xiii but in each of those segments much thinner and not so dark. Spermatheeal pores with rather irregular margins, not minute, centered about at B, two pairs, at 7/8-8/9 (6 specimens). Female pores minute, in setal annulus of xiv, sightly anteromedian to or just anterior to a (3). Male pores in seminal grooves, apparently at 17/18 (3, but location could not be confirmed by tracing male gonoduct through the musculature). Seminal erooves well lateral to B, pass mesially in xvii and xix onto conspicuous pro- tuberanees at apices of which are prostatie pores and follicle apertures. Genital markings unpaired, transversely placed, each with a single trans- verse row of circular areas, at 10/11, 11/12 and 20/21 (2 specimens), 11/12 and 20/21 (1), in BB posteriorly, reaching into BC anteriorly, unrecogniz- able and presumably not vet developed (3 specimens). Areas of translueence and of about the same size, slightly depressed, may be present, on some speci- mens at 16/17, 18/19 and 19/20. INTERNAL ANATOMY. Septa 5/6-11/12 funnel-shaped and posteriorly di- rected, funnels of 5/6—-9/10 especially large, 5/6 membranous but complete, 6/7-9/10 or 10/11 rather thickly muscular, 11/12-12/18 slightly streneth- ened. Special longitudinal muscle band at mD very distinet. Diagonal muscle bands broad, with median parietal insertions near nerve cord, pres- ent in xvi. VoL. XXX] GATES: EARTHWORMS OF EISEN’S COLLECTION 207 Gizzard large and strong, about at the level of vili—x as indicated by inter- segmental furrows externally but actually in v (5 specimens). Postgizzard portion of esophagus slender, fairly high up in coelom, deeply constricted at the septal insertions. Caleiferous lamellae fairly high and with free margins centrally, in vii—xii. Small, lateral sacs seemingly are present in each of vii-— x, the sacs opening widely into esophageal lumen. Sae¢s in xi—xii more dis- tinctly constricted off, though unstalked, and protruding slightly above and slightly below the median portion of gut. Intestinal origin in the region of the insertion of 14/15, almost certainly just behind 14/15 (1 specimen), ap- parently so but possibly not (the other four). Intestinal eaeca indistinet, in XXHI-xxiv (3 specimens), apparently just in front of the septa, and in one worm recognizable only on the left side, in (xxiii?) xxiv—xxvili (1), xxiv— xxvill (1). Typhlosole rudimentary till region of xxii—xxiy, then abruptly enlarged, widened ventrally and there with three low but lamelliform and longitudinal ridges, with obvious vertical ridges on the lateral faces dorsally for a number of segments, ending abruptly in 110th of 181 or 111th of 203 segments, still recognizable at 98th segment of one of the anterior fragments. Lateral typhlosoles low but lamelliform, beginning and ending abruptly, in XX111-xxvill. Supra-intestinal glands lacking. Dorsal blood vessel single throughout, bifurerting just behind or just under the brain. Ventral trunk complete, bifureating just over the sub- pharyngeal gangha. Extra-esophageal trunks median to segmental commis- sures, filled with blood anteriorly but empty and unrecognizable nosteriorly. Supra-esophageal trunk visible only in xi-xii. Subneural trunk unreeoeniz- able and presumably lacking. Lateroparietal trunks unrecognizable. Seg- mental commissures of vi-x lateral, none found in vy. Hearts of xi—xii, latero- esophageal (5 specimens). Exeretory system meronephric. A longitudinal band of closely crowded micronephridia is somewhat lateral to mV on each side of the body in iii-iy. Smaller clusters of nephridia are present on the anterior faces of septa 4/5— 11/12 or 12/13. Nephridia are on the parietes from xiii posteriorly and be- hind the eclitellum the loops are transversely placed. Nephridia almost cer- tainly are in three longitudinal ranks on each side posteriorly, one rank about in BC, the other two further laterally. The medianmost tubule on each side in post-typhlosolar segments has a preseptal funnel. Holandric. Seminal vesicles finely acinous, paired in xi and xii, in one worm smaller vesicles but with typical lobulation also present in ix. Male eonoducts of a side without epididymis, united just in front of 12/13, thence back apparently provided with a muscular sheath, zigzagged or shortly looped in xiv—xvi, shghtly slenderer in xvii where they are lateral to the pros- tatie ducts, passing down into the parietes in the region of the insertion of 17/18 (but not traceable through the musculature). Prostates tubular, 208 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH Ser. coiled in xvii and xix, the posterior pair sometimes extending into xx or XXi. Ducts muscular, 1-2 mm. long. Penial setae to 144 mm. long, slenderer than in the other species, shaft curved in an are like the parentheses but somewhat asymmetrically, narrowing rather gradually toward ectal end to a thin and flat band with both margins curved over on coneave side of shaft. Ornamen- tation of two longitudinal ranks of transverse serrations, one rank on the upper side of the shaft as it hes naturally on the slide and the other on the lower side. The serrations comprise 2—7 teeth of variable size and shape, some of a more or less triangular shape, others thorn-like. The functional setae are yellow and with each there is associated one small reserve seta. Spermathecae with a main axis that is 2-3 mm. lone, shehtly and gradu- ally narrowed entally, usually posteriorly directed on the ventral parietes and then upward on the anterior faces of the septa. One spermatheca turns mesially and anteriorly to pass into the segment next in front. The duct is much shorter than the ampulla which has on its inner wall numerous closely erowded annular ridges. Diverticulum a large, nearly circular dorsoven- trally flattened dise on the parietes of the segment next in front of that con- taining the main axis and reaching well toward or even to the anterior sep- tum, with a number of small seminal chambers. Ovaries fan-shaped and with several eee strings. REPRODUCTION. Iridescence is recognizable on the male funnels but is slight on those of the elitellate worm. Iridescence also is recognizable in the seminal chambers of the spermatheeal diverticula of aclitellate specimens. Reproduction, inasmuch as sperm are matured and exchanged in copulation, is assumed to be biparental. Remarks. A posterior portion had been torn off from three worms. Some intestinal caeca may have become unrecognizable because of softening and aleoholic browning in three of the dissected specimens. The penial setae usu- ally were broken, without ectal portions or with tips softened, and obviously more or less deformed. Only one caleiferous sae is present in x of one specimen, no slightest trace of a sae is recognizable on the other side. Hearts of xi (1 specimen) and xii (another worm) at first appeared to be absent. The associated septum, in each case, was adherent to the gut in front of the hearts as if normally in- serted there. The male gonoducts of one worm, though in contact in xii, obvi- ously do not unite until well into xiii. As the spermatheeal pores are not minute they may, when fully open, be large enough to permit insertion of protuberances of the male field function- ing as temporary intromittent organs. A testicular chamber was not recognized, possibly because the peripheral union of the bounding septa became unrecognizable as the worms were opened and pinned out. VoL. XXXT] GATES: EARTHWORMS OF EISEN’S COLLECTION 209 These specimens appear to differ from the type of R. eisent (cf. below) as follows: Saddle-shape of clitellum (annular but thinner ventrally) that reaches to xx/2 (instead of only slightly into xix). Absence of genital mark- ings at 14/15, 15/16, 21/22-25/26, and possibly also presence of a transverse row of circular areas within each marking. Presence of calciferous saes, espe- cially in xi-xl. Presence of intestinal caeca. Presence of seminal vesicles in xi and oceasionally in ix. Orientation and location of the spermatheeal diver- ticulum, 7.e., flattened on the parietes of the preceding segment (not verti- eally placed on the anterior face of the duct and perhaps in the same segment as the main axis of the organ). Individual variation as to number of genital markings is, of course, to be expected, but such very little data as now are available suggest specific uniformity as to shape and size of the elitellum in this genus. Intestinal caeca should have been recognizable if present in the type of R. eiseni, as it was well preserved, but they may have been overlooked or even considered to be unworthy of mention. Possibly some deformation or a peculiar reaction to preservation rendered the ealciferous saes unrecog- nizable. (For other comments see notes on FR. eisenr below.) Somatie characters show that R. strigosa belongs in the “quatemalana”’ group to which R. eisent probably will have to be added. Possibly only early stages in development of the testicular chamber, or none at all, are to be found in a holandrie section of the group. Ramiellona mexicana Gates, new species. Mexico, Dos Rios, Tehuantepec, January, 1900, 0-1-1. P. M. King (Eisen collection ). EXTERNAL CHARACTERISTICS. Length, 96 mm. (aclitellate worm), 116 mm. (elitellate specimen which lacks a posterior portion presumably torn off be- fore preservation). Diameter, 7 mm. (clitellate worm). Segments, 253 (aeli- tellate), 195 + (elitellate). Pigmentation not determinable (aleoholie pres- ervation, specimens browned). First segment soft, longitudinally furrowed, almost wholly withdrawn into the interior but intersegmental furrow 1/2 ree- ognizable just behind the anterior end of body. Prostomium presumably to be considered prolobous, also withdrawn out of sight but firm (not soft lke peristomium ), much wider than long, with a deep groove marking off a cireu- lar area at center of which is a deep pit. Secondary furrows two per segment behind clitellum. Setae unrecognizable on ii and probably lacking, small, very hard to see even when tips are protuberant from the epidermis, ventral setae paired and in regular ranks but dorsal setae in irregular ranks in pre- clitellar segments, posteriorly all ranks irregular and arrangement becoming quineunxial. Apertures of ventral follicles of vili-ix about at same level as spermatheeal pores and hence only slightly behind intersegmental furrows, 210 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. not in line with @ and b ranks of other segments. Nephropores unrecogniz- able and apparently microscopic. First dorsal pore at 11/12 (2 specimens). Clitellum saddle-shaped, dark red, reaching down to B, on xiv-xxii. A median region between the ventral borders of the clitellum is slightly de- pressed. The same region, in xv—xx of the aclitellate worm, is deeply de- pressed and so that the floor is invisible. Spermatheecal pores minute, superficial, obviously behind 7/8 and 8/9, unpaired and median. Female, male, and prostatic pores unrecognizable, the female pores possibly at A. Seminal groves unusually fine, shaped lke pa- rentheses, between equators of xvii and xix, at B. Slight tumescences at each end of the grooves may contain prostatic pores. Genital tumescenees areas of sheht epidermal thickening, without distinet boundaries, paired in vii around apertures of ab follicles, unpaired and reaching beyond B in viii and ix where the posterior intersegmental furrows are irregularly met. A tumescence in each of segments xxvi-xxxii (clitellate worm) reaches laterally on both sides to B and anteroposteriorly to the inter- seemental furrows. INTERNAL ANATOMY. Septum 4/5 membranous, 5/6—-9/10 rather thickly muscular and funnel-shaped, 10/11—11/12 sliehtly strengthened. Septal in- sertions on the parietes do not correspond to the intersegmental furrows, 9/10-14/15 crowded together so that the coelomic cavities of ix—xiv are short. Gizzard large and strong, with thick cuticular lining, in v. Four wide and strong muscle bands from the posterior end of the gizzard dorsally pass back at least to 12/13. Other and more numerous bands from the septa to the parietes are shorter. Hsophagus deeply constricted at the septal insertions and markedly moniliform through ix—xiv, in each of those segments short (especially so in xiv) and, relative to width of body, narrow. A thick typhlo- solar ridge is present at mV. Thin, vertical, and presumably ealeiferous lamellae with free median and dorsal margins are attached to floor and lat- eral walls but leave shghtly roughened roof of the gut free. Intestinal origin seemingly in xvi (but possibly in xv?). Typhlosole very rudimentary or lack- ing until about xxx, thence fairly high, lamelliform but with slight vertical ridges on lateral faces, ending effectively in 132d segment (clitellate worm) though a very small and round rudiment is recognizable in the next twenty metameres. Caeca and lateral typhlosoles were not recognized and may be absent. Supra-intestinal glands lacking. Dorsal blood vessel single throughout, traceable anteriorly to the brain and presumably complete (though bifureations to ventral trunk were not recognized). Ventral trunk not visible in front of ix. Supra-esophageal trunk double in part and apparently not continuous, recognizable only in xii—xiii. ixtra-esophageal trunks median to segmental commissures, recognizable only in vi-vill. No subneural trunk. Latere-parietal trunks unrecognizable. Seg- VoL. XXX] GATES: EARTHWORMS OF EISEN’S COLLECTION 211 mental commissures of v—x lateral (both specimens), those of v anterior to the gizzard, those of the other segments on anterior faces of the septa, trace- able to the ventral trunk only in ix—x. Hearts of xi—xiii (both specimens) rather small but obviously latero-esophageal. [xeretory system meronephric. A large horseshoe-shaped cluster of mi- eronephridia is present on the anterior face of 4/5. A fairly thick duct, ree- ognized only on one side of one specimen, passes forward on the parietes into segment 11. Very small clusters of micronephridia are present on the parietes in il, v, and posteriorly. Nephridia of postelitellar segments seemingly in six longitudinal ranks on each side. The median-most tubule on each side, in posterior segments of the body, is much thicker than the others and is pro- vided with a slightly elistening preseptal funnel. THolandrie, testes manicate. Male funnels large in aclitellate worm but smaller in the other, plicate. Male gonoducts very slender, looped or zig- zagged just behind the funnel septa, not traceable after reaching the parietes. Seminal vesicles vertical bodies on posterior faces of septa 10/11 and 11/12, medium-sized or smaller, acinous, and with many lobules. Prostates two pairs, coiled in xvii and xix. Duets slender but with muscular sheen, ea. 4 mm. long. Lumen in glands slit-like in eross section, much smaller and eir- cular in the duets. Penial setae and enlarged setal follicles were not found in Xvli-x1x. Parietes covered, in xv—xx, by numerous diagonal muscle bands presumably responsible for depression or grooving of ventrum in the clitel- lar region. Spermathecae long enough to reach up above gut, rather slenderly elub- shaped, without external or internal indieation of demareation into ampulla and duet, very much narrowed in the parietes. The anterior spermatheca (both specimens) reaches forward into vi and has two diverticula. Two sper- mathecae are present in ix or anteriorly, each with its own diverticulum, the ducts (both specimens) united within the parietes. The diverticulum which passes to the anterior face of the duct at the parietes is short and rather digitiform. The axial lumen, located on the side of the diverticulum next to duet, opens into six discrete seminal chambers. Ovaries small, fan-shaped, with numerous short ege strings. The ventral follicles of vii-ix are very thick but protrude only slightly into the coelomic cavities. Setal shafts without nodulus, curving slightly to one side near ental end. An ectal portion of each seta (clitellate specimen) is lacking as is ornamentation on the remaining portions. REPRODUCTION. Spermatozoal irideseence on the male funnels is limited to a peripheral region central to which the funnels are very dark, not black but with a reddish tinge. The color does not appear to be due to blood. Ivi- descence is unrecognizable in the spermathecae. Nevertheless, there are in the ampullary coagulum small bundles of fine threads that may be spermatozoa. 212 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4711 Sper. Small, spindle-shaped to ovoidal bodies of a definite red color are scattered through the coagulum that fills the seminal chambers of the spermatheeal diverticula. If, as seems possible, these red bodies contain sperm, then a mas- sive aggregation may have been present on the male funnels. Reproduction, in absence of evidence to the contrary, is assumed to be biparental. INGESTA. Soil. Remarks. These worms had been crowded into a too-short container. The epidermis of xiv, because of the folding over of the specimen, had been ereased and then cracked so that the females pores could not be found. The eut of one worm may have been ruptured during study of external charae- teristics. The second specimen was dissected before examination of external characters, but the gut already was ruptured in the region of xxx. Some sort of genital markings probably are present in BB of the clitellar region, but even after soaking in picrie acid solution optical differentiation was too poor to permit any attempt at characterization. Clitellar tumescenece is unrecognizable externally on the worm ealled aeli- tellate though some thickening of the epidermis is noticeable at the mid- dorsal incision. Sperm obviously had been matured and the worm almost certainly had copulated. Accordingly, one more must be added to the leneth- ening list of specimens that have matured sperm and copulated though aclitellate. No indieations of existence of a testicular chamber were recognized, but absence cannot be asserted. Some of the somatic anatomy obviously is like that of the “quatemalana” eroup: Retraction of the prostomium and derogation of the peristomium associated with loss of setae at least in ii. Small size and close pairing (ven- tral ranks only) of setae anteriorly. Location of the first dorsal pore. Pres- ence of all septa from 4/5, enlargement into posteriorly directed funnels of 5/6-9/10, marked museularity in 6/7-9/10. Primitive location of gizzard, in v. Postgizzard portion of esophagus short and slender. Presence of eal- ciferous lamellae in ix—xii. Presence of an intestinal typhlosole. Hearts of x lateral, of xi-xii latero-esophageal. Absence of a subneural vessel. Differences from the “guatemalana” group are as follows: Anterior dis- location of the apertures of the ventral follicles of vili-ix. Development of tumescences (associated with intraparietal glands?) around the apertures of the ventral follicles in vii-ix, enlargement of those follicles (and development of copulatory setae?), abortion of ventral follicles in xvii—xix. More pos- terior intestinal origin. Presence of hearts in xiii (and of sezmental commis- sures in v?). Union of the spermatheeae midventrally. Minute size of sper- matheeal pores. Some of those differences have been found elsewhere within generic limits and now seem likely to be of minor taxonomic importance. VoL. XXXI] GATES: EARTHWORMS OF EISEN’S COLLECTION 213 Evolution of the calciferous portion of the esophagus and of the excretory system in the anterior segments has been proceeding in ways that were not involved in the ancestry of the “gquwatemalana” group. Common origin with that group now appears to have been sufficiently remote to require generic distinction when adequate characterization of structure is possible. Ramiellona eiseni (Michaelsen). Eodrilus eiseni MICHAELSEN, 1911, Zool. Jahrb. Syst. 30, 559. This species was erected on a single specimen from Huehuetenango. The importance attached to somatie organization in the classical svstem is shown by the characterizations of two systems: excretory organs meganephrie, dor- sal blood vessel single, last hearts in xii. The meganephry required the spe- cies to go in an Aecanthodriline genus but micronephridia were later found (Pickford, 1937) in a small piece (now in the U.S.N.M.) of the body wall from the type. Pickford’s transfer to Howascolex was qualified by a ‘“?” because of absence of calciferous saes. A widening of the gut in xii-xili was attributed by Michaelsen to flattulence, but might have been due to disten- tion by ingesta that was passing through at time of preservation. Poor pres- ervation may have been responsible for failure to detect calciferous lamellae (the gut of earthworms sometimes is in very poor condition even though peripheral anatomy seems to be well preserved. ) The species obviously is holandrie, presence of sperm on male funnels fortunately having been recorded. Absence of seminal vesicles in ix and/or x1 1S unusual as their disappearance would be expected to follow rather than to precede metandry. Spermathecal pores were said to be small but what that means is unknown as size usually was not mentioned. Location of male pores also was not stated. Except for absence of calciferous sacs, P. ersenr now seems to be close to R. strigosa. If that is correct, both species probably share with others of the group certain unrecorded characters. Ramiellona irpex (Michaelsen). Eodrilus irpex MICHAELSEN, 1911, Zool. Jahrb. Syst., 30:555. ? Acanthodrilus irpex MICHAELSEN, 1925, Mitt. Zoll. Mus. Hamburg, 41:76. Eodrilus irpex was erected on a single specimen from Huehuetenango that was provided (like the type of the previous species) by Eisen. Size of spermatheeal pores, location of male pores, intestinal caeca, typhlosole and even the andry were not mentioned in the deseription. The species may be metandrie but with retention of anterior male funnels as in R. tecumumami and R. guatemalana, or holandrie but with only one pair of seminal vesicles as in R. etsent. A supposedly meganephridial excretory svstem, as in R. 214 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. eisent, was found (Pickford, 1987) to be micronephridial, 7.e., meronephric¢ but nothing is known about the tubules of the preclitellar region of the body where generic peculiarities are likely to be more obvious. Caleiferous lamel- lae are present in xii but in vili—xi are within paired, dorsally directed and diserete glands with slender stalks that open into the gut close to mD. The spermathecae are much like those of R. balantina. Acanthodrilus wrpex was deseribed from a specimen from Mexico which is larger than the Guatemalan worm and has paired genital markings. Seeg- ment number is more than twice that of the type, but a eclitellum had not been developed. No information as to internal organization was recorded though the worm presumably was opened to enable generic identification. In view of the external differences conspecificity with the type requires confirmation. Ramiellona tecumumami (Michaelsen). EKodrilus tecum-umami MICHAELSEN, 1911, Zool. Jahrb. Syst., 30:550. This species was erected on specimens from Huehuetenango that were provided by Eisen. Size of spermatheeal pores, location of first dorsal pore and of male pores, intestinal origin, and caeca were not mentioned. In the usual classical manner, location of the last pair of hearts is all that was re- corded about the vascular system. As in ease of the two preceding species, a supposedly meganephridial excretory svstem was later found (cf. R. vul- canica below) to be micronephridial. Unfortunately, no further information as to the type or other specimens was vouchsafed. Omissions in the list of differences between R. tecumumami and R. vulcanicus warrant assumptions that spermathecal pores are large, that the male pores are in seminal erooves at level 17/18, that caleiferous lamellae are present and especially large in xi-xil, that intestinal caeca are present, that there are vascular commis- sures or hearts in vill-xil. Spermathecae appear to be erect in coelomic eavi- ties. Spermatheeal pores are median to A, more so at 8/9. A continuation of the process that apparently is under way ean be expected to result in median union of the paired organs, first at 8/9 rather than at 7/8 as in R. mexicana. Ramiellona stadelmanni Michaelsen. Ramiellona stadelmanni MIcHAELSEN, 1934, Mitt. Zool. Mus. Hamburg, 45:53. This species is known only from the original deseription of two internally macerated specimens from Honduras. Reproductive apertures (except the female pores) and seminal grooves were not seen. Septum 5/6, the intestinal VoL. XXX] GATES: EARTHWORMS OF EISENS COLLECTION 2A origin, and intestinal caeca were not mentioned. One part of the setal for- mula appears to be incorrect, in AA= 3 —- AB, BC presumably should be read for AB as setae are closely paired anteriorly. Whether the spermathecal pores are minute or merely unrecognized be- cause of strong contraction is unknown but no porophores or special pro- tuberances are shown in a figure of the male field. Presence of seminal vesicles in xi-xli usually is associated with holandry which is assumed in order to include this species in the key below. Relationships indicated by the genital markings, caleiferous portion of the gut, the large hearts of xi—xil, and other structure, in absence of information about insertion of 5/6 (which may have been membranous and destroyed in pinning out or even in han- dling the specimen), is believed to warrant assuming that the gizzard is in v. Intestinal caeca (as in some of the author’s specimens), possibly even ealeif- erous sacs, may have been unrecognizable because of the internal maceration. Ramiellona americana (Cates). Ramiella americana GAteEs, 1957, Breviora, 75:1. This species was erected on a single, probably incomplete, specimen sup- posedly from Guatemala. The esophagus is slender behind the gizzard but if calciferous saes and lamellae are present in this species they had become un- recognizable because of internal maceration. Spermatheeal pores are large enough to admit protuberances functioning as temporary intromittent or- gans. The lateral typhlosoles, intestinal caeea, lateral hearts of x, latero- esophageal hearts of xi-xii, the testicular chamber and the spermathecae suggest relationships to a “guatemalana” group of species. From that group, however, R. americana is set apart by the nephridia of the anteriormost portion of the body. Ramiellona lasiura (Graff). Howascolex (Graceevelynia) lasiurus GRAFF, 1957, Senckenbergiana, 38:129. This species was erected on 3 specimens (2 juvenile) from Salvador. The description is in the best classical tradition. Accordingly, though the typhlo- sole was well characterized, information as to size of spermathecal pores, GM elands and lateral typhlosoles, kind of hearts, trunks of the cireulatory sys- tem, nephridia of the preclitellar region, testicular chambers, ete., is not to be expected. Gonads were not seen and perhaps also the gonodueal funnels as the latter were not mentioned. The number of seminal vesicles also is un- 2116 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. known. Possibly masses of coagulum (which have been mistaken for vesicles in the past) were what was seen in segment xi. A single pair of vesicles, in xli, usually is indicative of metandry but in R. eiseni is associated with holandry. Henee, the andry is unknown, but to enable inclusion of “lasiura” in the key, metandry is assumed. Inability to reeognize gonads and perhaps also the enteric insertion of septum 5/6 may have made recognition of axial location of certain organs difficult (though level of entrance of spermathecae into the parietes should have provided one good elue). If the gizzard is in v rather than in vi, other organs also will prove to be located as in R. guatemalana and a testicular chamber may well be present. Restriction of hearts to vii—xi and presence of caleiferous sacs in 1x—xili certainly require confirmation. Ramiellona sauerlandti (Graff). Howascolex (Graceevelynia) sauerlandti Grarr, 1957, Senckenbergiana, 38:131. This species was erected on 2 specimens (1 juvenile) from Salvador (cf. comment on deseription of R. lasturus). Male pores, gizzard, calciferous sacs, intestinal origin and hearts appear to be as in R. guatemalana. Perhaps, then, the typhlosole which was not characterized also is of the same sort and associated with intestinal caeeca. Spermatheeal pores were not recognized and may be minute—protuberanees in region of prostatic pores apparently lacking. Exeept for presence of hearts in xi—xii, nothing is known of the vascular system and the anterior nephridia are not characterized. Male fun- nels are said to be in xi—xii which in itself is very unlikely but becomes even more so when seminal vesicles are in xi-xil. A lapsus calami doubtless is responsible for an erroneous location of the testes. The holandry then sug- gests relationships with FR. strigosa and R. stadelmanni. Ramiellona vulcanica (Graff). Howascolex (Graceevelynia) tecumumami vulcanicus GRAFF, 1957, Senckenbergiana, 382 L27. This form was erected on 6 specimens (5 subadult) from Salvador. As usual, information as to the vascular system and nephridia of a preclitellar region is lacking. The level of insertion of septum 5/6 on the gut apparently was not determined as the gizzard was said to be in v or vi (v in R. tecumu- mamt). Graff examined the type of this species and found the excretory system to be micronephridial, 7.e., meronephrie but did not record other im- portant information that could have been obtained at the same time. Among the differences between the Salvador and Guatemala worms re- spectively, are the following: Leneth 140 to 180 mm.—440 to 1000 mm. Di- VoL. XXX} GATES: EARTHWORMS OF EISEN’S COLLECTION Zi ameter, 7 to 9 mm.—9 to 12 mm. Segments, 260—860 (+ ?). Spermatheeal pores, in AB (possibly not minute)—median to A (possibly minute). Geni- tal markings, present at 16/17 and 19/20—lacking. Septum 10/11, lacking —present. Intestinal caeca, present—lacking (?). Penial setae, 2.6 mm. lone—4 to 4.5 mm. lone. Spermathecae, with a large round diverticulum on the ampulla ( ?)—adiverticulate but with seminal chambers sometimes recog- nizable in a small, transverse protuberance near ental end of duct. More information is needed about important somatie anatomy to determine rela- tionships of the two forms not only to each other but also to the rest of the species whether holandrie or metandrie. DISCUSSION Certain species do or probably do share with “quatemalana” many if not all of the following characters: Peristomium, flaccid and usually more or less retracted into anterior end. Prostomium, prolobous, not flaccid thoueh usually retracted. (Associated with the retraction there seems to be a tend- ency to abort the setal follicles in the anteriormost segments.) Setae, small and closely paired, at least anteriorly, ventral setae of xviii retained, ventral couples of xvii and xix penial. First dorsal pore in reeion of 11/12-12/138. Spermatheeal pores, in AB at 7/8-8/9, not minute, large enough for insertion during copulation of temporary protrusions from the male field. Male pores, at 17/18 and in seminal grooves that are lateral to B. Genital markines, un- paired, median, at interseemental furrows, usually if not always with a single transverse row of small circular areas. Septa, present from 4/5, 5/6— 9/10 funnel-shaped, 6/7-9/10 thiekly museular, 10/11-11/12 peripherally fused except ventrally to form a testicular chamber. Gizzard, in v. Caleif- erous lamellae, in vili—xil, in saes (4 species) that are somewhat more dis- tinctly constricted off from the gut in xi-xii. Intestinal origin, near insertion of 14/15. Intestinal caeca, paired, from dorsum in some of xx—xxx. Typhlo- sole widened ventrally and with three longitudinal ridges. Lateral typhlo- soles present in region of xx—xxx. Hearts, of x lateral, of xi—xii lateroesopha- geal. Dorsal blood vessel, single throughout and complete. Extra-esophageal trunks median to hearts and segmental commissures. No subneural trunk. Nephridia of ii—iv (presumably astomate) closely crowded in two longitudi- nal bands parallel to the nerve cord. Astomate micronephridia on anterior faces of septa in next eight segments in each of which there is one pair of nephridial duets. Some such uniformity of structure must be shared by species so closely related as to belong in a monophyletic genus. However, some of the charae- ters just listed may not be available to define even the genus in which R. guatemalana belongs. Caleiferous saes of xi-xi, for instance, distinguish four species of a “gquatemalana” group from Ramiella but other species, obvi- 218 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH Ser. ously of the same group, seemingly have no saes. The exeretory system in the first few segments of R. americana and R. mexicana provides differences from the “quatemalana” group that may be significant at generic level but not necessarily so from the unknown structure in most species of Ramiella. The ealciferous section of the gut appears to deny close relationship between R. americana and R. mexicana as well as of both to the “guatemalana” group. Most divergent, possibly, is R. irpexr which certainly is generically distin- guishable by its caleiferous glands alone from Ramiella. Nothing is known, however, about an anterior portion of the excretory system in R. irpex. Re- lationships of the latter to other American species accordingly are uncertain. Ramuellona obviously is a congeries, as a whole morphologically indis- tinguishable from Ranvella and presently incapable of segregation into natu- ral generic units. The group may well prove to be a most important constitu- ent of the native earthworm fauna in a region from the isthmus of Tehuan- tepec to Nicaragua. Hisen long ago suggested that exotie species were replacing the natives in Mexico and Central America. If, as elsewhere, endemics are haemerophobie they should be sought in undisturbed soils, especially in jungles (which must exist even today in considerable areas) and preferably near or at end of the rainy season when mature individuals are most likely to be easily obtainable. Key TO SPECIES OF RAMIELLONA 1. Stalked calciferous glands lacking Slender stalks of calciferous glands pass to gut close to mD in viii—xi 25 Svermatheecal pores. paired)... eee 3 SOSIPONE NE YER) [OXONNS Wyo NPL, le WMS cece eee eee eer cs R. mexicana Sec HiOlamadiies Co. on ee ee I a kt es 4 Metandric? 2... eA reas Pen Sees No Ae ee iG 4° -Spermathecae adiverticulate 2.) eee eee 5 Spermathecae diverticulate {200 2) 2) 2) 6 es ee eee eee 6 On Clitellumvannular senital manrkines pained) see eee Clitellum saddle-shaped, genital markings unpaired _................... R. stadelmanni 6. Spermathecal diverticulum dorsoventrally flattened in preceding segment ...... BEL Te, | @ Topriz i = a 3e|- oe <= Meshed @ ry > ‘ Resi @ {ee Fad ave) | . 4 e 434 . 36-27 / @ Herat 2. Wi Tehron a 36}— “a e : . N > carr merce: 7 AFGHANISTAN ; 32 4 @ Kermonshah \ e 34|— x ¢ e @ Baghdad se @ sfohon a | e <4 @ ote EST @rverd / 30 Shustor@ @ Mosjd- i-Suleiman > o 13/4 0/5 aw 32 |RAQ 20 S07 -78 (sae @ Ahwar@ OF < Lats” Sf @ den bid iit PAKISTAN e O4 *, t) Abadon pro, 26 @ Bom -_: 128 . ~_—- £. Q a I 5 A ] £ © aaa @ Shiroz \ . ‘ ae | Se Bushir APOGEE 2 ON IP ta O3/-352 . Nv 33° o34-35 a ve Bondor Abbos @ @ Minob i 26 A 28 Rs, Ly’ i SAUDI ARABIA Ay, Josk . i le 24 GULF OF OMAN 26 \ \ \ \ 44 46 48 50 52 54 56 58 60 62 VoL. XXXII] ANDERSON: IRANIAN HERPETOLOGY 421 shrubs growing on the coastal plains and in the low coastal foothills (fig. 6). The primary agriculture is that of the date palm. In the south, .arge anticlinal structures rise abruptly out of the coastal plain. Inland from the gulf the plain is deeply dissected into rugged bad- lands. In these areas of severe erosion, vegetation is extremely scanty. Ficure 2. Map showing the localities in Iran where amphibians and reptiles were collected. Masjid-i-Suleiman, Khuzistan Province, 31° 57’ N., 49° 16’ E. Naftak, staff housing north of Masjid-i-Suleiman, Khuzistan Province, 31° 58’ Ne, 492 115" Masjid-i-Suleiman Airfield and vicinity, 31° 59’ N., 49° 15’ E. Sar-i-Gach, Khuzistan Province, 31° 57’ N., 49° 21’ E. Tul-i-Bazun, abandoned airfield, Khuzistan Province, 31°55’ N., 49° 25’ E. Bard-i-Nishunde, Khuzistan Province, 31° 57’ N., 49° 21’ E. Godar Landar, Khuzistan Province, 32° 01’ N., 49° 23’ EH. Zeloi, Khuzistan Province, 32° 13’ N., 49° 04’ E. Lali, Khuzistan Province, 32° 15’ N., 49° 05’ E. 10. Lali Well No. 1, Karun River gorge, Khuzistan Province, 32° 14’ N., 49° 11’ E. 11. Sar-i-Naftak, near Tang-i-Golestan, Khuzistan Province. 12. Culvert on road between Masjid-i-Suleiman and Batwand, Khuzistan Province, BULe Bysy/ ING, ZO 2S IDE 13. Lahabira Valley, Khuzistan Province, 31° 56’ N., 49° 09’ E. 14. Naft Sefid oil field, Khuzistan Province, 31° 40’ N., 49° 14’ E. 15. Yamaha, Khuzistan Province, 31° 47’ N., 49° 23’ E. 16. Haft Kel, Khuzistan Province, 31° 28’ N., 49° 30’ E. 17. Lake, 17 kilometers east of Haft Kel. 18. Champ Kure, Khuzistan Province, 31° 31’ N., 49° 50’ EK. 19. Agha Jari, Khuzistan Province, 30° 43’ N., 49° 49’ E. 20. Gach Saran, 30° 20’ N., 50° 48’ E. 21. Chah Sefid, 30° 00’ N., 50° 52’ E. 22. Dar-i-Khazineh, Khuzistan Province, 31° 45’ N., 49° 08’ E. 23. Sand dunes on Ahwaz Ridge, Khuzistan Province, 31° 18’ N., 48° 45’ E. 24. Kurait, east of Ahwaz, Khuzistan Province, 31° 17’ N., 48° 49’ E. 25. Sand dunes on road between Ahwaz and Haft Kel, Khuzistan Province, 31° 16’ IN, OS wal? 19¢ 26. Persepolis, 29° 56’ N., 52° 49’ EB. 5,500’. 27. Binak, at the foot of Kuh-i-Bang, 29° 44’ N., 50° 19’ E. 28. 35 miles north of Bandar Abbas, 27° 30’ N., 56° 18’ E. 29. Hast of Kuh-i-Ginau, 27° 33’ N., 56° 24’ EK. RO. IOs bl, AS Bay ING, NOS A? 1d, dle Road just north of Shaeu, 200 177 N., 562227 32. Dunes at Shagu, 27° 15’ N., 56° 25’ EH. 33. Dunes on road from Bandar Abbas to Kerman, northeast of Bandar Abbas, 27° WP IN, GOAL 1, 34. Date grove at Minab, 27° 09’ N., 57° 07’ E. 35. Dunes at Minab, 27° 09’ N., 57° 07’ E. 36. Foothills of Elburz Mountains, north of Tehran. 37. Hills east of Tehran on Ab Ali Road. bo eR OAD OT me oe 422 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 471 SER. Although there are many species of desert shrubs, there is no immediate eround-cover other than spring grasses, which complete their growing season within a very few weeks. There are a few small active dune areas inland from the coast, similar to other seattered dunes in southern Iran. Loealities 27-35 are in the coastal area. LOCALITIES The latitude and longitude of the localities were determined from U. 8. Air Foree Aeronautical Approach Charts and can only be regarded as approximations. Masjid-i-Suleiman (localities 1 and 3). Lat. 31° 57’ N., long. 49° 16’ EK. Field headquarters of the Iranian Oil Exploration and Producing Com- pany. On many maps it appears as Maidan-i-Naftun. Wall (1908) refers to it as Maidan Mihaftan. The terrain is one of valley and ridge, typical of the foothill belt. Incompetent folding of the gypsum rock of the lower Fars formation has led to an erosion pattern of ridges of resistant rocks and strike-valleys cut in the more readily eroded layers. Naftak (2). Lat. 31° 58’ N., long. 49° 15’ E. is a residence area about five kilometers to the north of Masjid-i-Suleiman proper, and is shghtly higher in elevation. The road from Masjid-i-Suleiman to Naftak winds through low hills composed chiefly of gypsum. Sar--Gach (4). Lat. 31° 57’ N., long. 49° 21’ EK. A ridge to the east of Masjid-i-Suleiman composed principally of gypsum. It is crossed by a well- maintained, surfaced road, which serves the wells of the Masjid-i-Suleiman oll field and the water pumping station at Godar Landar. Tul-Bazun (5). Jat. 31°.55’ N., long. 49° 25’ KE. Near the top ot the Sar-i-Gach ridge, a road runs south from the Sar-i-Gach road to an aban- doned airstrip. Along this road are sinkholes in the gypsum whieh contain water throughout the year. Bard-1-Nishunde (7). Lat. 81° 58’ N., long. 49° 21’ HE: Site of a pre- Achaemenian ruin lying north of the road near the crest of Sar-i-Gach. Godar Landar (7). lat. 382° 01’ N., long. 49° 23’ E. A pumping station supplying Masjid-i-Suleiman with water from the Karun River. Across the river from the pumping station rises a steep conglomerate wall, com- posed of limestone pebbles and cobbles, to the high ridge of Kuh-i-Landar. Zeloi (8). Lat. 82° 18’ N., long. 49° 04’ E. A plain west of the road between Masjid-i-Suleiman and Lali. There are streams and sinkholes in the gypsum. This was a former site of exploratory oil drilling and travers- able roads still exist to some of the well sites. Lali (9, 10). Lat. 82° 15’ N., long. 49° 05’ EK. This is the northernmost oil field in the foothill area of Khuzistan. It is located in the hills through VoL. XXXII] ANDERSON: IRANIAN HERPETOLOGY 423 whieh flows the Karun River. There are considerable salt deposits in this region. Sar-i-Naftak and Tang-i-Golestan (11). A surfaced road connecting Masjid-i-Suleiman with Batwand winds down the Sar-i-Naftak ridge, Ficure 38. Foothill region, March 1958. Zagros Mountains in the background. Foreground is typical habitat of Agama nupta, Mabuya aurata septemtaeniata, Colu- ber rhodorachis, and Psammophis schokari. 424 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4711 SER. through Tang-i-Golestan, a steep-sided canyon. A small stream passes under this road through a eulvert (12). Lat. 31° 55’ N., long. 49° 25’ BH. At Bat- wand this road meets the newer Masjid-i-Suleiman—Ahwaz road. Naft Safid (14). Lat. 31° 40’ N., long. 49° 14” E. An oil field on the Ficure 4. Stream in the foothill region near Station 12 (fig. 2), April, 1958. Habitat of Rana ridibunda and Hyla arborea savignyi. VoL. XXXI] ANDERSON: IRANIAN HERPETOLOGY 425 western slope of the foothill belt. The terrain is typieal of the foothill region. Yamaha (15). Lat. 31° 47’ N., long. 49° 23’ KE. An abandoned site of ex- ploratory oil drilling near the Tembi River in the foothills south of Masjid-i- Suleiman. A road runs from Yamaha, across the Tembi River, and through the Masjid-i-Suleiman oil field. Haft Kel (16). Lat. 31° 28’ N., long. 49° 30’ E. An oil field south of Masjid-i-Suleiman. The road from Masjid-1-Suleiman to Haft Kel runs through typical foothill terrain, traversing a part of the Tembi River drainage. At one point the road emerges onto the small Yamaha plain. It erosses the Ab-i-Lashear and minor intermittent streams. Haft Kel is near the edge of the foothills; the road from Haft Kel to Ahwaz soon drops onto the flat Ahwaz plain. Near Haft Kel a road runs east into the Zagros Mountains. Seventeen kilometers east of the Masjid-i-Suleiman—Haft Kel road is a small lake (17) in an undrained depression in the hills. Shrinking to a fraction of its winter size, it provides, even in summer, a refuge for frogs, turtles, and ducks. In summer there is no sign of vegetation near the lake, other than a stand of dry brown reeds at its periphery. The road into the Zagros leads ultimately to the village of Qal ’eh Tun, site of an old Bakhtiari fortress. From there a jeep track leads through stands of scrub oak to Malagha, a small village on a permanent stream, the Rud-i-Zard, at an elevation of 5,000 feet. Champ Kure (18), lat. 31° 31’ N., long. 49° 50’ E., is a government check station on this road. Agha Jari (19). Lat. 30° 43’ N., long. 49° 49’ E. An oil field in the foot- hills at the edge of the coastal plain. There is a small active dune area on a ridge in the foothills near the number 38 production unit of the field. Gach Saran (20). Lat 30° 20’ N., long. 50° 48’ EK. The southernmost oil field in the foothill region. The Dugumbadan airfield and the living areas are ona small plain at the foot of a high ridge. The oil field is located in the foothills. Chah Safid (21). Lat. 30° N., long. 50° 52’ E. A camp site on the road from Gach Saran to Ganaweh. It is approximately one mile from the june- tion of the Mishun road. The road traverses terrain of deeply incised gorges, most of which carry an intermittent flow of water. All of the gorges are sub- ject to flood torrents following heavy rains. Lahabira Valley (13). Lat. 31° 56’ N., long. 49° 09’ E. A valley in the foothills west of Masjid-i-Suleiman. Two to four miles wide, a thin alluvium covers the gypsum in much of the valley floor, through which flows Ab-i- Lashear, a permanent stream. This stream is fed by seasonal drainages, and here and there are sinkholes in the gypsum. Shustar. An old city on the Karun River in the relatively high eastern area of the Ahwaz Plain, and not far from the foothills. The road to Shustar passes through areas of low hillocks, ridges, and sandstone outerops. Much 426 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH SER. of the vegetation along this road is that characteristic of the foothill region. Annual rainfall is about 300 millimeters (Adams, 1962). Ahwaz. The eapital of Khuzistan Province, and the commercial and dis- tributional center. Ahwaz lies on the Karun River in the heart of the plain. Both the Iranian National Railway, running from Bandar Shapur and from Khorramshar to Tehran, and the road from Khorramshar to Tehran, pass through Ahwaz. The Karun River is navigable from the Shatt ’al Arab as far as Ahwaz. Small dune areas (23). Lat. 31° 18’ N., long. 48° 45’ E. are located along the Ahwaz Ridge on the road between Ahwaz and the Ahwaz number 6 well location at Kwramt (24), lat. 31° 17’ N., long. 48° 49’ E. Dar-i-Khazineh (22). Lat. 81° 45’ N., long. 49° 08’ E. A small village on the plain near the road from Masjid-i-Suleiman to Ahwaz. Khalafabad. A ferry crosses the Jarrahi River near the village of Khala- fabad on the summer road from Agha Jari to the Ahwaz—Haft Kel road. The road erosses a broad flat plain, where, except for small cultivated areas near villages, the vegetation is sparse and xerophytie. Sand dunes on the Ahwaz—Haft Kel road (25). Lat. 31° 16’ N., long. 49° 11’ E. The largest dune area visited was on the road between Ahwaz and Haft Kel. These dunes are composed of loose sand, transported and altered by the wind. A few shrubs grow on the dunes and on the sandy fringes. Some grasses and thorny shrubs are found in the “blow-out” depressions amone the dunes, and the root systems have some stabilizing effect. Persepolis (26). Lat. 29° 56’ N., long. 52° 49’ KE. Ruins of the palace of the Achaemenian kings. It is on a terrace above a plain, in a large valley in the Zagros Mountains. The elevation is approximately 5,500 feet. It les at the foot of high limestone hills. The hills are barren except for a few small shrubs. There is considerable agriculture on the plain. Binak (27). Lat. 29° 44’ N., long. 50° 19’ K. Site of an exploratory well at the foot of Kuh-i-Bang on the Persian Gulf. There is a narrow strip of coastal dunes, on which grow a few shrubs. Kuh-i-Bane is an anticlinal structure, approximately one mile in length, rising out of the coastal plain. The vegetation of the plain is sparse. There is some date culture at Ganaweh on the coast to the southeast. A few low hills lie at the foot of Kuh-i-Bang and along the coast. Bandar Abbas (28-33). An ancient seaport at the Strait of Hormuz. A road runs from Bandar Abbas to Kerman and Shiraz. East of the town, alone the road to Kerman, there is a small active dune area (33), lat. 27° 12’ N., long. 56° 21’ E., much like the dune areas on the Ahwaz Plain. Three prominent anticlinal structures rise out of the plain north of Ban- dar Abbas. These are Kuh-i-Ganau, Khureu, and Kush Kuh. Between Bandar Abbas and Kuh-i-Ganau, the road passes through areas of savannah, sandstone outerops, and low hills, emerging onto a higher, cobbled plain. VoL. XXXT] ANDERSON: IRANIAN HERPETOLOGY 427 Here there is true desert vegetation (fig. 5). Many of the thorny, xerophytie shrubs of the plain extend onto the slopes of the mountains. Although as a whole the area is quite arid, there are a few drainages where streams persist the vear round. There is considerable date culture alone these streams, even into the mountains to 3,000 feet. Shagu (31), lat. 27° 17’ N., long. 56° 22’ E., is a small village, primarily a truck stop, a few miles north of Bandar Abbas on the road to Kerman. Just east of the village, on the road to Minab, there is a small active dune Medmacn plataict loo Ne lone 56° 2a" Er: Minab (34, 35). Lat. 27° 09’ N., long. 57° 07’ E. The road from Shagu to Minab erosses a flat plain, many of the savannahs having numerous trees. Minab is a date-growing center. The Rud-i-Minab is the principal drain- age. Minab is located at the edge of the coastal plain, there being much folded strata to the east, continuing into Baluchistan. A small aetive dune area (35) hes north of the town, against a low ridge. The vegetation of the Minab area, other than the extensive date cultivation, consists of a few desert shrubs. Figure 5. Terrain near Kuh-i-Ginau, southern Iran (Station 28 in fig. 2), April, 1958. Habitat of Hremias guttulata watsonana. 428 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. SYSTEMATIC AND ECOLOGICAL DISCUSSION Class AMPHIBIA Order SALIENTIA Family BuFONIDAE Genus Bufo Laurenti Bufo viridis Laurenti. Bufo viridis LAURENTI, 1768, Synops. Rept., p. 27, pl. 1, fig. 1. KNOWN RANGE. Europe east of the Rhine and Rhone rivers; North Africa; southwest and central Asia, from the Mediterranean to Tibet and Mongolia; from below sea level near the Dead Sea to 15,000 feet in the Himalayas. MATERIAL EXAMINED (14). Station 1! (CAS 862622 [II/27/58; ¢]; 86263 [III/2/58, 2 |; 86285 [ITI/19/58, ¢]; 86330 [1V/13/58, 3&]; 86641— 86642 | 2 2 |; 86643-86645 [oo] [XI/30/58]). Stream 15 km. north of Station 1 (CAS 86284 [IIT/10/58, 3 ]). Station 2 (CAS 86324 [IIT/30/58, 3 |). Stream near Station 3 (CAS 86594 [IX/11/58, 9° ]). Station 10 (CAS 86289 [III/20/58, o]). Stream near Station 17 (CAS 86327 [III/28/58, ef IL). RemaArKs. The interorbital space is as wide as, or wider than, the upper eyelid in this series. There are numerous warts on the dorsum. The larger warts are surmounted by a prominent tubercle tipped with a black asperity, and this tuberele is surrounded by smaller, similar tubercles. The tubercles of nonbreeding females are reduced, and lack the asperity. The toads are numerous in Khuzistan Province along stream courses in early spring, usually in the grass, rarely actually in the water. They were found occasionally under rocks on dry hillsides several hundred yards from water. Active in cultivated gardens throughout the summer, they were often seen at night in the grass near an outdoor light, feeding on insects attracted to the hight. After rains in the late fall they were numerous on paved roads in the evening. Bufo viridis was seen, but not collected at Persepolis in August. On two oceasions males were heard to make a bird-like warbling eall along a stream in early March, several individuals participating in the chorus at dusk. Strings of eggs were found in the streams in late February and early March. By early April most of the larvae had metamorphosed, the water along seasonal drainages by this time limited to intermittent pools diminish- 1. Station numbers refer to localities described on p. 6 of ms., and fig. 2. 2. California Academy of Sciences catalog number. VoL. XXXII] ANDERSON: IRANIAN HERPETOLOGY 429 ing in size. Two females and three males collected November 30, at Masjid-i- Suleiman appear to be in breeding condition. Stomachs of toads collected in November contained ants and termites; CAS 86643 contained a centipede. The stomach of CAS 86285, collected March 19, contained terrestrial isopods, lepidopterous larvae, and other arthropods. There was also a calcareous mass, 6 mm. in diameter, in the stomach. Grass was present in the stomach of CAS 86289; this stomach con- tained no animal remains. Bufo olivaceus Blanford. (Figure 7.) Bufo olivaceus BLANForRD, 1874, Ann. Mag. Nat. Hist., ser. 4, vol. 14, p. 35. (Type lo- cality: Dasht River, Baluchistan [West Pakistan]. ) KNOWN RANGE. Southeast Iran and western Pakistan; to 3,000’. MATERIAL EXAMINED (33). Station 34 (CAS 86567-86568, 86570-86571, 86576, 86578, 86581-86582, 86584-86585, 86611-86613, 86615-86616, 86618 | oo |; 86560, 86569, 86572-86573, 86575, 86577, 86579-86580, 86583, 86595— 86599, 86610, 86614, 86617 [ 9 2] [X/21/58]). Remarks. These specimens do not show as pronounced an occipital fold as shown in Blanford’s illustration (1876, pl. 28, fig. 3). While Boulenger (1882) states that B. olivaceus lacks a tarsal fold, the present specimens pos- sess a well-defined tarsal ridge of closely aligned tubereles. The Minab series may be characterized as follows: Head and body de- pressed; snout projects slightly beyond mouth; crown lacks bony ridges; dis- tance between eye and nostril on canthus rostralis equal to at least 2% the diameter of the eye; interorbital space equal to at least 34 the leneth of the upper eyelid; distance between nostrils equal to 14, or only slightly greater than 14, the interorbital distance; tympanum distinet, the vertical diameter may be slightly greater than the horizontal diameter, the latter being 1% the diameter of the eve. Tongue in preserved specimens longer than wide. First finger longer than second; toes more than 44 webbed, with the ex- ception of the fourth, which is 13 webbed; toes with single subarticular tubercles; a well-defined tarsal ridge of closely aligned tubercles; inner meta- tarsal tubercle larger than outer metatarsal tubercle; palmar metacarpal tubercle considerably larger than thenar metacarpal tubercle. Tibio-tarsal articulation reaches angle of mouth or slightly beyond. A few small warts tipped with black asperities in the dorsal sacral and pelvic areas, slightly more numerous and more pronounced in females, and entirely lacking in CAS 86568 and 86613 (both males). Paratoids often twice as long or longer than broad, frequently extending to sacral region. 430 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Male with subegular vocal sae. Pale olive above, white below; immaculate. Male 56.5 mm. snout-vent leneth; female 58.0 mm. These toads are distinct from Boulenger’s (1891) deseription of B. surdus whieh lacks the tympanum (and the annulus typanicus, columella, and eusta- ehian tube as well, according to Parker, cited by Schmidt, 1955) and which has bony ridges on the crown. Chernov compared one of the animals of the Minab series with Nikolsky’s specimens of B. persicus (Carevsky, 1926, equated B. surdus with B. persicus, a view with which Schmidt, 1955, takes issue); he stated (personal communication) that the Minab toad is distinct from his specimens. It is clearly distinet from B. luwristanicus Schmidt, which has much smaller paratoids, a tubereulate dorsal skin, shorter hind limbs, and the interorbital space narrower than the upper eyelid. Blantford’s specimens came from the Dasht River, which flows into the Gulf of Oman some 15 km. east of the Iran—Pakistan border, Bahu Kalat, just west of the border, and Ghistigan, Bampusht, also on the Iranian side of the border. Blanford (1876) states that at elevations greater than 3,000 feet B. olivaceus is replaced by B. viridis. The present series was collected on a single October evening between 6 and 9 p.m. There were hundreds of these toads in and along irrigation ditches near a road running through a date grove (fig. 6). They were col- leeted in the grass, in the mud, on the banks, and in the water of the shal- low ditches, all within a distance of 100 yards. The majority were seen sit- ting or swimming in the water. Those on the banks and in the grass usually sought the water in their efforts to escape capture. These toads greatly out- numbered the representatives of Rana cyanophlyctis which were present in the same ditches. None were in breeding condition. Most of the stomachs examined contained ants primarily, and often beetle larvae and termites. Several individuals had flatworms in the urinary bladder. Family HYLIDAE Genus Hyla Laurenti Hyla arborea savignyi Audouin. Hyla savignyi Aupourn, 1827, Descr. Egypte, Rept., Suppl., p. 183, pl. 2, fig. 13. (Type locality: presumed to be Syria. ) Hyla arborea savignyi, MERTENS, 1924, Abh. Ber. Mus. Magdeburg, Bd. 3, p. 356. Ficure 6. Date grove at Minab (Station 34 in fig. 2), October, 1958. Bufo oliva- ceus and Rana cyanophlyctis were collected in irrigation ditches of this grove. FicurE 7. Bufo olivaceus. VoL. XXXT] ANDERSON: IRANIAN HERPETOLOGY 431 432 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. KNOWN RANGE. [Tyla arborea ranges from Europe through North Afriea and temperate Asia. The subspecies //. a. savignyi is the form of southwest Asia, meeting the typical form in Asia Minor and extending eastward through Iran, being found in the drainages of the Zagros and Elburz Mountains. MATERIAL EXAMINED (21). Station 1 (CAS 86286 [III/20/58, 9 |). Stream on road to Station 9, 15 km. north of Station 1 (CAS 86295-86296 [2 2]; 86297-86300 [vo] [III/10/58]). Stream on road to Station 9, 16.9 km. north of Station 1 (CAS 86312 [III/21/58, 9 ]). Vicinity of Sta- tion 1 (CAS 86355 [ @ |]; 86356 | #] [April, 1958] ). Stream between Station 1 and Station 16 (CAS 86446 [VI/14/58, ee ). Station 8 (CAS 86253- 86255, 86257, 86259-86260 [fo]; 86256, 86258 [2 9] [II/27/58]) Stream south of Station 9 (CAS 863 ae M2058 oule Remarks. These hylas are present in fair numbers along the seasonal stream courses in early spring. When collected, all were bright green, blend- ing well with the grasses in which they were found. When placed in the dark, some became a darker, olive green, some a steel gray or gray-brown, others showing no change, although all were in the same container. Some be- came much paler when nareotized with chlorotone. Just-metamorphosed in- dividuals were a hght gray-green or tan. In Khuzistan Provinee the eges seem to be laid in late February and early March, metamorphosis being completed in March and April. No hylas were seen after mid-June. No intensive search was made for them, however. Water remains throughout the year in sinkholes and small caves in the gyp- sum formations, as well as in permanent streams. The water temperature is 22°-26°C. during the hottest part of the summer. A retreat is thus pro- vided for the amphibians of the area. CAS 86304 has a supernumerary forelimb attached to the right side of the pectoral girdle and innervated from the right side. Family RANIDAE Genus Rana Linnaeus Rana ridibunda ridibunda Pallas. Rana ridibunda PAtias, 1771, Reise Rus. Reich., vol. 1, p. 458. (Type locality: Gurev, north coast of the Caspian Sea.) Rana ridibunda ridibunda, MERTENS, 1925, Abh. Senck. Ges., vol. 39, p. 55. KNowN RANGE. The whole of Hurope except northwest and central Italy; western Asia as far east as northern West Pakistan, Afghanistan, and eastern Turkestan (USSR); North Africa (Boulenger, 1891). MATERIAL EXAMINED (99). Vicinity of Station 1 (CAS 86353 [ & | ; 86354, 86359, 86364-86365 [| 2 2 | [| April, 1958] ). Stream 1.9 km. north of junetion VoL. XXXI] ANDERSON: IRANIAN HERPETOLOGY 433 of road between Station 1 and Station 9 and road between Station 1 and Ahwaz (CAS 86307-86311 [III/21/58, ¢o']). Stream 16.9 km. north of Station 1, on road to Station 9 (CAS 86313-86315 [III/21/58, juv.]). Fif- teen kilometers north of Station 1, on road to Station 9 (CAS 86290, 86292 [juv.]; 86291 [ 9]; 86293-86294 [io] [III/10/58]). Station 2 (CAS 86363 [III/28/58, 9]). Stream near Station 3 (CAS 86267 [III/14/58, 3&]). Pools near Station 3 (CAS 86619-86620, 86628 [ 2 2 |; 86621-86622, 86627 [ov ov] [Sept.—Oct., 1958] ). Stream on road to Station 5 (CAS 86264— 86265 [III/13/58, juv.]). Stream 9 km. above Station 7 (CAS 86261 [III/ 13/58, ¢@]). Station 8 (CAS 86268-86272, 86274-86278, 86280-86282 B22 86273; 86279) 86283" oo | [127/58]; 86287 (11/27/58, 2 |). Stream south of Station 9 (CAS 86301-86302, 86306 [ 2 2 |; 86303 [juv. | [11I/20/58]). Station 13 (CAS 86316-86319 [ITI/21/58, juv.]). Stream near Station 17 (CAS 86357 [¢]; 86358 [¢@] [III/28/58]). Station 21 (CAS 86411 [ 2 |]; 86412-86413 [juv.] [VI/8/58] ). Remarks. This frog is numerous in the foothill regions in all the per- manent streams throughout the year, and in the seasonal drainages in the spring. Sinkholes and small caves in the gypsum provide retreats from the extreme summer heat and contain the only permanent water in many loeali- ties. During the day these frogs are found sitting in the water or in the vegetation along the banks of pools and streams. Kees are laid in late February and Mareh, most metamorphosis being completed by early April. A few metamorphosine tadpoles were seen as late as May 23 in an area where the stream course was dry except for small, rapidly shrinking pools. Unsuecessful attempts were made to induce ovulation in mature females in September. Ovaries showed little response despite injections of both amphibian and mammalian anterior pituitary. These frogs may be an important part of the diet of many predators, since they are one of the few prey species available in large numbers throughout the vear. Seventeen of the 59 frogs in this series have a light vertebral stripe. In 40 of the 59 the tibiae do not overlap when the legs are at right angles to the body, and in several of these they do not even touch (a character inde- pendent of age or sex). Rana cyanophlyctis Schneider. Rana cyanophlyctis SCUNEIDER, 1799, Hist. Amph., vol. 1, p. 137. KNOWN RANGE. Widely distributed from southern Arabia and eastern Iran to the Malay Peninsula, and from the Himalayas (to 6,000 feet) to Ceylon. 434 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4111 SER. MATERIAL EXAMINED (6). Station 34 (CAS 86562 [¢%]; 86561, 86563— 86566 [2 2] [X/21/58]). Remarks. The frogs of this series have a single white streak on the hinder side of the thighs, the dark pigment below this streak giving a mottled appearance rather than forming a distinet second dark streak. The inter- orbital space is only slightly narrower than the upper evelid in these specimens. Stomach contents included beetles, beetle larvae, a mole cricket, spiders, and other arthropod remains. Class REPTILIA Order CHELONIA Family Emypmar Genus Clemmys Ritgen Clemmys caspica caspica ((imelin). Testudo caspica GMELIN, 1774, Reise durch Russland, vol. 3, p. 59, pls. 10-11. (Type locality: Hircania, northern Iran.) Clemmys caspica caspica, Scumipr, 1939, Field Mus. Nat. Hist., Zool. Ser., vol. 24, p. 89. KNOWN RANGE. Ivan, from the southern borders of the Caspian Sea to the Persian Gulf; Iraq; northern Arabia and Bahrein Island. MATERIAL EXAMINED (7). Station 3 (CAS 86634 [III/14/58, 9]) Sta- tion 17 (CAS 86629 | 2 |; 86639 [| ¢] | X/26/58]; 86635 | 9 |]; 86636-86637 , [juv.] [2IL/28/58]; 866388 [V/23/58, juv.)). Remarks. The carapace of CAS 86639 is misshapen, the left side much more depressed than the right, several shields having a shriveled appearance. These turtles were collected at only two localities. Turtles, apparently of this species, were seen on a stream bank near Istahan, and in streams near Persepolis. Presumably they are to be found wherever there is permanent water over successive years in the foothill regions. In the small lake (75-100 meters in diameter) where most of these speci- mens were collected, there were hundreds of these turtles of all ages, as well as numbers of Rana ridibunda. This appears to be a permanent lake, but in summer diminishes to half its sprine size. Reeds grow along its northern margin, and there is considerable aquatic vegetation in the lake itself. One of the females laid an ege on June 4, and another on June 26. These eges were about 40 mm. by 30 mm. The shells were brittle, and of a china- like texture. CAS 86629 and 86639 were found copulating on October 26, in the mud at the edee of the small lake. Man may be the principal predator on the adults of this species. Al- VoL. XX XT] ANDERSON: IRANIAN HERPETOLOGY 435 though the flesh is apparently not eaten, the eggs are considered a potent ingredient in eye medicine. It is possible many turtles are killed for the eges they contain, as eye ailments abound in southern Iran. Order SQUAMATA Suborder SAURIA Family GEKKONIDAE Genus Eublepharis Gray Eublepharis macularius (Blyth). (Figure 8.) Cyrtodactylus macularius BiyrH, 1854, Jour. Asiat. Soc. Bengal, vol. 23, p. 737. (Type locality: Salt Range, Punjab.) Eublepharis macularius, ANDERSON, 1871, Proc. Zool. Soc., p. 163. KNOWN RANGE. Northwestern India, Pakistan, and Afghanistan to south- ern Turkmen (USSR), Ivan, and Ivaq. MATERIAL EXAMINED (16). Road between Station 1 and Station 2 (CAS 86333 [1V/15/58, 3, DOR]; 86360 [| juv.]; 86361, 86366 [ 2 2 ]; 86362 [ov] [IV /19/58]; 86381 [IV /28/58, o']; 86383 [V/13/58, juv.]; 86385 [V/21/ 58, 2 |]; 86396, 86398 [vo vo]; 86397 | 2 | [V/22/58]; 86416 [V/26/58, ov]; Ssoa0t—-V 11/20/58, 2 | ). Lise d between Station 1 and Batwand (CAS 86337 [IV/18/58, &, DOR]; 86382 [V/13/58, 2 ]; 86384 [V/20/58, oJ). Remarks. Each dorsal tubercle is encircled by a single rine of slightly enlarged, juxtaposed dorsal seales. In those specimens in which the tail has been rejuvenated, the tail is swollen just distal to the break, the rejuvenated portion lacking tubercles, and having smaller, less regular ventral seales. Preanal pores are present in the females, although much less distinet than in the males. The postanal saes and deep axial pouches contain many small mites in all specimens. All specimens were collected on surtaced roads between 8 p.m. and mid- night. They were first seen in mid-April on roads along which collecting had been done during the same hours for the preceding six weeks. The lizards were fairly numerous on these roads until the end of May. None were seen from June until August 20, when a single specimen was collected. These lizards were never encountered during the day, although many rocks were overturned in the area. Deep crevices in the gypsum probably provide a retreat during the day. During the period that these lizards were seen on the roads, grasshoppers were also present on the roads in large numbers. Also on the roads at the 436 CALIFORNIA ACADEMY OF SCIENCES [Proc. 471 SER. same hours were scorpions, solpugids, and large spiders. Stomach contents included these animals as well as beetles and other arthropods. They have been reported to eat other lizards (Smith, 1935) and may prey on the vari- ous small geckoes of this reeion. The foxes which abound in this area were observed eating lizards of this species which had been hit by automobiles, and probably prey regularly on them. When captured, these lizards give a loud, prolonged squeek and attempt to bite. They often defecate, wrapping the short tail around the hand of their captor. This may oceasionally enable them to drop the tail. CAS 86361, 86366, 86385, 86397, 86507 contain eges. Air temperature recorded when these lizards were collected varied from 32.0° to 34.4°C. The road surface temperatures were 32.6° to 36.4°C., and were usually at least 2°C. higher than the surrounding soil temperatures, which ranged from 30:2° to 35.0°C. The critical maximum temperature (as defined by Cowles and Bogert, 1944) was recorded for one individual (CAS 86507) and found to be 42.6°C. The lizard was placed on an asphalt surface which had a temperature of Fiaure 8. Hublepharis macularius. VoL. XXXII] ANDERSON: IRANIAN HERPETOLOGY 437 48.6°C., in the direct sunlight. The air temperature was 44.6°C. The anal temperature at the start of the experiment was 35.6°C. The lizard was tethered so that it could not reach the shade which was visible a short dis- tance away. The anal temperature at which all efforts to reach the shade became uncoordinated and ineffective was considered the critical maximum. Anal temperatures were taken with a Schulthize rapid-readine thermometer. The eritical maximum temperature was reached four minutes after the ini- tiation of the experiment. Efforts to reach the shade began immediately upon exposure of the animal to direct sunlight, and were unceasing until collapse. Considerable moisture was passed from the vent, and the mouth opened only when the animal was on the point of collapse. After the anal temperature was recorded, the animal was removed to the shade where it subsequently recovered. No paralysis was noted after recovery, but what, if any, other functional impairments resulted from exposure to high tem- perature was not determined. Genus Cyrtodactylus Gray Cyrtodactylus scaber (Hayden). Stenodactylus scaber HAYDEN, 1827, in Riippell, Atlas Reise Nord. Africa, Rept., p. 15, pl. 4, fig. 2. (Type leoality: vicinity of Tor, Sinai.) Cyrtodactylus scaber, UNpERWOOD, 1954, Proc. Zool. Soc., London, pp. 469-492. KNowN RANGE. From Egypt to northwest India; Arabia, along the Persian Gulf. MATERIAL EXAMINED (9). Station 1 (CAS 86481 [VI/23/58, do, on a wall] ; 86513 [IX/7/58, juv.] ; 86626 [Nov., 1958, juv., in a house] ). On wall on a house near Station 3 (CAS 86532 [IX /10/58, juv.]; 86533 [IX /11/58, juv.]). Station 19, on wall of I.O.E.P.C. guest house (CAS 86481 [VITI/14/ 58, 2 |; 86482 [ 2 ]; 86483 [of] [VIII/15/58]). 1O.E.P.C. guest house at Station 24 (CAS 86558 [X/5/58, juv.]|). Remarks. There is a vertebral series of tubercles, smaller than the 10—12 rows ot longitudinally and transversely arranged tubercles. This median row usually extends from the scapular to the pelvic region. This species is the most common house-gecko in the foothill region. It was quite common on walls of houses from late June through November, becoming active during the late daylight hours. It was frequently seen at night just beyond the cirele of light thrown by an outdoor fixture, darting forward occasionally to capture a moth or other insect attracted to the heht. Hatehlings were common in mid-August. One female laid two eges, ovoid, white, and with brittle shells, in late August. These were 10.5 mm. lone, the greatest width being 7 mm. 438 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Cyrtodactylus agamuroides (Nikolsky). (Figure 9.) Gymnodactylus agamuroides Nixousky, 1899, Ann. Mus. Zool. Imp. Acad. Sci. St. Pétersbourg, ser. 4, pp. 384-385. KNOWN RANGE. Eastern Iran. MATERIAL EXAMINED (1). Station 29 (CAS 86370 [IV /26/58, 3 ]). Remarks. The single specimen was found under a rock by village boys. Nikolsky (1899) designates no holotype specimen, nor type locality. The three syntypes are from Neizar in Seistan, Pendseh-Sara, and Duz Abad, eastern Kerman Provinee, Iran. The only reeord of this form subsequent to that of Nikolsky is by Wettstein (1951) who lists it from Kerman. The present specimen differs from C. kotschyi in that the limbs are longer, the dorsal tubercles are approximately as broad as long, the abdomi- nal seales are about 22-24 at mid-body (30 in C. kotschyi), and it has sub- quadrangular spots in three longitudinal series, with a less distinct lateral series, while C. kotschyi has angular transverse bars. It differs from C. fedtschenkov and C. longipes in having the snout longer than the distance between eve and ear, fewer longitudinal rows of abdominal seales (28-30 in C. fedtschenkov) ; scales of limbs are not keeled, dorsal tubereles are small, rounded, smooth to weakly keeled, subeonieal, rather than large and sub- trihedral; there are but 4 preanal pores (28-36 preanal and femoral pores in C. fedtschenkor) ; C. fedtschenkoi has dorsal crossbars. It is distinguished from C. kermanensis in that the diameter of the eye goes into the leneth of the snout only 144 times (2-214 in C. kermanensis), in that the mental shield is triangular rather than pentagonal, and in that it lacks strong keels on the dorsal tubercles. It is different from C. zarudnyi in having the first pair of chin shields in contact behind the mental; C. zarudnyi has triangular, strongly keeled tubercles, and lacks dark dorsal markines. It differs from C. kachhensis in having smaller, less strongly keeled tubercles, having reeu- lar infracaudal plates, and fewer longitudinal series of abdominal scales (30-40 in C. kachhensis). It is distinet in C. caspius, which has large, strongly keeled, trihedral tubercles, about 20 femoral and preanal pores, and indistinet dorsal crossbars. Cyrtodactylus brevipes has shorter limbs and imperfect longitudinal dusky bands on the dorsum, formed of arrow- head-shaped marks. It does not differ significantly from Nikolsky’s deseription of Gymnodac- tylus agamuroides. At least eight species of Cyrtodactylus have been recorded from eastern Tran (C. scaber, C. fedtschenkoi, C. kirmanensis, C. zarudnyi, C. longipes, C. caspius, C. brevipes, and C. agamuroides). The types of these species should be examined to determine whether or not these are all specifically distinct forms. VOL. XXXII] ANDERSON: IRANIAN HERPETOLOGY 439 Figure 9. Cyrtodactylus agamuroides. (ienus Bunopus Blanford Bunopus tuberculatus Ilanford. Bunopus tuberculatus BLANrorD, 1874, Ann. Mag. Nat. Hist., ser. 4, vol. 13. p. 454. (Syntypes from Bahu Kalat, Mand, and near Bampur, southeastern Iran.) KNOWN RANGE. Syria, lraq, eastern Arabia, southern Iran, Afghanistan, Pakistan; to 3,000 fect. MATERIAL EXAMINED (6). Station 25 (CAS 86492 [VIII/17/58, juv.]; S6n22 (VII /22/58, juv. |; S65386 | 2) 86537 |] |IX/13/58]). Station 35 (CAS 86590-86591 [X/20/58, oo']). ReMARKS. Leviton and Anderson (1963) have demonstrated that Blan- ford’s genus (Bunopus) of tuberculate geckoes is distinet from the non- tuberculate Alsophylax of Fitzinger. In the present specimens the imbricate seales of the belly and lower sur- faces of the imbs and tail have their posterior borders distinetly denticulated. During the day these lizards were found in burrows among the roots of shrubs in the active sand-dune areas. They were active about 9:00 P.M. on the surface of the sand, particularly near the base of shrubs, on October 20 at Minab. Blanford (1876) collected this species in Baluchistan in houses and under rocks. This species may be distinguished from Cyrtodactylus heterocercus (Blanford) and Bunopus persicus (Nikolsky) by the lack of postmental shields. The scales of the belly are smooth in Bunopus tuberculatus, but de- seribed as keeled in B. persicus. 440 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Coloration in life: sandy gray above, the juveniles distinctly barred with chocolate-brown; six crossbars, broader than the interspaces, between the shoulders and pelvis, becoming broken and less distinct in adults. A brown crescent curves around the nape and passes through the eyes. There is dark pigment in the temporal areas, and in front of the eyes, the lips banded or spotted with brown. The hind limbs are crossbarred with brown, the forelimbs mottled. The tail is erossbarred above, flecked with brown below. The throat and belly are immaculate white. Genus Microgecko Nikolsky Microgecko helenae Nikolsky. (Figure 10.) Microgecko helenae Nixousky, 1907, Ann. Mus. Zool. Acad. Imp. Sci. St. Pétersbourg, vol. 10, pp. 265-268, pl. 1, figs. 4, 4a. (Type locality: Arabistan [= Khuzistan], Iran.) KNOWN RANGE. Khuzistan Province, southwest Iran. A doubtful reeord from southeast Iran (Mertens, 1956), and recorded from west Pakistan (Minton, 1962). MATERIAL EXAMINED (1). Station 4 (CAS 86408 [V/13/58, ¢@]). Remarks. This small gecko was first deseribed from southwestern Iran by Nikolsky (1907). Mertens (1956) tentatively assigned a single, poorly preserved specimen from southeastern Iran to this species, regarding this specimen as congeneri¢ with Tropiocolotes Peters. His specimen lacked dark crossbars on the dorsum. Minton (1962) includes Tropicalotes helenae (sic) in the fauna of west Pakistan, and states that it 1s common. The present specimen agrees with Nikolsky’s description of JMicrogecko helenae in all particulars except that the tail is slender and crossbarred throughout. Nikolsky’s specimens were reported as having thick tails, the distal third being solid blaek. His illustration clearly shows that the tail is regenerated, and it is possible that all nine of his specimens lacked the original tail. CAS 86408 has been deseribed in detail in a previous paper (Anderson, 1961), in whieh I coneurred with Mertens in placing Microgecko in the synonymy of Tropiocolotes. I now feel that while MW. helenae Nikolsky and T. steudneri Peters are congeneric, they are probably generically distinet from 7’. tripolitanus Peters (the type species of T'ropiocolotes ). This tiny gecko was collected on a surfaced road at 8:00 p.m. The road surface was 37.6°C., the surrounding soil 35.2°C., and the air 34.2°C. It was exceedingly agile, able to Jump several inches. VoL. XXXII] ANDERSON: IRANIAN HERPETOLOGY 441 Figure 10. Microgecko helenae. Genus Hemidactylus Oken Hemidactylus persicus Anderson. Hemidactylus persicus ANDERSON, 1872, Proc. Zool. Soc. London, p. 378, fig. 2. (Type locality: probably Bushire, Iran, according to Blanford, 1876.) KNOWN RANGE. Pakistan, Iran, Iraq, Arabia. MATERIAL EXAMINED (9). Road between Station 1 and Station 2 (CAS 86499-86500 [| VIII/20/58, 2 2 |). Road between Station 1 and Station 16 (CAS 86877 [V/12/58, @ ]; 86421 [V1I/14/58, 9 ]). Station 4 (CAS 86414 [ov]; 86415 [9] [V/26/58]; 86424 [VI/24/58, ¢ |; 86454-86455 [VI/13/ Bano ©): Remarks. The females in this series contain ovarian eges; CAS 86377 contains a large ege in each oviduct, the largest 13 mm. The stomach of CAS 86455 contains grasshopper remains almost ex- clusively. These lizards were found at night from May through August on the roads of the foothill region. The air temperatures were 31.4°-36.4°C., the road surfaces 35.4°-38.0°C., and the soil at the road edges 30°-35°C., the road surface usually 3°-5° higher than the surrounding’ soil. This species occurs oceasionally as a house gecko in this region, but less frequently so than Cyrtodactylus scaber. 442 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 41H SER. Genus Phyllodactylus Gray Phyllodactylus elisae Werner. Phyllodactylus elisae Wrenner, 1895, Verh. Zool.-Bot. Ges. Wien, vol. 45, p. 14, pl. 3. fig. 1. (Type locality: ruins of Ninevah, near Mosul, Iraq.) KNOWN RANGE. Western Iran and eastern Iraq. MATERIAL EXAMINED (18). Station 4 (CAS 86432 [V1I/24/58, |). Sta- tion 12 (CAS 86339-86340, 86352 |IV/18/58, oo]; 86435, 86437-86438 [oo]; 86436, 86489-86443 [| VII/6/58] ; 86525-86529 [ VIII/22/58, ¢ dv ]). REMARKS. Stomach contents of this series seem to be exelusively spiders. There were many spiders in the culvert where these lizards were collected on several occasions. These geckoes are quite agile, and proved adept in avoiding capture in the dark culvert. The tails are extremely fragile, and although care was taken in collecting them, few were captured with complete tails. Several individuals were present on the stone walls of the culvert each time it was visited. A small triekle of water ran through the culvert in the sumimer. The lizards were usually in the middle, or darkest area of the culvert. Family AGAMIDAE Genus Agama Daudin Agama nupta de [ilippi. Agama nupta bE Finiprt, 1843, Giorn. Ist. Lomb. e Bib. Ital., vol. 6, p. 407. (Type lo- cality: Persepolis, Iran.) KNOWN RANGE. Pakistan, Afghanistan, Iran, Iraq. MATERIAL EXAMINED (23). Station 1 (CAS 86250 [II/27/57, juv.]; 86321 [III/26/58, juv. & |; 86332 [IV /6/58, juv. &); 86334-86335 [juv. oo]; 86336 [juv. 2] [IV /16/58] ; 86512 [1 X/7/58, juv.]). School build- ing above road between Station 1 and Station 2 (CAS 86508 [ VIII/20/58, juv.|). Road between Station 1 and Station 16 (CAS 86372 [V/12/58, o']). Road between Station 1 and Batwand (CAS 86434 [VII/6/58, 2 |). Station 3 (CAS 86531 [IX/4/58, &]). Station 6 (CAS 86502, 86510 | 2 2 ]; 86504, 86509 [| oo] | VILI/21/58]). Station 7 (CAS 86252 [juv. oi]; S6266n nom) [111/13/58]). Station 9, above Karun River (CAS 86559 [X/11/58, 9 ]). Station 19 (CAS 86623 [XI/4/58, ¢ ]). Station 26 (CAS 86474 | VIII/9/ 58, juv.|; 86476 [juv.]; 86477 [ ¢] | VIII/10/58]). Station 27 (CAS 86511 Xe 58, juve |) Remarks. The spines on the neck, around the ear, and the posterior tem- poral region are absent in the very young specimens. The scales become much more strongly keeled and mucronate with age, and the spines become VoL. XX XT] ANDERSON: IRANIAN HERPETOLOGY 443 increasingly pronounced. The occipital seales are relatively much larger in the voung specimens than in older individuals. The young have distinet dark crossbars in the area between the pectoral and pelvie girdles, these breaking up into less distinct reticulations as the animal matures. Mites cluster in the folds on the neck, and under the imbrieate seales of the adults. These lizards were seen to eat both insects and herbaceous vegetation. The identifiable stomach contents were primarily orthopterous and coleop- terous insects. CAS 86266 and 86559 contained ovarian eges, the largest measuring 3.5 mm. CAS 86502 and 86510 had eggs in the oviducts, measuring up to 26 mm. Thus gravid females were collected in March, August, and October, indicating that eges are laid at least in the spring and in the autumn in this region. The smallest juveniles (44 mm. snout—vent length) were seen in early September, but half-grown individuals were regularly observed from Mareh through November. This species is abundant in the foothill region, inhabiting rock outcrops where there are deep crevices. It is common on walls, cemetery monuments, and buildings. Almost always, one or more of these lizards are to be found living on and about each oil company staff bungalow at Masjid-i-Suleiman. These hang head down on the walls or sereens and make brief forays onto the ground to capture insects. At night and during the hottest part of the day they retreat under the eaves. These lizards were noted occasionally on the trunks of the few trees in the foothills. One was seen in a small evypsum cave. They are present in considerable numbers on the terrace of Persepolis, living in and about the ruins. This is an extremely wary species, retreating quickly into a crevice (al- ways close at hand) when alarmed. Observations (particularly of those liv- ing near buildings) indieate that the area of activity of an individual generally has a radius of less than fifty feet. The basking area, as indicated by the distribution of feeal pellets, is usually not more than five to ten feet in diameter. From February through the summer a very definite shift in the periods of activity was observed. In late February and March this species was seen only at midday and early afternoon, when the air temperature was about 30°C. They were seen then basking on rocks, exposing the maximum surface area to the direct sunlight. In the afternoon they were raised on their fore- legs, facing the sun so that the heht rays would strike the head and belly, and the angle of reflection from the surface of the rock exposed them to the maximum reradiation. Later in the spring basking was initiated progres- 444 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. sively earlier in the day and continued later into the afternoon. In the sum- mer the lizards retired during the hottest hours, their activity restricted to the earliest daylight hours and to late afternoon in July and August. The anal temperature of a basking individual at 6:35 a.m. on August 20 was 27.2°C. when the air was 22.4°C. and the substrate 25.5°C. By mid-September they were again active until later in the morning and retired only during the midday heat, becoming active again early in the afternoon, when air temperatures were still over 40°C., and the rocks on which they were seen considerably higher. In mid-October they were active at midday. Observations to determine the critical maximum temperature for this species were carried out with seven individuals (see p. 436). t ls ‘ina Bio! oe 3] I Bal iar le D Ci -¢ y | ees A I Y > n (05° i i LANHSC Ln] £ NA if ge W OODS HULE, MASO. - — aa 500 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. pupil horizontal; maxillary teeth present; no odontoids on mandible; vo- merine teeth in two small, distinct patches lying well behind the small choanae; tongue elongate, entire and free behind; fingers free of web, toes with but the merest vestige of web at the base; cranial derm free of skull; root of skull not exostosed. Niceforonia nana Goin and Cochran, new species. (Figure 1.) Typr. Museo de La Salle 142, adult female, Colombia, Santander, Paramo de La Rusia, collected by Hno. Nicéforo Maria. —rS>) FIGURE 1. Niceforonia nana, type, Museo de La Salle 142, Colombia, Santander, Paramo de La Rusia. Actual length, 20.5 mm. x 8. VoL. XXXT] GOIN AND COCHRAN: NEW GENERA OF FROGS 501 DESCRIPTION OF TYPE. No bony ridges on top of head. Vomerine teeth in 2 short, slanting, narrowly separated series far behind the choanae; tongue 14 as wide as mouth opening, elongate, broadly oval, its posterior border free and unnoteched; snout moderate in length, rounded when viewed from above, slanting forward in profile, the upper jaw extending very slightly beyond the lower. Nostrils small, scarcely projecting, their dis- tance from end of snout a trifle greater than their distance from eye. Canthus rostralis strong, curved; loreal region weakly coneave, slanting outward to the upper lip. Eye rather small, but with very thick lids to give it additional prominence, its diameter about +5 its distance from tip of snout; interorbital diameter 114 times that of upper eyelid, equal to interval between nostrils. Tympanum concealed. Fingers moderately long, with faint lateral ridges, free, their tips scarcely enlarged, first a little shorter than second, which equals the fourth, both of these extending to base of terminal phalanx of third; a distinet oval thumb pad present, also one at base of second finger; a rounded palmar callus; metacarpal tubercles well developed. Toes fairly long, with little trace of web, third longer than fifth, its tips reaching to middle of antepenultimate phalanx of fourth, the tips of all the toes slightly enlarged and flattened above; a distinct, oval, inner metatarsal tubercle, and a smaller, rounded, outer one; no apparent tarsal ridge; a series of heavy skinfolds on heel and knee. Body very stout, in postaxillary region much wider than greatest width of head. When hind leg is adpressed, heel reaches front of shoulder; when limbs are laid along the side, knee and elbow are widely separated; when hind legs are bent at right angles to body, heels are narrowly separated. Skin of upper parts finely granular, with narrow, interrupted, lateral, glandular line between posterior border of eye and sacrum; some heavy tubercles seattered over posterior third of back; sides heavily granular; venter smooth on throat, chest and anterior half of belly, becoming granular on posterior part of belly, on posterior femur and below vent; a short, thick, glandular ridge from lower posterior corner of eye, dropping behind corner of mouth, and continuing as a series of 2 or 3 small glands along side of neck, ending on proximal anterior part of humerus; a strong skinfold across the chest; a very distinct ventral disk. Dimensions. Head and body, 20.5 mm.; head length (to end of upper jaw), 7 mm.; head width, 7.5 mm.; femur, 8 mm.; tibia, 7.5 mm.; foot, 8 mm.; hand, 5.5 mm. CoLor (in aleohol). Dorsum seal brown to walnut brown, lightening on sides to wood brown; venter wood brown, immaculate except for some 502 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. darker shadings between the granules on posterior belly; side of head im- maculate fawn color, becoming a little darker near canthus; anterior femur seal brown on upper half, light wood brown on lower half; posterior thigh Mars brown with one or two paler spots; soles of feet and palms of hands pale wood brown, their tubercles and webs similar in color. Discussion. Just what the relationships of Niceforonia are is at pres- ent a moot question. In many ways it is structurally more like the forms of Borborocoetes which lack the tympanum. When first examined the type specimen gives the impression of being immature, but there can be no doubt that it is an adult for it is a female packed with large ripe eggs. The large unpigmented eges (the individual eggs are approximately 2.00 mm. in diameter) probably indicate that Niceforonia is a terrestrial breeder. This would not be unexpected or inappropriate for a species liv- ing in the paramos. It may be that this species, the type of which was collected in the Paramo de La Rusia, is restricted to the paramos. The other new form was collected 20 miles east of Pasto, Narifo. While the collector, E. S. Ross, gave no more precise data, this locality is cer- tainly in a mountainous district and may well be on a paramos. Trachyphrynus Goin and Cochran, new genus. TYPE SPECIES. T'rachyphrynus myersi, new species. Dracnosis. { \ uy \, na a ; re , : aS = = ~ “4 { ay Eh e8s | . vos, 19 Jag jv al , ‘s f & -, = hy ] . 4 ; ; » a sen . § at ki AP leery ‘ rf 1 , \ * 5 » i : 7 : ar} y oy (é a es. i Ao kc: J a es i : : Mf, = Poe " aes: . - { % : . 7 ; ® i text fig: 3; 1905, pp. 207-208, pl. 1, figs. 14, 14a, 15, Iba; text fig. 32; 1940, pp. 279-280, pl. 30, fig. 40; 1944, pp. 29. Corr®a, 1953a, pp. 130-134, pl. 1, figs. 1-2; 1955, pp. 68-72, pl. 1, figs. 1-3; pl. 2, figs. 4-6. GRIFFIN, 1898, pp. 207-209, figs. 17-18, as Emplectonema viride. IstER, 1901, p. 274. Iwata, 1954, p. 15. Jousrn, 1890, pp. 585-586, pl. 31, figs. 15-16, 21, as Nemertes gracilis. RICKETTS and CALVIN, 1956, pp. 43, 170, 206. StreEPHENSON, 1913, pp. 12-138, fis. 6. STIMPSON, 1857, p. 163, as Hmplectonema viride. YAMAOKA, 1940, pp. 237-238, pl. 4, figs. 1-2; text fig. 14. MATERIAL. Several worms from March to August, 1958. Loealities 1, 2. Very common. DistrIBuTION. Northern coasts of Europe to Madeira; Mediterranean; Aleutian Islands and coast of Alaska to Ensenada, Mexico; Chile; Kam- chatka to Japan. Intertidal zone to 100 m. The largest worm measured in life up to 20 em. in length and 1 mm. in width. The color of the back is uniform dark green. The ventral side is pale yellow or whitish. The color and aspect of the Mediterranean and Cali- fornia worms are not completely alike. The former are stronger and their color is paler than in the California worms (Birger, 1895, pl. 2, fig. 1; Coe, 1905, pl. 1, fig. 14). The color is still present after preservation in hot “Susa.”’ The head flattened dorsoventrally, wider and lghter than the trunk, contains numerous eyes distributed in two rows on each side: one antero- lateral with 12-14 eyes and the other posterolateral with the same number. 536 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Young worms have fewer eyes distributed irregularly. Cephalic furrows are not distinct. The worms have a tendency to coil into a spiral in life and during preservation. The epidermis is high and rich in mucous glands. The worms make a mucous tube with the fine particles of mud existing between the mussel beds, their most common habitat. The basement membrane is thick and at certain levels thicker than the external circular muscle layer. The longitudinal muscles are high. Both layers reach the tip of the head. Anteriorly to the brain there is a dense muscular net where abundant dorsoventral muscular fibers oceur. The cephalic glands are voluminous, distributed dorsally, ven- trally, and laterally, reaching as far back as the posterior level of the stomach. The esophagus opens into the rhynechodaeum. The stomach is large, lone, and villous, and its ciliated epithelium is rich in eyanophilous glands. There are two long anterolateral intestinal pouches which join at the begin- ning of the gut on its sides. The rhynchodaeum is surrounded by the cephalic glands. The proboscis septum belongs to the closed type. The rhynchocoel has about 1/3 of the body length. The rhynehocoel wall is composed of a flat epithelium, a layer of longitudinal muscles and a layer of circular muscles. There is a central, curved stylet surrounded by abundant diaphragmatic glands. The wall of the anterior proboscis chamber is composed of a high external epithelium, a layer of circular muscles, a thick layer of longitudinal muscles, and the flat internal epithelium. In front of the brain there are two blood vessels united by a large com- missure. Posterior to the brain there are three blood vessels, one dorsal, situated between the rhynchocoel, and the intestine, and two lateral ones. The dorsal vessel originates from the right lateral vessel. The nephridia are well developed at the posterior level of the brain and the first part of the stomach. There are two pairs of cerebral ganglia united by the commissures. The lateral nerve cords have only one core of fibers. The cerebral sense organs are situated ventrolaterally and far anteriorly to the brain. Besides EF. gracile, another species of Emplectonema, E. biirgeri Coe, 1901, is expected to occur on the mussel beds and other growths in the inter- tidal zone of the localities 1 and 2 where I have collected E. gracile. Emplec- tonema biirgeri can be separated trom E. gracile by the following char- acters: it is up to 1 m. long; dark velvety brown above and creamy-white below; the base of the stylet is short, only a little longer than the central stylet; the latter is weak and straight. Occurrences: Alaska to Monterey Bay, California; Japan. Intertidal zone to 500 m. VoL. XXXII] CORREA: NEMERTEANS; CALIFORNIA and OREGON 537 Genus Paranemertes Coe, 1901. REFERENCE: FRIEDRICH, 1955, p. 178. Up to 50 em. long, more or less stout; two to numerous eyes; subepithe- lial glands present, also behind the brain, sometimes reaching region of median gut; longitudinal musculature double layered in anterior part of body, internal and external layer separated from each other by a layer of parenchyma, farther behind the parenchyma disappears and the lonei- tudinal museular layer is uniform; cephale glands large but not reaching beyond brain; esophagus opens into rhynchodaeum; caecum of median gut short but with paired pouches; rhynehocoel 1/2 or 1/3 of body leneth; lateral nerve cords with only one core of fibers; cerebral sense organs small, anterolaterally to brain. Paranemertes peregrina Coe, 1901. PRINCIPAL REFERENCES: Cor, 1901, 1904, pp. 33-36, pl. 2, fig. 6; pl. 3, fig. 5; pl. 7, fig. 7; text fig. 7; 1905, pp. 220-224, pl. 1, figs. 7-9; pl. 16, figs. 95-96; pl. 17, fig. 103; pl. 24, fig. 192; pl. 25, figs. 189-199; text figs. 37-38; 1940, p. 286; 1944, p. 29. Twata, 1954, p. 15. Ricketts and Cavin, 1956, pp. 146-147, 206. YAMAOKA, 1940, pp. 240-243, pl. 17 (in text 4), figs. 3-6; text figs. 17-19. MATERIAL. Several worms from March to August, 1958. Loealities 1, 5. This is one of the commonest and most widely distributed species of nemer- teans on the Pacific coast. Very restless, frequently it was found on cloudy days crawling about on the beaches between tides. It occurs from low-tide level well up toward high-water in every habitat, under stones, among seaweeds, barnacles, mussel beds, and on seagrass in mudflats. DisTRIBUTION. Commander Islands; Aleutian Islands; Alaska to Gulf of California, Mexico; Kamehatka to Japan. Intertidal zone and farther below. The worms measured in life up to 20 em. in length and 4 mm. in maxi- mum width. The head is very variable in shape, commonly fan-shaped, flattened dorsoventrally, and a little wider than the adjoining portion of the trunk. The color is also very variable, but the commonest is uniform purplish or dark violet on the back and whitish-vellow on the ventral sur- face as well as on the margins. The width of the ventral yellow surface is in some worms only a narrow longitudinal stripe, the margins being also purple colored. The color is well preserved after fixation in hot ‘“Susa.” The purple head is bordered in front and laterally by the lighter color of the ventral surface. The same color occurs at the posterior border of the head as a small angular spot on each side. Numerous small eyes are arranged in groups on the head. Two groups are situated on the anterolateral mar- eins and the other two are disposed in clusters just in front of the brain. 538 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. There is one transverse cephalic furrow, V-shaped, with its apex directed backward. The glandular epidermis shows the purple pigment of the body in its deepest zone. Before the brain there is a dense muscular net besides the normal muscular layers underneath the epidermis. The cephalie glands are very abundant, scattered among the muscular net fibers and reaching the nephridial region backward. In the anterior part of the body the longi- tudinal muscles form an internal and an external layer separated by parenchyma. Farther behind the parenchyma disappears. The esophagus opens into the rhynchodaeum. There are two anterolateral intestinal pouches which reach the posterior border of the brain. They unite under the stomach forming a short median caecum. The rhynehodaeum opening is a subterminal slit situated ventrally to the frontal sense organ. The rhynchodaeum has a large lumen lined by a high epithelium. The strong proboscis septum, connected with the anterior muscular net, belongs to the closed type. The rhynechocoel reaches about 3/4 the length of the body. Its wall is composed of a flat epithelium, a layer of longitudinal muscles, and a layer of circular muscles. The wall of the anterior proboscis chamber is composed of a high external epithelium, a layer of circular muscles, a layer of longitudinal muscles, and a flat internal epithelium. The 14 longitudinal proboscidial nerves are located on the outer border of the longitudinal muscular layer. The central stylet is small, slender, and sharply pointed as are the accessory ones. All stylets have a braided appearance. The base is very small and pear-shaped. The number of accessory stylet pouches is 2, and each contains from 4 to 6 stylets. There are a pair of long and large cephalic blood lacunae in the head joined anteriorly by an anastomosis. The three longitudinal posteerebral vessels are well developed, being two lateral and one dorsal, situated between the rhynchocoel and the gut. The nephridia occupy the region behind the brain and are situated above, below, or lateral to the lateral nerve cords. They are provided with numerous branches. The brain is very simple. It is composed of two pairs of ganglia. The lateral nerve cords have only one core of fibers. The cerebral sense organs are small, simple, and lie well in front of the brain. They open into a slight furrow on the sides of the head, a little in front of them. A frontal sense organ is present. KEY FOR THE SPECIES OF Paranemertes a1 ey O tO GO ieee ae 2 Sea A 2 a, ee «ee ee P. biocellata Coe, 1944. No record is available of coloration in life. Occurrence: Biloxi, Mississippi. Burrowing in intertidal sand flats and in shallow water. VoL. XXXT] CORREA: NEMERTEANS; CALIFORNIA and OREGON 539 EEA NT OOH Chia es oO CCl ier ee ee aed se gt ee ee ce ek) oe eee eee 7 Dd NYA El OU gt brake aS ECON cre tein (pl 01,2 oS He Mages eens se NN ee eee eee ee eee 3 EVV EOC TEL ATISIVET SC pill Gusto em teen ce Sie saree eee ee ee ee eee 4 3. Transverse rings as very minute dark brown spots; 12 proboscidial nerves; number of ocelli not known. One can infer from the description of the species (Iwata, 1952, pp. 142-143) that it has more than 2 eyes as it resembles P. CALC Oe 1U ea Wel CEG TY DTA: CO ees eae re ta ere re tee a eee een gee we Skene cerns nee ee P. incola Iwata, 1952. Bright chestnut brown anteriorly, yellow ventrally and marginally. Occurrence: Tomioka, Japan. Under stones near low water-mark. eran sVvierseurines, swihitishi) tene prOWOSCIGMal me TVS ese rere pare Bree i.e on Pee SF P. plana Iwata, 1957. Pale brown with numerous whitish dots besides the rings. Occurrence: Sagami Bay, Japan. From 250 to 300 m. Collected by His Majesty the Emperor of Japan. 4. Stylets with braided appearance; 14 proboscidial nerves; 2 pouches of accessory SHEN AU] ASF eS ee Saree ee ee er et ee ee P. peregrini Coe, 1901. — Stylets not braided in appearance: less than 14 proboscidial nerves; more EH anerom OMCs = Ol maCCESSOI yee trys CUS eee ee ee ee 5 5. Opaque white dorsally and ventrally; 9 to 10 proboscidial nerves; 4 pouches Olea CCESSORYVaS tLe t Ses ae ee IN ce ce se ee ee ce 175 Sasa 5 P. pallida Coe, 1901. Occurrence: Alaska. Intertidal zone. — Not opaque white dorsally and ventrally; variable number of proboscidial nerves ING SSD OL CIES Ota CCOSS OT Vi eS UV US rer ee ee eee ae en 6 6. Pale red dorsally and ventrally; 11 or 12 or 13 proboscidial nerves; 6 to 12 POUCHES HOMMACCESSORY: Sty letse swe. mA k oe ee ee ee re ee eee eae _P. carnea Coe, 1901. Occurrences: Alaska to Puget Sound. Intertidal zone. — Rosy or pinkish anteriorly, often tinged with yellow or orange, intestinal region gray or salmon; 10 to 13 proboscidial nerves; 2, 4, or 6 pouches of ACCESSOnYs StUYIECUS (2 ee eet 2 eee P. californica Coe, 1904. Occurrences: Monterey Bay, California to Ensenada, Mexico. In sandy and muddy flats. BIOLOGICAL OBSERVATIONS AND EXPERIMENTS. The worms were maintained in sea-water at a temperature between 12 to 15° C. They endure better if some fragments of algae are added. These serve as shady shelter and sub- stratum. In general the worms lasted for about 15 to 20 days under these conditions without food. Their calm and uniform eliding is accompanied by movements of the tip of the snout to both sides or hfting it. Sometimes they contract rapidly, distend, and indulge in corkscrew and antiperistaltie movements. After about 30 minutes, all worms were sheltered under the pieces of algae. Acephalous fragments (Friedrich, 1933; Eggers, 1924, 1936; Corréa, 1953, 1953a) decapitated 5 em. behind the anterior extremity, do not show any locomotion beyond muscular contractions which could produce a ehange of place. Only very strong stimuli could bring the peripheral 540 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. nervous elements to give impulses to the cilia. Cephalic fragments, whieh contain the brain, behave like complete animals in regard to locomotion. Immediately after the cut, as the cilia continue to beat, they start a calm and uniform locomotion in a rectilinear path. After 24 hours they still are able to present the same sort of locomotion if the water is changed. The ae- cumulation of mucus on the bottom of the dishes impedes the spontaneous oliding. Paranemertes peregrina belongs to the locomotory type a (Corréa, 1953, p. 548, 554). Spontaneous eliding depends on the brain. The threshold of stimulation of the nervous elements in the posteephalie part is much higher than that of the brain. The worms are negatively phototactic. When in the nearest corner to the light souree of square dishes they elide to the farthest corner. If they have the anterior end directed against the light source they turn round before they glide. The path is very seldom diagonal, as the worms are posi- tively tigmotactie to the borders of the dish. Both paths, the diagonal and the bordering, took the same time to be performed. The same reaction was shown by complete worms and cephalic fragments. Paranemertes peregrina represents excellent material for classes experi- ments on food intake in nemerteans, as was already described and illustrated by MaeGinitie (1949, p. 162, f. 43). Besides being one of the commonest intertidal species of nemerteans on the California coast, they are voracious feeders (Coe, 1901, 1904, p. 36) and are easily fed in the laboratory with fragments of polychaetes. Genus Zygonemertes Montgomery, 1897. REFERENCE: FRIEDRICH, 1955, p. 154. Many eyes, extended backward beyond brain, over and beside the lateral nerve cords, and, specially behind the brain more or less serially disposed; anterior part of body with numerous subepithelial glands (?); esophagus opens into rhynehodaeum; median gut without unpaired caecum directed forward, substituted by two long paired pouches (evidently Z. capensis Wheeler, 1934 and Z. glandulosa Yamaoka, 1940, with caecum) ; rhynechocoel reaching the posterior end, its wall double-layered; proboscis thick and short, anterior part of it with only one layer of circular muscles; base of central stylet long, cylindrical, posteriorly truncate, in front of the posterior end generally an annular furrow; lateral nerve cords with one fiber core; anal commissure dorsal to gut; cerebral sense organs large, imme- diately in front of or near the brain (?), clearly separated from it. Zygonemertes virescens (Verrill, 1879). PRINCIPAL REFERENCES: CoE, 1905, pp. 214-216, pl. 22, figs. 141-144; 1940, pp. 295- VoL. XXXT] CORREA: NEMERTEANS; CALIFORNIA and OREGON 541 296, pl. 30, fig. 39; 19438, pp. 270-273; text figs. 63-64; 1944, p. 30; 1951, p. 329; 1951la, pp. 170-171, text figs. 16a—e. Corriéa, 1961, pp. 25-28. Monracomery, 1897, pp. 2-4, pl. 1, figs. 14-15, 238-24, 28. VerRrixy, 1892, p. 400, pl. 33, figs. 4-4e as Amphiporus virescens; l.c., pp. 400-401, pl. 35, fig. 4; text fig. 4, as A. agilis. MATERIAL. Three worms in September, 1958. Locality 10. Not common. DisTRIBUTION. Bay of Fundy, New England, and southward to Miami and Key Largo, Florida, and along the Gulf coast at least as far as Pensa- cola, Florida; Puget Sound to Gulf of California. Intertidal zone to 120 m. The living mature worms are up to 3 em. long and 1 mm. wide. The color is variable from white to yellow and green. There are numerous precerebral eyes disposed in irregular rows and postcerebral ones disposed in only one row on each side. In living worms, flattened between slide and cover-slide, numerous gen- erally sickle-shaped bodies appear, first described in Emplectonema echino- derma (Marion, 1830) (Biirger, 1895, p. 124, 216). At a first glance they seem to be calcareous corpuscles. Probably they are extra-cellular secretions, colorless or pigmented, originated from epidermal cells (Coe, 1943, p. 217). The sickle-shaped bodies ean also be seen in clarified worms and in sections, as they do not dissolve in the liquids commonly used for preservation. The large rhynchodaeal pore leads to a very narrow rhynechodaeum. The proboscis septum belongs to the closed type. The rhynechocoel reaches the posterior tip of the body. The proboscis is much shorter. The rhynchocoel wall is composed of an epithelium, a thick layer of longitudinal muscles, and a thin laver of cireular ones. The proboscis wall is composed of a high external epithelium, circular muscles, a double layer of longitudinal muscles, circular muscles, and internal epithelium. There are ten proboscidial nerves. The central stylet is shorter than its base which is truneate and posteriorly lobate. OTHER SPECIES OF THE GENUS. The genus Zygonemertes is quite uniform in its features. Besides size and color the best character to separate its eleven known species (Corréa 1961, p. 26) is the number of proboscidial nerves. Two more species occur on the West coast of North America. They can be separated from virescens as follows: 1. Zygonemertes thalassina Coe, 1904. Closely related to Z. virescens; 12 proboseidial nerves. Occurrence: Alaska. Intertidal zone. 2. Zygonemertes albida Cor, 1904. Number of proboscidial nerves unknown; whitish, small, only up to 25 mm. long. Occurences: British Columbia to Ensenada, Mexico. Intertidal zone. 542 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Genus Amphiporus Ehrenberg, 1831. REFERENCE: FRIEDRICH, 1955, p. 154. Generally large, stout forms; seldom none, 2 or 4 eyes, generally many to very many, distributed in groups; cephalic and subepithelial glands pres- ent or lacking; cutaneous-muscular tube with or without diagonal layers; dorsoventral muscles generally present; esophagus opens into rhyneho- daeum; caecum with pouches of different length; without diverticula of esophagus, stomach or pylorus; rhynehocoel as long as body; nephridial apparatus probably always limited to a short streteh behind the cerebral ganglia; cerebral sense organs anterior to brain, maximally reaching under its anterior border, sometimes small, in the tip of the head, sometimes larger and nearer to the brain (the A. pulcher-group differs essentially in this point). Amphiporus cruentatus Verrill, 1879. PRINCIPAL REFERENCES: Cor, 1904, pp. 154-155, pl. 20, figs. 1-6; 1905, pp. 246-247, pl. 1, fig. 13; text figs. 12, 20, 51-52; pp. 279-280, pl. 22, fig. 161, as A. leptacanthus ; 1940, p. 299; 1943, pp. 279-281, pl. 3, fig. 1, text figs. 66-67; 1944, p. 30. VERRILL, 1892, pp. 399-400, pl. 33, figs. 7-8a; pl. 35, fig. 3, pl. 39, fig. 9. MareriaL. Two worms in April, 1958. Loeality 1. Uncommon. Distrisution. New England to Florida; Puget Sound to San Diego. California. Intertidal zone to 80 m. The living, mature worms are up to 25 mm. in length. The head is slen- der, not wider than the rest of the body, with indistinet cephalie furrows. The color is whitish or pale yellow. There are only a few eyes, about 15, distributed irregularly in two rows, one on each lateral margin of the head. The size of the eyes is variable, the most anterior being the largest ones. The rhynchocoel is almost as long as the body. The proboscis is also very long and thick. The slender and pointed central stylet is as long as the base. The base is irregularly eylindrical in shape. There are two pouches of accessory stylets each having two to four stylets. The most striking feature of this small species of Amphiporus is the deep red color of the blood, easily seen by transparence through the large vessels in living animals. There are two precerebral vessels united anteri- orly by a commissure and three posteerebral ones, two lateral and a dorsal one originated from the right lateral vessel at the level of the brain. Amphiporus imparispinosus (Griffin, 1898. PRINCIPAL REFERENCES: Cor, 1901, 1904, pp. 51-53, pl. 7, fig. 6; text fig. 13, as A. leuciodus; 1904, p. 115; 1905, pp. 247-249, pl. 16, figs. 99-100; pl. 25, figs. 195- VoL. XXXT] CORREA: NEMERTEANS,; CALIFORNIA and OREGON 543 197; text fig. 53; 1940, p. 300; 1944, p. 30. Grirrin, 1898, p. 210, text figs. 19-20. PUNNETT, 1901, p. 95, as A. leuciodus. MATERIAL. Several worms from March to August, 1958. Locality 1. It is one of the commonest nemerteans in the Dillon Beach area. DISTRIBUTION. Coast of Siberia; Bering Sea; Alaska to San Diego, Cali- fornia and Ensenada, Mexico. Intertidal zone to 50 m. The slender, elongated worms measured in life up to 20 em. in length. The head is narrower than the following part of the body which is flattened posteriorly. The color is uniformly opaque white, sometimes with a pinkish or yellowish tinge. There are two groups of eyes on each side of the head. The anteromarginal groups form elongated rows of 10-12 eyes each, and the posterior groups, nearer the median line, have about the same number of eves. The glandular epidermis is moderately high. The basement membrane is thick. Both layers of muscles, the external cireular and the thicker internal longitudinal one, attain the tip of the head. Before the brain there is a dense net of muscular fibers. The cephalic glands are not very abundant. They reach backward only the posterior level of the brain. The large esophagus opens into the rhynechodaeum at the level of the cerebral sense organs. There are two laree, branched, anterolateral intestinal pouches which reach forward as far as the brain region. They unite beneath the stomach and form a median caecum provided with lateral diverticula. Also the main gut is provided with diverticula. The rhynechodaeum opens anteriorly by a large opening. It is lined by a high epithelium and contains a thick muscular coat in its most internal part. The strong proboseidial septum belongs to the closed type. The rhynechocoel is long, reaching almost the posterior end of the body. Its wall is composed of an epithelium, a laver of longitudinal muscles and a layer of circular muscles. The proboscis wall is composed of a high ex- ternal epithelium, a laver of cireular museles, a thick layer of longitudinal muscles and a flat internal epithelium. There are 16 longitudinal probos- cidial nerves, situated in the longitudinal muscular layer. The central stylet is almost as long as the very wide base. The number of pouches of accessory stylets was three in all specimens examined. The number of accessory stylets was 2-3 in each pouch. There are two precerebral vessels united anteriorly by a commissure and lined by a high epithelium. Posterior to the brain there are three ves- sels, two lateral and a dorsal one. The nephridia are well developed. The brain is composed of two pairs of ganglia united by a broad and short ventral commissure and a long and narrow dorsal commissure. The lateral nerve cords have only a core of fibers. The cerebral sense organs 544 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. are composed of a series of chambers which open to the exterior by one lateroventral cerebral canal and aperture. They he far in front of the brain. A frontal sense organ is present. Amphiporus formidabilis Griffin, 1898. PRINCIPAL REFERENCES: Cok, 1901, 1904, pp. 54-56, pl. 3, fig. 1; pl. 7, fig. 5; pl. 11, fig. 3; text fig. 14, as A. exilis; 1904, p. 115; 1905, pp. 250-252, pl. 17, fig. 101— 102; text figs. 13, 15, 23, 54, 1940, p. 300, 1944, p. 30. GrirFIN, 1898, p. 211, text figs. 21-23. Iwara, 1952, pp. 144-146, text figs. 15-16. MATERIAL. Several worms from March to August, 1958. Localities 1, 8. Very common. DistrisuTION. Bering Island, Aleutian Islands, coast of Alaska and southward to Monterey Bay, California; Japan. Intertidal zone and below. The slender worms measured in life up to 30 em. in length and 2-3 mm. in maximum width. The commonest color is uniform opaque-white, some- times pale yellowish-white. The intestinal canal, brain, and lateral nerve cords can be seen by transparence. The eyes, very small but extremely nu- merous, up to 250, are distributed in four clusters, two anterior and two posterior in the head. Before the brain there is a dense muscular net. The cephalic glands are very well developed all around the rhynchodaeum. I could not see their posterior limits because there occur abundant submuscular glands at the brain level which show the same aspect as the cephalic ones. The large esophagus opens into the rhynchodaeum. There are two branched anterolateral intestinal pouches which reach forward to the brain region. They unite beneath the stomach to form a branched and large median caecum. The large rhynechodaeum is lined by a high epithelium. At its internal end it is coated by a thick muscular laver. The extremely strong probosecidial septum belongs to the closed type. The rhynchocoel is long, reaching almost the posterior end of the body. Its wall is composed of an epithelium, a longi- tudinal muscular layer, and a circular one. The proboscis is of moderate size. The wall of its anterior chamber is composed of an external epithelium, a layer of circular muscles, a thick layer of longitudinal muscles and a flat internal epithelium. There are about 25-30 proboscidial nerves situated within the longitudinal musculature. The central stylet is slender but its base is very massive. There are usually 6-12 pouches of accessory stylets each bearing 1-2 stylets. The nephridia are remarkably long and well developed. The brain is large. The lateral nerve cords have only one core of fibers. VoL. XXXII] CORREA: NEMERTEANS; CALIFORNIA and OREGON 545 The cerebral sense organs are situated far in front of the brain, ventro- laterally as well as their canal and opening. A frontal sense organ is present. OTHER SPECIES OF THE GENUS. Eighteen species and one variety of Amphiporus are known on the Pacifie coast of North America (Coe, 1944, pp. 30-31). Besides the three described here, eight more and the variety could oceur in the intertidal zone of the localities where I collected. They can be separated by the following characteristics: He Amphiporus angulatus (Fabricius, 1774). Length 20 em. or more; dark brown, reddish-brown or purplish-brown on dorsal surface, with paler margins and conspicuous angular whitish spots on each side of head, continuous with whitish ventral surface; a pair of elongated clusters of ocelli situated on the anterolateral borders of the head and a posterior eroup lies in or near the angular white spot on each lateral margin of head. Occurrences: Circumpolar; Greenland; Davis Strait; Labrador to Cape Cod; Bering Strait; Bering Sea; Aleutian Islands and Kamchatka to Japan; Alaska; British Columbia and southward to Point Conception, California. Intertidal zone to 150 m. or more. . Amphiporus tigrinus Coe, 1901. Length 10 em.; color of various shades of lemon, yellow, and vellowish-brown; ocelli irregular in shape and variable in size arranged in two irregular and indistinctly separated clusters on each side of head. Occurrences: British Columbia and Puget Sound. Intertidal zone. . Amphiporus bimaculatus Coe, 1901. Up to 15 em. long; dorsal surface, behind the head, deep red, brownish-red or brownish-orange, sometimes yellowish; head whitish with two angular or oval black or very dark brown spots placed symmetrically on dorsal surface; ventral surface pale reddish, or pale orange; ocelli arranged in an irregular, elongated cluster on each anterolateral margin. Occurrences: Sea of Okhotsky; Alaska to Ensenada, Mexico. Intertidal zone to 250 m. Amphiporus fulvus VCoe, 1905. Leneth 15 to 25 mm. or more; pale brownish anteriorly, deep brown in intestinal region, much paler beneath; head pale brown; color of body consists of innumerable minute dark brown spots scattered thickly over the ground color. Occurrence: south- ern California. Intertidal zone to 85 m. Amphiporus californicus Coe, 1905. Length 10 to 25 mm. or more; deep red orange with an opaque white tinge, dark, dull yellowish red, or bright red; ventral surface duller and with more whitish. Occurrence : southern California. Intertidal zone to 80 m. or more. 546 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 6. Amphiporus punctatulus Coe, 1905. Between 35 and 50 mm. in length; in preserved specimens the whole surface of the body is dark mottled brown, the color consisting of innumerable dark dots which run together, forming irregular blotches; the mottling is less complete on the ventral surface. Occurrence: Catalina Island, California. Intertidal zone. =I Amphiporus rubellus Coe, 1905. Length 25 mm. or more; pale orange, or pale red; much paler and usually grayish beneath. Occurrence: south- ern California. Intertidal zone to 200 m. 8. Amphiporus flavescens Coe, 1905. Up to 20 mm.; color very variable, usually whitish, or pale yellow; there are commonly 12 to 25 ocelli ar- ranged in two groups on each side of head. Occurrences: Monterey Bay, California to Ensenada, Mexico. Intertidal zone. 9. Amphiporus imparispinosus Griffin, 1898 var. similis (Coe, 1905). Differs from the species in having two instead of three pouches of accessory sty- lets. Occurrences: Puget Sound to Ensenada, Mexico. Intertidal zone and below. Genus Tetrastemma Ehrenberg, 1831. REFERENCE: FRIEDRICH, 1955, p. 170. Generally small, slender worms; tip of head with circular and longi- tudinal muscles as well as retractors; cephalic glands present, very variable in size, sometimes reaching far beyond brain; esophagus opens into rhyn- chodaeum; caecum of midgut present, with lateral pouches and two more pouches directed forward; pouches of midgut shallow, generally not branched; rhynehocoel as long as body (except 7. hansi Biirger, 1893), with- out diverticula; precerebral septum (as far as known) closed; nephridia short, generally with one or two pores; lateral nerve cords with one fiber core; cerebral sense organs generally large situated in front of brain, ex- ceptionally in cerebral region; generally dioecious (except T. marioni Joubin, 1890 and 7. caecum Coe, 1901). Tetrastemma nigrifrons Coe, 1904. PRINCIPAL REFERENCES: Cok, 1904, pp. 159-164, pl. 15, fig. 7; pl. 16, figs. 6-9; pl. I, es. Ls ply 20; fies 165 pl 2, fies: 15=29)) 1905, pps 289-293) la hoe oops 18, figs. 111-115, text figs. 57-59; 1940, pp. 305-306: pl. 31, fig. 42; 1944, p. 31. IwaTa, 1954, pp. 30-32, fig. 8 B 1-8; 1957, p. 27, pl. 1, fig. 14. Yamaoxa, 1940, pp. 249-251, pl. 3, fig. 14; figs. 9-12; textfigs. 26-29, as Prostoma nigrifons. MaTeErRIAL. Two worms in May, 1958. Locality 1. Uncommon. VoL. XXXII] CORREA: NEMERTEANS; CALIFORNIA and OREGON 547 DISTRIBUTION. Puget Sound to coasts of Mexico and Costa Rica; Japan. Intertidal zone. Both worms were 30 mm. long. Variations in color and markings are so striking that several species appear to be represented by individuals which actually present all degrees of intergradation (Coe, 1905, p. 290). My worms had a whitish head with a large, quadrangular dark-brown dorsal marking and a purplish-brown trunk. Their coloration is close to that given by Coe (1904, 1905, 1940) for the variety “purpuratum.” Four eyes are situated one on each corner of the head marking. OTHER SPECIES OF THE GENUS. Besides 7’. nigrifrons, five more species of Tetrastemma could occur in the intertidal zone of the localities where I have collected. They can be separated by the following external character- istics: 1. Testrastemma candidum (Miiller, 1774). Recently redeseribed by Corréa (1961, p. 40) is up to 20 mm. long and uniformly whitish or pinkish- white in color. Occurrences: Circumpolar; Greenland and Norway to Madeira; South Africa; Labrador to New England and southward to Miami and Key Largo, Florida; Alaska to Ensenada, Mexico. Inter- tidal zone. i) Tetrastemma bilineatum Coe, 1904. Only 5 to 10 mm. long; general color creamy or grayish with two sharply defined brown stripes extending nearly along whole leneth of dorsal surface. Occurrence: San Diego, California. Intertidal zone. 3. Tetrastemma quadrilineatum Coe, 1904. Long 8 to 12 mm.; general color of body whitish with four longitudinal brown stripes, two of which are situated near lateral margins of body and the other two placed on dorsal surface. Occurrences: Monterey Bay, California to Ensenada, Mexico. Intertidal zone. 4. Tetrastemma reticulatum Coe, 1904. About 8 to 15 mm. long; ground color white with rectangular and longitudinal brown markings; head provided with a brown marking. Occurrence: southern California. Inter- tidal zone. 5. Tetrastemma signifer Coe, 1904. About 15 to 25 mm. long; general color of body deep reddish-brown or purplish except for the head which is white with a dark brown marking formed by a transverse, basal portion, from which two semicircular branches pass anteriorly. Occurrences: Monterey Bay, California to San Diego, California. Intertidal zone to 10 m. 548 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Genus Prostoma Dugés, 1828. REFERENCE: FRIEDRICH, 1955, p. 162. In general small, up to 3.5 em. long; none to six eyes; muscular-cutane- ous tube without diagonal layer; tip of head with longitudinal muscles; cephahe glands short, only precerebral; midgut without or with very short unpaired blind pouch; only with two pouches directed froward; special pylorus tube not developed; true esophagus absent; rhynchocoel generally not reaching the posterior end, its wall musculature not crossed; blood vessels without brain commissures, the dorsal vessel arises from one of the lateral vessels; nephridia well developed, with many pores; cerebral sense organs before the brain, relatively simple; hermaphrodite. ? Prostoma rubrum (Leidy, 1850). PRINCIPAL REFERENCES: Cok, 1940, p. 308; 1943, pp. 299-301, text fig. 75; 1959, pp. 366-367, text fig. 14. CorpERO, 1943, pp. 125-134, figs. 1-2. Corré&a, 1951, pp. 257— 264, pl. 1, figs. 1-5; pl. 2, figs. 6-11. Monrcomery, 1896, pp. 436-438. Riova, 1941, pp. 663-668, text figs. 1-4. WisjNuoFF, 1938, pp. 219-230. MarTertau. Three worms in August, 1958. Locality 3. The species occurs in large numbers at that season (Coe, 1948, p. 299). Distripution. New England to Georgia and Florida and westward to Ohio, Nebraska, Washington, and California; Xochimileo, Mexico and per- haps Caracas, Venezuela. The living worms were up to 18 mm. long and 1.5 mm. wide. The anterior end is rounded and the posterior is pointed. The color of the three specimens was pale vellow or whitish. There are three pairs of small eves disposed in two rows in front of the brain. An indistinet transverse cephahe furrow is present at the posterior level of the eyes. The rhynchocoel is long as is also the proboscis. The central stylet is short. The base is pear-shaped and as long as the stylet. Two pouches are present, each containing 2-4 accessory stylets. The unfavorable conditions of collecting and preserving made it im- possible to obtain more information and a larger number of specimens. In her synopsis of the genus Prostoma, Stiasny-Wijnhoff (1938) records 16 different names applied to freshwater nemerteans and gives their bibli- ography. Six species are recognized as valid in her revision, viz., P. lumbri- cordeum Duges, 1828; P. graecense (Bohmig, 1892); P. etlhardi (Mont- gomery, 1895); P. grande (Ikeda, 1913); P. padanum Pierantoni. 1926; and P. puteale Beauchamp, 1932. Strangely enough, P. rubrum (Leidy, 1850) does not figure in Wijnhoff’s list, although Coe (1940, 1943, 1959) revalidated this name for the specimens described from North America VoL. XXXT] CORREA: NEMERTEANS, CALIFORNIA and OREGON 549 (P. aquarum-dulcium Silliman, 1885, from New York State and P. asen- soruatum Montgomery, 1896, from Pennsylvania) and Rioja, 1941, for specimens found in Xochimileo, in the vicinity of Mexico City, the most meridional known occurrence of the species. The South American specimens from Venezuela (Cordero, 1943) were not determined; they could belong to P. rubrum as the frontal sense organ was stated (p. 129) as absent. Friedrich (1955, p. 163) records P. rubrum in his list of species of Prostoma. Wijnhoff (p. 6) considers the North American species and P. graecense as identical, but hesitated to introduce the corresponding change of the name because she had not seen North American material. As the name P. asensorvatum indicates, the American species has no supra-oral or frontal sense organ. In Montgomery’s opinion (1896, p. 436) the cephalic glands of P. asensoriatum open individually, scattered on the frontal side. Pro- stoma graecense, however, has a frontal organ toward which the duets of the cephahe glands converge (Bohmig, 1898, pp. 481, 536). Even Reisinger (1926, pp. 2-3), who unites all central European species under the name of P. graecense, maintains the North American species separated. The living worms from California examined with regard to the presence of a frontal sense organ have shown it, and I confirmed it in preserved and cleared state as well as in sections. However, the sections were not good enough to reveal other very delicate characteristics of this species, e.g., the ciliated epithelium in the esophagus and true layer of thick longitudinal muscles of the rhynchodaeum (Correa, 1951, pp. 259, 262). In the sections of the California worms I did not see the caleareous corpuscles in the parenchyma characteristic of P. graecense. Only further study of the Cali- fornia species of Prostoma could elucidate its exact systematic status. Malacobdella blainville, 1827. REFERENCE: FRIEDRICH, 1936, p. 44. Posterior end of body with a sucker; buccal cavity villous, serving for food intake; intestine sinuous; midgut without lateral pouches; rhyneho- daeum absent; proboscis without stylet. Malacobdella grossa (Miller, 1776). PRINCIPAL REFERENCES: BURGER, 1895, p. 597, pl. 18, figs. 1-5; pl. 23, fig. 39; pl. 27, figs. 21-23; pl. 28, figs. 25, 28, 39. Cor, 1940, p. 310; 1943, pp. 309-310, text fig. 79; 1944, p. 32. GrrING, 1911, pp. 673-720, pl. 32, figs. 1-13; text fig. 1. GUBERLET, 1925, pp. 1-13, pl. 1. HAmMaRsreNn, 1918, pp. 1-95, figs. 1-26, text figs. 1-18. RIcCKETTS and CaLvin, 1956, pp. 195-196. Riepen, 1933, pp. 323-496; text figs. 1-63 (the most complete work done on the species). VERRILL, 1892, pp. 444-445, pl. 39, fig. 23, text fig. 9, as M. obesa and M. mercenaria. 550 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4vH SER. MATERIAL. One worm in July, 1958. Locality 4+. A second specimen was received from Dr. Liliana Forneris, Oceanographic Institute, Sao Paulo, (Brazil), collected during her stay at the University of Kiel (Germany), 1959. The latter worm was found in the mantle cavity of the lamellibranech Arctica (Cyprina) islandica (Linnaeus). Occurrence: Stoller Grund, Bay of Kiel, Germany. Riepen (1933) studied material from the same locality. DistriBpuTION. Northern coasts of Kurope, Mediterranean; Nova Scotia to Chesapeake Bay, Atlantic coast of North America and Puget Sound to California, Pacific coast. The length of both present specimens, in preserved state, was about 15 mm. The maximum width, situated at the posterior half of body, was 5 mm. The body is short, broad, and thick. The anterior end is rounded and contains a large and excavated mouth, which serves as aperture for the atrium and proboscis. The posterior end is provided with a sucker. The worms cleared in clove oil showed atrium, esophagus, stomach, midgut, rectum, anus, brain, nervous commissures, lateral nerve cords, proboscis, rhynchocoel, and gonads. The broad, villous atrium extends from near the tip of the head to the brain commissures where the proboscis opens into its dorsal wall. Posterior to the proboscis opening: the digestive canal continues as esophagus without any change in size of lumen or in character of the lining epithelium of short ciliated cells. The esophagus is followed by the stomach. Its lumen is narrower, the ciliated epithelium is higher and the subepithelial glands in the parenchyma become more numerous. There is only a gradual tran- sition in the lumen and epithelium of the esophagus and stomach and of the stomach and midgut. The midgut bends alternately to the left and right, forming at least six loops on the right and seven on the left side. The mid- out terminates in the reetum which opens by the anus on the dorsal side of the sucker. The rhynechocoel extends nearly along the whole length of the body, while the proboscis is much shorter reaching backward less than half that leneth. The unarmed proboscis fills most of the rhynchocoel and follows its coiling. The unusually long proboscidial retractor attains the posterior end of the rhynchocoel. There are two much branched and convoluted lateral vessels united by three commissures, one anterior cephalic, one esophageal, situated on the dorsal side of the esophagus, from where the convoluted dorsal vessel arises, and one anal which unites the three vessels behind. The dorsal vessel lies between rhynehocoel and gut; the lateral vessels run on the sides of the body, close to the lateral nerve cords. A single pair of nephridia extend from near the brain to the region VoL. XXXT] CORREA: NEMERTEANS; CALIFORNIA and OREGON 551 of the stomach. The branched tubules lie in the parenchyma close to lateral nerve cords and lateral vessels. They unite posteriorly to form a pair ot large efferent ducts which lead to the exterior on the ventrolateral surface of the body. The long cerebral ganglia are widely separated by the broad atrium and connected by a small dorsal and a large ventral commissure. The ventral commissure passes between rhynehocoel and atrium and the dorsal one between rhynehocoel and body wall. Anterior, lateral, and posterior nerves arise from the ganglia. The lateral nerve cords are united posteriorly by a slender suprarectal commissure. Some other commissures and nerves arise from the cords along their course. Eyes and sense organs are wanting. Both specimens, sexually mature females, had many ovaries situated on both sides of the body, irregularly scattered from the limit between esophagus and stomach to the rectal region. Each gonad has a dorsal opening. The Order Bdellomorpha consists of a single genus, Malacobdella. Be- sides the here-deseribed species, three more are known which can be sepa- rated by the somewhat modified Coe’s key (1945, p. 67). 1. Commensal in marine lamellibranchs 2 — Commensal in fresh-water gastropods............................ M. auriculae Blanchard, 1847. Occurrence: Chile. In the pallial cavity of Chilina dombeyana Brug. bo Proboscis sheath nearly as long as body; posterior commissure of nerve cords ONMGOTES ale SUG nO fre CEU ee er ee cee ee eee eee oe 3 — Proboscis sheath only two thirds as long as body; posterior commissure of nerve cords in sucker.......................M. japonica Takakura, 1897. Occurrence: Coasts of Japan. Commensal in the mantle cavity of Spisula sachalinensis. 3. Nephridiopores on dorso-lateral surfaces; gonads large, situated in a single TLOwe On each: Sid exOb, WOM yee ee eee eee ee es ee eee caine Vee a tee Ss gn ee ee M. minuta Coe, 1945. Occurrence: Off Point Loma, Southern California, at a depth of 40 m. Commensal in the mantle cavity of Yoldia cooperi Gabb. — Nephridiopores on ventro-lateral surfaces; gonads relatively small and numer- Ousteirresularly, -scattere Ge. 2.-. Se M. grossa (Miller, 1776). As the main purpose of this paper is to determine the intertidal species of nemerteans occurring along the California coast, I add a list of 11 more species not collected by me but recorded by Coe (1944, pp 27-32). 1. Carinomella lactea Coe, 1905. Recently redeseribed by Corréa, (1961, pp. 8-11, fig. 8). Length 50-100 mm.; general color of body milk-white, more or less translucent; after preservation a brown band appears about 5 mm. back from the tip of the head, which fades backward. Occurrences: Biscayne Bay, Miami, Florida; Monterey Bay, California to San Diego, California. Intertidal zone to 20 m. On 6. 10. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Baseodiscus punnetti (Coe, 1904+). Large species, 40-60 cm. long; gen- eral color deep brownish-red; anterior tip is much deeper brown, marked off from a narrow terminal and lateral white border; ocelli as an irregular longitudinal row of 40-60 or more on the anterolateral margins of head. Occurrences: Monterey Bay, California to Gulf of California, Mexico. Intertidal zone to 380 m. . Zygeupolia rubens (Coe, 1895). Rather slender, 5-8 cm., long; head pure white; esophageal region white or pale yellow; intestinal region from rose to pale yellow, light brown or chocolate-brown; white caudal cirrus present; cephalic furrows absent. Occurrences: New England and southward to North Carolina; Monterey Bay, California to En- senada, Mexico. Intertidal zone to 50 m. Euborlasia nigrocincta Coe, 1940. Up to 50-70 em. long; there are two color varieties: the darker is rich purplish brown dorsally and ventrally; head white with fine red or brown dets except the tip; the paler is rosy with reddish brown or purplish dots dorsally; head white or pink; ocelli absent. Occurrences: San Francisco Bay, California; Monterey Bay, California to Ensenada, Mexico. Intertidal zone to 30 m. Nemertopsis gracilis Coe, 1904. Commonly 10-15 em. long; whitish with two longitudinal bands of deep brown along the whole length of body; four ocelli. Occurrences: Puget Sound to Ensenada, Mexico. Intertidal zone and below. Nemertopsis gracilis var. bullocki, 1940. Differs from the typical form in having the longitudinal brown bands connected anteriorly and some- times interrupted at intestinal region. Occurrence: coast of central California. Intertidal zone. . Dichonemertes hartmanae Coe, 1938. Small, 10-15 mm. long; pale red or yellowish with deep red blood; four ocelli; hermaphrodite. Occeur- rence: San Diego, California. Intertidal zone. Carcinonemertes epialti Coe, 1902. Commensal on crabs of the genera Portunus, Pugettia, and Euphylax. Occurrences: Monterey Bay, Cali- fornia, to San Diego, California; Peru. Ototyphlonemertes spiralis Coe, 1940. Minute, 3-5 mm. long; white, eray, straw color or yellow; ocelli absent; statocysts containing statolith with 8, 12, or 16 globules. Occurrence: San Diego, California. On sandy shores of bays. Prosorhochmus albidus (Coe, 1895). Usually 10-15 mm. long; milk- white or creamy; four ocelli; two ocelli of same side united by a fine line VoL. XXXT] CORREA: NEMERTEANS,; CALIFORNIA and OREGON 553 of dark pigment. Occurrences: Monterey Bay, California, to Ensenada, Mexico. Intertidal zone. 11. Oerstedia dorsalis (Abildgaard, 1806). Recently redescribed by Corréa (1961, pp. 23-24). Up to 15 mm. long; color light cream irregularly spotted with brown of various shades and with considerable variations in shape and distribution; four ocelli. Ocurrences. Circumpolar; Nor- way to Mediterranean; Madeira; Nova Seotia to Miami, Florida and Key Largo, Florida; Puget Sound to Gulf of California. Intertidal zone and below. BIBLIOGRAPHY Boumia, LupWwIG 1898. Beitrage zur Anatomie und Histologie der Nemertinen. Zeitschrift fiir Wissenschaftliche Zoologie, vol. 64, no. 3, pp. 479-564, pls. 13-17, one text fig. 1929. Nemertini, W. Kiikenthal & Th. Krumbach, Handbuch der Zoologie, vol. 2, first half, pp. 1-110, text figs. 1-75. Bork, HEINZ-J URGEN 1952. Die Paketdriisenzone von Lineus ruber O. F. Miiller (Nemertini). Zeitschrift fiir Morphologie und Oekologie der Tiere, vol. 41, pp. 188— 222, text figs. 1-17. BURGER, Oro 1892. Zur Systematik der Nemertinenfauna des Golfs von Neapel. Nachrichten von der Gesellschaft der Wissenschaften zu GO6ttingen, no. 5, pp. 137— 178. 1895. Die Nemertinen. Fauna und Flora des Golfes von Neapel, Monogr. 22, pp. xvi + 743, pls. 1-31. 1897-1907. Nemertini. Dr. H. G. Bronn’s Klassen und Ordnungen des Tier- Reichs, vol. 4 (suppl.) pp. viii + 542, pls. 1-22, text figs. 1-63. 1904. Nemertini. Das Tierreich, Preussische Akademie der Wissenschaften zu Berlin, vol. 20, pp. xvii + 151, text figs. 1-15. Cor, WESLEY ROSWELL 1901. The Nemerteans. Papers from the Harriman Alaska Expedition. XX. Proceedings of the Washington Academy of Sciences, vol. 3, pp. 1-111, pls. 1-13, text figs. 1-16. 1904. Nemerteans of the Pacific coast of North America. Harriman Alaska Expedition, vol. 11 (part 1), pp. 1-111, pls. 1-13, text figs. 1-16 (re- printed from 1901); (part 2), pp. 112-220, pls. 14-22, text figs. 17-23. 1905. Nemerteans of the west and northwest coasts of North America. Bulle- tin of the Museum of Comparative Zoology, Harvard, vol. 47, pp. 1-318, pls. 1-25, text figs. 1-62. 1930. Two new species of nemerteans belonging to the family Cephalothri- chidae. Zoologischer Anzeiger, vol. 89, pp. 97-103, text figs. 1-8. 554 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. Cor, WESLEY ROoSwELL—Cont. 1940. Revision of the nemertean fauna of the Pacific coasts of north, central, and northern South America. Allan Hancock Pacific Expedition, vol. 2, no. 13, pp. 247-323, pls. 24-31. 1948. Biology of the nemerteans of the Atlantic coast of North America. Transactions of the Connecticut Academy of Arts and Sciences, vol. 35, pp. 129-328, pls. 1—4, text figs. 1-79. 1944. Geographical distribution of the nemerteans of the Pacific coast of North America, with descriptions of two new species. Journal of the Washington Academy of Sciences, vol. 34, no. 1, pp. 27-32. 1944a. A new species of Hoplonemertean (Paranemertes biocellatus) from the Gulf of Mexico. Journal of the Washington Academy of Sciences, vol. 34, no. 12, pp. 407-409, text figs. 1—4. 1945. Malacobdella minuta, a new commensal nemertean. Journal of the Washington Academy of Sciences, vol. 35, no. 2, pp. 65-67, text figs. 1-3. 1951. Geographical distribution of the nemerteans of the northern coast of the Gulf of Mexico, etc. Journal of the Washington Academy of Sciences, vol. 41, no. 10, pp. 328-331, text figs. 1-2. 1951la. The Nemertean faunas of the Gulf of Mexico and of southern Florida. Bulletin of Marine Science of the Gulf and Caribbean, vol. 1, no. 3, pp. 149-186, text figs. 1-28. 1952. Geographical distribution of the species of nemerteans of the Arctic Ocean near Point Barrow, Alaska. Journal of the Washington Academy of Sciences, vol. 42, no. 2, pp. 55—58. 1959. Nemertea. In Ward, H. B. & G. C. Whipple. Fresh-water biology, 2nd ed., pp. 366-367, text fig. 14. 1. CoRDERO, ERGASTO H. 1943. Hallazgos en diversos paises de Sud America de nemertinos de agua dulce del Genero Prostoma. Anais da Academia Brasileira de Ciéncias, vol. 15, no. 2, pp. 125-135, figs. 1-2 a-f. Correia, Diva DINIz 1951. Freshwater nemertines from the Amazon region and from South Africa. Boletim da Faculdade de Filosofia, Ciéncias e Letras da Universidade de Sao Paulo, Zoologia, no. 16, pp. 257-269, pls. 1-2. 1953. Sobre a neurofisiologia locomotora de Hoplonemertinos e a taxonomia de Ototyphlonemertes. Anais da Academia Brasileira de Ciéncias, vol. 25, no. 4, pp. 545-555, text figs. 1-7. 1953a. Sdbre a locomocao e a neurofisiologia de nemertinos. Boletim da Facul- dade de Filosofia, Ciéncias e Letras da Universidade de Sao Paulo, Zoologia no. 18, pp. 129-147, pls. 1-2. 1954. Nemertinos do litoral brasileiro. Boletim da Faculdade de Filosofia, Ciéncias e Letras da Universidade de Sao Paulo, Zoologia no. 19, pp. 1-90, pls. 1-14. VoL. XXXT] CORREA: NEMERTEANS, CALIFORNIA and OREGON 555 Correa, Diva Diniz—Cont. 1955. Os géneros Emplectonema Stimpson e Nemertopsis Birger (Hoplo- nemertini Monostilifera). Boletim da Faculdade de Filosofia, Ciéncias e Letras da Universidade de Sao Paulo, Zoologia no. 20, pp. 67-78, pls. 1-4. 1961. Nemerteans from Florida and Virgin Islands. Bulletin of Marine Science of the Gulf and Caribbean, vol. 11, no. 1, pp. 1—44, text figs. 1-34. EGGERS, FRIEDRICH 1924. Zur Bewegungsphysiologie der Nemertinen I. Emplectonema. Zeitschrift fiir vergleichende Physiologie, vol. 1, no. 3-4, pp. 579-589, text figs. 1-5. 1936. Zur Bewegungsphysiologie von Malacobdella grossa Miill. Zeitschrift fir Wissenschaftliche Zoologie, vol. 147, no. 1, pp. 101-131, text figs. 1-11. FRIEDRICH, HERMANN 1933. Vergleichende Studien zur Bewegungs- und Nervenphysiologie bei Nemertinen. Zoologische Jahrbticher Allgemeine Zoologie und Physi- ologie der Tiere, vol. 52, no. 3, pp. 537-560, text figs. 1-6. 1935. Studien zur Morphologie, Systematik und Oekologie der Nemertinen der Kieler Bucht. Archiv fiir Naturgeschichte, N. F., vol. 4, no. 3, pp. 293-375, text figs. 1-34. 1936. Nemertini. Grimpe & Wagler, Tierwelt der Nord- und Ostsee, vol. IV d, pp. 1-69, text figs. 1-18. 1936a. Eine Bemerkungen zur Anatomie von Tubulanus borealis n. sp., ete. Zoologischer Anzeiger, vol. 116, no. 3-4, pp. 101-108, text figs, 1—5. 1955. Beitrage zu einer Synopsis der Gattungen der Nemertini monostilifera nebst Bestimmungsschlissel. Zeitschrift flr Wissenschaftliche Zoologie, vol. 158, no. 2—4, pp. 133-192. 1956. Zur Morphologie des Vorderdarmes der monostiliferen Hoplonemertinen. Veroffentlichungen des Instituts fiir Meeresforschung in Bremerhaven, vol. 4, pp. 45—53, pls. 1-2. 1956a. Die Palaeonemertine Carinoma Oudemans 1885 an der Chilenischen Kiuste. Reports of the Lund University Chile Expedition 1948—49. Lunds Universitets Arsskrift, N. F. 2, vol. 52, no. 1, pp. 3-5. 1958. Nemertini. The zoology of Iceland, vol. 2, part 18, pp. 1-24, text figs. 1-8. 1960. Bemerkungen tiber die Gattung Micrura Ehrenberg 1831 und zur Klas- sification der Heteronemertinen nebst vorlaufigem Bestimmungs- schliissel. Veroffentlichungen des Instituts fiir Meeresforschung in Bremerhaven, vol. 7, pp. 48-62. GERING, GUSTAF 1911. Beitrage zur Kenntnis von Malacobdella grossa (Miull.). Zeitschrift fiir Wissenschaftliche Zoologie, vol. 97, no. 4, pp. 673-720, pl. 32, one text fig. 556 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH SER. GONTCHAROFF, MARIE 1951. Biologie de la régénération et de la reproduction chez quelques Lineidae de France. Annales des Sciences Naturelles, Zoologie, sér. 11, vol. 13, pp. 149-235, pls. 1-7, text figs. 1-29. 1955. Nemertes-Tuniciers. Inventaire de la faune marine de Roscoff, suppl. 7 aux Travaux de la Station Biologique de Roscoff, pp. 1-15. GRIFFIN, BRADNEY B. 1898. Descriptions of some marine nemerteans of Puget Sound and Alaska. Annals of the New York Academy of Sciences, vol. 11, no. 10, pp. 193- 218, text figs. 15-24. GUBERLET, JOHN E. 1925. Malacobdella grossa from the Pacific coast of North America. Publi- cations of the Puget Sound Biological Station, vol. 5, pp. 1-13, pl. 1. HAMMARSTEN, OLOF D. 1918. Beitrag zur embrionalentwiclung der Malacobdella grossa (Miull.). Zootomischen Institut der Hochschule zu Stockholm, vol. 1, pp. 1—95, figs. 1-26, text figs. 1-18. HEDGPETH, JOEL W. 1951. Pyenogonids from Dillon Beach and vicinity, California, etc. Wasmann Journal of Biology, vol. 9, no. 1, pp. 105-117, pls. 1-3. HyLBpom, RICHARD 1957. Studies on palaeonemerteans of the Gullmar Fiord area (west coast of Sweden). Arkiv for Zoologi, ser. 2, vol. 10, no. 17, pp. 539-582, pls. 1-9, text figs. 1-6. HYMAN, Lispie HENRIETTA 1951. The Invertebrates: Platyhelminthes and Rhynchocoela, vol. 2, pp. vii + 550, text figs. 1-208. McGraw-Hill Book Comp. Inc., New York. ISLER, ERNST 1901. Die Nemertinen der Sammlung Plate. Zoologische Jahrbiicher Suppl. 5, Fauna Chilensis, vol. 2, no. 2, pp. 273-280. IWATA, FUMIO 1952. Nemertini from the coasts of Kyusyu. Journal of the Faculty of Science, Hokkaido University, ser. 6, Zoology, vol. 11, no. 1, pp. 126-148, text figs. 1-18. 1954. The fauna of Akkeshi Bay XX. Nemertini in Hokkaido. Journal of the Faculty of Science, Hokkaido University, ser. 6, Zoology, vol. 12, no. 1-2, pp. 1-39, text figs. 1-9. 1957. Nemerteans from Sagami Bay. Publications from the Akkeshi Marine Biological Station, vol. 7, pp. 1-31, pls. 1-7. JOUBIN, Louis M. 1890. Recherches sur les Turbellariés des cotes de France (Némertes). Archives de Zoologie Expérimentale et Générale, ser. 2, vol. 8, pp. 461-602, pls. 25-31, text figs. 1-6. VoL. XXXII] CORREA: NEMERTEANS; CALIFORNIA and OREGON 55 -~1 JOUBIN, Louis M.—Cont. 1905. Note sur quelques némertiens recueillis en Basse-California par M. Di- guet. Bulletin du Museum National d’Histoire Naturelle de Paris, vol. 11, pp. 309-315, text figs. 1-5. MAcGInItTig£, G. E., and Nerric MAcGINITIE 1949. Natural history of marine animals. pp. xii + 473, McGraw-Hill Book Comp. Inc., New York. McINTOSH, WILLIAM CARMICHAEL 1873-74 A monograph of the British Annelids. Part I. The Nemerteans. Ray Society, pp. 1-214, pls. 1-23. MONTGOMERY, THOMAS H. 1896. Stichostemma asensoriatum, n. sp., a freshwater nemertean from Penn- sylvania. Zoologischer Anzeiger, vol. 19, pp. 436-438. 1898. Descriptions of new Metanemerteans, with notes on other species. Zoologisches Jahrbiicher Systematik, Geographie und Biologie der Tiere, vol. 10, no. 1, pp. 1-14, pl. 1. PUNNET, R. C. 1901. On some Arctic nemerteans. Proceedings of the Zoological Society of London, vol. 2, pp. 90-107, pls. 6—7, text figs. 1-6. 1903. On the nemerteans of Norway. Bergens Museums Aarbog, no. 2, pp. 1-35, pls. 1-2. REISINGER, ERICH 1926. Nemertini. Dr. Paul Schulze, Biologie der Tiere Deutschlands, part 7, pp. 1—24, text figs. 1-8. RICKETTS, Epwarp F., and JACK CALVIN 1956. Between Pacific tides, 3rd ed. rev. by Joel W. Hedgpeth. pp. xiii + 502, pls. 1-46, text figs. 1-134. Stanford University Press, Stanford, Cali- fornia. RIEPEN, OTTO 1933. Anatomie und Histologie von Malacobdella grossa (Miill.). Zeitschrift fiir Wissenschaftliche Zoologie, parts 1-2, vol. 148, no. 4, pp. 324— 496, text figs. 1-63. RiosJA, ENRIQUE 1941. Hallazgo en Xochimilco de Stichostemma rubrum (Leidy), nemerte de agua dulce. Anales del Instituto de Biologia, vol. 12, no. 2, pp. 663-— 668, text figs. 1-4. Snaw, F., C. FiscHer, and G. H. HARLAN 1949. Oil lamps and iron ponies. San Francisco: Bay Books, Ltd., pp. 187, illus. (copied from Hedgpeth, 1951). SOUTHERN, R. 1913. Nemertinea. Proceedings of the Royal Irish Academy, Clare Island Sur- vey 55, vol. 31, no. 3, pp. 1-20, pl. 1. r 558 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. STEPHENSON, J. 1913. The nemertines of Millport and its vicinity. Transactions of the Royal Society of Edinburgh, vol. 48, pp. 1-29, text figs. 1-18. STIMPSON, W. 1858. Prodromus descriptionis animalium evertebratorum, quae in expedi- tione ad Oceanum, Pacificum Septentrionalem a Republica Federata, ete. Proceedings of the Academy of Natural Sciences of Philadelphia 1857, part 2, pp. 159-165. VERRILL, ADDISON E. 1892. The marine nemerteans of New England and adjacent waters. Trans- actions of the Connecticut Academy of Arts and Sciences, vol. 8, pp. 382-456, pls. 33-39, text figs. 1-9. WHEELER, J. F. G. 1934. Nemerteans from the South Atlantic and Southern Oceans. Discovery Reports, vol. 9, pp. 215-294, pls. 15-16, text figs. 1-66. 1940. Some nemerteans from South Africa and a note on Lineus corrugatus McIntosh. The Linnean Society’s Journal-Zoology, vol. 41, no. 276, pp. 20-49, text figs. 1-13. WIJNHOFF, GERARDA 1912. List of nemerteans collected in the neighbourhood of Plymouth from May-—September, 1910. Journal of the Marine Biological Association of the United Kingdom (new ser.), vol. 9, no. 3, pp. 407—434, text fig. 1. WIJNHOFF, GERARDA STIASNY 1938. Das Genus Prostoma Dugés, eine Gattung von Siisswasser-Nemertinen. Archives Néerlandaises de Zoologie, vol. 3, suppl., pp. 219-230. YAMAOKA, TEICHI 1940. The fauna of Akkeshi Bay 9. Nemertini. Journal of the Faculty of Science, Hokkaido University, ser. 6, vol. 7, no. 3, pp. 205-263, pls. 14-17, text figs. 1-33. Marine Biological Laboratory LIBRARY JAN 2 71965 WOODS HOLE, MASS. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXI, No. 20, pp. 559-562, 7 figs. January 15, 1965 PHYLLOPHAGA SAYLORI, N. SP., FROM NUEVO LEON, MEXICO (COLEOPTERA: SCARABAEIDAE) By Milton W. Sanderson Illinois Natural History Survey, Urbana, Illinois Jean Mathieu, Instituto Technologico y de Estudios Superiores in Mon- terrey, Nuevo Leon, Mexico, made a special search for Phyllophaga galeanea Saylor (1943: 28) on May 12 and 13, 1961, at its type locality, Galeana, Nuevo Leon. He examined various plants, including Pinus and Juniperus, suspected as possible hosts, but without suecess. Mr. Mathieu slept under the pines that night, and early next morning he noted a large species of Phyllophaga dropping from the high pine foliage to the forest floor. It is a new species in the “ignava” group, as defined by Sanderson (1958: 160, 172), and it is unique in several features among some 30 known species in that oroup. This species is named in honor of Mr. L. W. Saylor, Head of the Ma- terial Department, Administrative Branch, Naval Supply Depot, Oakland, Califorma. From 1934 to 1948, Mr. Saylor published some 35 papers on New World Phyllophaga in which he deseribed many new species. He is especially to be credited, however, with breaking the taxonomic impasse which existed in the Central American and Mexican literature since the [ 559 J 560 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. publication of the Biologia portion treating this genus. Until he began his studies for that area, it was virtually impossible to identify species from the literature. He illustrated male genitalia and prepared keys to several groups of species, making it possible to recognize a large proportion of the species in Mexico and Central America. The Saylor collection of Phyl- lophaga and other Scarabaeidae is now located at the California Academy of Sciences. I am indebted to Jean Mathieu for the specific determination of the pine. Phyllophaga saylori Sanderson, new species. (Figures 1-7.) Houotyre. Male. Illinois Natural History Survey, collected 4 km. south of Galeana in Nuevo Leon, Mexico, at 6000 feet, May 13, 1961, by Jean Mathieu. Dropped from Pinus teocote about 5 o’clock in the morning. Gen- eral appearance. A large plump brownish beetle, the posterior margin of the elytra (fig. 7) reflexed. Length 24 mm.; width 12.5 mm. Head. Antenna 10-segmented, 3-segmented club short, approximately equal in leneth to five preceding segments combined, and approximately equal in length to elypeus at middle. Clypeus evenly rounded to the slightly emarginate apex, apical margin strongly reflexed in anterior one-third, elypeal punctures Shallow, generally closely placed, separated on the average by less than their own widths; frons similarly punctured but with irregular impunetate areas, especially adjacent to clypeo-frontal suture. Labrum exposed when viewed from above. Pronotum 1.6 times as wide as long, finely and very irregularly punctured, some punctures separated from others by from one to six times their widths; lateral margins broadly and faintly crenulate, nearly parallel in basal one-half; basal angle distinet, approximately 135 degrees, anterior angle not produced. Elytra rather strongly flared posteriorly, widest at about apical one-third, sutural costa very broad at middle, narrowed toward apex and seutellum, with irregularly seattered punctures; a large diagonal nearly impunctate costa arising near middle of base of elytron, expanded and extended toward sutural apex; apical margin of each elytron (fig. 7) strongly reflexed; entire lateral margin of elytron, except near sutural apex, with a conspicuous fringe of long hairs. Abdomen. Pygidium nearly evenly convex for most of its length, more strongly convex before apex, very irregularly and shallowly punctured, glabrous. Abdomen in lateral view (fig. 7) strongly convex to the narrowly transversely im- pressed posterior margin of the penultimate sternite; surface shining, nearly impunctate on disc, punetures dense in a median pateh on posterior one-half of penultimate sternite; three abdominal sutures on dise slightly pro- duced anteriorly at middle; last sternite flattened, about one-half length of penultimate. J/estasternum. Densely punctured toward sides, more sparsely VoL. XXXT] SANDERSON: PHYLLOPHAGA SAYLORI 561 punctured at middle, with long hairs not quite as long as greatest width of middle femur. Legs. Anterior tibia tridentate; each anterior tarsal seg- ment with a ventral longitudinal carina, carinae of first four segments each terminating in a strong internal tooth; each tarsal claw with a long nearly median tooth (fig. 6); apical margin of posterior tibiae each with 22 and 24 strong setae; apical tibial spurs slender, longer spur a little longer than first posterior tarsal segment; posterior tibia without special modifications. Genitalia (figs. 1-3) asymmetrical, apical lobe of each paramere free, slender; one of the two lobes of paramere base (fig. 3) more expanded at apex than the other; aedeagus (fig. 2) bearing two stronely curved processes one one side, one longer than the other. PaRATYPES. Eleven males and two females, all collected at the type locality, and except three males collected at light, under the same conditions. California Academy of Sciences; Instituto Technologico y de Estudios Be oA Figure 1. Left lateral view of male genitalia, Phyllophaga saylori. FicgurE 2. Right lateral view of aedeagus of male genitalia, Phyllophaga saylori. Figure 3. Dorsal view of asymmetrical base of parameres of male genitalia, Phyllophaga saylori. Figure 4. Anal plates of female, Phyllophaga saylori. FIGURE 5. Pubic processes of female genitalia, Phyllophaga saylori. FiGuRE 6. Tarsal claw of right foreleg of male, Phyllophaga saylori. Figure 7. Lateral view of elytron and abdomen of male, Phyllophaga saylori. Arrow indicates reflexed elytral margin. 562 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Sen. Superiores in Monterrey, Nuevo Leon; Illinois Natural History Survey. Males vary in color from light brown to dark brownish red; length 21 to 25 mm.; width 11 to 12.4 mm. The pronotum ranges from 1.67 to 1.90 times as wide as long, and the posterior tibial setae from 17 to 24. The female is similar to the male except as follows: antennal club shorter than clypeal leneth; pygidium deeply, narrowly grooved at apex; abdomen more pointed, last sternite longer, about two-thirds leneth of penultimate sternite. Pubie process of female genitalia divided (fig. 5); anal plates as in figure 4. Discussion. The genitalia of one dissected male are paler in color and less heavily sclerotized than other males, suggesting recent transformation from the pupal stage. The reflexed apical elvtral margin, and the asym- metrical male genitalia will distinguish this species from all other known members of the “tgnava” group. In size and appearance, P. saylort resembles P. torta LeConte (Mexico, United States), and P. pleroma Reinhard (Texas), both belonging to the “ignava” group. The symmetrical male genitalia will at once distinguish the two species from P. saylori. LITERATURE CITED SANDERSON, M. W. 1958. Faunal affinities of Arizona Phyllophaga, with notes and descriptions of new species. Journal of the Kansas Entomological Society, vol. 31, pp. 158-1738. Saytor, L. W. 1943. Seis nuevos Coleopteros Lamelicornios de Mexico. Revista de la Sociedad Mexicana de Historia Natural, vol. 4, pp. 25-81. Marine Biological Laboratory LIBRARY JAN 2 7 1965 WOODS HOLE, MASS, PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXI, No. 21, pp. 563-600, 3 figs. January 15, 1965 OSTEOLOGICAL CHARACTERISTICS AND AFFINITIES OF THE HEXAGRAMMID FISHES, WITH A SYNOPSIS’ By Jay C. Quast U.S. Bureau of Commercial Fisheries Biological Laboratory, Auke Bay, Alaska ABSTRACT Representatives of hexagrammid genera are analyzed for skeletal fea- tures, and the findings compared with original and published data on other mail-cheeked fishes (suborder Cottoidei). Pereciform groups suggested in the literature as possible mail-cheeked allies are also examined. Results indicate that the fish skeleton has numerous characteristics of taxonomic importance that have been used little or not at all. Examples inelude caudal structure, dorsal ptervgiophore patterns, and ratios between dorsal fin spines, pterygiophores, and vertebrae. Taxonomic findings indicate that the present classification of mail-cheeked fishes, based on Regan’s work of 1915, is in need of revision. The Hexagrammidae and Zaniolepididae are probably primitive offshoots of the cottid evolutionary line and the three groups should be placed in the same superfamily. The zaniolepidids are generally intermediate between the hexagrammids and cottids. The Anoplo- JA revised portion of the dissertation submitted in partial fulfillment of the requirements for Doctor of Philosophy at the University of California at Los Angeles, California. [565 J 564 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. pomatidae (Anoplopoma, Erilepis) are sufficiently distinct from the seor- paenids and hexagrammid-cottids to warrant a separate superfamily. The mail-cheeked fishes may be an artificial assemblage containing at least three distinct evolutionary lines, scorpaenid, anoplopomatid, and hexagrammid- cottid. The findings are summarized in a synopsis. CONTENTS arto cuetiOme ao we aewce lhe one is oarouee oe etaue) Gretel eo. o.0 ore oa eee 565 Materials: and \Methods.2.0--de:c 26 acces. pe clon Gore seek ee 565 Osteolooy ated. etic Rey cles ee Ble Pe Se See EES ake dino eo Bee a ee ee D68 Solace ng Grey ee cce ed, gle secre arte eat are cae ones er 568 Toothed Upper Pharyneeals:. os \ secs lee. eee 568 Biranehiost€cale 854s eee oe oe hears Wet en eee D69 INGmmoeraial Wii y.132)Gs Bae oa os oye tee ae RR eee 569 Suborbital shones.-.4.. si). dase SS ne Se 569 Supratenmporal Canales i< 2.) o 0.5 Soe See tee ee 570 C)ISPIO FICS a8 ae Secrest suse ay eaves, ae eee ear al HP XOCCIPIGAIS “So-36. cele og oan banca. & Paes Comat Oe eee MP4 Parasphenoid-pterosphenoid: jumetion = 052... see 52 BASIS PIG I OVE | ice ier deat gelig 3 elo /arcael a dap tert pepe Vee eee oT4 Lateral ethimoids 02a 7. Bee ne oe ee ee 574 Vertebraevand Associated Structures 2... 4:06... 20 eee ble NiCTtEDPAG” i. ee ee eee Sh RUT gH ee a To, cue Arenas cek rites ine banana eNO 576 Dorsal Pteryeiophores, sci... 4e.cc ee oe eee 2 oe 5976 Dorsal Ray Totals: «2. jscpieeeie coere ee eee ee oe eee 578 Caudal. skeleton, is. 2c1apy adden Li hes eee 580 Appendicular Skeleton‘. 7 aus. A8) es oon re ee ee ee 581 Peélvie-eirdiie “Se tac ke cee ae ae eee 581 Pectoral oimdle 0.3.55 oc ce Spe ees ok ee ee ee 581 DISCUSSION CAs Ske ed SIA RR ee 583 SIVMMO SIS pid airy fe hl abs us Sul ge dieealeg Med & 2 Leta Pa 586 NGO WILEGEMMEIULS Ts Bos as Pe YS oa de eR Ve es 597 Literatures @ived a. si teat Sher wm ie, Meret hod > oe ee oe Se 597 Vou. XXXII] QUAST: HEXAGRAMMID FISHES ol 7) on INTRODUCTION The nine extant species of the family Hexagrammidae form an im- portant element in the mail-cheeked fish fauna of the North Pacifie Ocean. The phyletice position of the family within the mail-cheeked fishes (suborder Cottoidei) has been uncertain with considerable difference of opinion con- cerning where the family lines should be drawn. To investigate these and related questions, I undertook an osteological study of the family and its purported members and allies as part of my doctoral problem at the Uni- versity of California at Los Angeles. The mail-cheeked families listed by Berg (1940) and the Serranidae and Cirrhitidae are compared for features of cranium, axial skeleton, and appendicular skeleton. The Serranidae and Cirrhitidae are included because one or both of these families are usually mentioned by students of the phylogeny of the mail-cheeked fishes, e.g. Gill (1889), Jordan and Ever- mann (1898), and Regan (1913). The findings, as brought out in the dis- cussion, indicate that considerable changes should be made in the arrange- ment of the families in the suborder and that the suborder is probably poly- phyletic. To facilitate comparisons, the findings are summarized in a synopsis, with special emphasis on comparisons with the Hexagrammidae. Taxonomic changes in several species follow the recommendations of Quast (1960): Oxylebius pictus and Ophiodon elongatus are in monotypic subfamilies under the Hexagraminidae; the Japanese Agrammus agrammus is placed in the genus Hexagrammos; and the nominal species Heragrammos superciliosus and H. lagocephalus ave synonymized under H. lagocephalus. MATERIALS AND METHODS Skeletal material included X-ray photographs, boiled fresh specimens, Clorox- or Purex-treated preserved material, and dissections (table 1). Dissection and study of skeletal material were sometimes facilitated by alizarin staining, with the specimens being dissected to the vicinity of the skeletal elements to aid stain penetration. Techniques of Green (1952) generally were followed except for storage of specimens in a 50 per cent isopropyl alcohol solution before fine dissections. Clearing in glycerine, was avoided when further examination was contemplated because of re- sultant slipperiness and irregularities of light refraction. In skeletal preparations the entire fish was immersed in very hot, but not boiling, water with a detergent. Skeletal elements were removed as they loosened and were cleaned with a toothbrush. Care was taken not to cook the fish so extensively that neurocranial bones became separated or vertebral column and hypural fan beeame disarticulated. Jets of water and air were useful for removing soft tissues after cooking. When dry, some fatty bones were degreased by soaking for a week or more in toluene or xylene. 566 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr. No one method of preparation is entirely satisfactory for material used in osteological studies, and two or more methods should be used on separate specimens of the same species when possible. Although dry skeletal prepa- rations are superior for neuroecranial study, dissections and alizarin stain- TABLE 1. Number of specimens examined Symbols for the columns represent method of examination: X, X-ray; S, alizarin staining; C, cooked or Clorox-treated material. x Ss Cc Holocentridae ET OUOCETUGUS SAU OO TO UE CLUS ree ee WED ROIPOSEOS: (GUC PU OIA GLOSS OSecene nee peace fe 2 AS eee Serranidae PAT DIUCSECS nO CULO) GC TUS TS teseeee ee eee 2s eee aoe be a anes eee 1 PAUL DIVESIGC SHITE LL CEL OLUTIECLL LES omen mentee eet ae ee eee EE 2 ID ERP OCR DUIDOS 7 J OWN CORIO a oa ono ee eee SESE il DA DlCGEALT AI LGELODONUG = ee eee ee 1 BT TIUIUC DIUGUILS mam LTE CULO OLS ene ee VEU CU TOD CIRC Cay O17; C010 een nee enn ene ee EZ CUUGO TO OUTS O OUT CCG TES ree ee ee 1 EX ORCL CGO ROAR UC GTUIG CL GLU eee ene 1 ZORCGULO TECTIA CDALUU] Clyne eee eee 1 POGOIUEN TUS RC OUONALS eee er ee See iL 1 ISECPODUG DUS: Of KORS ass eaten ey ate mE MeUeU Ns Patel Ly Centrarchidae VTCT.O DIG CTU Same S CULTTU OU, CS eases ee eae 1 Sciaenidae MECH ODO GOT mi CIUGUDUTTUTS ses eeen eee ee eee ee Bares il Cichlidae CAChlGSOTUCRA OU ee eee il Pomacentridae ED ONUOCETUUTAULS OC CU. ene nee aaa eee eee if Cirrhitidae GUUS ITE CTY OG OCULS eeaeenee 1 Ca WS VOU VOLS -- TP) Ns aici } PoPOF SVO LYIDAY DUIAD|SITYydu yy yisuoy] S}JUSULSIS S]USULSaS SsTeiqe'y quo A /S10Q IeL(yny ) +7e J+ Apog xo9 Jequnu [eovo[p uryo SLNANWDHS ITONNYV pure sateds suawmdrads a1gpjwpan s40f DIDp [0 21QD]L f HIEVL VoL. XXXII] GANS: NEW AMPHISBAENA 619 FIGuRE 4. Amphisbaena heathi. Dorsal, lateral and ventral views of the head of the paratype, CAS 49424, from Ceara Mirim, RGN, Brazil. 620 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. Fieure 5. Amphisbaena heathi. Dorsal and ventral views at midbody of CAS 49424, to show segment proportions. side. They are paralleled by the sutures between ocular and third infralabial, which continue in an unbroken straight line as the post-ocular-temporal sutures and as those between the first parietals and the segments ventral to them. All of this gives the seale pattern a very characteristic regular veometrical appearance. There are three subequal supralabials, the second the largest, the third occupying the shortest distance along the labial edge. The posterior edge of the third supralabia! always lies slightly anterior to that of the third infralabial. The interlabial sutures run anteriorly at angles (to the labial edge) of 30°, 45°, 90°, and 90° respectively. The ocular is quadrilateral, in contact with the dorsal edges of the second and third supralabials, in broad contact with the prefrontals, in point contact with the frontal, and mareined posteriorly by the equal-sized postocular. The mental is elongate and triangular, and in point contact with the small rhomboidal postmental, the sides of which are approximately equal in width to the elongate trapezoidal first infralabials whose medial edges contact it. The second infralabials are huge and in point contact with the postmental. Two first large postgenials lie between the medial edges of the second infralabials, and the posterolateral edges of the postmental. These are followed by the three small second postgenials. The third infralabial is FIGURE 6. Amphisbaena heathi. Lateral (top), and ventral (middle) views of the cloaca and tail of the holotype, CAS 49374, and ventral view (bottom) of the tail of the paratype, CAS 49424, to show expression of the precloacal pores, and healing of the autotomized stump. 622 CALIFORNIA ACADEMY OF SCIENCES [Proc. 411m SER. small. Medial to it lies a pair of large shields on each side, occupying the position normally held by the malars. Neither pair contacts the postmental. The posterior ones are here considered as the enlarged extreme lateral segments of the postmalar row, which then numbers six segments. Enlarged postsupralabials (postinfralabials) larger than the preceding segment (s) in line, lie caudad to the angulus oris. Dorsally the first body annulus includes two large shields back of the first supralabial and the trapezoidal postocular. The second body annulus also sweeps forward to include two segments plus the enlarged first parietals. The second parietals represent a remnant of an intercalated dorsal half annulus. The anterior edge of the third body annulus sweeps shehtly anterior and its middorsal elements are slightly lengthened. The midventral elements of the first through fourth annuli are somewhat nar- rower but otherwise regular. This narrowing accounts for the anterior curvature of the ventral portions of these and the four succeeding annuli. More posterior annuli show no anterior curvature and lie normal to the lone axis of the trunk. There are 183 to 187 body annuli from the back of the third infra- labial, up to and ineluding the precloacal “pore bearing” row. The pectoral region is very short and not complexed. There are neither intercalated dorsal half annuli nor irregularities along the trunk. Only the paratype has an irregularity in the immediate precloacal region. There are 12 dorsal and 18 to 20 ventral segments to a midbody annulus. The cloaeal region of the male is characterized by four large round precloacal pores which are completely lacking in the female. Both speei- mens are clearly adult and have mature gonads. The female has the oviduet filled with a poorly preserved yolk mass in an early stage of development. There are eight pre- and 11 to 12 posteloacal segments, with the two lateral precloaeals very much the smallest, and three to four lateral half annuli. There are seven or eight caudal annuli up to and including the autotomy annulus after which the tail is freshly autotomized in the holotype and autotomized and healed in the paratype (which accounts for Sehmidt’s remarks). The lateral sulci are faintly indicated by elaboration of the aligned intersegmental sutures after approximately the thirty-fifth body annulus and until the eighth precloacal annulus. Dorsal and ventral sulei are, if present, expressed only by alignment of intersegmental sutures. The middorsal segments are approximately 1.5 times as long as wide and the midventral 1.2 to 1 as long as wide, so that there are no segments wider than long. There is relatively little change in segment proportions along the leneth of the trunk. Rance. Brazil. Rio Grande do Norte, lowland coastal region. Trunk Diameter mm eo A. heathi o A. carvalhoi Length b O Tail Oo 3) 8 12 Snout-vent Length - cm FIGURE 7. Amphisbaena. Scatter diagram of body diameter and of tail length, versus snout-vent length of specimens mentioned in the paper. Ficure 8. Amphisbaena carvalhoi. Dorsal, lateral and ventral views of the head of the paratype, MN R-—2094, from Pocéo, Pernambuco, Brazil. The line equals 1 mm. to scale. (Mrs. M. McKinney, del.) or) bo | VoL. XXXT] GANS: NEHW AMPHISBAENA DISTRIBUTION RECORDS. BRAZIL: Rio Grande do Norte: Ceara Mirim, CAS 49424 (paratype). Baixa Verde, CAS 49374 (holotype). Amphisbaena carvalhoi Gans, new species. DraGnosis. A very small form of Amphisbaena without major fusions of head shields, with slightly enlarged first parietals, with three supra- and three infralabials; and with two postgenial and no postmalar rows. The head is slightly compressed dorsoventrally, noticeably set off from the neck and bears a regular segmentation pattern. The form has 231 to 245 body annuli; 7 to 8 caudal annuli up to the autotomy constriction (all tails are complete); 19 to 22 eaudal annuli from the cloaca to the conical caudal spine; 12 to 14 dorsal and 16 to 18, generally 18, ventral segments to a midbody annulus; and four clearly expressed, small, round precloacal pores in both sexes. The color of preserved specimens is brown dorsally, fading out ventrally on the trunk. The nape and parietal aspects of the head and the dorsal surface of the tail are slightly darker. The lightened area extends across the supralabials and the rostral region. Pigmentation consists of general coloration of the dorsal segments, emphasized on the rectangular segmental centers. Laterally the margins fade, the rectangles decrease in size, and segmental drop out occurs. Ilototyrr. MN R2095, an adult male, collected by Antenor Leitao de Carvalho at Poecao, Municipio de Pesqueira, Pernambuco, Brazil. Pocao lies on the Serra de Acahy at 1035 m elevation, and the specimen was collected in broken up granite gravels near the top of the mountain range (cf. Carvalho, 19387). ParRATyPes. MN R1759, R2093—R2094, R2096—R2098; SU 17289-17290 taken with the holotype). DEsScRIPTION. Figure 8 of the present paper shows views of the head, figure 9 shows ventral and lateral views of the tail and figures 10 through AST srt) \SBBEBAL ERE SREEREREERER 32 ae re HEE S EEE aaa RE 9. eam isbaena carvalhoi. Ventral view of cloaca and tail of the para- type, MN R-2094, from Pocio, Pernambuco, Brazil. The line equals 1 mm. to scale. (Mrs. M. McKinney, del.) 626 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. 12 are photographs of the coloration and other aspects. Figure 7 compares body proportions with those of Amphisbaena heathi. Meristie data are listed in table 1. This is a small-sized species of Amphisbaena with an even brown dorsal color, emphasized on the parietal region and on the dorsal surface of the tail. The ventral surface is h¢eht colored from the tip of the tail up to and including the infralabials, first (and seecend) supralabials and the rostral tip. Some specimens (including the holotype) show a fainter version of the dorsal color on the ventral surface of the tail. Pigmentation is uniform across many of the dorsal segments; others show a rectangular darkened area. The countershading is achieved by a gradual fading of the segmental margins, proceeding more rapidly below the lateral line, by a decrease of the area occupied by the central rectangle, and by marked segmental drop-outs. Head segmentation is characterized by lack of major fusions, by large pretrontals, by postoculars almost as large as the frontals, and by medium- sized first and no enlarged second parietals. The head is relatively short, more or less pointed and dorso-ventrally compressed. The temporal muscles bulge noticeably and the middorsal sutures are faintly sunk from the level of the frontals to the third body annulus. The nuchal region is shehtly constricted. After this the trunk continues at constant diameter alone most of its leneth. The rostral is approximately of the size of the first supralabial and only its tip is visible in dorsal view. Pairs of sizeable nasals, large pretrontals, small frontals, and medium-sized first parietals form a sequence of large shields alone the dorsal surface of the head. The posterior edge of the frontals lies slightly anterior to the level of the angulus oris. There are three supralabials, the second the largest; the third the smallest. The pos- terior edge of the third supralabial generally lies slightly anterior to that of the third infralabial. The interlabial sutures run anteriorly at angles (to the labial edge) of 30°, 50°, 60°, and 75° respectively. The ocular is quadri- lateral, in contact with part of the dorsal edges of the second and third supralabials, in broad eontaet with the prefrontal, in point contact with the frontal, and margined posteriorly by the postoecular and possibly by the segment ventral to it. The mental is a small, rectangular segment only slightly larger than the small first infralabials, and approximately one-half the size of the septagonal postmental. The sceond intralabials are very large and are medially in full contaet with the postmental. The malars are large, and le immediately posterior to the large second and medial to the small third infralabials. They never contact the postmental. Two subtriangular first postgenials enclose the posterior tip of the postmental and are followed Figure 10. Amphisbaena carvalhoi. Dorsal, lateral and ventral views of the head of the holotype. MN R—2095. 628 CALIFORNIA ACADEMY OF SCIENCES [Proc. 41T1r Ser. Ficure 11. Amphisbaena carvalhoi. Dorsal and ventral views at midbody, of MN R-2095, to show segment proportions and pigmentation pattern. by three postgenial segments of the second row. One specimen has a tiny median segment midway between the two large first postgenials. There are no postmalars. The lateralmost postmalar segments of the first body annulus are widened.There is always a pair of enlarged postsupralabials and post- infralabials. Dorsally the first body annulus ineludes two large shields back of the third supralabial and the large postocular. The second body annulus gen- erally splits one segment dorsal to the angulus oris, giving rise to two dorsal half annuli. The anterior of these is normally an interealated one and the posterior one the dorsal continuation of the second body annulus, though irregularities occur and some specimens have two rather than one inter- ealated half annuli. The anterior dorsal half annulus includes two medium- sized, pentagonal first parietals as well as two to three smaller segments on each side. The posterior dorsal half annulus culminates in the second parietals which are secareely enlarged. The midventral elements of the second through sixth body annuli are somewhat narrowed. The posterior annul show no curvature and lie normal to the long axis of the trunk. There are 231 to 245 body annuli from the back of the third infralabial up to and including the precloacal porebearing row. The pectoral region VoL. XXXT] GANS: NEW AMPHISBAENA 629 Figure 12. Amphisbaena carvalhoi. Lateral (top) and ventral (bottom) views of the cloacal region and tail of MN R—2095, to show proportions, segment and pore arrangement, and color pattern. is not complexed. There are no irregularities or intercalated dorsal half- annuli along the trunk, though there generally is some irregularity in the ten precloacal annuli. There are 12 to 14 dorsal and 16 to 18, generally 18, ventral segments to a midbody annulus. The cloacal region is characterized by four medium-sized, but clearly expressed, round precloacal pores, present in both sexes and in juveniles. Six larger precloacal segments cover a half-moon-shaped precloacal shield, that is extended laterally by a small straight segment on each side, so that there generally are eight precloacals. Twelve to 14 radially arranged postcloacal segments, with the two midventral ones slightly enlarged, border the posterior lip of the cloaca. Lateral half annuli number three to four. There are 7 to 8 caudal annuli up to and including the clearly nar- rowed and constricted autotomy annulus, and 19 to 22 from the ecloaea up to and including the caudal tip. The tail swells slightly posterior to the autotomy constriction, and then becomes gradually conical, reducing down to the parabolic, vertically oval tip. The lateral sulei are clearly indicated after the fiftieth body annulus 630 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. and run up to the level of the cloaca. At midbody they are expressed as erooves half as wide as one of the fringing segments and filled with broken seoments. The dorsal suleus is barely indicated by alignment of interseg- mental sutures. It is clearly apparent only at the base of the tail and in the nuchal region. The ventral is indicated only by alignment of inter- segmental sutures. The middorsal segments are approximately 1.3 to 1.5 times as long as wide. The midventral segments vary from 0.9 to 1.2 times as wide as long. There is some minor variation in segmental proportions along the length of the trunk. Rance. Brazil. Pernambuco, Serra do Acahy. DISTRIBUTION RECORDS. BRAZIL: Pernambuco: Pocao, Munieipio de Pesqueira, elev. 1035 m., MN 1759, 2093-2094, 2095 (holotype), 2096-2098; SU 17289-17290. LITERATURE CITED CARVALHO, ANTENOR LEITAO DE 1937. Notas oecologicas e zoogeographicas sObre vertebrados do nordeste bra- sileiro. O Campo, pp. 12-14. GANS, CARL 1963a. Notes on amphisbaenids (Amphisbaenia: Reptilia). 7. Redescription and redefinition of Amphisbaena mitchelli Proctor and A. slevini Schmidt from the middle and lower Amazon, Brazil. American Mu- seum Novitates, no. 2127, 22 pp. 1963b. Notes on amphisbaenids (Amphisbaenia: Reptilia). 8. A redescription of Amphisbaena stejnegeri and description of a new species of Amphis- baena from British Guiana. American Museum Novitates, no. 2128, 18 pp. GANS, CARL and A. ALLAN ALEXANDER 1962. Studies on amphisbaenids (Amphisbaenia: Reptilia). 2. On the amphis- baenids of the Antilles. Bulletin of the Museum of Comparative Zoology, vol. 128, no. 3, pp. 65-158. KLAPPENBACH, MIGUEL A. 1960. Notas herpetologicas, I. Amphisbaenad munoai n. sp. (Amphisbaenidae). Communicationes Zoologicas del Museu do Historia Natural, Monte- video, vol. 4, no. 84, pp. 1-12. ScuMiptT, Kart PATTERSON 1936. Notes on Brazilian Amphisbaenians. Herpetologica, vol. 1, pp. 28-382. Marine Biological Laboratory LIBRARY JAN 2 7 1965 WOODS HOLE, MASS. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXI, No. 24, pp. 631-692, 11 figs. January 15, 1965 BEHAVIOR AND NATURAL REACTIONS OF THE NORTHERN ANCHOVY, ENGRAULIS MORDAX GIRARD, UNDER THE INFLUENCE OF LIGHT OF DIFFERENT WAVE LENGTHS AND INTENSITIES AND TOTAL DARKNESS By Anatole S. Loukashkin and Norman Grant California Academy of Sciences San Francisco I, INERODUCTION It has been known from times immemorial that certain fishes respond positively to artificial light and ageregate within illuminated zones. This peculiar behavior of fish has long been extensively exploited by fishermen. Torches and bonfires (still in use in some areas) were the first sources of arti- ficial light for attracting the schools of fishes into nets and fish traps. With advancing technology, these light sources gave way to petrol and acetylene lamps and electricity, especially to the latter because of its applicability for underwater illumination (Verheijen, 1958). With underwater illumination possible, a new trend in commercial fishing has been developing since the end of World War II (Ellison, 1953), particularly in the Soviet Union [ 651 J 632 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. (Borisov, 1950; Borisov and Protasov, 1959; Leskutkin, Nikonorovy and Patéev, 1955; Nikonorov, 1955, 1956, 1958, 1959a, 1959b; Terentiev, 1957). Instead of using conventional gear such as nets or traps, new, so-called “netless” fishing equipment has been introduced in certain fisheries. It con- sists of submerged electric lamps and the “fish pump.” The fish attracted by the light at night are sucked into the pump funnel and pumped directly into the vessel’s hold. In this technique, experiments have been made aiso to apply an electrical field within the illuminated zone so that the agere- gated fish would be forced to swim toward the pump funnel, which is made the positive pole (Nikonoroy and Patéev, 1959; Smith, 1955). More and more species of fishes and other aquatic organisms have been reported in the literature as reacting positively to sources of artificial ight under laboratory conditions or in the natural environments. Considerable research has been done on the structure and funetion of the fish eye (Baburina, 1955, 1958;\ Brett, 1959; Tamura, 1959; Vilter, 1950), on the ability of the fish to discriminate colors, and on innate preferential selee- tivity of monochromatic hehts (Arora and Sperry, 1958; Breder, 1959; Bull, 1957; Kawamoto, 1959; Loukashkin and Grant, 1959), on the ability of fish to respond differently to different intensities of artificial leht (Breder, 1959; Privolnev, 1956, 1958), and on many other specifie prob- lems related to fish behavior as it is affected by natural and artificial lights. Out of the voluminous literature on the subject published in recent years and of special interest to the writers, only a few papers are selected and mentioned below. Borisov (1950) recorded 42 species and subspecies of fishes which responded positively to electric light. His list includes marine, anadromous and freshwater fishes found in the USSR; in 1955, he listed more than 60 forms. In 1954, Radovich and Gibbs reported 44 species of marine fishes from the waters of California and western Mexico which re- sponded positively to eleetrie ght under natural conditions.t Baranoy (1955) listed 17 species for the northwestern Pacific, and Parin (1958) mentioned 54 marine fishes collected at night light stations during oceanic exploration of the Pacifie in 1954—55.? Among pelagic fishes of commercial importance, the clupeids, or herrine- like fishes, have been found the most responsive to artificial light, and 1 Since the date of Radovich and Gibbs’ report (1954), many more species of the fishes from the same area have been found to respond positively to electric light in the open sea (a continuously expanding unpublished list has been maintained by the California State Fisheries Laboratory at Terminal Island). While on research cruises of the California Fish and Game M/V Alaska in Mexican territorial waters in 1958 and 1961, the senior author recorded 20 species as supplementary to Radovich and Gibbs’ list of 1954. These fishes are as follows: Astroscopus zephyreus, Auxis sp., Carcharhinus lamiella, Cetengraulis mysticetus, Chloroscombrus orqueta, Cynoscion parvipinnis, Harengula thrissina, Menidia starksi, Mugil cephalus, Mugil sp., Nectarges nepenthe, Oligoplites sp., Polynemus sp., Pseudophallus starksii, Raja sp., Sphyraena sp., Sphyrna sygaena, Synodus lucioceps, Trachurops crumenophthalmus, and Upeneus sp. 2 A complete list of the fishes collected at night light stations by Parin in the Pacific Ocean during the 1954-1960 oceanological expeditions aboard the research vessel Vitiaz will be published by him and is in press. VoL. XXXI] LOUKASHKIN & GRANT: THE NORTHERN ANCHOVY 633 most of the references herein cited refer to this family. Species displaying < strong positive taxis to artificial light, readily aggregating in masses within illuminated zones, are as follows: sardines—Sardinops caerulea (Radovich and Gibbs, 1954: Rasalon, 1959), Sardinops sagax melanosticta (Borisov, 1955: Yudovieh and Kolegov, 1956), Sardina pilchardus sardina (Ver- heijen, 1957, 1958; Nikonorov, 1959), Sardinella macrophthalma (Breder, 1959), Sardinella aurita (Verheijen, 1958); herrings—Clupea pallasu (Gristehenko, 1951: Radovieh and Gibbs, 1954; Baranov, 1955; Borisov, 1955: Nikolaev, 1957), Clupea harengus harengus (Craig and Baxter, 1952; Borisov, 1955; Blaxter and Parrish, 1958; Radakov and Soloviev, 1959, Tihonoy, 1959; Zaitsev and Azhazha, 1959), Clupea harengus membras (Borisov, 1950, 1955); Caspian shads—Alosa brashnikovi brashnikovi, Alosa brashnikovi agrachanica, Alosa caspia caspia, Alosa kessleri kessleri, and Alosa kessleri volgensis (Borisov, 1955; Chugunova, 1955); Caspian sprats “kil’ka”—Clupeonella delicatula caspia, Clupeonella engrauliformas, and Clupeonella grimmi (Kremtstovy and Nikonova, 1949; Tokarev, 1949; Borisov, 1950, 1955; Bondarenko, 1951; Prihodko, 1951, 1957a, b; Leskutkin and Prihodko, 1951; Safronov, 1952; Evtéev, 1953; Leskutkin, Nikonorov and Patéev, 1955; Lovetskaya, 1955, 1958; Nikonorov, 1955, 1956a, b, 1958, 1959a, b; Chugunova, 1955; Terentiev, 1957; Borisov and Protasov, 1959) ; sprats—Sprattus sprattus sprattus (Blaxter and Parrish, 1958), Sprattus sprattus balticus, and Sprattus sprattus phalericus (Borisov, 1950, 1955) ; Pacific round herring—Ktrumeus acuminatus (Radovich and_ Gibbs, 1954); Pacifie thread herrine—Opisthonema libertate (Radovich and Gibbs, 1954); Atlantic dwarf herrine—Jenkinsia lamprotaena (Breder, 1959); and zunasi herrine—Harengula zunasi (Sasaki, 1959). Among other commercially important pelagic fishes which are known to respond strongly to artificial light are the following: anchovies—Engraulis mordax, Anchoa delicatissima, and Anchoa compressa (Radovich and Gibbs, 1954), Engraulis japonica (Borisov, 1950, 1955; Baranov, 1955; Parin, 1958), Engraulis encrasicholus (Verheijen, 1958), Engraulis encrasicholus pontica and Engraulis encrasicholus maeotica (Borisoy, 1950, 1955; Safia- nova, 1952, 1958; Kirillov, 1955; Radakov, 1956) ; mackerels—Scomber scom- brus (Blaxter and Parrish, 1958), Pnewmatophorus diego (Radovich and Gibbs, 1954), and Pneumatophorus japonicus (Borisov, 1950, 1955; Baranoy, 1955; Parin, 1958) jack-mackerels or horse-mackerels—Trachurus symmetri- cus (Radovich and Gibbs, 1954), Trachurus japonicus (Parin, 1958, Sasaki, 1959), and Trachurus trachurus (Borisov, 1950, 1955; Safianova, 1952, 1958: Radakov, 1956; Protasov, 1957; Blaxter and Parrish, 1958; Borisov and Protasov, 1959); saury—Cololabis saira (Pochekaey, 1949; Radovich and Gibbs, 1954; Baranov, 1955; Borisov, 1955; Yudovich, 1956; Parin, 1956, 1958; Gristehenko, 1957; Pokrovsky, 1957; Fukuhara, 1959); tunas— 634 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. Neothunnus macropterus and Euthynnus yaito (Hsiao, 1952; Tester, 1959) ; eod-like fishes—Gadus morhua morhua (Borisov, 1950, 1955; Lagunov, 1955), Gadus morhua macrocephalus (Baranov, 1955), Melanogrammus aeglefinus, Odontogadus merlangus ewrinus, and Boreogadus saida (Borisov, 1955).* The behavioral studies conducted at the California Academy of Sei- ences have been confined to four species of marine pelagic fishes: Pacifie sardine, Sardinops caerulea (Girard); northern anchovy, Engraulis mordax Girard; Pacific mackerel, Pnewmatophorus diego (Ayres); and Pacifie jack mackerel, Trachurus symmetricus (Ayres). The behavior and reactions of the sardine under the influence of white and colored hghts and darkness have already been explored (Loukashkin and Grant, 1959). The present paper sums up the results of the study of the behavior and reactions of the northern anchovy stimulated by artificial light of different wave lengths and intensities and by darkness. In essence, it is a continuation of the earlier experimental work on sardines. The equipment, facilities, and methods (fig. 1) used in the laboratory experiments for the larger part of the study were exactly the same as described earlier for the sardine; therefore, to avoid unnecessary repetition the reader is referred to that report. However, Light fixtures incosed in shades Plywood partitions Plywood top 32" wide a_i ———» +— 50° + 50° —_____> Fixture's support eee ———_s 4 > Adjustable scaled supports a Fluorescent tube Z Z Z Z Z Zz sm = Z Z Z = A Se eae error erere es eee Z A gat Sen (SZ 2 Z Zan 8 aR BS A Z Aaa 3 Sen BS Z Z aoc TTA Zee ZeSss SS aS Zee ae SET * eseeez AA AA > Se —=— SS SSS = eau _ZAZZZZZZZ-ZZZZZZZZZZZZZZZDEDEEEEEEEEEEEE \UCZ2ZZZ2ZZz « I2'6° > 21%" > Clamps LONGITUDINAL CUT CROSS-SECTIONAL CUT Figure 1. Sketch drawing of the experimental tank divided into three light zones for testing the anchovy’s ability to discriminate colors of the light and intensities of white light. (After Loukashkin and Grant, 1959.) changes in technique or equipment are noted and full information is pre- sented in appropriate sections below. For measuring light intensities, a Weston Illumination Meter, model 756, was used. This model is visual and cosine corrected, with direct dial read- 3% The size of the present report excludes the possibility of listing all the other marine and freshwater fishes whose phototactic responses to artificial light have been tested in recent years. Readers interested in this subject will find more information in the accounts by Baranov (1955), Blaxter and Parrish (1958), Borisov (1950, 1955), Parin (1958), Pochekaev (1949), Privolnev (1958), Protasov (1957, 1958), Radovich and Gibbs (1954), Sasaki (1959), and especially in the Verheijen report (1958) in which a review of the literature on fish responses to light is included. VoL. XXXII] LOUKASHKIN & GRANT: THE NORTHERN ANCHOVY 635 ines on the scales ranging from 0 to 500 foot-candles. The illumination meter was manufactured by the Weston Electrical Instrument Corporation of Newark, New Jersey. This account is based on the experiments carried out on two large schools of adult anchovies kept in the display tanks of the Steinhart Aquarium, California Academy of Sciences, at different times (approxi- mately two years apart). The majority of the experiments were devoted to the investigation of the ability of the anchovy to discriminate the same monochromatic lights, white light and darkness, which had been successfully applied in the experiments with the Pacific sardine (fig. 2). The second por- tion of the study involved the use of ultraviolet and infrared wave lengths and observations on the reactions of the anchovy to different intensities of the white light. As with the stock of the Pacific sardine used in earlier ex- periments, the northern anchovy schools were kept in a 1,000-gallon display tank illuminated with an ordinary 300-watt incandescent lamp which was suspended two feet above the water surface. Therefore, the fish used in the study can be considered “‘light-adapted” animals. The scientific names of most of the fishes mentioned in the text are based on Roedel (1953) for the California and Mexican species, and on Berg (1932-33, 1949), Borisov and Ovsiannikov (1951), and Svetovidov (1952) for the fishes of the USSR. Il. REACTIONS OF NORTHERN ANCHOVY TO LIGHT WAVE LENGTHS AND INTENSITIES (1) Preferential reactions to monochromatic lights, white light, and dark- ness. The ability of the anchovy to react differently to different light wave lengths was tested in a tank which could be divided into two, three, or four SPECTRAL ENERGY DISTRIBUTION BLUE "F" LAMP + BLUE FILTER “37 GREEN "F" LAMP +GREEN FILTER “40 PINK "F"LAMP+RED FILTER “I5 = > a = 30 - = = = | = | ~ 20 =! iL a ! e) & | x KS ” i= } . | Bees =20 (leas? 4 = 4000 4500 5000 5500 6000 6500 7000 = ANGSTROMS Figure 2. Spectral energy distribution of the monochromatic light sources used in the present study. (After E. A. Lindsay, 1948.) 636 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. zones and equipped with electric light sources of contrasting illumination. Results of the experiments of two-zone tests are presented in tables I—XI, those of the three-zone tests in tables XII and XIV, and of the four-zone tests in table XIII. In the two-zone tests different groups of six fish were subjected to the effect of a given pair of lghts or of ight and darkness. Each combination was used in two experiments consisting of six tests with 100 recorded observations, totalling 7,200 fish. Altogether, 79,200 fish are erouped in eleven tables for comparison of natural preference reactions to one type of illumination, or another. In testing the ability of the anchovy to distinguish green light from other colors, it was found that when this light ras paired with white leght, 5,424 fish out of 7,200 moved to or remained in the green-lght zone, displaying definite preference for this hght (74.34 per cent) over the white light (24.66 per cent), as seen from table I. When green and red lights were paired, this preference for green light rose to 97.86 per cent. The highest degree of negative reactions to red light in tests 2, 6, and 12 was manifested by total avoidance of the red light zone, as shown in table Il. When green light was presented along with blue light, anchovies were able to differentiate these two lights in contrast to the Pacific sardine, which was unable to do so (Loukashkin and Grant, 1959). As seen from table III, anchovies reacted preferentially to green light; 73.18 per eent of the individuals which were tested selected the “green zone,” com- pared to 26.82 per cent which showed preference for the “blue zone.” When paired with a darkened zone, the green-light zone was frequented by 6,918 fish (96.08 per cent), while only 282 (3.92 per cent) made occasional move- ments of short duration into the darkened zone. In tests 1 and 5, avoidance of the darkened zone was total (table IV). When testing the blue light paired with white hght or red light or darkness, fish responded favorably to the blue light. Table V shows a sheht preference for blue (52.15 per cent) over the white light (47.85 per cent), and amarked preference for blue (81.60 per cent) over the red (18.40 per cent) and (97.03 per cent) over darkness with four examples of total avoidance of the darkened zone in tests 1, 6, 7, and 12 (tables VI and VII). Red heht, when paired with white light (table VIII), as in the trials with green and blue lights, elicited negative responses on the part of the fish tested (in tests 1 and 9 only one fish entered the red light zone each time). Preferential reaction for the white light was as high as 88.39 per eent. The red hght attracted anchovies only when it was opposed by total darkness (92.97 per cent) as seen from table LX. When testing white light versus darkness, anchovies responded posi- tively to the former (97.88 per cent) and negatively to the latter (2.12 per cent) with total avoidance of that zone in tests, 4, 5, 6, 8, 10, 11, and 12 (table X). This is in full accord with other experiments in which an illumi- VoL. XXXI] LOUKASHKIN &€ GRANT: THE NORTHERN ANCHOVY 637 nated zone was presented with the darkened one (tables IV, VII, and IX). Diagrammatic interpretation of the relationship in the effects of dif- ferent lights on the anchovy’s discriminating ability tested in pairs is shown in figure 3. To evaluate the significance of the apparent preference responses of the fish to monochromatic lights, the same eroups of anchovies, either before or after experiment, were kept in a two-zone tank under a white light of the same intensity. The results obtained are presented in table XI, and they clearly display a normal distribution of 7,200 fish very close to a 50:50 ratio; however, the relationship varied from test to test. The average dis- tribution of anchovies for 12 tests was found to be 50.06 per cent for one zone, and 49.94 per cent for the other. These tests were considered as controls. After completing the series of experiments in a two-zone tank, anchovies were subjected to experiments in three-zone and four-zone tanks. In these experiments light intensities were maintained at a uniform level for all lights as in the two-zone experiments, or they were presented in different values. The latter modification was intended to see if the increment in light intensity would elicit a change in response because of brightness of illami- nation regardless of the color of light. The results of these experiments are presented in tables XIJ and XIII. The first four experiments in a three- 18.40 2.14 Hy61 88.)39 47.85 24.66 81.60 97.86 52.15 75.34 26.82 73.18 Figure 3. Diagrammatic interpretation of the relationships between the effects of different lights on the anchovy’s discriminating ability tested in pairs in the two- zone tank. Positive and negative reactions are expressed in per cent. All sources of light were maintained at 9 foot-candle intensity. 638 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. zone tank illuminated with white, green, and red lights, regardless of varia- tion in intensities, demonstrated overwhelming preference of the anchovies for the ereen light. The positive preference responses in these experiments for the green light averaged as high as 66.33 per cent (14,400 fish), though in separate cases this preference varied from 56.0 per cent to 72.0 per cent. Negative responses to the other two lights were as follow: 19.65 per cent for white light and 14.04 per cent for red light. These results are in full agreement with those obtained for the green light when tested in pairs with the others in a two-zone tank. In experiment WGD (table XII) the in- tensity of the green light was reduced to 6 foot-candles, while the white- light intensity was increased up to 30 foot-candles. The third zone was dark- ened. Again, anchovies responded in favor of the green light (69.35 per cent). In another experiment RDW (table XII) red and white hehts were presented in intensities of 30 foot-candles with the middle zone darkened. As anticipated, the white-light zone was frequented most of all (50.34 per cent), and red-light zone least of all (19.33 per cent). The reason why more fish were found in the darkened zone than in the red-light zone may have been that the white light penetrated the darkened zone. In other experi- ments, DBR-1 and DBR-2 (table XIII), the blue and red lights were applied, the third zone having been darkened. In both experiments with uniform intensities of 9 foot-eandles and with contrasting intensities (4 foot-eandles for the blue light, and 30 foot-eandles for the red heht) an- chovies displayed extremely high preference for the blue light (95.16 per cent and 98.67 per cent respectively ). Table XIII presents the results of experiments in a four-zone tank in which ereen, blue, and red lights of 9 foot-eandle intensity, and darkness were tested. Out of 9,600 fish, 80.19 per cent were found in the green-light zone, 15.34 per cent in the blue, 2.56 per cent in the red, and 1.91 per cent in the darkened zone. This preference of the anchovies for the green light perfectly agrees with all previous results. The last experiments in the present series were made to duplicate approximately the natural vertical distribution of the sunlight spectrum in water. The tank was divided into three zones as follow: daylight (white light with normal percentage of red light in it)* to imitate surface and near-surface illumination; green light for a deeper horizon of water mass; and blue light to represent the deepest horizon of the water medium in which the anchovy is found. In experiment DGB-1 the intensities of lights were maintained at 16, 7.8, and 0.5 foot-candles respectively; for DGB-2 these intensities were reduced to 6.0, 3.0, and 0.25 foot-eandles respectively (table XIV). The results of 12 tests with a group of nine anchovies in each of the two experiments show the same preferential tendency of the fish ‘ General Electric 20-watt ‘‘Daylight’’ fluorescent tube 24 inches long, ordering symbol F20T12/D. VoL. XXXII] LOUKASHKIN & GRANT: THE NORTHERN ANCHOVY 639 toward the ereen light as in all other experiments in which various com- binations of monochromatic and white lights were applied. This preference for the ereen light was found to be 48.57 per cent, in the experiment DGB-1, and 44.79 per cent in DGB-2 compared to 30.01 per cent and 30.24 per cent respectively for the dayheght and 21.42 per cent and 25.40 per cent for the blue heht. (2) Responses to ultraviolet wave length. In this series of experiments, low and high intensity sources of ultra- violet radiation were used. In the first set of experiments a “black light” 20-watt fluorescent tube (24 inehes long) manufactured by the General Electric Company (trade symbol F20T12/BLB) was used. Its spectro- eraphie characteristics are shown by the curve in figure 4, from which it is seen that this lamp emits a certain amount of visible light, too. This source 357 pS = a = ke = OF i 3 = ox lJ a n = 25- [o) [ea = 2) oO = et oie fo) 2 ec W a (ep) (Siz | al | a = = =! =e t ' ac WwW = fo) a a 5 2 =< (=) <= c 2500 3000 3500 4000 4500 5000 WAVELENGTH - ANGSTROMS Figure 4. Spectral energy distribution of the “Black Light Integral Filter Fluo- rescent Lamp” manufactured by the General Electric Company. Official drawing on file with the Company based on 40-watt lamp is reproduced here with written permission of the manufacturer. The curve is also typical for the 20-watt lamp (F20T12/BLB) used in the present study. 640 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. of ultraviolet radiation was first tested paired with monochromatic leghts, using the same colored fluorescent tubes and filters as in the preceding series. Because of the extremely low intensity of the “black light,” the other lamps were masked to reduce the intensity of colored lights to the level of the former, which was as low as 0.2 foot-candle. The results of six experi- ments covering the distribution of 57,600 fish are tabulated in tables XV— XVII. Paired with blue light, the ultraviolet wave length had no specific effect upon behavior of the anchovies. The average figures show a 50:50 distri- bution ratio (table XV). The ultraviolet-green combination revealed sheht preferential reactions toward the green light (54.79 per cent). This tend- ency was observed in all of the 24 tests (table XVI), while in ultraviolet- blue combination fish responses varied considerably from test to test, espe- cially in experiment UL-1. In experiments using ultraviolet light and red light, anchovies at first displayed very sheht but constant preference for the ultraviolet zone (52.25 per cent in experiment UL-5.). In the next experiment (UL-6) this prefer- enee rose to 91.70 pev cent varying between 80.50 per cent to 100.0 per cent from test to test, and averaging 71.79 per cent and 28.03 per cent for ultra- violet and red light respectively (table XVII). However, in this ease it can be assumed that it was not the attractive value of the ultraviolet rays that resulted in greater frequenting of the “black light” zone, but rather the repelling effect of the red light as revealed in previous experiments when monochromatic lghts were used and the anehovies avoided the red-light zone unless the alternative was darkness. The same avoidance reactions to- ward the red light were demonstrated earlier on the Pacific sardine (lou- kashkin and Grant, 1959). The next two experiments, with application of higher light intensity, were made in a two-zone tank. It was illuminated with clear light, and a souree of ultraviolet radiation alternately added to one of these zones. The white light was produced by the General Eleetrie 15-watt incandescent lamp (‘‘frosted’’), one in each zone, and the ultraviolet source was the same 20-watt “black hght” deseribed above. A light intensity of 10.5 foot-candles was maintained in both zones. The results of 24 tests involving the distribu- tion of 19,200 fish are shown in table XVIII. The averages for the white- light zone and white-light plus ultraviolet zone are almost identical: 49.84 per cent for the former, and 50.16 per cent for the latter. The fish seemed to be unable to differentiate one zone from the other, and the numbers of fish frequenting one zone or the other varied considerably from test to test, especially in experiment UL-7. Following this, an ultraviolet source of very high intensity was tested. For this purpose a “New Black-Ray Model B-100 (3660A)” equipped with 100-watt mereury spotlight bulb, ballast, and ultraviolet-transmitting Kopp 641 RN ANCHOVY ~ y NORTHE E LOUKASHKIN &€ GRANT: TH VoL. XXXT] AVY-HoRIG MON,, 94] UO posn se , TP ‘OU ddoy,, 10I[Y SUIQWIWUISuRA, JO[OTARAZ[N OY} JO Sotjtodoad [BAyedG “G ANNOY ‘T9INJOVJNUBU ay} Jo ASoVINOD “BIUAOFI[VO ‘Jolaqey uRg “OUT ‘syoNpoOrd JOTA-BAYM 94} AQ poanjzoejnuem duel ..(VO99E) OOT-A TePOIN SNOYDIWITTIW NI HLONAT SAVM oos O8b O9b Orr O@b O0b ose oge Ove oze [-S9 me TRANSMISSION PER CENT Ol- SI- Oc o¢4 ces Ov- St os+ 642 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. 41 filter was used. This source of ultraviolet radiation was manufactured by the Ultra-Violet Products, Inc., San Gabriel, California. The speetral- energy distribution of this lamp, with filter attached, is shown in figure 5. In addition, an extra filter (Corning Glass Works, no. 5840) was aequired in order to filter out most of the visible rays. Its properties are shown in figure 6. L. se 660 680 700 720 740 640 300 320 340 360 380 400 ieee e: Waar ae WAVE LENGTH MILLIMICRONS 280 70 60 a se See ent ae TRANSMITTANCE PER CENT FiGURE 6. Spectral properties of the ultraviolet-transmitting filter no. 5840 (7-60) of the Corning Glass Works. Courtesy of the manufacturer. VoL. XXXI] LOUKASHKIN € GRANT: THE NORTHERN ANCHOVY 643 At first, attempts were made to test the effect of ultraviolet wave lengths in total darkness by placing this source in one zone and keeping the other zone darkened. These, as all other tests herein reported, were carried out in a specially built dark room in the Steinhart Aquarium. Despite all pos- sible efforts, the investigators had to abandon this experiment because the use of both filters together failed to entirely filter out visible leht rays. Though of extremely low intensity and detectable by the human eye only after prolonged stay in the dark room, these rays, fortified by the ultraviolet wave leneth, created fluorescence in water. Reflections from the bottom and tank sides dimly illuminated the entire tank though a much brighter glow- ing spot appeared direetly under the lamp. Under this meager illumination the fish were able to still orientate and swim ina loose school formation and to continue their typical counter-clockwise movement in the tank. Ilowever, the speed of swimming slowed to one-half of normal. The intensity of light was far below 0.01 foot-candle. In the next trial, the 100-watt mereury spothght lamp (General Elec- trie H-100-SP4) was suspended over the center of the experimental tank. Its spectrographic characteristics are shown in figure 7. A dividing shield was removed. Light intensity at the surface of water directly under the lamp was 500-++ foot-candles with a rapid decrease toward the tank’s ends. One-half of the tank was covered with a clear glass plate to filter out ultra- violet rays. The other half remained open to allow ultraviolet radiation to enter. In this experiment (UL-9, table XIX) 64 per cent of 8,000 fish re- sponded positively to the zone covered with the glass plate, while 36 per cent entered the ultraviolet zone. The glass plate was then removed and both halves of the tank were subjected to ultraviolet radiation. In this experi- ment (UL-10, table XIX) 52.20 per cent of the fish entered one zone, and 47.80 per cent the other, which is close to a 50:50 ratio. After this, in order to evaluate the role of the clear-glass plate as a filter and its effect upon the numbers of fish gathering under it, an ultraviolet source was replaced by the KEN-RAD 300-watt reflector flood lamp emitting clear heht of the same intensity as the mercury spotlight lamp. One-half of the tank was again covered with the glass plate. This time (UL-11, table XIX) the fish distribu- ted themselves evenly (50.45 per cent and 49.55 per cent). Thus, it seems reasonable to assume that the 64.0 per cent response of the fish to the ultra- violet-free zone in experiment UL-9 was not incidental, and that the fish displaved a normal ‘avoidance reaction” toward the ultraviolet zone. In the last set of experiments with ultraviolet radiation, the light inten- sitv was reduced by half, and the procedure was different. In the experi- ment UL-12 (table XX) the tank was divided again into two zones by in- stalling a separating shield in the center. In each zone one KEN-RAD 500- watt reflector flood lamp emitting white light of 225 foot-candle intensity 47it SER. [ Proc. u 7 vay CALIFORNIA ACADEMY OF SCIENCE 644 ‘AuBdWOHN O11}D9[W [BloUer) oY} JO Aso NOD [‘scg~ Aaevnuere poiep ‘surjutad puodes ‘EQT-G7T ,,{Sdomuojsuviy, puv sdwey Aanoss,, Jorydued syy] “soanjory -nueu 34} Aq peyst[qnd vyep oy} UO peseq (IYSIT YoRI_) FdS—-O00T-H ‘ON (uinaA}0edS 9Y} JO UOTSa1 Jo[OIARAQ[N -IB9U) AW] IYSITIOdS 10JD9Yod AINIIOU WVM-YNOT ILAQO9[G [VIOUS 9Y} JO UOTINGIAWSIP ASious [eaqQOedS *) aANNDIY SWOULSONV ololey loleley a oos9s 0009 0OSS 0005 OOS O000b oose ooo¢e RADIANT POWER - WATTS VoL. XXXI] LOUKASHKIN &€ GRANT: THE NORTHERN ANCHOVY 645 was installed. In ten tests involving 6,000 fish, as anticipated, there resulted a more or less even distribution of fish (48.47 per cent and 51.53 per cent). In experiment UL-13 (table XX), one of the white lights was replaced with 100-watt mercury spotlight lamp (ultraviolet), and the positions of these two sources were alternated during the experiment. The intensity of light in both zones remained the same as in the previous experiment. Throughout all ten tests, the anchovies consistently preferred the white-light zone. Their responses for the white-light zone varied between 60.0 per cent to 100.0 per cent from test to test, averaging 72.1 per cent and displaying negative or avoidance reaction toward the ultraviolet zone (27.9 per cent) once again. Diagrammatic interpretation of the anchovy reactions toward the ultra- violet wave length is shown in figure 8. (3) Responses to infrared wave length. In this series of experiments the first tests were made in a two-zone tank; ene zone was exposed to infrared radiation, the other remained in total darkness. Instead of being six inches deep, as in all other experiments, the Figure 8. Diagrammatic interpretation of the anchovy’s reactions toward the ultraviolet wave length in relation to opposing monochromatic and white lights. Positive and negative reactions are expressed in per cent. 646 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4m Ser. water level was lowered to three inches and the lamp was suspended six inches above its surface. A G.H. 250-watt reflector heat lamp with red coating provided the source of the infrared radiation (its spectrographie features are shown in figure 9). A Corning filter no. 2540 was used to absorb all visible rays, transmitting infrared rays alone. Figure 10 shows the spectrographie properties of this filter. Bight anchovies were placed in an experimental tank two hours prior to testing and were kept there in total darkness. A recording of fish distribu- tion was made every ten minutes with the aid of dimmed ruby-red flash- light; this operation required not more than two or three seconds and only the fish in one zone were counted at a time. Altogether ten tests each of 30 reeorded observations were made covering the distribution of 2,400 fish. The results of experiment INF-1 are shown in table XXI. From the very start, it was clearly evident that the fish did not respond to infrared radia- tion. In both darkened and infrared zones, they behaved in exactly the same manner as did the Pacific sardine in total darkness (Loukashkin and Grant, 1959). The school was broken up; fish were scattered throughout the tank; swimming speed was slowed almost to a “stand still”; orientation was com- pletely lost, individual fish moving randomly, and ail the fish moved so close to the surface of the water that their dorsal fins and backs projected above the water. The average distribution of fish in ten tests was found to be about even: 51.33 per cent of the fish were recorded in the infrared zone, and 48.67 per cent in the darkened zone. To check the results, the infrared lamp was turned off, and the fish were kept in total darkness in both zones. Following the same procedure as in experiment INF -1, the investigators obtained exactly the same results: 48.58 per cent and 51.42 per cent (exp. INF-2, table X XI). After this, an infrared source was turned on again, and to the surprise of the observers, the fish began to concentrate under the lamp, though there was no visible change in the over-all situation. The mir- ror, placed under the lamp, revealed a tiny crack in the filter, through which just a pin point of red leht was reflected by the mirror. Intensity of this heht was about 0.001 foot-candle. The human eye, adapted to the darkness of the dark room, was unable to see this heht without the use of a mirror, but the anchovies were able to perceive such a meager light value and to re- spond to it very readily. The averages for ten tests (exp. INF-3, table X XI) show a definite preference by the fish for this zone (74.17 per cent) over the zone of darkness (25.83 per cent). In the next two experiments (INF-4 and INF-5, table XXII), one of the two zones was illuminated by white hght using a KEN-RAD 300-watt reflector floodlight lamp; light intensity at the surface of the water mea- sured 500 foot-ecandles. The other zone was illuminated with a G.E. 250-watt reflector heat lamp without red coating, which emitted both white light and 250 WATT R40/10 INFRARED (RED) WAVELENGTH BAND WATTS MICRONS 40- .76 4.92 .76 -3.0 171.99 3.0 -7.0 17.75 -40-7.0 194.66 INPUT EFFICIENCY = 78.75 % RADIANT POWER-MILLIWATTS PER 100 ANGSTROMS PER WATT &) 5 41.0 LS 20 25 3.0 10-27-55 WAVELENGTH - MICRONS GENERAL GB ELECTRIC ETL-GHZ LARGE LAMP DEPARTMENT APPLICATION ENGINEERING 250 WATT R40/I0 INFRARED (RED) INPUT RADIANT POWER-MILLIWATTS PER 100 ANGSTROMS PER WATT 3.0 3.5 4.0 4.5 5.0 55 6.0 @5 70 10-27-55 WAVELENGTH- MICRONS GENERAL @@ ELECTRIC ETL-GHZ LARGE LAMP DEPARTMENT APPLICATION ENGINEERING Figure 9. Spectral energy distribution of the 250-watt reflector heat lamp with red coating (infrared) manufactured by the General Electric Company. These graphs are official manufacturer’s copies reproduced here with the Company’s written permission. 648 CALIFORNIA ACADEMY OF SCIENCES [Proc. 41H SER. TRANSMISSION PERCENT WAVELENGTH MICRONS Figure 10. Spectrographic properties of the infrared transmitting filter of the Corning Glass Works’ manufacture no. 2540 (7-56). Courtesy of the manufacturer. infrared rays, as shown in figure 11. The light intensity of this lamp was also 500 foot-eandles. Observations were made every ten seconds. In 24 tests the fish behaved normally, maintaining typical school-forma- tion, and circled at normal speed in counter-clockwise direction. They dis- played preference for neither zone; of 19,200 fish 49.52 per cent were found in the white light zone, 50.48 per cent in the infrared zone. Two more experiments concluded the infrared studies (INF-6 and INF-7, table XXIII). In these, one zone was illuminated with white heht produced by the G.E. “soft white” fluorescent tube. The other zone was illu- minated with a similar source of light to which the G.E. 600-watt electric heater was added as a source of infrared radiation. The intensity of light in both zones was equal to 25 foot-candles at the water’s surface. As in the preceding case, the fish behaved normally and maintained typieal school- formation and circular path of movement. They showed no marked prefer- ence for either of the zones. The average figures for 24 tests in the two ex- periments involving 19,000 fish are as follows: 51.79 per cent for the white- light zone and 48.21 per cent for the white-light-plus-infrared-wave-leneth zone. For all practical purposes these figures show an even distribution of the fish, and as in all other experiments with application of infrared radia- 250 WATT ~~ R 40/1 INFRARED CLEAR WAVELENGTH BAND WATTS MICRONS 34- 40 .08 40 - .76 13.95 .76 -3.0 179.12 3.0 - 7.0 22.51 34-70 215.66 EFFICIENCY = 86.02 % — > a Zz ba ee ar = ee Bees ot RADIANT POWER-MILLIWATTS PER 100 ANGSTROMS PER WATT 3 ss 1.0 1S 20 rai) 3.0 10-27-55 WAVELENGTH - MICRONS GENERAL ® ELECTRIC ETL-GHZ LARGE LAMP DEPARTMENT APPLICATION ENGINEERING 250 WATT R 40/1 INFRARED CLEAR) 0.6 04 0.2) RADIANT POWER -MILLIWATTS PER 100 ANGSTROMS PER WATT INPUT 3.0 3.5 4.0 4.5 5.0 55 6.0 45 70 10-27-55 WAVELENGTH - MICRONS GENERAL Q@@ ELECTRIC ETL-GHZ f LARGE LAMP DEPARTMENT APPLICATION ENGINEERING FIGuRE 11. Spectral energy distribution of the 250-watt reflector clear heat lamp (infrared) manufactured by the General Electric Company. These graphs are offi- cial copies of the manufacturer. Courtesy of the General Electric Company. 650 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. tion they manifest very clearly the inability of the anchovy to perceive infrared radiation. (4) Responses to different values of intensity of white light. In an attempt to determine the ability of the northern anchovy to re- spond differently to different values of light intensity, the experimental tank was divided first into five zones, then into four, and finally into two zones. Lllumination was provided by G.E. incandescent (“frosted”) lamps emitting white light. In the five-zone arrangement, light intensities were as follows: 2, 10, 20, 50, and 100 foot-candles. After five tests in each of two experiments (INT-1 and INT-2, table XXIV), the positions of the hight sourees were reversed in order to avoid conditioning responses. Average response percentages for the gradient values in both experiments were close to each other, especially for the first three intensity gradients. The 20,000 fish used in 20 tests were distributed with respect to the five different light intensities as follows: 0.53 per cent — 2 foot-candles 6.03 per cent — 10 foot-candles 29.23 per cent — 20 foot-candles 41.94 per cent — 50 foot-candles 22.27 per cent — 100 foot-candles In the four-zone tank, illumination was provided by incandescent lamps of the same type and manufacture, which emitted white hght in 75, 125, 250, and 500 foot-candle intensities. As in the preceding experiment, the positions of light sources were reversed after the first five tests. The average response percentages in experiment INT-3 (table XXV) were: 14.20 per cent — 75 foot-candles 30.21 per cent — 125 foot-candles 37.16 per cent — 250 foot-candles 18.43 per cent — 500 foot-candles In these two arrangements of five and four intensity values, anchovies seemed to keep within the region of moderate light intensity; they shied away from the extremes. In each instance, most of the fish responded more positively to the lights of moderate intensities in centrally located zones and displayed an avoidance reaction to lights of the highest and lowest in- tensities. In the first arrangement, 71.17 per cent of 20,000 fish were found to frequent the two adjacent zones of 20 and 50 foot-candles; in the second—67.37 per cent of 10,000 fish frequented adjacent zones of 125 and 250 foot-candle intensities. The most striking example of avoidance by anchovies of the brighter zone was demonstrated in experiments INT-4, INT-5, INT-6, and INT-7, when intensities of white light were presented in sharply contrasting pairs, Vou. XXXII] LOUKASHKIN & GRANT: THE NORTHERN ANCHOVY 651 in which the higher value remained constant throughout the four experi- ments. An intensity of 500 foot-candles was opposed by intensities of 20, 10, 5, and 2 foot-candles. Each of the above-mentioned experiments con- sisted of three tests of 100 recorded observations of the behavior of eight anchovies. The preference responses to the lower values of light intensity over the 500 foot-ecandle intensity were found to be 65.50 per cent for 20 foot-eandles, 64.71 per cent for 10 foot-candles, 60.42 per cent for 5 foot- candles, and 83.71 per cent for 2 foot-candles. Of the 9,600 fish involved in these experiments, the average percentage in favor of all the lower inten- sities taken together equalled 68.52 per cent; that for the 500-foot-candle intensity, 31.48 per cent. ie TABLES TABLES J—-XI Records of experiments using the tivo-zone tests for determining the preference reactions of the northern anchovy (Engraulis mordax Girard) for monochromatic lights, white light, and darkness when presented in contrasting pairs. Light in- tensity was maintained at 9 foot-candles for all light sources. Each experiment con- sisted of six tests with 100 recorded observations made every ten seconds for six anchovies subjected to the effect of the light. Fluorescent tubes, manufactured by General Electric, and gelatine filters, made by Rascoe Laboratories, used in the present study were described by Loukashkin and Grant (1959). Table I Green Light Soft White Light Total Frequency of Occurrence Exp. Test Number Per cent Number Per cent Number Per cent GR-1 1 468 78.00 132 22.00 600 100 s 2 404 67.33 196 32.67 600 100 a 3 443 78.83 iby 7 26.17 600 100 4 399 66.50 201 33.50 600 100 5 453 75.50 147 24.50 600 100 3 6 425 70.83 1L0(S) 29.17 600 100 Total 6 2,592 72.00 1,008 28.00 3,600 100 GR-2 7 551 91.83 49 8.17 600 100 “A 8 468 78.00 132 22.00 600 100 ‘7 9 582 97.00 18 3.00 600 100 ie 10 523 87.17 ca 12.83 600 100 s 131 396 66.00 204 34.00 600 100 12 312 52.00 288 48.00 600 100 Total 6 2,832 78.67 768 21.33 3,600 100 Grand Total 12 5,424 75.34 EEG 24.66 7,200 100 652 CALIFORNIA ACADHMY OF SCIENCES [Proc. 4TH SER. Table II Green Light Red Light Total Frequency of Occurrence Exp. Test Number Per cent Number Per cent Number) Boreas GR-3 1 597 99.50 3 0.50 600 100 = "4 600 100.00 0 0.00 600 100 3 561 93.50 39 6.50 600 100 4 593 98.83 0 Hilts 600 100 5 597 99°50 3 0.50 600 100 6 600 100.00 0 0.00 600 100 Total 6 3,548 98.56 52 1.44 3,600 100 GR-4 7 598 99.67 2 0.33 600 100 a 8 599 99.83 1 0.17 600 100 = 9 591 98.50 9 1.50 600 100 s 10 514 85.67 86 14.33 600 100 Wal 596 99.33 4 0.67 600 100 12 600 100.00 0 0.00 600 100 Total 6 3,498 9717 102 2.83 3,600 100 Grand Total 12 7,046 97.86 154 2.14 7,200 100 Table III Green Light Blue Light Total Frequency of Occurrence Exp. Test Number Per cent Number Per cent Number Per cent GR-5 il 433 M2 167 27.83 600 100 4 2 429 71.50 1b7/at 28.50 600 100 x 3 383 63.83 PAILT( 36.17 600 100 4 381 63.50 219 36.50 600 100 4 5 480 80.00 120 20.00 600 100 6 465 UD) 135 22.50 600 100 Total 6 2,0 0 71.42 1,029 28.58 3,600 100 GR-6 i 563 93.82 37 6.17 600 100 e 8 419 68.33 181 31.67 600 100 9 280 46.67 320 53.33 600 100 10 505 84.17 95 15.838 600 100 ial 441 73.50 159 26.50 600 100 ws 12 490 81.67 110 18.33 600 100 Total 6 2,698 74.94 902 25.06 3,600 100 Grand Total 12 5,269 73.18 1,931 26.82 7,200 100 VoL. XXXII] LOUKASHKIN &€ GRANT: THE NORTHERN ANCHOVY 653 Table IV Green Light Darkness Total Frequency of Occurrence bBue. Test Number Per cent Number Per cent Number Per cent GR-7 af 600 100.00 0 0.00 600 100 ie 2 454 75.67 146 24.33 600 100 3 557 92.83 43 Tule 600 100 4 597 99.50 3 0.50 600 100 5 600 100.00 0 0.00 600 100 6 594 99.00 6 1.00 600 100 Total 6 3,402 94.50 198 5.50 3,600 100 GR-8§ G 594 99.00 6 1.00 600 100 < 8 559 aI 41 6.83 600 100 ra 9 592 98.67 8 Ise} 600 100 10 583 Siedler, ake 2.83 600 100 fet 594 99.00 6 1.00 600 100 1, 594 99.00 6 1.00 600 100 Total 6 3,516 97.67 84 2:00 3,600 100 Grand Total 12 6,918 96.08 282 3.92 7,200 100 Table V Blue Light Soft White Tah Total Frequency of Occurrence Exp. Test Number Per cent Number Per cent Number Per cent BL-1 1 180 30.00 420 70.00 600 100 os 2 464 ese 136 22.67 600 100 3 373 (HALL 227 37.83 600 100 4 333 55.50 267 44.50 600 100 5 367 Gulag 233 38.83 600 100 6 318 53.00 282 47.00 600 100 Total 6 2,035 56.53 1,565 43.47 3,600 100 BL-2 i 379 63.17 221 36.83 600 100 - 8 289 48.17 Sie 51.83 600 100 9 23K 39.50 363 60.50 600 100 10 280 46.67 320 Horao 600 100 ial 234 39.00 366 61.00 600 100 12 303 50.50 297 49.50 600 100 Total 6 122 47.83 1,878 Bye Ur 3,600 100 Grand Total 12 3,757 52.15 3,443 47.85 7,200 100 654 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. Table VI Blue Light Red Light Total Frequency of Occurrence Exp Test Number Per cent Number Per cent Number Per cent BL-3 1 474 79.00 126 21.00 600 100 oe 2 591 98.50 9 1.50 600 100 3 353 58.83 247 41.17 600 100 y 4 390 65.00 210 35.00 600 100 5 438 73.00 162 27.00 600 100 2 365 45.63 435 54.37 800 100 3 327 40.88 473 5912 800 106 1 360 45.00 440 55.00 800 = 100 5 386 48.25 414 Bylo 7its) 800 100 6 337 42.13 463 57.87 800 100 i 375 46.88 425 5oelzZ 800 = 100 8 351 43.88 449 56.12 800 100 9 358 44.75 442 55.25 800 100 ry > — 471 58.87 329 41.13 800 =100 Vou. XXXI] LOUKASHKIN ¢€ GRANT: THE NORTHERN ANCHOVY 661 Table XVII — Cont. Red Light Ultraviolet Radiation Total Frequency of Occurrence Exp. Test Number Per cent Number Per cent Number Per cent ial 444 56.50 356 44.50 800 100 12 423 52.87 377 47.13 800 100 Total 12 4,584 47.75 5,016 52:25 9,600 100 UL-6 tS 89 UG als3 (Alal 88.87 800 100 i 14 93 11.62 707 88.38 800 100 ss 15 136 17.00 664 83.00 800 100 rs 16 Te 14.62 683 85.38 800 100 ss au 156 19.50 644 80.50 800 100 18 109 13.62 691 86.38 800 100 19 35 4.38 765 95.62 800 100 " 20 25 SulliZ 1) 96.88 800 100 sf 21 6 0.75 794 99.25 800 100 Ze, 10 125 790 98.75 800 = 100 23 0 0.00 800 100.00 800 100 24 ill 2.62 779 97.38 800 100 Total 12 797 8.30 8,803 91.70 9,600 100 Grand a ae Total 24 5,381 28.03 13,819 71.97 19,200 100 TaBLE XVIIT Records of preference reactions of the northern anchovy (Engraulis mordax Girard) to white light and ultraviolet rays presented simultaneously in a two-zone tank. Eight fish were used in each test. White light zone Ultraviolet zone Two G.E. 15-watt Two G.E. 15-watt incandescent lamps imcandescent lamps. and 1 fluorescent Intensity 10.5 foot- G.E. “Black light’ Total candles tube. Intensity 10.5 f.-c. Frequency of Occurrence Exp. Test Number Per cent Number Per cent Number Per cent UL-7 il 282 35.25 518 64.75 800 100 : 2 240 30.00 560 70.00 800 8100 rn 3 316 39.50 484 60.50 800 100 y 4 325 40.63 475 59.39 800 100 sy 5 328 41.00 472 59.00 800 100 m=) 6 302 31.05 498 62.25 800 100 ee 7 482 60.25 318 39.75 800 100 8 520 65.00 280 35.00 800 100 d 9 546 68.25 254 ales 800 100 662 CALIFORNIA ACADEMY OF SCIENCES [Proc. 411m Ser. TABLE XVIII — Cont. White light zone Ultraviolet zone Two G.E. 15-watt Two G.E. 15-watt incandescent lamps incandescent lamps. and 1 fluorescent Intensity 10.5 foot- G.E. “Black light” candles tube. Intensity 10.5 Frequency of Occurrence Exp. Test Number Per cent Number Per cent Number Per cent 10 480 60.00 320 40.00 800 100 ial 484 60.50 316 39.50 800 100 12 434 54.25 366 45.75 800 100 Total 12 4,739 49.37 4,861 50.63 9,600 100 UL-8 15% 463 57.87 337 42.13 800 100 < 14 427 Boron 373 46.63 800 100 ie 15 416 52.00 384 48.00 800 100 = 16 474 59.25 326 40.75 800 100 i i 459 Deo 341 42.63 800 100 " 18 436 54.50 364 45.50 800 100 5 19 366 45.75 434 54.25 800 100 a 20 373 46.63 427 53.37 800 100 21 aS 7b 46.38 429 53.62 800 100 22 358 44.75 442 55.25 800 100 ij 23 Sol 41.38 469 58.62 800 100 is 24 378 47.25 422 S2ad 800 100 Total 12 4,852 50.54 4,748 49.46 9,600 100 Total 24 OES Oil! 49.84 9,609 50.16 19,200 100 TABLE XIX Records of preference reactions of the northern anchovy (Engraulis mordax Girard) to white light and ultraviolet rays presented in pairs in a two-zone tank. Hight fish were used in each test. General Electric 100-watt mercury spot- light lamp suspended over the middle of the tank. Intensity 500+ foot-candles Zone “A” covered with clear glass to filter out ultravio- let rays Zone “B” free for Total ultraviolet radiation Frequency of Occurrence Exp. Test Number Per cent Number Per cent Number Per cent UL-9 1 663 82.87 137 IPS} 800 100 oe 2 704 88.00 96 12.00 800 100 : 3 639 79.85 161 20.15 800 100 nf 4 569 Cledi2 231 28.88 800 100 oe 5 347 43.38 453 56.62 800 100 VoL. XXXI] LOUKASHKIN ¢€ GRANT: THE NORTHERN ANCHOVY 663 TABLE X LX — Cont. General Electric 100-watt mercury spot- light lamp suspended over the middle of the tank. Intensity 500+ foot-candles Zone “A” covered Zone “‘B’ free for Total with clear glass to ultraviolet radiation filter out ultravio- let rays Frequency of Occurrence Exp. Test Number — Per cent Number — Per cent Number Per cent 6 421 52.62 379 47.38 800 100 " Ul 418 52.25 382 47.75 800 100 e 8 402 50.25 398 49.75 800 =100 9 388 48.50 412 51.50 800 100 * 10 569 71.12 231 28.88 800 =100 Total 10 5,120 64.00 2,880 36.00 8,000 100 Clear glass filter removed, both zones under effect of ultraviolet radiation UL-10 iL 361 45.13 439 54.87 800 =100 es 2 375 46.88 425 53.12 800 =100 A 3 361 45.13 439 54.87 800 =—-100 * 4 448 56.00 352 44.00 800 =—-100 * 5 428 53.37 372 46.63 800 =—100 “ 6 440 55.00 360 45.00 800 =©100 = tf 441 55.12 359 44.88 800 =—100 5 8 450 56.25 350 43.75 800 =—100 9 416 52.00 384 48.00 800 =100 10 456 57.00 344 43.00 800 =100 Total 10 4,176 52.20 3,824 47.80 8,000 100 To check the role of the glass filter in experiment UL-9, it was introduced again, but instead of 100-watt mercury spotlight (ultraviolet) KEN-Rad 300-watt reflector flood light (white) lamps were installed in each zone. Light intensity was 500+ foot-candles. The results are shown below. Zone “A” covered with glass Zone “‘B” open free Total UL-11 al 432 54.00 368 46.00 800 100 is 2 431 53.87 369 46.13 800 100 3 398 49.75 402 50.25 800 100 4 393 49.12 407 50.88 800 100 5 408 51.00 392 49.00 800 100 6 404 50.50 396 49.50 800 100 7 371 46.38 429 53.62 800 100 8 420 50.25 380 49.75 800 100 9 Sil! 46.38 429 53.62 800 100 10 408 51.00 392 49.00 800 100 Total 10 4,036 50.45 3,964 49.55 8,000 100 664 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TABLE XX Records of preference reactions of the northern anchovy (Engraulis mordax Girard) to white light and ultraviolet rays presented in pairs in a two-zone tank. Six fish were used in each test. Experiment UL—-12 shows typical distribution of the fish when white light alone was applied, and experiment UL—13 shows change in dis- tribution after replacing the white-light lamp in one of the zones with a lamp producing ultraviolet radiation. Exp. Test Zone ‘‘A”’ ORC Bas Total Frequency of Occurrence Number Per cent Number Per cent Number Per cent KEN-RAD 300-watt KEN-RAD 300-watt reflector flood light reflector flood light lamp (white) In- lamp (white) In- tensity 225 f.-c. tensity 225 f.-c. UL-12 1 240 40.00 360 60.00 600 100 2 263 43.84 337 56.16 600 100 3 270 45.00 330 55.00 600 100 4 280 46.67 320 53.37 600 100 5 300 50.00 300 50.00 600 100 6 240 40.00 360 60.00 600 100 7 280 46.67 320 53.37 600 100 8 340 56.67 260 43.33 600 100 9 335 55.83 265 44.17 600 100 10 360 60.00 240 40.00 600 100 Total 10 2,908 48.47 3,092 51.53 6,000 100 KEN-RAD 300-watt G.E. 100-watt mer- reflector flood light cury spotlight lamp lamp (white) In- (ultraviolet). In- tensity 225 f.-c. tensity 225 f.-c. UL-13 1 600 =100.00 0 0.00 600 100 2 477 79.50 123 20.50 600 §=100 3 387 64.50 213 35.20 600 = 100 4 404 67.33 196 32.67 600 100 5 372 62.00 228 38.00 600 100 6 380 63.33 220 36.67 600 100 U 400 66.67 200 33.33 600 100 8 420 70.00 180 30.00 600 100 9 480 80.00 120 20.00 600 100 10 406 67.67 194 32.37 600 = 100 Total 10 4,326 72.10 1,674 27.90 6,000 100 VoL. XXXII] LOUKASHKIN & GRANT: THE NORTHERN ANCHOVY 665 TABLE X XI Records of preference responses of the northern anchovy (Engraulis mordax Girard) to infrared radiation and total darkness in a two-zone tank. Hight fish were used in each test. Exp. Test Infrared Radiation Total Darkness Total G.E. 250-watt re- Absolute Darkness flector heat lamp with red coating and Corning infra- red transmitting fil- ter. Frequency of Occurrence Number Per cent Number Percent. Number Per cent INF-1 il 167 69.58 83 30.42 240 100 a 2 135 56.25 105 43.75 240 100 3 135 56.25 105 43.75 240 100 4 106 44.17 134 55.85 240 100 5 162 67.50 78 32.50 240 100 6 96 40.00 144 60.00 240 100 rl 98 40.83 142 5 Om 240 100 8 74 30.84 166 69.16 240 100 9 153 63.75 87 36.25 240 100 10 106 44.17 134 55.83 240 100 Total 10 1232 Hilko3 1,168 48.67 2,400 100 Total Darkness Total Darkness INF-2 1 170 70.83 70 29.19 240 100 os 2 124 SEO 116 48.33 240 100 3 105 43.75 135 56.25 240 100 4 116 48.33 124 51.67 240 100 ‘ 5 120 50.00 120 50.00 240 100 6 95 39.58 145 60.42 240 100 ul 96 40.00 144 60.00 240 100 S 8 86 35.83 154 64.17 240 100 ss 9 150 62.50 90 37.50 240 100 10 104 43.33 136 56.67 240 100 Total 10 1,166 48.58 1,234" 542 2,400 100 Infrared: Visible Red. Total Darkness Corning filter cracked and began to transmit visible red in the in- tensity about 0.001 f.-c. INF-3 1 124 51.67 116 48.33 240 110 3 2 183 76.25 57 24.75 240 110 3 170 70.83 70 29.17 240 100 f 166 69.17 74 30.83 240 100 5 166 69.17 74 30.83 240 100 6 192 80.00 48 20.00 240 100 7 196 81.67 44 18.33 240 100 8 201 83.75 336) 16.25 240 100 a 9 SZ 80.00 48 20.00 240 100 : 10 190 TAIT 50 20.83 240 100 Nene ee Total 10 1,780 74.17 620 25.83 2,400 100 666 CALIFORNIA ACADEMY OF SCIENCES TABLE X Noll [Proc. 4TH Ser. Records of preference reactions of the northern anchovy (Engraulis mordax Girard) to white light and infrared radiation in a two-zone tank. Hight fish were used in each test. Exp. Test Clear Light KEN-RAD 300-watt reflector flood light lamp. Light inten- sity - 500 f.-c. Infrared Radiation Total G.E 250-watt reflec- tor infrared (heat) industrial lamp (white bulb). Light intensity - 500 f.-c. Frequency of Occurrence Number Per cent Number Per cent Number Per cent INF-4 il 377 47.13 423 52.87 800 §=100 ‘ 2 352 44.00 448 56.00 800 100 f 3 370 46.25 430 53.75 800 §=100 as 4 363 45.38 437 54.62 800 =100 = 5 365 45.63 435 54.37 800 §=100 : 6 352 44.00 448 56.00 800 =100 “ C 406 50.75 394 49.25 800 100 % 8 399 49.88 401 50.12 800 =100 i 9 393 49.13 407 50.97 800 =100 - 10 480 60.00 320 40.00 800 =100 : ile 441 55.12 359 44.88 800 §=100 12 413 51.62 387 48.38 800 =100 Total 12 4,711 49.07 4,889 50.93 9,600 100 INF-5 13 415 51.87 385 48.13 800 =100 : 14 405 50.62 395 49.38 800 =—100 * 15 413 51.62 387 48.38 800 §=100 $ 16 471 58.87 329 41.13 800 =100 ‘ 17 401 50.12 399 49.88 800 =100 s 18 442 55.25 358 44.75 800 =100 mi ils) 388 48.50 412 51.50 800 =100 : 20 379 47.38 421 52.62 800 ~=—100 ; 21 344 43.00 456 57.00 800 =100 : 22 385 48.13 415 51.87 800 ~=100 23 364 45.50 436 54.50 800 ~=100 24 389 48.63 411 51.37 800 =100 Total 12 4,796 49.96 4,804 50.04 9,600 100 Grand Total 24 9,507 49.52 9,693 50.48 19,200 100 VoL. XXXII] LOUKASHKIN € GRANT: THE NORTHERN ANCHOVY 667 TABLE XXIII Records of preference reactions of the northern anchovy (Engraulis mordax Girard) to white light and infrared radiation plus visible light in a two-zone tank. Hight fish were used in each test. Exp. Test White Light Infrared Radiation Total Fluorescent tube Fluorescent tube “Soft White’ light; “Soft White” light, Intensity 25 f.-c. and G.E. 600-watt electric heater. In- tensity 25 f.-c. Frequency of Occurrence Number Per cent Number Per cent Number Per cent INF-6 1 374 46.75 426 53.25 800 =100 2 390 48.75 410 51.25 800 100 3 373 46.63 427 53.37 800 = 100 4 406 50.75 394 49.25 800 =100 5) 389 48.63 411 51.37 800 =100 6 404 50.50 396 49.50 800 = 100 7 438 54.75 362 45.25 800 =100 8 459 57.37 341 42.63 800 =100 9 410 51.50 390 48.50 800 =100 10 434 54.25 366 45.75 800 =100 ial 429 53.62 371 46.38 800 =100 12 425 53.12 375 46.88 800 100 Total 12 4,931 51.36 4,669 48.64 9,600 100 INF-7 13 419 52.37 381 47.63 800 =100 4s 14 458 57.25 342 42.75 800 =100 15 420 52.50 380 47.50 800 =100 16 387 48.38 413 51.62 800 =100 ; 17 415 51.87 385 48.13 800 =100 18 388 48.50 412 51.50 800 =100 i) 394 49.25 406 50.75 800 =100 20 449 56.12 351 43.88 800 =100 21 402 50.25 398 49.75 800 =100 22 435 54.37 365 45.63 800 =100 23 416 52.00 384 48.00 800 =100 24 429 53.62 asa 46.38 800 =100 Total 12 5,012 52.21 4.588 47.79 9,600 100 Grand Total 24 9,943 51.79 9,257 48.21 19,200 100 rH SER. 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T €LNI 19) 49g AOQUUNAT qU9) 49g AOQUUN NY U9) 49g 4OQUNN’ 109) dq 49QUNN U9) 49g 49QUNN OL dxy DIUIAANIIQ, {0 Kuanbasy ‘2-"f 00 PAINO Se GEE WGA 1710.1, y, 9U0Z Ds, 2U0Z i, 2U0Z «Vy 9U0Z UL pasn asan YSY UAT, “Sa~pund-JOOL WOE 0} CG) -™ I NOXOXG MTSE “Y89] Yona wouf fijisuajur spr uw Hurhuns (sdwon) JUBISIPUDIUL) VYAY apLyNn pup Yup}, auoz -INOf HWIsn $}89a}] JUatpDAH YAY OF (panary XeBp1OW ST[NVIsUG) AAOYIUD UsaYyLOU ay) {O SUOLJIDIA AIUAlafaId {O SpLOIIY 670 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. TABLE XX VI Records of preference reactions of the northern anchovy (Engraulis mordax Girard) to different intensities of white light presented in sharply contrasting pairs in a two-zone tank. Hight fish were used in each test. Zone “A” Zone ‘‘B” Frequency of Occurrence Exp. Test ae Per cent Number Per cent Number Per cent (A) 500 f.-c. 20 f.-c. INT-4 1 256 32.00 544 68.00 800 100 ee 2 297 Oneal: 503 62.89 800 100 3 275 34.38 525 65.62 800 100 Total 3 828 34.50 1,572 65.50 2,400 100— (B) 500 f.-c. 10 f.-c. INT-5 1 319 39.88 481 60.12 800 100 * 2 302 BUCS 498 62.25 800 100 3 226 28.25 574 TASS) 800 100 Total 3 847 35.29 1,553 64.71 2,400 100 (C) 500 f.-c. SiGe INT-6 il 335 41.88 465 58.12 800 100 = 2 334 41.75 466 58.25 800 100 33 285 35.63 515 64.37 800 100 Total 3 954 39.58 1,446 60.42 2,400 100 (D) 500 f.-c. 2ife=Ge INT-7 1 210 26.25 590 135053 800 100 a 2 PAY 15.88 673 84.12 800 100 “ 3 54 6.75 746 93.25 800 100 Total 3 391 16.29 2,009 83.71 2,400 100 Grand Total 12 3,020 31.48 6,580 68.52 9,600 100 IV. DISCUSSION (1) On responses of the northern anchovy to monochromatic lghts in re- lation to reactions of other species. As stated in the introduction, the experiments deseribed in the preceding pages were carried out as a part of the general study on color vision in cer- tain species of the marine pelagic fishes of the Pacifie Ocean. The first stage of this study was published in 1959 by the present investigators. At that time they studied the Pacific sardine to determine the influence of monochromatic and white lights and darkness as environmental stimuli for elucidation of behavioral changes in schooling patterns and conversely to VoL. XXXII] LOUKASHKIN & GRANT: THE NORTHERN ANCHOVY 671 determine the ability of the sardine to discriminate colored and white lights qualitatively. Those experiments demonstrated clearly that different lights and darkness do affect school behavior and schooling patterns; also shown was the ability of the sardine to discriminate between lights on the basis of wave length. The sardines were attracted most of all by green light; they were repelled by both red light and total darkness (Loukashkin and Grant, 1959)/5 The results of recent experiments with the northern anchovy, to deter- mine their ability to discriminate among differently colored lights and dark- ness, are strikingly similar to those obtained in the experiments with the Pacific sardine. The anchovy, however, was able to differentiate green light from the blue, while the sardine failed to do so. In choice experiments in which the blue and white lights were presented responses of the anchovy in favor of the blue light (52.15 per cent) were lower than those of the sardine (73.05 per cent). Comparative data on the responses of these two species are assembled in the table XXVIT. TABLE X XVII Comparison of preference reactions of the northern anchovy and Pacific sardine to monochromatic and white lights in a two-zone tank. Responses in per cent Description Light source Light Source Green White ANSTO A See OO eS ee ee 75.34 24.66 Sangin w s0s-7: eee ee Re cee eee ee ea eee 78.63 Ze Sit Green Red PANT CHO Vay Neen ee eee cee ak EE See he 97.86 2.14 Sandinlewae ee meee mere oes aeeeer Sak. | lo DRS, he oe 95.25 4.17 Green Blue JNST\GL MONA eee hg hon Dee ea ee 73.18 26.82 AS Gai CUTTS aes ei ee 6 Ta ea 49.17 50.83 Blue White INTE NON AT See ae ee ee ees 52.15 47.85 Sain Cm st ee Cement cree nae ee 73.05 26.95 Blue Red PARITIC ING VV ioe eae ee nn ee nd Fe 81.60 18.40 Sars Cin gee eee eee ee ye a tw oS oe Sf S 97.26 2.74 Red White PANTIC HN O\VaV agree ee ee SE a 2 Be IL Oil 88.39 SNE IUELG NTS OS Ss a ee ate ela a) Sal I ae gh Sn ee 12.43 Seon 5 Verheijen (1956, 1958, 1959), speaking of the mass gathering phenomena of certain clupeids under the light at night at sea, disqualifies the interpretation of these phenomena in terms of “‘positive phototaxis,” “being attracted,” “intensity preferendum,”’ or “light optimum.’’ He considers all of them unsatisfactory and he attributes the above phenomena merely to a ‘‘mass photic disorientation’’ of the fish. 672 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. In the series of experiments comparing the effects of green, blue, red, and white lights in a two-zone tank (tables I, II, III, V, VI, VIII) only 2.315 fish or 10.72 per cent out of 21,600 fish responded positively to red light, and 19,285 fish or 89.28 per cent responded positively to the other lights. The negative reaction of the sardine toward red light was stronger: of 36,000 fish, 2.300 or 6.67 per cent were found in the red-light zone, and 33,610 fish or 93.33 per cent in the other ight zones. Comparative data are shown below to better illustrate the preferential responses of the anchovy and sardine to colored and white lights. TABLE XXVIII Responses in per cent Description Anchovy Sardine (CAS) 2Green Are oe a ee A ee 82.13 74.35 Bluey Reds awihiitemtosethie reese eens 17.87 25.65 (GB ABC yea a ee LE CS ee eee 53.52 Gomme Greeny ede awWihtitemtos ethic ines nanan 46.48 26.29 GO) tg CRG Gio ec rie oe a ey ee aes Ae eee ee 10.72 6.67 Green Blues Wihitestorether =e 89.28 93.33 (GBP We ee ee cee ne cy PIL CORRE 53.63 45.30 Green Blue mived sto ret hierar 46.37 54.70 In the three- and four-zone tank (tables XII, XIII, and XIV) green light was found to have the same effect as in the two-zone tank. The anchovies consistently responded in favor of the green light regardless of the intensities of the opposing lights. As was true for sardines (Loukashkin and Grant, 1959), anchovies were attracted mostly by the blue-green region of the spectrum. They showed a preference for green light over blue, for green over red, and for green over white. A preference for blue, in the ab- sence of green, over red and white was also evident. Similar results were obtained by Breder (1959) in his experiments using monochromatic lights of low intensities (2 foot-eandles) on Sardinella macrophthalma, Jenkinsia lamprotaenia, and some other fishes. He observes that, when contrasting colored lights are presented in pairs, “a general tendeney is evident for fishes to respond more definitely toward the shorter wave lengths (the blue and greens) and much less toward the longer wave lengths (reds).” The attractive value of the blue-green region of the spectrum was dem- onstrated in experimental studies by several Japanese behaviorists on young marine fishes, such as Oplegnathus fasciatus, Stephanolepis cirrhifer, Scomberomorus niphonius, Fugu niphobles, Fugu rubripes, Mugil cephalus, Girella punctata, Pempheris japonica, Trachurus japonicus, and the fresh- VoL. XXXI] LOUKASHKIN & GRANT: THE NORTHERN ANCHOVY 673 water Oryzias latipes (Kawamoto, 1959; Kawamoto and Konishi, 1952; Kawamoto and Takeda, 1950, 1951; Ozaki, 1951). Protasov (1957) investigated the responses of several species of the Black Sea fishes to monochromatic and white lights in the seaquaria of the Sebastopol Biological Station. He found that the ombre, Corvina wmbra, responded positively to violet, blue, light blue, green and white lights, and even to ultraviolet rays. The juvenile sturgeon “sevriuga,” Acipenser stellatus, was found to be phototatic to all types of lights applied but the responses were rather quantitative in character. The fish reacted positively only to higher intensities of the light, regardless of color. The horsemackerel “stavrida,” Trachurus trachurus, displayed indifference to the blue-green region of the spectrum, but when the intensity of the light was increased ( <5) the fish reacted negatively. This fish responded less positively to white light than to red, especially when water temperature was lowered.® A year later, Protasov (1958) published the results of his studies on the sensitivity of the fish eve to different wave lengths of light, establishing boundaries of the visible spectrum for certain marine and freshwater fishes, as shown in the following table: Species Limits in millimacrons OOM POSUINMCOM 4 nie o. Ga cde ohne = 420 = “620 PUCUDCTUSER (SLCULOLUS © 0 nase. os ons oo eS oe 2420) 9 100 TEMES te ny ME Son ek es os = A350) e100 SCORPTONU POTCUS ine ook se. sah) os bade bs =~ 400 = 600 MP AUTRESMOUENTS 4 AL Sis as Gk hs eed 500 see OU CUPMNUSACALDIO Frias en tlee ed oka =~ 480 — = 700 RU MOUSMICORGNUSIS Ret Lois oh ae sloe: = 400 °:—: = 620 He also tested the ability of the fish to discriminate monochromatic lights regardless of their intensities, applying the electrophysiological method suggested by Bongard (1955) and Bonegard and Smirnova (1959). This study revealed that Mugil auratus could distinguish blue, green, red, and orange lights from one another, but failed to distinguish blue from the violet and “extreme red” from red. Scorpaena porcus could discriminate red, vellow, orange, green, blue, and light-blue lights, but was unable to diserim- inate violet from the blue and “extreme red” from the red. The Black Sea turbot, Rhombus macoticus, could distinguish blue, light blue, green, yellow, orange, and red lights, but could not differentiate violet from the blue.’ ® The rather unusual reaction of the Black Sea horsemackerel (in view of the Kawamoto experiments with the Japanese horsemackerel) had been reported earlier by Safianova (1952), who demonstrated preferen- tial reactions of this fish to the orange-red illumination. 7 Protasov’s studies would have been more complete had he determined the ability of his fishes to react preferentially to certain wave lengths as well. 674 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. As to the responses of marine fishes to monochromatic lights in experi- ments under natural conditions in the open sea, there are several reports of interest to be mentioned in connection with the present study. Pochekaev (1949), testing the effects of overhead and submerged electric lights in the inshore waters of Sakhalin Island as possible attractants in local fisheries, obtained positive phototactic reactions as follows: (1) of the pond smelt, Hypomesus olidus; saury, Cololabis saira; and Eastern dace Leucascus brandti (all three in juvenile stage) to white, yellow, and violet lights; (2) adult dace, to white and yellow (violet light was not used); and (3) trout “kundzha,” Salvelinus leuwcomaenis, pond smelt and saury (all adult), to white light (the other two sources were not used ).* In addition to Pochekaev’s data on pond smelt, Baranov (1955) found this fish also responded readily to and aggregated in quantities around sub- merged electric lamps emitting blue, red, and white light, the latter ap- pearing to be the more effective attractant. As to the saury, Yudovieh (1956) and Gristechenko et al. (1957) deseribed the effectiveness of the blue and red lights in experimental saury catches in the northwestern Pacific. The blue light was used to attract the fish to the vessel (up to forty 500-watt incandescent lamps were installed alone one side of the vessel) ; the red light (not more than four 500-watt incandescent lamps on the op- posite side of the vessel) was used for operational purposes. When an aggre- gation would form in the blue light zone, the light would be extinguished and the red lamps would be turned on. The fish aggregation then would move from the darkened zone into the new dimly illuminated red zone where conical lift nets or blanket nets were installed. Upon lifting the nets, the red lights would be turned off and the blue lights turned on. This pro- cedure would be repeated several times at one night hght station.® Experiments carried out by Japanese fishery biologists in the open sea revealed the effectiveness of other monochromatic lights in attracting saury. Light of 4,000 angstroms (violet) wave length was found to be most ef- feetive, and that of 6,000 angstroms (red) the least effective (Takayama, 1956). * Pochekaev indicated that violet light attracted the squid, Ommastrepes sloani pacificus, in great masses. A marked preference for violet light over both yellow and white lights was displayed by an instant phototaxis following switching on of the violet lamp and in a short time by the mass aggregation of the squid schools within the illuminated zone. The other two lights were found to be good attractants too, but to a much lesser degree. Positive phototaxis toward white light was recorded for the California squid, Loligo opalescens, by Radovich and Gibbs (1954) and for the Mediterrancan squid, Loligo vulgaris, by Verheijen (1958). ® The use of a two-light arrangement as described by Yudovich (1956) and Gristchenko (1957) was introduced in the saury fisheries industry of Japan in the years following the end of World War TI; it has been highly appreciated by the fishermen whose catches have rapidly increased (Parin, 1956; Pokrovsky, 1957). The total annual landings of seury in prewar years (1936-1939) in Japan, before the use of artificial lights, amounted to less than 10,000 metric tons. With introduction of light attractants, the catch in 1947 reached 22,900 metric tons; in 1950, 126,400 metric tons: and in 1954, 292,700 metric tons (Rass, 1956). By 1957 the number of fishing vessels with electric-light equipment employed in saury fisheries exceeded 2,000; the annual catch for the same year reached 375,000 metric tons (Fukuhara, 1959). > VoL. XXXI] LOUKASHKIN &€ GRANT: THE NORTHERN ANCHOVY ( ~] ol In experimental studies of natural visual responses of the yellowfin tuna, Neothunnus macropterus, and little tunny, Euthynnus yaito, in the Hawaii Marine Laboratory of the University of Hawaii, electrie lights of white, blue, green, orange, red, and yellow colors were applied. The fish responded to colored and white lights, but “green light appeared to attract tuna” (Hsiao, 1952; Tester, 1959). According to Nikonorov (1956a), the Caspian anchovy-like sprat “kilka,” Clupeonella engrauliformis, in its natural environment “prefers” white light emitted by the submerged electric lamp when this light was presented paired with green or red lights. When green and red lights were presented together, the fish concentrated near the green lamp. In studying another Caspian sprat, Clupeonella delicatula caspia, under identical en- vironmental conditions and using monochromatic and white lights, Borisov (1955) found out that the most effective attractant was ordinary white light. The results of his trials, expressed in per cent, are shown below: white light i ers Oe yellow light... 27.6 orange light .:.. 5.9 ; | blue light peta oe) Catches made with application of : < ereen light SH Qre red leht see AOS no light 2. OL Totals eee ee 100.0 In evaluating the results of his exploration, Borisov observed, ‘‘Here, apparently, is reaction to the intensity of hight but not to the color of light.” From this remark it could be assumed that light intensities of the lamps used by Borisov and his associates were not uniform, and therefore the results he obtained were not conclusive.'° During the present investigation, two other species of schooling marine fishes, occasionally available for comparative study, were subjected to the influence of lights. One of them, the topsmelt, Atherinops affinis (Ayres), was kept in captivity for quite a long time; the second, the Pacific herring, Clupeaa pallasti Valenciennes, had been captured at the end of the spawning season and was used in tests following the fish’s initial adaptation to an artificial environment in the 1000-gallon display tanks of the Steinhart Aquarium. The results obtained with these fishes are shown in tables below: 1° Borisov never mentioned either light intensity figures or spectrographic values of his monochromatic lights in his report. This is also true for most of the Russian works cited in the present paper. 676 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. TABLE X XIX Records of responses of the topsmelt to monochromatic and white lights and darkness. Each experiment consisted of five tests of 100 recorded observations on a group of eight fish. Light intensity was maintained at the 5 foot-candle level. Pair of contrasting light zones Een ETTTRCHE One Ae Zone “‘B” Total number Frequency of distribution Number Per cent Number Per cent Number Per cent Top-l Green light Red light 3,970 99.25 30 0.75 4,000 100 Top-2 Blue light Red light 3,610 90.25 390 9.75 4,000 100 Top-3 Blue light White light 3,400 85.25 590 14.75 4,000 100 Top-4 Red light Darkness 2,536 63.40 1,464 36.60 4,000 100 As seen from this table, the preference responses of the topsmelt were toward the green and blue lights. As in the case of the Pacifie sardine and northern anchovy, the red light had no attractive value, except when it was opposed by darkness. MAB Linke No Records of responses of the Pacific herring to monochromatic and white lights. Hach experiment consisted of five tests of 100 recorded observations on a group of sia fish. Light intensity was maintained at the 10 foot-candle level. Pair of Contrasting light zones Experiment Lone Aw Zone ‘‘B” Total number Frequency of distribution Number Per cent aon Per cent Number Per cent Hirg-1 Blue light Red light pales) 70.50 885 29.50 3,000 100 Hrg-2 Green light Red light 1590 Eyer) 1,409 46.90 3,000 100 Hrg-3 White light Red light 1,524 50.80 1,476 49.20 3,000 100 Possibly, because of the physical and physiological condition of the her- ring captured during spawning season (in fact, a few females spawned on the tank’s walls soon after delivery of the captured fish), their responses to monochromatic and white lights are quite different from those of the an- VoL. XXXI] LOUKASHKIN &€ GRANT: THE NORTHERN ANCHOVY 677 chovy and topsmelt. Only in one of the three experiments did the herring display strongly negative reactions to red light and preferentially positive reaction to the contrasting light (blue). Gristehenko (1951) and Nikolaev (1957), speaking of the Pacific herring, and Tihonov (1959)—of the At- lantic herring, state that in experiments conducted in the open sea they found seasonal changes in phototactical behavior of this fish to the artificial lights. During the fattening period, both herrings reacted positively to light, and readily aggregated in masses in the illuminated zone. During the spawning season they became phototactically negative.tt This may well ex- plain the confusing results shown in the table XXX. In elosing this discussion on color vision in fishes, a few words should be said about the results of certain experiments in which “training” tech- niques have been successfully applied (e.g., feeding responses associated with a stimulus of restricted wave leneth). In the classical work of Reeves (1919) the sunfish, Lepomis gibbosus, and horned dace, Semotilus atro- maculatus, were trained to discriminate heht of longer wave lengths from light of shorter wave lengths and from clear light. Blennius pholis, used in experiments reported by Bull (1957) in which he applied differential con- ditioning, displayed unusual ability to qualitatively discriminate mono- chromatie hehts. One of the most interesting studies on color vision in fishes recently published is that of Arora and Sperry (1958). These investigators applied training techniques too. Astronotus ocellatus was used as an experi- mental animal. They found that this fish was able to distinguish red, blue, vellow, and green lights, and painted objects from each other and from various shades of grey. After training, the optic nerve was sectioned; the fish became blind. Regeneration of the sectioned optie nerve and restoration of vision took from 36 to 40 days; upon recovery of vision the fish displayed an ability to discriminate among the colors without further training. A fish which had not been trained prior to the blinding, by sectioning of the optic nerve, learned color discrimination as fast as normal fish. In the opinion of Arora and Sperry, the fish were able to discriminate between colors qualitatively rather than merely because of variation in intensity. In the much earlier work of Brown (1937), who worked with large-mouth black bass, it was concluded that, “in general, and excepting the violet, the degree of difference of different colors to bass is a funetion of difference in wave length.” Puchkoy (1954) states, “the ability of the fish to distinguish colors undoubtedly exists.” Discussing the results of von Frisch’s (1933) experiments, Puchhov observed, “if the fish were color blind, it would per- 11 Similar seasonal peculiarities in the behavior of certain marine fishes were recently reported by several Russian investigators: Parin (1956) in regard to saury; Safianova (1952, 1958) and Radakoy (1956) con- cerning the Black Sea anchovy, Engraulis encrasicholus pontica, and horsemackerel, Trachurus trachurus; and Lovetskaya (1958) about the Caspian sprat, Clupeonella delicatula caspia. Of the freshwater fishes, adult bream, Alburnus alburnus, in experimental studies in the laboratory carried out by Privolnev (1956) displayed phototactical periodicity with a change four times a year. 678 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER. ceive the red color as grey, and thus it would mistake red cups for the grey enes of corresponding brightness. However the fish always distinguished red cups from the grey ones of different degrees of brightness.” Walls (1942) flatly concluded that ‘no reasonable student of the problem [of the eolor vision in fishes| any longer doubts that fishes—all duplex teleosts at least—can experience hue as a sensation-quality apart from brightness.” Fifteen years later, Brett (1957) recognized Walls’ statement as the best formulated conclusion to the problem. As to the present study of the innate ability of the northern anchovy to react differently to light of different wave lengths, the authors are in- clined to consider the anchovy’s perception of the applhed leghts strictly as a function of wave length apart from the intensity of the light, in accord- ance with their earlier report on color vision in the Pacific sardine (Lou- kashkin and Grant, 1959). (2) On responses of the northern anchovy to ultraviolet wave length in relation to reactions of other species. Illumination of the aquatic media differs from that of the aerial environ- ments both quantitatively and qualitatively. Clark (1954) said that the sunlight upon entering water undergoes many changes. First of all, about 10 per cent or more of the light is lost because of reflection at the surface or beneath it. Traveling downward, the light is further modified not only in its intensity but also in its spectral and other properties.’” Baburina (1955) states that infrared rays are absorbed in the first meter layer of water. Ninety per cent of the red rays disappear within a depth of five meters; and ninety per cent of the green region of sunlight speetrum is absorbed before reaching thirteen meters of depth. Only violet and ultraviolet rays reach a depth of five hundred meters. The ultraviolet rays were detected 1,000 and more meters below the ocean surface. In conformity with this she maintains that “the eye of the fish is less sensitive to the red and more sensitive to the yellow, green, blue, and violet rays than the hu- man eye, but in contrast with the human eye it is also sensitive to the ultra- violet region of the spectrum.” Craig and Baxter (1952), speaking of the physiological importance of the ultraviolet component of natural ght in aquatic environments, observed that “in the sea water there is differential absorption so that the centre of maximum intensity is displaced somewhere towards shorter wave leneths, the precise effect depending upon depth and the nature of the sea water. We should not, therefore, be surprised to find marine creatures sensitive to a range including a portion of ultraviolet spectrum.” These theoretical reasonines concerning the ability of the fish 1° For instance, Boden ef al. (1960) found that in the Bay of Biscay sunlight passing through water “becomes steadily bluer with depth until at 400 meters the spectrum peaks sharply between 475 and 480 millimicrons.”’ VoL. XXXI] LOUKASHKIN &€ GRANT: THE NORTHERN ANCHOVY 679 eye to perceive ultraviolet wave lengths appear to be well founded and correct as has been demonstrated by recent experiments in both the open sea and in the laboratory. Protasov (1957), who apphed an electrophysiological method in the investigation of vision in a number of marine fishes, obtained definite proof that the Black Sea ombre, Corvina umbra, could respond to ultraviolet rays as positively as to the rays of the visible speetrum."® With facilities, sources of radiation, and techniques used in the present study, natural responses of the anchovy to ultraviolet rays seemed to be mis- leading because the fish responded inconsistently to ultraviolet light in various combinations with opposing wave leneths of light. These responses were found to vary from indifferent and negative to highly positive. Be- cause of this seemingly individualistic and confusing behavior of the anchovy in response to the ultraviolet radiation, further experimentation is neces- sary, especially in total darkness with the application of better filters totally isolating the wave lengths of the visible region of the spectrum. Breder (1959), who experienced the same difficulties with his experimental fishes, in his very earefully worded conclusion states “there is some evidence to support the view that some fishes show a positive reaction toward ultraviolet wave leneth, but this requires extended analysis...” He found out that males of Gambusia sp. were ultraviolet positive, the females negative. In his experiments Anoptichthys hubbsi reacted positively, and Anoptichthys jordani negatively in one ease; both species were slightly negative in another case. Jenkinsia lamprotaeni was found to be “ultraviolet positive to a very marked extent,” and Atherina stipes showed an individualistic behavior toward the ultraviolet, being either attracted, or repelled, or in- different. Brachydanio rerio displayed a strong positive reaction to the ultraviolet radiation. As to the use of sourees of ultraviolet radiation in tests in the open sea, only a few attempts have been made. A Netherlands research vessel carried cut experiments alone the Belgian coast but without success (de Boer, 1950). Craig and Baxter (1952), however, obtained immediate reactions of several species of marine fishes and other marine organisms to a sub- merged source of ultraviolet radiation (125-watt “black” ultraviolet lamp). They list the following fishes as influenced by ultraviolet rays: herring, mackerel, horsemackerel, dogfish, and whiting. Blaxter and Parrish (1958) also obtained positive aggregation of fish around the same source of radia- tion as used by Craig and Baxter, but they assumed that the reactions to the ultraviolet light might have been “due to the fluorescence from micro- organisms in the water” rather than to the ‘black light” itself. The inconclusive results of the experiments herein discussed prompt 13 Of freshwater fishes, the trout and pike have been known to perceive ultraviolet wave lengths of light (“Reflector,’’? 1949). 680 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SEr. the authors to consider the data obtained as a preliminary step toward further experimentation using improved sources of radiation and applying perfected techniques in the study of behavioral responses of the marine fishes toward ultraviolet radiation. (3) On responses of the northern anchovy to infrared radiation with refer- ence to experiments of other behaviorists. There was no evidence that they were attracted, repelled, or frightened by the radiation, which suggests that they did not perceive the infrared wave length. This conforms with work of Dunean (1956) who found that fingerling silver salmon, Oncorhynchus kisutch, failed to respond in any manner to intrared radiation. Breder (1959) has found no indicative evi- dence that fishes would respond differently to radiant heat (infrared radia- tion) and ambient temperature. (4) On responses of the northern anchovy to different values of intensity of white light. The experiments using white light of varying intensities in two separate arrangements (in one these intensities ranged from 2 to 100 foot-candles, in the other from 75 to 500 foot-eandles) revealed a natural ability of the anchovy to respond positively to intensities of moderate values regardless cf the order of light arrangement, and to react negatively to both the highest and the lowest intensities in the arrangement. On the other hand, in a series of experiments utilizing sharply contrasting light intensities pre- sented in pairs, the anchovies always responded positively to the lower values and displayed a marked avoidance reaction toward the brighter illumination. The results obtained in the present preliminary study suggest other tests, to be made in near future, might disclose the degree of sensi- tivity of the fish eve to the changes in the intensity of illumination, as well as the specific adaptation of the eve to certain intensity values as earlier demonstrated by Privolney (1958) on samples of young earp, Cyprinus carpio, and young teneh, Tinca tinca. He had found both were able to dif- ferentiate intensities of white light when these intensities were 75 per cent to 85 per cent greater than those to which the experimental fish were orig- inally adapted. As with other species found suitable for training, the northern anchovy and Pacific sardine should not present any difficulty in trainine: studies. Usually, the newly delivered wild anchovies and sardines began to take food after 5 to 7 days of acclimation to the artificial environment of the Stein- hart Aquarium. Following this, the fish were trained to break up the school, to ascend to the surface, and to swim close to the position oeeupied by the feeding person. The training consisted of propelling a tablespoon in the VoL. XXXI] LOUKASHKIN &€ GRANT: THE NORTHERN ANCHOVY 681 water for 10-15 seconds prior to dropping live food (brine shrimp) into water. Both the sardines and anchovies became conditioned to the sound of propelling the spoon, developing a feeding reaction within three to five trials (once a day), and they retained this response permanently. This con- ditioned response was of great help to the investigators at times when they had to pick up a few live specimens from the 1,000-gallon tank. V. SUMMARY 1. The present investigation was conducted in order to study experimen- tally the effects of various types of illumination on the northern anchovy, Engraulis mordax Girard, from the point of view of its ability to diserim- inate between different wave leneths of the light spectrum and different intensity values of the white heht. 2. The discriminating ability of the anchovy in regard to different types of visible and non-visible light radiation was explored in the specially con- structed dark room and an experimental wooden tank which was divisible into a number of light zones [in accordance with the nature of the experi- ment to be carried out]. 3. In the two-zone experiments the following paired lights were tested: ereen-blue, green-red, green-white, green-darkness, blue-red, blue-white, blue-darkness, red-white, red-darkness, and white-darkness. Ultraviolet was tested in pairs with green, blue, red, white, and darkness; infrared with darkness or with white light. 4. In the three-zone experiments the following combinations of lights were tested: green-red-white, green-white-darkness, red-white-darkness, blue-red- darkness, and blue-green-white (“daylight’’). 5. In the four-zone experiments the ereen-blue-red-darkness combination was tried. 6. In the two-zone experiments with monochromatie and white lights, the intensity was maintained uniformly at the 9 foot-candle level; in experi- ments with monochromatic Hghts and ultraviolet rays the intensity was adjusted to the maximum intensity of the ‘black lamp” which was equiva- lent to 0.2 foot-candle. In other experiments using ultraviolet or infrared wave leneths and white light or darkness, the intensities varied from almost zero to 500+ foot-candles. 7. In the three- and four-zone experiments, the intensities of monochro- matic and white lights tested were either uniform or of different values. 8. In all combinations of monochromatic and white lights, the effect of red light on the anchovy remained invariably negative in contrast to the sharply positive reaction of these fish toward other lights tested. 9. In two-zone choice experiments the positive reaction of the anchovy for eveen light was found to be 97.86 per cent over the red (2.14 per cent) ; 682 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4711 Ser. 75.34 per cent over white (24.66 per cent); 73.18 per cent over blue (26.82 per cent). The preference tor other lights, tested in pairs, was as follows: 81.60 per cent for blue hght over red (18.40 per cent), and 52.15 per cent over white (47.85 per cent); and 88.39 per cent for white light over red (ueoll per cent). 10. In the three- and four-zone experiments, the anchovies consistently dem- onstrated positive responses toward the green light as they did in the two- zone experiments. Hven a considerable increase in the intensities of the opposing lights could not alter the positive reaction to green light. 11. In the two-zone experiments using ultraviolet light paired alternately with ereen, blue, or red heht, the anchovies displayed three conflicting responses. These responses were “indifference” in ultraviolet versus blue light (50.15 per cent — 49.85 per cent), “shehtly negative” (45.21 per cent) when ultraviolet was contrasted with green light (54.79 per cent), and “highly positive” (71.97 per cent) when it was paired with red (28.03 per cent). 12. In the other two-zone experiments, when ultraviolet and white lehts of mueh higher intensities were tested, the results were confusing as des- eribed above. With respect to the ultraviolet light, the responses of the anchovies varied from negative or avoidance (36:64), through indifference (50:50), to positive (72:28). 13. In experiments utilizing infrared radiation, the anchovies seemed totally unable to perceive it. 14. In experiments intended to test the ability of anchovies to differentiate among different white light intensity values they seemed able to do so as evidenced in the tests with four and five intensity zones, and even more markedly in the two-zone experiments. 15. In the five-zone test arrangement in which intensities of heht ranged from 2 to 100 foot-candles, the fish responded preferentially to the moderate intensities of the central zones (29.23 per cent for the 20 foot-candle zone, and 41.94 per cent for the 50 foot-eandle zone, or 71.17 per cent for both). 16. In the four-zone test arrangement of white light used in intensities of 75, 125, 250 and 500 foot-ecandles, the anchovies reacted toward the mod- erate intensities of 125 foot-candles (30.21 per cent) and 250 foot-candles (37.16 per cent). 17. In the two-zone test experiments involving sharply contrasting inten- sities of 500 foot-candles, as a constant value, paired with much lower values ranging from 2 to 20 foot-candles, the reaction of the anchovies was always in favor (60.42 per cent to 83.71 per cent, averaging 68.52 per cent) of the lower intensity values. 18. The experiments herein deseribed and discussed reveal a few important factors in the reactions of the anchovy to light and darkness: (1) the an- VoL. XXXI] LOUKASHKIN &€ GRANT: THE NORTHERN ANCHOVY 683 chovy is a phototactic animal; (2) it is capable of discriminating qualita- tively between monochromatic (green, blue, red} and white lights; (3) it is able to distinguish green light from blue (the Pacifie sardine failed to do so); (+) it shows a preference for green and blue lights over white; (5) it proved to react strongly negatively to red light. However, the fish tolerated this type of illumination when it was tested as an alternative to total dark- ness, and showed a highly positive response in such a ease to the red light; (6) in its reaction toward the ultraviolet wave lengths it displayed a rather individualistic pattern of behavior; (7) it is unable to perceive infrared radiation; (8) it is capable of reacting differently to different intensities of white hght ranging from 2 foot-candles to 500 foot-candles. VI. ACKNOWLEDGMENTS This work has been carried on with funds provided by the Marine Re- search Committee as a part of the California Cooperative Oceanic Fisheries Investigations. The authors express their appreciation to Dr. Robert C. Miller, Senior Scientist of the California Academy of Sciences, for his keen interest in the studies and for encouragement and valuable suggestions during the progress of the research; to Dr. Earl S. Herald, Curator of Aquatie Biology, California Academy of Sciences, for his willing coopera- tion and valuable counsel; and to Dr. Alan E. Leviton, Curator of the De- partment of Amphibians and Reptiles, and Mrs. Lilian Dempster, Associate Curator of Fishes, California Academy of Sciences, for their extremely helpful assistance in preparing the manuscript. Acknowledgment is made also to Mr. Garth I. Murphy, Coordinator of the California Cooperative Oceanic Fisheries Investigations, and to Miss Carol Bumgardner for help- ful and constructive criticism of the manuscript; and to the management of the General Electrie Company, the Ultra-Violet Products, and the Corning Glass Works for providing important data and for their permission to reproduce official charts in the present study. LITERATURE CITED Arora, H. L., and R. W. SPERRY 1958. Studies on color discrimination following optic nerve regeneration in the cichlid fish, Astronotus ocellatus. Anatomical Record, vol. 131, no. 3, DS 029) BABURINA, E. A. 1955. Structural and functional properties of the fish eye. Trudy Sovestchania po voprosam povedenia i razvedki ryb 1953 g. (Transactions of the Conference on the problems of fish behavior and reconnaissance of the fish, 1953.) Edited by E. N. Pavlovsky, Academy of Sciences of the U.S.S.R., Ichthyological Commission, issue 5, pp. 90-103, 12 text figs., Moscow, Academy of Sciences Press, in Russian. 684 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. BABURINA, HE. A. — Cont. 1958. Characteristic properties of development of the eyes and their functions in fishes. Trudy Sovestchania po fiziologii ryb 1956 g. (Transactions of the Conference on physiology of the fishes, 1956.) Edited by E. N. Pav- lovsky, Academy of Sciences of the U.S.S.R., Ichthyological Commis- sion, issue 8, pp. 101-110, 10 text figs., Moscow, Academy of Sciences Press, in Russian. BARANOV, J. B. 1955. Experimental catch of fish with the aid of electric light in waters of Primorie. Izvestia (Bulletin) of the Pacific Research Institute of Fisheries and Oceanography, vol. 43, pp. 139-166, 7 text figs., Vladi- vostok, in Russian. Bere, L. S. 1932-33. 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Visserij-Nieuws, 2(11)127 (Summarized in the World Fisheries Abstracts, vol. 1, no. 4, July-August, 1950). BONDARENKO, A. F. 1951. The mass catch of the sprat ‘kil’ka’’ with electric light. Rybnoie Ho- ziaistvo, no. 12, pp. 8-10 (Fisheries Economy), Moscow, in Russian. BonGaArp, M. M. 1955. Colorimetry on animals. Doklady Akademii Nauk S.S.S.R. (Compte Rendus of the Academy of Sciences of the U.S.S.R.), vol. 103, no. 2. [After V. P. Protasov, Electrophysiological study of vision in fishes, 1958, as listed below], in Russian. BONGARD, M. M., and M. S. SMIRNOVA 1959. Color vision in man and animals. Priroda (Nature), no. 5. [After I. V. Nikonorovy, 1959, Application of light blocks in sprat fishing with fish pump, Rybnoie Hoziaistvo, no. 12, pp. 42-47, 5 text figs., in Russian. | Vou. XXXI] LOUKASHKIN € GRANT: THE NORTHERN ANCHOVY 685 Borisov, P. G. 1950. Fishing with the aid of electric light, 24 pp., 10 text figs., Moscow, in Russian. 1955. The behavior of fishes under the influence of artificial light. Trudy Sovestchania po voprosam povedenia i razvedki ryb 1953 g. (Trans- actions of the Conference on the problems of fish behavior and recon- naissance of fish, 1953.) Edited by E. N. Pavlovsky, Academy of Sciences of the U.S.S.R., Ichthyological Commission, issue 5, pp. 121— 143, 5 text figs., Moscow, Academy of Sciences Press, in Russian. Borisov, P. G., and N. S. OvSIANNIKOV 1951. A key to the commercial fishes of the U.S.S.R., 178 pp., 267 text figs., Moscow, in Russian. Borisov, P. G., and V. P. Prorasov 1959. The employment of incandescent lamps and selective sources of light for catching the fish. Rybnoie Hoziaistvo (Fisheries Economy), no. 12, pp. 48—49, Moscow, in Russian. BREDER, C. M., JR. 1959. Studies on social groupings in fishes. Bulletin of the American Museum of Natural History, vol. 117, no. 6, pp. 397-481, 28 text figs., 20 tables, and plates Nos. 70-81 with 23 photos., New York. Beer Je Re 1957. The sense organs: the eye. Chapter II, part 1, in The Physiology of Fishes, vol. I1, Behavior, Edited by Margaret E. Brown. Pp. 121-154, 10 text figs., and 1 table, New York. Brown, F. A., JR. 1937. Responses of the large-mouth black bass to colors. Illinois Natural His- tory Survey Bulletin, vol. 21, no. 2, pp. 33-55, 10 text figs., 11 tables and 1 color plate. 1digieit, 16 (Oy 1957. Behavior: Conditioned responses. Chapter III, part 1, in The Physiology of Fishes, vol. II, Behavior, Edited by Margaret E. Brown. Pp. 211-228, 1 text fig., 1 table, New York. CuucuNnova, N. I. 1955. Discussion (on responses of the Caspian sprat and shads to the under- water electric light). Trudy Sovestchania po voprosam povedenia i razvedki ryb 1953 g. (Transactions of the Conference on the problems of fish behavior and reconnaissance of fish, 1953.) Edited by E. N. Pav- lovsky, Academy of Sciences of the U.S.S.R., Ichthyological Commis- sion, issue 5, pp. 206-212, 6 text figs., Moscow, in Russian. CLARKE, G. L. 1954. Elements of ecology. Contribution no. 704 from the Woods Hole Ocean- ographic Institution, 534 pp., many text figs., New York — London, 686 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. CRAIG, R. E., and I. G. BAXTER 1952. Observations in the sea on the reaction to ultraviolet light of certain sound scatterers. Journal of the Marine Biological Association of the United Kingdom, vol. 31, no. 2, pp. 223-227, 1 plate with 2 photos., Cambridge University Press. DUNCAN, R. EK. 1956. Use of infrared radiation in the study of fish behavior. Special Scientific Report — Fisheries, no. 170, U.S. Department of the Interior, Fish and Wildlife Service, 16 pp., 9 text figs. ELLSON, J. G. 1953. Experiments with a “fish pump.” Commercial Fisheries Review, vol. 15, no. 2, pp. 27-29, 4 text figs., Washington, D.C. EREMTstTov, F.. Z., and M. G. NIKONOVA 1949. Catching the sprat ‘“kil’ka” with electric light. Rybnoie Hoziaistvo (Fish- eries Economy), no. 3, pp. 19-20, Moscow, in Russian. EXVTEEV, I. A. 1953. The influence of the moonlight on catching the sprat ‘kil’ka” with the aid of electric light. Rybnoie Hoziaistvo (Fisheries Economy), no. 4, pp. 16, Moscow, in Russian. FRISCH, K. 1933. Sind der Fische farbenblind? Zoologische Jahrbiicher, Abteilung fiir Allgemeine Zoologie, Bd, 33. [After Prof. N. V. Puchkov, 1954, listed below. ] FUKUHARA, A. 1959. Some improvements in the stick-held dipnet for saury fishing. An article in Modern Fishing Gear of the World, edited by Hilmar Kristjonsson, pp. 422-425, 11 text figs., London, England. GRISTCHENKO, G. I. 1951. Experimental herring fishing with application of the electric light. Izvestia (Bulletin) of the Pacific Research Institute of Fisheries and Oceanography, no. 34, Vladivostok. [After T. I. Privolney, 1956, as listed below], in Russian. GRISTCHENKO, G. I., V. V. LANDYSHEVSKy, and A. J. YUGANSON 1957. The results of the saury fishing in the Pacific Ocean using electric light. Rybnoie Hoziaistvo (Fisheries Economy), no. 3, pp. 15-17, 1 text fig., Moscow, in Russian. lela, Sj. (Ob 1952. Observations on the reaction of tuna to artificial light. (Contribution no. 24 of the Hawaii Marine Laboratory, University of Hawaii, Honolulu, Hawaii.) Special Scientific Report — Fisheries, no. 91, U.S. Department of the Interior, Fish and Wildlife Service, Washington, D.C., pp. 36-58, 3 text figs. and 8 tables. Hurst, P. M., Jr: 1952. Can fish see color? Pennsylvania Angler, vol. 21, no. 12, p. 5. Vout. XXXII] LOUKASHKIN € GRANT: THE NORTHERN ANCHOVY 687 KAWAMOTO, N. Y. 1959. The significance of the quality of light for the attraction of fish. An article in Modern Fishing Gear of the World, edited by Hilmar Krist- jonsson, pp. 553-555, 1 table, London, England. Kawamoto, N. Y., and J. Konisii 1952. The correlation between wavelength and radiant energy affecting photo- taxis. 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Commercial fishes of the U.S.S.R. Description of fishes (text to the atlas of colored plates of fishes), 787 pp., 10 color plates and many text figs., edited by Leo S. Berg in association with A. S. Bogdanov, N. I. Kozhin, and T. S. Rass; All-Union Research Institute of Marine Fisheries and Oceanography (VNIRO), Moscow, in Russian. NIKOLAEV, A. P. 1957. On the fishing of fattening herring in the inshore waters of Sakhalin island with employment of electric light. Rybnoie Hoziaistvo (Fish- eries Economy), no. 3, pp. 12-14, 3 text figs., Moscow, in Russian. NIKONoROV, I. V. 1955. Catching the Caspian sprat ‘“kil’ka” with the aid of underwater electric illumination and the fish pump. 32 pp., Moscow, in Russian. 1956a. The observations on the behavior of the Caspian sprats. Voprosy Ihtiologii (The Problems of Ichthyology), published by the Academy of Sciences of the U.S.S.R., Division of Biological Sciences, Ichthyo- logical Commission, issue 6, pp. 21-36, 10 text figs., Moscow, in Russian. 1956b. The theoretical foundation and experimental investigation of the fishing of the Caspian sprat with fish pump and underwater illumination. Rybnoie Hoziaistvo (Fisheries Economy), no. 1, pp. 73-77, 8 text figs., Moscow, in Russian. 1958. The catching Caspian sprat “kil’ka’’ with the fish pump and underwater illumination. 96 pp., Moscow, in Russian. 1959a. Application of the light blocks in sprat fishing with the fish pump. Rybnoie Hoziaistvo (Fisheries Economy), no. 12, pp. 42-47, 5 text figs., Moscow, in Russian. 1959b. The basic principles of fishing for the Caspian “‘kil’ka’”’ by underwater light. An article in Modern Fishing Gear of the World, edited by Hil- mar Kristjonsson, pp. 559-566, 15 text figs., London, England. Nrixkonorov, I. V., and A. H. PATEEV 1959. Sprat fishing with pump and underwater illumination with application of pulsating electrical current. Rybnoie Hoziaistvo (Fisheries HEcon- omy), no. 7, pp. 53-58, 4 text figs., Moscow, in Russian. VoL. XXXII] LOUKASHKIN € GRANT: THE NORTHERN ANCHOVY 689 OzAKI, HSAo 1951. On the relation between the phototaxis and the aggregation of young marine fishes. Report, Faculty of Fisheries, Prefectural University of Mie, vol. 1, no. 1, pp. 55-66, 10 text figs., 5 tables. Parin, N. V. 1956. Distribution of concentrations of saury and modern methods of saury fishing in the north-western part of the Pacific Ocean. Rybnoie Hoziaistvo (Fisheries Econemy), no. 7, pp. 57-61, 3 text figs., Moscow, in Russian. 1958. The ichthyofauna of the pelagial of the north-western part of the Pacific Ocean. Priroda (Nature), no. 5, pp. 60-66, 7 text figs., Moscow, in Russian. PARRISH, B. B., and R. HE. Craie 1951. Sonie layers in the sea. Nature, no. 168, p. 472, London. POCHEKAEY, V. M. 1949. Reactions of certain aquatic organisms to the electric light. Rybnoie Hoziaistvo (Fisheries Economy), no. 3, pp. 36-39, 3 text figs., Moscow, in Russian. Poxrovsky, A. N. 1957. On the Japanese saury fishing with the blanket net and light. Rybnoie Hoziaistvo (Fisheries Economy), no. 3, pp. 75-78, 2 text figs., Moscow, in Russian. PrIHopko, B. I. 1951. On sprat Caspian “kil’ka” fishing with electric light in winter. Rybnoie Hoziaistvo (Fisheries Economy), no. 8, pp. 43-46, Moscow, in Russian. 1957. Some new data on the behavior of the Caspian sprat “kil’ka” within electric light zone. Rybnoie Hoziaistvo (Fisheries Economy), no. 10, pp. 47-52, 5 text figs., 3 tables, Moscow, in Russian. PRIVOLNEV, T. I. 1956. The reaction of fish to the light. Voprosy Ihtiologii (The Problems of Ichthyology), published by the Academy of Sciences of the U.S.S.R., Division of Biological Sciences, Ichthyological Commission, issue 6, pp. 3-20, 5 text figs., Moscow, in Russian. 1958. The reactions of fish to different intensities of the light. Trudy Sovestchania po fiziologii ryb 1956 g. (Transactions of the Conference on physiology of the fishes, 1956.) Edited by E. N. Pavlovsky, Academy of Sciences of the U.S.S.R., Ichthyological Commission, issue 8, pp. 93-96, 1 text fig., Academy of Sciences Press, Moscow, in Russian. Provrasovy, V. R. 1957. Reactions of certain Black-Sea fishes to the light. Voprosy Ihtiologii (The Problems of Ichthyology), published by the Academy of Sciences of the U.S.S.R., Division of Biological Sciences, Ichthyological Com- mission, issue 9, pp. 144-146, 2 tables, Moscow, in Russian. 690 CALIFORNIA ACADEMY OF SCIENCES {Proc. 4TH SER. Prorasov, V. R. — Cont. 1958. Electrophysiological study of vision in fishes. Trudy Sovestchania po fiziologii ryb 1956 g. (Transactions of the Conference on physiology of the fishes, 1956), edited by EH. N. Pavlovsky, Academy of Sciences of the U.S.S.R., Ichthyological Commission, issue 8, pp. 111-114, 2 text figs., 2 tables, Academy of Sciences Press, Moscow, in Russian. PucHKov, N. V. 1954. The physiology of fishes, 369 pp., 108 text figs., Moscow, in Russian. RADAKOV, D. V. 1956. On studying behavior of the fishes during the fishing operations. Voprosy Ihtiologii (The Problems of Ichthyology), published by the Academy of Sciences of the U.S.S.R., Division of Biological Sciences, Ichthyo- logical Commission, issue 6, pp. 37-46, Moscow, in Russian. RapaAkovy, D. V., and B. S. SOLOVIEV 1959. The first experimental employment of the submarine for observation of the herring’s behavior. Rybnoie Hoziaistvo (Fisheries Economy), no. 7, pp. 16-21, 4 text figs., Moscow, in Russian. RapovicH, J., and HK. D. GiBss 1954. The use of a blanket net in sampling fish populations. California Fish and Game, vol. 40, no. 4, pp. 353-365, 7 text figs., 4 tables, Sacramento, California. RASALON, S. B. 1959. 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Reactions of the Black-Sea fishes to the electric light. Materials of the Conference held at the Ministry of Fisheries Industry of the U.S.S.R. [After V. R. Protasov, 1957, as listed below.] Moscow, in Russian. VoL. XXXI] LOUKASHKIN & GRANT: THE NORTHERN ANCHOVY 691 SAFIANOVA, T. E. — Cont. 1958. On certain correlations in the changes of reaction to the light in the anchovy “hamsa” and horsemackerel “stavrida” throughout the year. Trudy Sovestchania po fiziologii ryb 1956 g. (Transactions of the Con- ference on physiology of the fishes, 1956.) Edited by E. N. Pavlovsky, Academy of Sciences of the U.S.S.R., Ichthyological Commission, issue 8, pp. 97-100, Moscow, Academy of Sciences Press, in Russian. SAFRONOV, P. A. 1952. My experimental catch of the sprat ‘“kil’ka” with electric light. Rybnoie Hoziaistvo (Fisheries Economy), no. 4, pp. 53-54, Moscow, in Russian. SASAKI, T. 1959. The use of light attraction for traps and setnets. An article in Modern Fishing Gear of the World, edited by Hilmar Kristjonsson, pp. 556— 558, 5 text figs., 2 tables, London, England. SmitTH, K. A. 1955. Use of an electric attraction and guiding device in experiments with a “fishpump.” Commercial Fisheries Review, vol. 17, no. 2, pp. 1-7, 4 text figs., Washington, D.C. Sverovipov, A. N. 1952. Herring-like fishes (Clupeidae). Fauna S.S.S.R. (The Fauna of the U.S.S.R.), new series no. 48, vol. 2, no. 1, Fishes, 331 pp., 54 text figs., 53 plates on separate pages; Zoological Institute of the Academy of Sciences of the U.S.S.R., Academy Press, Moscow - Leningrad, in Russian. TAKAYAMA, S. 1956(?) Saury lift net fishing with light. Fisheries Statistics and Technological Development (in Pokrovsky, 1957, On the Japanese saury fishing with the blanket net and light, in Russian). TAMURA, T. 1959. Fundamental studies on the visual sense in fish. An article in Modern Fishing Gear of the World, edited by Hilmar Kristjonsson, pp. 543— 547, 1 text fig., 3 tables, London, England. TERENTIEV, A. V. 1957. Technical development in the fisheries industry of the U.S.S.R., in the past forty years. Rybnoie Hoziaistvo (Fisheries Economy), no. 11, pp. 31-39, 3 text figs., Moscow, in Russian. TESTER, A. L. 1959. Summing of experiments on the response of tuna to stimuli. An article in Modern Fishing Gear of the World, edited by Hilmar Kristjonsson, pp. 538-542, London, England. TitONOV., D>) 2 1959. Some supplementary data on the reaction of herring to the electric light. Rybnoie Hoziaistvo (Fisheries Economy), no. 4, pp. 13-14, Moscow, in Russian. 692 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. TOKAREV, A. K. 1949. The anchovyform sprat “kil’ka” and catching it with the aid of light. Rybnoie Hoziaistvo (isheries Economy), no. 6, pp. 33-86, Moscow, in Russian. VERHEWEN, F. I. 1956. On a method for collecting and keeping clupeids for experimental pur- poses, toget.er with some remarks on fishery with light-sources and a short description of free cupulae of the lateral line organ on the trunk of the sardine, Clupea pilchardus Wabl. Reprint from Pubblicazioni della Stazione Zoologica di Napoli, vol. 28, pp. 225-240, 1 text fig., 1 table with 4 photos. 1958. The mechanisms of the trapping effect of artificial light sources upon animals. Reprint from the Archives Néerlandaises de Zoologie, vol. 13, no. 1, pp. 1-107, 6 text figs., 1 plate. 1959. Attraction of fish by the use of light. An article in Modern Fishing Gear of the World, edited by Hilmar Kristjonsson, pp. 548-549, 1 text fig., London, England. VILTER, V. 1950. Adaptation biologique de l’appareil visuel et les structures rétiniennes de la sardine. Compte Rendus de Sceances de la Societe Biologique, vol. 144, nos. 3—4, pp. 200-208. WALLS, G. L. 1942. The vertebrate eye and its adaptive radiation. Cranbrook Institute of Sciences Bulletin, no. 19, XIV + 785 pp., 195 text figs., 1 plate. YUuDOVICH, J. V. 1956. Experimental saury fishing with the aid of electric light. Rybnoie Hoziaistvo (Fisheries Economy), no. 4, pp. 8-11, 2 text figs., Moscow, in Russian. YupovicH, J. V., and P. A. KoLEGOv 1956. Scouting of the sardine and experimental sardine fishing in the Sea of Japan. Rybnoie Hoziaistvo (Fisheries Economy), no. 2, pp. 17-19, 2 text figs., Moscow, in Russian. ZAITSEV, V. P., and V. G. AZHAZHA 1959. Submarine scientific expeditions on the “Severianka.’ Rybnoie Hoziaistvo (Fisheries Economy), no. 7, pp. 7-16, 7 text figs., Moscow, in Russian. INDEX TO VOLUME XXXI FOURTH SERIES New names and principal reference in boldface type abata, Esenbeckia, 74 Agama, 442, 482 abdominalis, Plenoculus, 32, 38 43, 44 agilis, 444, 445, 446, 447, 448, 449, 450, Stypommia, 88, 100 451, 452, 453, 475, 480, 484, 485 aberrans, Atractomorphia, 246 agilis agilis, 445 Abelpharus bivittatus, 476 agilis isolepis, 445 grayanus, 476, 481 agilis sanguinolenta, 445 pannonicus, 329, 337, 476, 480 aralensis, 446 persicus, 476 caucasica, 329, 336, 337, 475 Absidia scabra, 260 erythrogaster, 475, 482 acanthias, Squalus, 673 kirmanensis, 475, 482 Acanthodactylus, 456, 482 megalonyx, 475, 481 cantoris, 456, 458, 459, 480 melanura, 475, 481 cantoris blanfordi, 456, 457, 475 microlepis, 475 cantoris schmidti, 456, 457, 475 microtympanum, 475 micropholis, 475, 481 nupta, 336, 423, 442, 444, 446, 447, Acanthodrilidae, 187, 224 448, 475, 480, 484 Acanthodrilus, 194 persica, 451, 452, 453, 474, 475, 481 irpex, 213, 214 rubrigularis, 475, 481 Achnanthes prava, 351, 354 ruderata, 453, 475, 480 Acipenser stellatus, 673 sanguinolenta, 445, 446, 451 Acmaea, 106, 107, 114, 118 Agamidae, 442, 475 mitra, 343 Agamura persica, 474, 482 (Patelloida), 108 agamuroides, Cyrtodactylus, 438, 439, 474 (Patelloida) tessulata, 103 Gymnodactylus, 438 persona, 344 Agaricaceae, 260 tessulata, 107 agilis, Agama, 444, 445, 446, 447, 448, 449, virginea, 103, 107 450, 451, 452, 453, 475, 480, 484, 485 Acrochordus, 373, 376 Agama agilis, 445 Acrodectes philopagus, 503 agilis, Agama, 445 acuminatus, Etrumeus, 633 Amphiporus, 541 acutipennis gerstaeckeri, Atractomorpha, 246 isolepis, Agama, 445 Addisonia, 108 sanguinolenta, Agama, 445 adenanthus, Phaseolus, 297 Agkistrodon, 483 aeglefinus, Melanogrammus, 634 halys, 478, 480 aegrotus, Phymateus (Phymateus), 233 Agnostokasia, 504, 506, 508, 511 aegyptia, Walterinnesia, 470, 471, 478, 481 sublima, 504, 505, 506, 507, 508, 509, aegyptium, Dactyloctenius, 271 Salil, Sly aeruginosa, Anacystis, 264 Agonidae, 596 aestuans, Corchorus, 276, 294 agrachanica, Alosa brashnikovi, 633 aestuarii, Lyngbya, 266 Agrammus, 589 affinis, Atherinops, 675 agrammus, Agrammus, 565 Ceramodactylus, 474 Hexagrammos, 567, 576, 579 Afrosphena rhodesiensis, 241 Agrophis, 394 [693 | 694 CALIFORNIA ACADEMY OF SCIENCES Ahaetulla prasina, 373 prasina prasina, 378, 382, 385, 400, 405 prasina preocularis, 374, 379, 380, 383, 384, 387, 388, 389, 390, 393, 394, 398, 402, 404, 410 alascanus, Sebastolobus, 567, 568, 576, 579 alaskensis, Micrura, 533 alba, Amphisbaena, 615 Eclipta, 280 albibarbis, Chelommia, 98, 99 Tabanus, 98 albida, Zygonemertes, 541 albidum, Octoblepharum, 269, 29 albidus, Prosorhochmus, 552 albifrons, Cerebratulus, 516, 518, 519, 531 albilinearis, Stenotabanus, 78, 100 albini, Pyrgomorphella, 242 albiostris, Prionotus, 567, 579 albipes, Solierella, 3 albithorax, Scaptia, 70 alboater, Tabanus, 98 albocinctus, Tubulanus, 525 albocirculus, Tabanus, 93, 94 albomarginata, Euphorbia, 27, albonotatus, Tabanus, 94 albonuchalis, Pseudorabdion, 409 albopruinosus, Tabanus, 94 Alburnus alburnus, 677 alburnus, Alburnus, 677 alcalai, Oligodon, 409 Opisthotropis, 374, 393, 409 Allolobophora chlorotica, 224 bo 36, 46 trapezoides, 224 turgida, 224 Alosa brashnikovi agrachanica, 633 brashnikovi brashnikovi, 633 caspia caspia, 633 kessleri kessleri, 633 kessleri volgensis, 633 alpestris, Myersophis, 373, 374, 391 Alphestes galapagensis, 566, 579 multiguttatus, 566, 579 Alpsophylax, 334, 335, 336, 439, 482 crassicauda, 474 laevis, 334, 336 pipiens, 329, 334, 336, 474 alternata, Siphonaria, 111, 115, 116, 118 altipinnis, Micropogon, 556, 579 altivelis, Sebastolobus, 567, 579 [Proc. 4TH SER. amabilinus, Tabanus quadripunctatus, 89, 100 amabilis, Eragrostis, 271, 285 Hypopelma, 89 Phaeotabanus punctatus, 90 Tabanus, 89 Tabanus quadripunctatus, 100 amaranthina, Taphronota, 230 amarus, Phyllanthus, 275 amazonensis, Chelommia, 98 ameghinoi, Tabanus, 91 americana, Ramiella, 215 Ramiellona, 215, 218 americanum, Solanum nigrum, 278 Amphibia, 428, 473 Amphibians and_ Reptiles Steven C. Anderson, Amphiporidae, 517 Amphiporus, 517, 542, 545 agilis, 541 angulatus, 545 bimaculatus, 545 californicus, 545 cruentatus, 517, 518, 54.2 flavescens, 546 formidabilis, 517, 518, 519, 544: fulvus, 545 imparispinosus, 517, 518, 542, 546 imparispinosus similis, 546 leptacanthus, 542 leuciodus, 542, 543 pulcher, 542 punctatulus, 54.6 rubellus, 546 tigrinus, 545 virescens, 541 Amphisbaena, 614, 615, 617, 623, 624, 625, 626 alba, 615 earvalhoi, 613, 615, 617, 618, 623, 624, 625, 627, 628, 629 heathi, 613, 615, 617, 618, 619, 621, 623 mitchelli, 613 munoal, 613 from Iran, by 417-498 620, pretrei, 615 slevini, 613 vanzolinii, 613 vermicularis, 615 Amphisbaenidae, 456, 475 Vout. XXXII] Amphithrix violacea, 265, 302 Anacystis, 302 aeruginosa, 264 montana, 264, 302 analis, Clinocottus, 567, 579 analogus, Epinephelus, 566 Anchoa compressa, 633 delicatissima, 633 ancorus, Oligodon, 374, 391, 392, 394, 395, 404, 405, 408, 409 Anderson, Steven C., see Leviton, Alan E. Anderson, Steven C., Amphibians and Rep- tiles from Iran, 417-498 andreana, Coluber, 477 A new spine-throated grasshopper from the White Mountains of California (Orthoptera: Acrididae: Cyratcan- thacridinae), by Ashley B. Gurney, and David C. Rentz, 503-513 Anguidae, 475 Anguis, 483 fragilis, 480 fragilis cochicus, 475 angulata, Boiga, 373, 383, 389, 390, 393, 398, 402 angulatus, Amphiporus, 545 angustifrons, Tabanus, 98 angustirostris, Mirounga, 602 angustivittus, Tabanus, 91, 100 annulicornis, Tabanus, 71 annulifer, Oligodon, 409 annuus, Helianthus, 36 Anophrys bison, 567 Anoplopoma, 564, 570, 571, 572, 580, 582, 586, 590 fimbria, 567, 569, 579, 581 Anoplopomatidae, 567, 590, 591 Anoptichthys hubbsi, 679 jordani, 679 antennata, Maura, 228 Pterois, 566, 579 antiquatus, Hipponix, 111, 112, 113, 114, 118 Apathya cappadocica urmiana, 475, 481 aphanopterus obscuripilus, Phaeotabanus, 81 Phaeotabanus, 81 apicalis, Plenoculus, 13, 32, 40, 44, 47, 48 Aploactidae, 588 Aploactis, 569 INDEX 695 Aplopeltura, 405 boa, 373, 378, 379, 393, 400 apodus, Ophisaurus, 475, 479 aponina, Gomphosphaeria, 264 appendiculatus rus, 386 Rhacophorus appendiculatus, 386 aquarum-dulcium, Prostoma, 549 arabicus, Bufo viridis, 473 arachidis, Pyrgomorphella, 241 aralensis, Agama, 446 arborea, Hyla, 432, 479 savignyi, Hyla, 424, 430, 432, 473 arcatus, Paracirrhites, 566, 579 Archoplites interruptus, 577, 580 Arctica (Cyprina) islandica, 550 arenosus, Heliofugus, 125, 151, 152, 153, 156, 160, 166 Heliofugus (Heliofugus), 151 argenta, Tournefortia, 284 argenteum lanatum, Bryum, 269, 300 Argiacris, 503, 504, 506, 508 Argilophilus, 188 arguta, Eremias, 476 argyrophora, Dicladocera, 83, 84 armatus, Leptocottus, 567, 579 arrhizus, Rhizopus, 260 Artedius notospilotus, 567, 579 Ascalabotes pipiens, 334, 336 Ascomycetes, 260 asenoriatum, Prostoma, 549 asianus, Coluber jugularis, 477 appendiculatus, Rhacopho- asmussi, Uromastix, 475, 481 Aspergillus flavus, 261 flavus-oryzae, 261 fumigatus, 261 micro-virido-citrinus, 261 niger, 261 phoenicis, 261 sydowl, 261 terreus, 261, 262 versicolor, 261, 262 violaceo-fuscus, 262 Astrea gibberosa, 118 Astronotus ocellatus, 677 Astroscopus zephyreus, 632 ater, Pseudorabdion, 374, 393, 409 Atherina stipes, 679 Atherinops affinis, 675 696 CALIFORNIA ACADEMY OF SCIENCES atlanticus, Plenoculus, 43 Plenoculus davisi, 3, 37, 39, 43, 47 atra leucothorax, Scaptia, 70 Scaptia, 70, 100 Atractaspis, wilsoni, 470 Atractomorpha, 246 aberrans, 246 acutipennis gerstaeckeri, 246 atricornis, Tabanus, 98 atromaculatus, Semotilus, 677 aulicus capucinus, Lycodon, 378, 384, 386, 390, 392, 393, 395, 398, 401, 402, 404 Lycodon, 374, 377, 386 aurata, Mabuya, 464, 479 septemtaeniata, Mabuya, 423, 463, 476, 484 auratus, Mugil, 673 auribarbis, Dicladocera, 84 auriculae, Malacobdella, 551 auriculata, Natrix, 374, 379, 380, 389, 394, 404, 409 auripilis, Dichelacera 100 aurita, Sardinella, 633 auroguttatus, Paralabrax, 566, 579 australis, Echeneis, 170, 171, 172, 177 Remilegia, 169) 170) Wik 172, 173) WA. WIS UV, Lao) Remora, 170, 171 austriaca, Coronella, 477, 480 464, (Catachlorops), 86, baccatus, Phymateus (Maphyteus), 233 Baccharis salicina, 8 badia, Dicladocera, 86 bairdii, Cottus, 567 Delphinus, 178 balantina, Ramiellona, 202, 205, 214, 218 Balanus (Balanus) nubilis, 344 nubilis, 114 (Balanus) nubilis, Balanus, 344 balticus, Sprattus sprattus, 633 Banta, Benjamin H., and Alan E. Leviton, Remarks on the Colubrid genus Chil- omeniscus (Serpentes: Colubridae), 309-327 barbouri, Natrix dentrophiops, 391, 395, 399, 408 Barbourula, 385, 410 busuangensis, 382 [Proc. 4TH SER. barrosi, Heliofugus, 138, 144, 160, 166 Heliofugus (Heliofugus), 143 Baseodiscus punnetti, 552 Basidiomycetes, 260 basivittus, Tabanus, 95, 98 Bdellomorpha, 517 Bdellonemertini, 517 beadensis, Surirella, 356, 365, 366 beaumonti, Plenoculus, 3, 4.4. bedriagai, Teratoscincus, 474, 482 beebei, Eupomacentrus, 566, 579 Behavior and natural reactions of the north- ern anchovy, Engraulis mordax Gir- ard, under the influence of light of different wave lengths and intensities and total darkness, by Anatole S. Lou- kashkin, and Norman Grant, 631-692 bella, Nitzschia, 356, 364 Bellardia rubrofemorata, 92 berndti, Myripristis, 580 bicolor, Lycodon striatus, 478, 482 bilineata, Maticora intestinalis, 385, 401 bilineatum, Tetrastemma, 54.7 bilineatus, Lineus, 528 bimaculatus, Amphiporus, 545 Bimastos, 221 parvus, 221, 224 biobioensis, Heliofugus, 149, 155, 160, 166 Heliofugus (Heliofugus), 149 biocellata, Paranemertes, 538 bipartita, Oligodon, 409 bipartitus, Tabanus, 94 bison, Enophrys, 567 bitinctus, Tabanus, 95 bitorques, Calamaria, 374, 390, 391 bivittatus, Ablepharus, 476 Sibynophis, 374, 378, 382, 385, 401, 410 blandfordi, Acanthodactylus cantoris, 456, 457,475 Ophiomorus, 476, 481 Ophisops, 476, 481 Blennius pholis, 677 3741382. Blind limpet, Lepeta concentrica Midden- dorff, mantle cavity, habits, and habi- tat in the, by C. M. Yonge, 103-110 boa, Aplopeltura, 373, 378, 379, 393, 400 boharti, Plenoculus, 2, 6, 7, 28, 30, 34, 41 Boidae, 477 VoL. XXXII] Boiga, 482 angulata, 373, 383, 389, 390, 393, 398, 402 cynodon, 373, 378, 379, 385, 389, 390, 393, 400, 403 dendrophila, 373 dendrophila divergens, 374, 390, 391, 403 dendrophila latifasciata, 374, 393, 394, 404, 409 dendrophila multicincta, 374, 378, 400 drapizei, 373, 378, 400 philippina, 374, 390, 391 trigonata, 480, 482 trigonata melanocephala, 477 bolaui, Dichogaster, 224 bonariensis, Conyza, 280, 296, 297 bonduc, Caesalpinia, 274, 281, 282 Boraginaceae, 278 Borborocoetes, 502 borealis, Tubulanus, 522, 523 boregensis boregensis, Plenoculus, 21 perniger, Plenoculus, 2, 21, 48 Plenoculus, 2, 4, 5, 6, 8, 20, 21, 22 Plenoculus boregensis, 21 Boreogadus saida, 634 Borlasia camillea, 534 boulengeri, Rana, 334 Brachydanio rerio, 679 brachyura, Calamaria, 409 Bradynotes, 511 braminus, Typhlops, 476, 480 brandti, Lacerta, 476, 481 Leuciscus, 674 brannani, Siphonaria, 115 brashnikovi agrachanica, Alosa, 633 Alosa brashnikovi, 633 brashnikovi, Alosa, 633 brasiliensis, Ipomoea pes-carpae, 276 Brassica, 274, 279, 285, 290, 294, 295 juncea, 274, 297 brevipennis, Heliofugus, 143 Heliofugus (Heliofugus) 143 Heliofugus sulcipennis, 143, 159, 166 brevipes, Crytodactylus, 438, 474 Ophiomorus, 464, 465, 476, 481 Zygnopsis, 464 sulcipennis, brevirostris, Eremias, 480 brunnea, Tegula, 118 Bryaceae, 269 INDEX 697 Bryophyta, 268 Bryum, 268, 300, 301 argenteum lanatum, 269, 300 cuspidatum, 269 Bubalus, 396 mindorensis, 396 Buelliaceae, 263 Bufo, 405, 428 luristanicus, 430, 473, 481, 482 oblongus, 473 olivaceus, 429, 430, 472, 481 persicus, 430, 473, 481, 482 surdus, 430, 473, 481 viridis, 329, 330, 331, 428, 430, 479, 481 viridis arabicus, 473 viridis viridis, 473 Bufonidae, 428, 473 bullocki, Nemertopsis gracilis, 552 Bunopus, 336, 439, 482 persicus, 439, 474 tuberculata, 336 tuberculatus, 439, 473, 480 birgeri, Emplectonema, 536 busuangensis, Barbourula, 382 caeca, Lepeta, 104 caecum, Tetrastemma, 546 caerulea, Sardinops, 633, 634 Caesalpinia, 283 bonduc, 274, 281, 282 major, 281 Calamaria, 388 bitorques, 374, 390, 391 brachyura, 409 everetti, 373, 378, 400 gervaisi, 374 gervaisi gervaisi, 390, 391, 394, 399, 403, 407 gervaisi hollandi, 379, 393, 394, 403, 409 gervaisi iridescens, 384, 398, 399, 402, 409 gervaisi polillensis, 403 joloensis, 374, 387, 388, 410 mearnsi, 374, 393, 394, 409 suluensis, 373, 383 vermiformis, 373 vermiformis grayi, 374, 379, 389, 393, 394, 404, 409 zamboangensis, 374, 393, 394, 401, 409, 411 698 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. calcaetus, Dictyophorus, 228 capucinus, Lycodon aulicus, 378, 384, 386, californica, Paranemertes, 539 390, 392, 393, 395, 398, 401, 402, 404 Pheretima, 224 Capulus, 112 californianus, Mytilus, 343, 344 Caracanthidae, 588 Zalophus, 603 carbo, Mesomyia, 71, 72, 100 californicus, Amphiporus, 545 Mesomyia (Coracella), 71 Croton, 15, 16, 21, 27, 32, 36, 46 Tabanus, 71 californiensis, Cerebratulus, 532, 632 Carcharhinus lamiella, 632 calligaster calligaster, Calliophis, 390, 391, Carcinonemertes epialti, 552 394, 399, 404, 405, 408, 409 carinata, Pseudoboa, 471 Calliophis, 374 carinatus, Echis, 471, 472, 478, 480 Calliophis calligaster, 390, 391, 394, 399, Zaocys, 374, 378, 401 404, 405, 408, 409 Carinella cingulata, 524 gemianulis, Calliophis, 384, 398, 399, 402, dinema, 521 408, 409 exlineata, 521 mcclungi, Calliophis, 403, 408 polymorpha, 520 Calliophis, 411 rubra, 520 calligaster, 374 sclineata, 521 calligaster calligaster, 390, 391, 394, 399, sexlineata, 521 404, 405, 408, 409 speciosa, 520 calligaster gemianulis, 384, 398, 399, 402, | Carinoma, 516, 525, 529 408, 409 griffini, 526 calligaster mcclungi, 403, 408 mutabilis, 516, 519, 526 gracilis, 411 Carinomella lactea, 551 calliparea, Taphronota, 230, 231 Carinomidae, 516 Poecilocerus, 230 carnea, Paranemertes, 539 callosus, Tabanus, 91 carneus, Tabanus, 90 calogastra, Chrysops, 75, 76 Tabanus (Taeniotabanus), 90 caloptera, Dichelacera, 87 carpio, Cyprinus, 673, 680 Calotes, 482 earvalhoi, Amphisbaena, 613, 615, 617, 618, versicolor, 475, 480, 482 623, 624, 625, 627, 628, 629 Calothrix, 302 caspia, Alosa caspia, 633 crustacea, 265, 302 caspia, Alosa, 633 Clupeonella delicatula, 633, 675, 677 caspica caspica, Clemmys, 434, 473 Clemmys, 480 Clemmys caspica, 434, 473 Testudo, 434 caspius, Cyrtodactylus, 438, 474, 481 castanea, Dicladocera, 84 (Catachlorops) auripilis, Dichelacera, 86, 100 parietina, 265 stellaris, 265 camillea, Borlasia, 534 Canavalia, 283 rosea, 281 candidum, Testrastemma, 547 cantoris, Acanthodactylus, 456, 458, 459, 480 blandfordi, Acanthodactylus, 456, 457, 475 caucasica, Agama, 329, 336, 337, 475 schmidti, Acanthodactylus, 456, 457, caudaelineata, Coluber, 477 475 Caudata, 473 capax, Schizothaerus, 343, 344 caudolineatus caudolineatus, Dendrelaphis, capensis, Phymella, 243 378, 382, 401 Zygonemertes, 540 Dendrelaphis, 374, 379, 403 capistratus, Tubulanus, 525 Dendrelaphis caudolineatus, 378, 382, cappadocica urmiana, Apathya, 475, 481 401 VoL. XXXIT] luzonensis, Dendrelaphis, 374, 383, 390, 391, 395, 403, 404, 407 terrificus, Dendrelaphis, 374, 378, 379, 380, 382, 388, 389, 393, 394, 398, 399, 403, 404, 406, 411 Cawendia glabrata, 240 grossa, 240 Cellana, 103 Cenchrus, 256, 258, 285, 290, 293, 295, 296, 297 echinatus, 271, 294, 296 Centrarchidae, 566 Centrotrachelis loricatus, 453 Cephalothricidae, 516 Cephalothrix linearis, 526 cephalus, Mugil, 632, 672 Ceramodactylus, affinis, 474 doriae, 474, 481 Cerastes persicus, 472 Cerebratulus, 516, 529, 551 albifrons, 516, 518, 519, 531 californiensis, 532, 632 hercules, 532 lineolatus, 532 longiceps, 516, 519, 531 marginatus, 531 montgomeryi, 516, 519, 529, 531 occidentalis, 516, 518, 519, 530 Cetengraulis mysticetus, 632 Chaetopalpus, 71 coracinus, 71, 100 chagresensis, Esenbeckia, 73 Chalcides ocellatus, 479 ocellatus ocellatus, 476 Chama pellucida, 114 Chamaesiphonaceae, 264 Chaperina, 405 Chara, 268, 271, 285 kenoyeri, 268 zeylanica, 268, 303 Characeae, 268 Chelommia, 98 albibarbis, 98, 99 amazonensis, 98 crassicornis, 97, 98 Chelonia, 434, 473 Chelotabanus, 98 chevalieri, Eurotium, 260 Chilina dombeyana, 551 INDEX 699 Chilomeniscus, 309, 310, 317, 320, 321 cinctus; 309% SIs SIVA SIS si5a0S lisse 319, 321 ephippicus, 313, 317, 321, 322 punctatissimus, 309, 313, 315, 317, 322 savagei, 309, 313, 315, 317, 322 stramineus, 309, 310, 313, 315, 321, 323 stramineus esterensis, 315, 317, 323 stramineus fasciatus, 313, 317, 321, 322 stramineus stramineus, 310, 317, 323 Chirindites marshalli, 241 odendaali, 241 oldendaali, 241 Chitonotus pugetensis, 567 chloreus, Dendrochirus, 566 chlorogaster, Lacerta, 476 Chlorophyceae, 267 Chloroscombrus orqueta, 632 chlorotica, Allolobophora, 224 chlorostictus, Sebastodes, 566, 579 Chorizanthe parryi, 36 Chroococcaceae, 264 Chroococcus turgidus, 264 chrysarga, Natrix, 374, 378, 382, 385, 401 Chrysioekea paradisi, 373 chrysogenum, Penicillium, 262 Chrysopelea paradisi, 378, 379, 382, 383, 384, 387, 388, 389, 390, 393, 395, 398, 401, 403, 404, 405, 406 Chrysops calogastra, 75, 76 cuttula, 75 laeta sublaeta, 75 leucospila, 75 rossi, 75, 100 Chrysothamnus, 36 paniculatus, 15 Cichlosoma dovii, 566, 579 Cichlidae, 566 ciliaris, Eragrostis, 271, 285 cincta, Poecilocera, 230 cinctus, Chilomeniscus, 309, 311, 312, 313, 315. Sif, «31880319. es2i Gryllus, 230 Rutidoderes, 231 cingulata, Carinella, 524 Herse, 258, 299 cingulatus, Tubulanus, 516, 519, 524, 525 cirrhifer, Stephanolepis, 672 Cirrhitidae, 566 700 CALIFORNIA ACADEMY OF SCIENCES Cirrhitus, 570, 571, 572 marmoratus, 566, 579 rivulatus, 566, 569, 575, 584 citrinum, Penicillium, 262 clarionensis, Myripristis, 566, 579 claripennis, Tabanus (Taeniotabanus), 91 clathratus, Paralabrax, 566 Clemmys, 434, 483 caspica, 480 caspica caspica, 434, 473 Clinocottus analis, 567, 579 clippertonensis, Cosmarium, 267 Closterium parvulum, 267 parvulum majus, 267 Clupea harengus harengus, 633 harengus membras, 633 pallasii, 633, 675 Clupeonella delicatula caspia, 633, 675, 677 engrauliformis, 633, 675 grimmi, 633 Cobitus, 396 cochicus, Anguis fragilis, 475 Cochran, Doris M., see Goin, Coleman J. cockerellii, Plenoculus, 2, 4, 5, 7, 12, 15, 17, 18, 21, 28, 29, 40, 45, 46, 47 Cocoid palms, 281 Cocos, 279, 285 nucifera, 273 Cocotropus, 569 colasi, Heliofugus, 159, 166 Heliofugus (Heliofugus), 14.2 Collariheliofugus, 125, 126, 166 (Collariheliofugus) collaris, Heliofugus, 126 cryptocephalus curicoensis, Heliofugus, 128 cryptocephalus, Heliofugus, 126 collaris, Eirenis, 477, 479, 481 Euschatia, 126 Heliofugus, 124, 125, 126, 127, 128, 158, 166 Heliofugus (Collariheliofugus) , 126 collinus, Plutellus, 224 Cololabis saira, 633, 674 colonus, Parathias, 566, 579 Coluber, 465, 483 andreana, 477 caudaelineata, 477 dahlii, 477, 479 gemonensis, 477, 479 jugularis asianus, 477 [Proc. 4TH Serr. karelini, 466, 477, 481 ravergieri, 477, 480 rhodorachis, 329, 338, 423, 465, 466, 477, 480 schokari, 470 tyria, 477 ventromaculatus, 466, 477, 480 Colubrid genus Chilomeniscus (Serpentes: Colubridae), Remarks on the, by Ben- jamin H. Banta, and Alan E. Leviton 309-327 Colubrid snake Natrix pryeri from the Riu- kiu Islands, with description of a new subspecies, Interpopulation variation in the, by Edmond V. Malnate, and Harold E. Munsterman, 51-67 Colubridae, 465, 477 Comephoridae, 595 commune, Penicillium, 262 Compositae, 280 compressa, Anchoa, 633 concentrica, Lepeta, 103, 105, 106, 107, 109, 344 Lepeta (Cryptoctenidia) , 104 concolor, Rutidoderes, 231, 232 confervoides, Lyngbya, 266, 302 confirmatus, Tabanus, 83 Congiopodidae, 594 conirostris, Scincus, 462, 463, 476, 481, 485 conjuncta, Kaloula, 400 contortula, Navicula, 352, 353, 354 Conus, 283 convexa, Pinnularia, 354, 359 Convolvulaceae, 276, 283 Conyza, 285, 297 bonariensis, 280, 296, 297 cooperi, Yoldia, 551 coquimboensis, Heliofugus, 150, 160, 166 Heliofugus (Heliofugus), 152 Coracella, 100 (Coracella) carbo, Mesomyia, 71 Mesomyia, 71, 72 rubricornis, Mesomyia, 72 rufopilosus, Mesomyia, 72 Veprius, 72 coracinus, Chaetopalpus, 71, 100 corallipes, Tapronota, 229 Corchorus, 258, 285, 294, 295, 296 aestuans, 276, 294 corethrurus, Pontoscolex, 205, 219 VoLt. XXXI] Cornufer hazelae, 400 polillensis, 403 Coronella austriaca, 477, 480 coronella, Eirenis, 477, 481 coronelloides, Eirenis, 477 Corréa, Diva Diniz, Nemerteans from Cali- fornia and Oregon, 515-558 Corticium lactescens, 260 Corvina umbra, 579, 673 Cosmarium clippertonensis, 267 subprotumidum, 267 Cottidae, 567, 594, 595 Cottocomephoridae, 595 Cottunculidae, 596 Cottus bairdii, 567 couleensis, Cymbella, 356, 363 craspedogaster, Natrix, 65 crassicauda, Alsophylax, 474 crassicornis, Chelommia, 97, 98 cribricephalus, Heliofugus 137, 139, 160, 166 cribriceps, Heliofugus, 136, 140, 159, 166 Heliofugus (Heliofugus), 139 cristatus karelini, Triturus, 473 longipes, Triturus, 473 Triturus, 480 crocatus crocatus, Neurergus, 473 kaiseri, Neurergus, 473 Neurergus, 481 Neurergus crocatus, 473 Crossobamon, 336 eversmanni, 474 Crossodactylodes, 502 Crossodactylus, 502 Croton californicus, 15, 16, 21, 27, 32, 36, 46 Cruciferae, 274 cruentatus, Amphiporus, 517, 518, 542 crumenophthalmus, Trachurops, 632 crustacea, Calothrix, 265, 302 cryptocephalus curicoensis, Heliofugus, 127, 158, 166 curicoensis, Heliofugus (Collariheliofu- gus), 128 cryptocephalus, Heliofugus, 127, 158, 166 Heliofugus, 124, 125, 126, 128, 158 Heliofugus (Collariheliofugus) , 126 Heliofugus cryptocephalus, 127, 158, 166 Heliofugus (Euschatia), 126 (Cryptoctenidia) concentrica, Lepeta, 104 impressus, 136, INDEX 701 cuneatus, Planoculus, 2, 4, 5, 7, 12, 13, 15, 18, 19 Cunninghamella echinulata, 259 elegans, 260 Cunninghamellaceae, 259 curicoensis, Heliofugus (Collariheliofugus) cryptocephalus, 128 Heliofugus cryptocephalus, 127, 158, 166 curvassavicum, Heliotropium, 278, 284, 296 curvassavicus, Heliotropus, 36 cuspidatum, Bryum, 269 cyanophlyctis, Rana, 430, 433, 473, 480 Cyclocorus, 373, 374, 375, 392, 412 lineatus, 374 lineatus lineatus, 384, 390, 392, 394, 395, 398, 403 407 lineatus nuchalis, 379, 394, 404, 409 Cyclophis persicus, 466 cyclopium, Penicillium, 262 Cyclopteridae, 567, 596, 597 Cyclotella gamma, 350, 354 cylindrica, Pyrgomorpha, 245 Cymbella couleensis, 356, 363 rainierensis, 356, 363 cynodon, Boiga, 373, 378, 379, 385, 390, 393, 398, 400, 403 Cynoscion parvipinnis, 632 Cyperaceae, 272 (Cyprina) islandica, Arctica, 550 Cyprinus carpio, 673, 680 Cyrtodactylus, 334, 336, 437, 481 agamuroides, 438, 439, 474 brevipes, 438, 474 caspius, 438, 474, 481 fedtschenkoi, 438, 474, 481 heterocercus, 439, 474, 482 kachhensis, 438 kirmanensis, 438, 474 kotschyi, 438, 474, 479 longipes, 438, 474, 482 macularius, 435 russowl, 474 scaber, 437, 438, 441, 474, 480 zarudnyl, 438, 474, 482 Dactyloctenium, 285 aegyptium, 271 dahlii, Coluber, 477, 479 Dalea spinosa, 12, 16 Dasybasis, 82 702 CALIFORNIA ACADEMY OF SCIENCES dasyphrytina, Tabanus, 89 Dasyrhamphis minos, 97 davisi atlanticus, Plenoculus, 3, 37, 39, 43, 47 davisi, Penoculus, 2, 30, 37, 43, 44, 47 gracilis, Plenoculus, 3, 29, 39, 42 mojavensis, Plenoculus, 3, 38, 39, 45 Plenoculus, 2,356, 7 13, 265 27, 28. 29 80) 31, 32> 33, 34 35: 68s 39s 40143) 44, 46, 47, 48 Plenoculus davisi, 2, 30, 37, 43, 44, 47 transversus, Plenoculus, 3, 13, 34, 37, 38, 40, 42 decagrammus, Hexagrammos, 567, 570, 575, 576, 579 decemlineata, Eirenis, 477, 481 defillipi, Lacerta saxicola, 476 delicatissima, Anchoa, 633 delicatula caspia, Clupeonella, 633, 675, 677 Delphinus bairdii, 178 delphis, 178 delphis, Delphinus, 178 Dematiaceae, 261 Dempster, Lillian J., see Follett, W. I. Dendrelaphis caudolineatus, 374, 379, 403 caudolineatus caudolineatus, 378, 382, 401 caudolineatus luzonensis, 374, 383, 390, 391, 395, 403, 404, 407 caudolineatus terrificus, 378, 379, 380 pictus, 374, 378 pictus pictus, 382, 383, 384, 385, 387, 388, 390, 393, 395, 398, 401, 402, 403, 404 Dendrobaena, 221 octaedra, 224 rubida, 221, 223, 224 Dendrochirus chloreus, 566 dendrophila, Boiga, 373 divergens, Boiga, 374, 390, 391, 403 latifasciata, Boiga, 374, 393, 394, 404, 409 multicincta, Boiga, 374, 378, 400 dendrophiops barbouri, Natrix, 391, 395, 399, 408 dendrophiops, Natrix, 379, 393, 394, 399, 409 Natrix, 374, 394 Natrix dendrophiops, 379, 393, 394, 399, 409 [Proc. 4TH SER. negrosensis, Natrix 384, 392, 398, 399, 402, 409 dennyi, Liparis, 567, 579 Denticula rainierensis, 356, 364 Dermatolepis punctata, 566, 572 deserti, Plenoculus, 2, 6, 7, 26, 27, 45 desertus, Tabanus, 91, 100 Desmidiaceae, 267 deusta, Entophysalis, 264, 302 diadema diadema, Spalerosophis, 478 Lytorhynchus, 478, 479 Spalerosophis, 480 Spalerosophis diadema, 478 diaphana, Esenbeckia, 73 Diatomineura leucothorax, 100 Diatoms from Oregon and Washington, new and rare, by H. E. Sovereign, 349- 368 Dichelacera, 88 caloptera, 87 (Catachlorops) auripilis, 86, 100 ecuadoriensis, 87 Dichogaster, 194 bolaui, 224 saliens, 224 Dichonemertes hartmanae, 552 Dicladocera argyrophora, 83, 84 auribarbis, 84 badia, 86 castanea, 84 hoppi, 83 macula, 83, 84 neosubmacula, 83, 84 peruviana, 86 satanica, 84 scutellata, 84 submacula, 83, 84 tinctipennis, 84, 100 unicolor, 85 Dictydiaethalium plumbeum, 259, 260 Dictyophorus calcaetus, 228 (Dictyophorus) spumans, 228 (Dictyophorus) spumans, Dictyophorus, 228 Dictyophorous (Tapesiella) griseus, 229 (Tapesiella) griseus fuscoroseus, 229 (Tapesiella) griseus intermedius, 229 (Tapesiella) laticinctus, 229 diego, Pneumatophorus, 633, 634 diffringens, Pheretima, 219, 224 dinema, Carinella, 521 VoL. XXXI] Dinodon, 412 Dioclea, 281, 282 magacarpa, 281 reflexa, 282 dione, Elaphe, 477 Diplectrum macropoma, 566, 579 Diplometopon, 456, 482 zarudnyi, 456, 475, 481 discus, Tabanus, 99 dispar semlikiana, Pyrogomorpha, 245 dissapata undulata, Nitzschia, 356, 365 divaricata, Larrea, 25 divergens, Bioga dendrophila, 374, 390, 391, 403 elliptica, Pinnularia, 359 Pinnularia, 359 dombeyana, Chilina, 551 doriae, Ceramodactylus, 474, 481 dorsalis, Oerstedia, 553 Xylophaga, 112 dovii, Cichlasoma, 566, 579 drapizei, Boiga, 373, 378, 400 Dryocalamus subannulata, 374, 401 tristrigatus, 374, 378, 401 philippina, 374, 390, 395, 398, 405, 406 rubescens, 374, 385 dubia, Parasphena, 242 Parasphenella, 242 dubius, Microcolex, 224 dumerili, Lycodon, 374, 375, 379, 393, 394 Earthworms of Eisen’s collection, On some, by G. E. Gates, 185-225 Echeneis, 171, 176, 177 australis, 170; 171) 1172, 17:7 naucrates, 172 scutata, 170, 171, 176, 177 echinatus, Cenchrus, 271, 294, 296 echinoderma, Emplectonema, 541 echinulata, Cunninghamella, 259 Echis, 471, 482 carinatus, 471, 472, 478, 480 Eclipta, 285, 294 alba, 280 ecuadoriensis, Dichelacera, 87 Esenbeckia, 73 Egregia menziesii, 518 ehrenbergi, Ophisops elegans, 476 eilhardi, Prostoma, 548 INDEX 703 Eirenis, 466, 482 collaris, 477, 479, 481 coronella, 477, 481 coronelloides, 477 decemlineata, 477, 481 frenatus, 477, 480 iranica, 477 meda, 477 modesta, 477 persica, 466, 467, 477, 480 punctatolineata, 477 rothi, 477, 481 eiseni, Eodrilus, 213 Ramiellona, 209, 213, 216, 218 Sparganophilus, 220 Eisenia feotida, 224 hortensis, 224 rosea, 224 Eiseniella tetraedra, 224 Elaphe dione, 477 erythrura, 374, 382, 411 erythrura erythrura, 384, 387, 388, 389, 390, 393, 395, 398, 402, 403, 404 erythrura philippina, 378, 381, 382, 385, 401 hohenackeri, 477 longissimus, 478, 480 melanura, 411 quatuorlineata, 478 Elaphidae, 470, 478 elegans, Cunninghamella, 260 ehrenbergi, Ophisops, 476 elegans, Ophisops, 462, 476 Erys, 477 Ophisops, 462, 480 Ophisops elegans, 462, 476 Pelecorrhynchus, 99 Tabanus, 89 Zonocerus, 234. Eleocharis, 300 geniculata, 255, 272, 285, 300 mutata, 255, 272, 285, 298, 299, 300 elisae, Phyllodactylus, 442, 472, 481 elliptica, Pinnularia divergens, 359 elongatus, Ophiodon, 565, 567, 576, 577, 578, 579, 582 Emplectonema, 517, 534, 535, 536 biirgeri, 536 echinoderma, 541 gracile, 517, 518, 535, 536 704 CALIFORNIA ACADEMY OF SCIENCES neesii, 534, 535 viride, 534, 535 Emplectonematidae, 517 Emydidae, 434, 473 Emys orbicularis, 473, 480 encrasicholus, Engraulis, 633 maeotica, Engraulis, 633 pontica, Engraulis, 633, 677 engrauliformis, Clupeonella, 633, 675 Engraulis encrasicholus, 633, 675 encrasicholus maeotica, 633 encrasicholus pontica, 633, 677 japonica, 633 mordax, 631, 633, 634, 658, 659, 661, 662, 664, 665, 666, 667, 668, 669, 670, 681 Entada, 283 gigas, 282 Entophysalis, 302 deusta, 264, 302 granulosa, 265 Eodrilus, 194 eiseni, 213 irpex, 213 tecumumami, 214 ephippicus, Chilomeniscus, 313, 317, 321, 322 epialti, Carcinonemertes, 552 Epinephelus analogus, 566 Eragrostis amabilis, 271, 285 ciliaris, 271, 285 Eremias, 460 arguta, 476 brevirostris, 480 fasciata, 476 guttulata, 460, 480 guttulata guttulata, 476 guttulata watsonana, 329, 337, 427, 460, 461, 462, 476, 484 intermedia nigrocellata, 476 lineolata, 476, 481 (Mesalina) watsonana, 460 scripta, 476 velox persica, 476 velox strauchi, 476 Ereunias, 594 Erilepis, 564, 575, 580, 582, 586, 590 zonifer, 567, 569, 572, 579, 581, 597 Eriogonum, 25, 36 fasciculatum, 36 gracile, 25, 34 inflatum, 33 [Proc. 47TH Serr. polifolium, 34 reniforme, 36 thomasi, 16 trichopes, 8 trichopodum, 12, 32 wrighti, 36 Erisphex, 569 Eristocophis memahoni, 472 erythraea, Rana, 406 erythrogaster, Agama, 475, 482 erythrura, Elaphe, 374, 382, 411 Elaphe erythrura, 384, 387, 388, 389, 390, 393, 395, 398, 402, 403, 404 erythrura, Elaphe, 384, 387, 388, 389, 390, 393, 395, 398, 402, 403, 404 philippina, Elaphe, 378, 381, 382, 385, 401 Eryx, 482 elegans, 477 jaculus, 477, 481 johni, 477, 480 miliaris, 477, 480 tataricus, 477 Esenbeckia, 70 abata, 74 chagresensis, 73 diaphana, 73 ecuadorensis, 73 gertschi, 73 nigronotata, 74 schlingeri, 72, 100 seminuda, 74 tepicana, 74 translucens, 73 esterensis, Chilomeniscus Binge Etrumeus acuminatus, 633 Eublepharis, 435 macularius, 435, 436, 474, 480 nigrocincta, 552 Eudichogaster, 186 Eudrilidae, 224 Eudrilus eugeniae, 224 stramineus, 315, eugeniae, Eudrilus, 224 Eumeces, 483 schneideri, 479 schneideri variegatus, 476 taeniolatus, 476 zarudnyi, 476, 482 Eumetopias jubata, 604 Vor. XXXI] Eunemertes gracilis, 535 Euphorbia, 8, 32, 46 albomarginata, 27, 36, 46 marginata, 25 polycarpa, 16, 24, 32 polycarpa hirtella, 15, 24, 33, 46 Euphorbiaceae, 275 eupharticus, Trionyx, 474, 481, 483 Euphylax, 552 Eupomacentrus beebei, 566, 579 Euprepis septemtaeniata, 463 Euritium chevalieri, 260 Eurotiaceae, 260 Euscatia, 124 Euschatia, 124 collaris, 126 (Euschatia) cryptocephalus, Heliofugus, 126 Euschatia laticollis, 137 parva, 152 proxima, 154 punctata, 135 punctato-striata, 136 sulcata, 128 sulcipennis, 142, 143 Euthynnus yaito, 634, 675 Eutyphoeus, 186 euxinus, Odontogadus merlangus, 634 everetii, Calamaria, 373, 378, 400 eversmanni, Crossobamon, 474 evexa, Navicula, 353, 354 exaestuans, Leucotabanus, 91] exilissima, Navicula, 353 exlineata, Carinella, 521 exogyra, Pseudochama, 114 fairchildi, Proboscoides, 74 fairmairei, Heliofugus, 150, 158, 166 Heliofugus (Heliofugus), 150 fallax iberus, Telescopus, 468, 478, 481 fasciata, Eremias, 476 fasciatus, Chilomeniscus stramineus, 313, 317, Sail, ee) fasciculatum, Eriogonum, 36 fascinatus, Oplegnathus, 672 Fauna from the Upper Pleistocene Battery Formation near Crescent City, Cali- fornia, Interpretation of the In- vertebrate, by Warren O. Addicott, 341-347 fedtschenkoi, Cyrtodactylus, 438, 474, 481 INDEX 705 femurrubrum, Melanoplus, 506, 508 fenderi fenderi, Plutellus, 224 Plutellus fenderi, 224 feotida, Eisenia, 224 ferruginea, Taphronota, 229, 230 Fidena, 74 fimbria, Anoplopoma, 567, 569, 579, 581 First records of the echeneidid fish Remi- legia australis (Bennett) from Cali- fornia, with meristic data, by W. I. Follett, and Lillian J. Dempster, 169- 184 fitzgeraldi, Maura, 228 flavescens, Amphiporus, 546 Lineus, 529 Stypommia, 88 Tabanus, 88 flavicorpus, Tabanus (Lophotabanus), 94, 100 flavifrons, Maura, 228 flaviviridis, Hemidactylus, 474, 480 flavofemoratus, Stenotabanus, 77 flavomaculatus flavomaculatus, Trimeresurus, 380, 383, 387, 388, 389, 391, 393, 394, 395, 398, 410 halieus, Trimeresurus, 403 mcegregori, Trimeresurus, 380 Trimeresurus, 374, 376 Trimeresurus flavomaculatus, 380, 383, 387, 388, 389, 391, 393, 394, 395, 398, 410 flavus, Aspergillus, 261 flavus-oryzae, Aspergillus, 261 Flora and vegetation of Clipperton Island, by Marie Hélene Sachet, 249-307 florae, Liparis, 567, 579 flos-aquae, Microcystis, 264 Follett, W. I., and Lillian J. Dempster, First records of the echeneidid fish Remi- legia australis (Bennett) from Cali- fornia, with meristic data, 169-184 formidabilis, Amphiporus, 517, 518, 519, 544 fragilis, Anguis, 480 cochicus, Anguis, 475 frenata, Zaniolepis, 567, 579 frenatus, Eirenis, 477, 480 Tubulanus, 525 Freude, Misolampini: Guerin Und Myrmecodema_ Gebien Chilenischen Gattungen Heliofugus Heinz, Revision der 706 CALIFORNIA ACADEMY OF SCIENCES (= Myrmecosma Germain) Coleo- ptera: Tenebrionidae), 121-168 fritzei, Liopeltis semicrinata, 54 frontalis, Mesomyia (Vepriella) , 72 Frustulia rhomboides occidentalis, 351 Fucus, 116, 119 Fuga niphobles, 672 rubripes, 672 fulvistriatus, Stenotabanus, 77 fulvus, Amphiporus, 545 fumigata, Sartorya, 260 fumigatus, Aspergillus, 261 fumomarginatus, Tabanus, 94, 95 funebralis, Tegula, 343 funiculosum, Penecillium, 262 Further observations on Hipponix antiqua- tus with notes on north Pacific pul- monate limpets, by C. M. Yonge, 111-119 Further records of neotropical Tabanidae (Diptera) mostly from Peru, by Cor- nelius B. Philip, 69-102 fuscoroseus, Dictyophorus griseus, 229 fuscus, Pelobates, 473 (Tapesiella ) gaddi, Lytorhynchus, 478 (Gadinia) reticulata, Trimusculus, 111 Gadus morhua macrocephalus, 634 morhua morhua, 634 galapagensis, Alphestes, 566, 579 galeanea, Phyllophaga, 559 gamma, Cyclotella, 350, 354 Gans, Carl, On Amphisbaena heathi Schmidt and A. carvalhoi, new species, small forms from the northeast of Brazil (Amphisbaenia: Reptilia), 613-630 Gates, G. E., On some Earthworms of Eisen’s collection, 185-225 Gecarcinus plantaus, 258 Gehyra, 377 Gekkondae, 435, 474 gemianulis, Calliophis calligaster, 384, 398, 399, 408, 409 geminatus, Sibynophis, 378, 411 gemonensis, Coluber, 477, 479 geniculata, Eleocharis, 255, 272, 285, 300 germaini, Heliofugus, 156, 157, 158, 159, 167 Heliofugus (Heliofugus), 157 [Proc. 4TH Ser. gerstaeckeri, Atractomorpha acutipennis, 246 gertschi, Esenbeckia, 73 gervaisi, Calamaria, 374 Calamaria gervaisi, 390, 391, 394, 399, 403, 407 gervaisi, Calamaria, 390, 391, 394, 399, 403, 407 hollandi, Calamaria, 379, 393, 394, 403, 409 iridescens, Calamaria, 384, 398, 399, 402, 409 polillensis, Calamaria, 403 gibberosa, Astrea, 118 gibbosus, Lepomis, 677 gigantea, Lottia, 104, 118 giganteus, Saxidomus, 344 gigas, Entada, 282 Stereolepis, 566 gillaspyi, Plenoculus, 2, 4, 5, 8, 10, 23, 47 Girella punctata, 672 glabrata, Cawendia, 240 glandulosa, Zygonemertes, 540 glanis, Silurus, 673 glauca, Nicotiana, 278, 279 Glenodinium, 268 Glossoscolecidae, 219 Goin, Coleman J., and Doris M. Cochran, Two new Genera of Leptodactylid frogs from Colombia, 499-505 Gomphonema grovei, 357, 367 hedini, 356, 366 Gomphosphaeria aponina, 264 Gonyosoma oxycephala, 374, 378, 380, 390, 398, 401 (Graceevelynia) lasiurus, Howascolex, 215 sauerlandti, Howascolex, 216 tecumumami_ vulcanicus, 216 gracile, Emplectonema, 517, 518, 535, 536 Eriogonus, 25, 34 gracilis bullocki, Nemertopsis, 552 Calliophis, 411 Eunemertes, 535 Nemertes, 534, 535, 552 Nemertopsis, 552 gracilis, Plenoculus davisi, 3, 29, 39, 42 graeca, Testudo, 479 ibera, Testudo, 474, 481 graecense, Prostoma, 548, 549 Gramineae, 271 Howascolex, Vor. XXXI] grande, Prostoma, 548 grandis, Parasphenula, 240 Pisonia, 284 Grant, Norman, see Loukashkin, Anatole S. granulata, Pyrgomorpha, 245 granulosa, Entophysalis, 265 Grasshopper from the White Mountains of California (Orthoptera: Acrididae: Cyrtacanthacridinae), A new spine- throated, by Ashley B. Gurney, and David C. Rentz, 503-513 grayanus, Ablepharus, 476, 481 Calamaria veriformis, 374, 379, 389, 393, 394, 404, 409 grevillei, Protococcus, 268, 302 griffini, Carinoma, 526 grimmi, Clupeonella, 633 grisea juncundus, Scaptia, 70 Scaptia, 70 griseus, Dictyophorus (Tapesiella) , 229 fuscoroseus, Dictypohorus (Tapesiella), 229 intermedius, Dictyophorus (Tapesiella), 229 Tupinambis, 455 Varanus, 455, 475, 480 grossa, Cawendia, 240 Malacobdella, 517, 518, 549, 551 grovei, Gomphonema, 357, 367 Gryllus cinctus, 230 thaelephorus, 230 guatemalana, Ramiellona, 195, 205, PQ 213,.2 Wsp216.2lia218s 219 guaymasensis, Lyngbya, 266 Gurney, Ashley B., and David C. Rentz, A new spine-throated grasshopper from the White Mountains of Cali- fornia (Orthoptera: Acrididae: Cyr- tacanthacridinae) , 503-513 guttata, Scorpaena, 566, 579 guttula, Chrysops, 75 guttulata, Eremias, 460, 480 Eremias guttulata, 476 guttulata, Eremias, 476 watsonana, Eremias, 329, 337, 427, 460, 461, 462, 476, 484 Gymnodactylus, 334, 336 agamuroides, 438 microtus, 336 209, pipiens, 336 INDEX 707 halieus, Trimeresurus flavomaculatus, 403 halys, Agkistrodon, 478, 480 Hampala, 386 hamulirostris, Leptotyphlops, 477 hannah, Ophiophagus, 374, 378, 387, 391, 393, 395, 398, 401 Phyiophagus, 378 hansi, Tetrastemma, 546 Harengula thrissina, 632 zunasi, 633 harengus, Clupea harengus, 633 harengus, Clupea, 633 membras, Clupea, 633 hartmanae, Dichonermertes, 552 hasselquisti, Ptyodactylus, 474, 479 hawayana, Pheretima, 224 hazelae, Cornufer, 400 heathi, Amphisbaena, 613, 615, 616, 617, 618, 619, 620, 621, 623 Hebardacris, 511 hecticus, Rhacophorus, 405 hedini, Gomphonema, 356, 366 (Helcion), Patina, 103 helenae, Microgecko, 440, 441, 474, 481 Tropiocolotes, 336, 440 Helianthus annuus, 36 Helicolenus, 580 Heliofuge, 123 Heliofugus, 121, 123, 124, 125, 126, 131, 158, 166 arenosus, 125,, USile W529 15381565160; 166 (Heliofugus) arenosus, Heliofugus, 151 Heliofugus barrosi, 138, 144, 160, 166 (Heliofugus) barrosi, Heliofugus, 143 Heliofugus biobiensis, 149, 155, 160, 166 (Heliofugus) biobiensis, Heliofugus, 149 Heliofugus brevipennis, 143 colasi, 159, 166 (Heliofugus) colasi, Heliofugus, 142 Heliofugus (Collariheliofugus), collaris, 126 (Collariheliofugus) cryptocephalus, 126 (Collariheliofugus) cryptocephalus euri- coensis, 128 collaris, 124, 125, 126, 127, 128, 158, 166 coquimboensis, 150, 160, 166 (Heliofugus) coquimboensis, Heliofugus, 152 Heliofugus cribriceps, 136, 140, 159, 166 (Heliofugus) cribriceps, Heliofugus, 139 708 CALIFORNIA ACADEMY OF SCIENCES Heliofugus cryptocephalus, 124, 125, 126, 128, 158 cryptocephalus curiocensis, 127, 158, 166 cryptocephalus cryptocephalus, 127, 158, 166 (Euschatia) cryptocephalus, 126 fairmairei, 150, 158, 166 (Heliofugus) fairmairei, Heliofugus, 150 Heliofugus germaini, 156, 157, 158, 159, 167 (Heliofugus) germaini, Heliofugus, 157 Heliofugus (Heliofugus) arenosus, 151 (Heliofugus) barrosi, 143 (Heliofugus) biobioensis, 149 (Heliofugus) colasi, 142 (Heliofugus) coquimboensis, 152 (Heliofugus) cribriceps, 139 (Heliofugus) fairmairei, 150 (Heliofugus) germaini, 157 (Heliofugus) impressus, 132 (Heliofugus) laticollis, 137 (Heliofugus) leeehi, 147 (Heliofugus) leechi maulensis, 148 (Heliofugus) penai 141 (Heliofugus) penai penai, 140 (Heliofugus) proximoides, 146 (Heliofugus) proximoides rotundangu- ohigginsi, lus, 147 (Heliofugus) proximus, 154 (Heliofugus) proximus punctatosulca- tus, 155 (Heliofugus) punctatostriatus, 136 (Heliofugus) quillotaensis, 148 (Heliofugus) rossi, 153 (Heliofugus) sulcatus, 131 (Heliofugus) sulcipennis, 142 (Heliofugus) sulcipennis brevipennis, 143 (Heliofugus) tenuipunctatus, 136 (Heliofugus) ventriosus, 144 impressus, 132, 134, 135, 139, 141 impressus cribricephalus, 136, 137, 139, 160, 166 (Heliofugus) impressus, Heliofugus, 132 Heliofugus impressus impressus, 130, 135, 136, 138, 140, 159, 166 impressus punctatus, 133, 136, 139, 140, 159, 166 (Inseutoheliofugus) kuscheli, 130 kuscheli, 126, 158, 166 laticollis, 137, 138, 139, 141, 144, 159, 166 134, [Proc. 4TH Ser. (Heliofugus) laticollis, Heliofugus, 137 Heliofugus leechi, 145, 148, 159, 160 (Heliofugus) leechi, Heliofugus, 14.7 Heliofugus leechi leechi, 146, 160, 166 leechi maulensis, 160, 166 (Helofugus) 148 Heliofugus neuqueni, 126, 130, 158, 166 parvus, 151, 152 penai, 141, 151, 159 penai ohigginsi, 159, 166 (Heliofugus) penai ohigginsi, Heliofugus, 141 Heliofugus penai penai, 159, 166 (Heliofugus) penai penai, Heliofugus, 140 (Heliofugus) proximoides, 145, 147, 149, 160 (Heliofugus) proximoides, Heliofugus, 146 Heliofugus proximoides proximoides, 160, 166 proximoides rotundangulus, 160, 166 (Heliofugus) proximoides rotundangulus, Heliofugus, 14.7 Heliofugus proximus, 132, 134, 135, 139, 141, 1465, 11525) 154) 1565 160 (Heliofugus) proximus, Heliofugus, 154 leechi maulensis, Heliofugus, Heliofugus proximus proximus, 146, 157, 160 proximus punctatosulcatus, 156, 158, 167 (Heliofugus) proximus punctatosulcatus, Heliofugus, 155 Heliofugus punctatostriatus, 135 (Heliofugus) punctatostriatus, 136 Heliofugus punctatosulcatus, 155 punctatus, 135 quillotaensis, 148, 149, 155, 160, 166 (Heliofugus) quillotaensis, Heliofugus, 148 Heliofugus rossi, 154, 156, 160, 167 (Heliofugus) rossi, Heliofugus, 153 Heliofugus, Heliofugus rufitarsis, 158 (Rugosiheliofugus) neuqueni, 129 (Rugosiheliofugus) sulcatulus, 128 striatus, 151, 152, 157 sulcatulus, 125, 126, 130, 132, 158, 166 sulcatus) 1235) 128) 129 eS ie Zee 151, 156, 158, 159, 166 (Heliofugus) sulcatus, Heliofugus, 131 Heliofugus sulcipennis, 140, 142, 143, 150, 156, 159 sulcipennis brevipennis, 143, 149, 166 VoL. XXXT] (Heliofugus) sulcipennis brevipennis, Helio- fugus, 143 sulcipennis, Heliofugus, 142 sulcipennis sulcipennis, 143, 159, 166 tenuipunctatus, 135, 136 (Heliofugus) — tenuipunctatus, 136 Heliofugus ventriosus, 160 ventriosus nancaguensis, 145 (Heliofugus) ventriosus, Heliofugus, 144 Heliofugus ventriosus ventriosus, 144, 166 Heliophygus, 124, 155 helioscopus, Phrynocephalus, 475 Heliotropium, 258, 285, 290, 295, 296, 299 curassavicum, 278, 284, 296 Heliotropus curassavicus, 36 Hemibungarus, 411 Hemicarpha, 285, 300 micrantha, 255, 273, 300 hemictenus, Puntius, 396, 397 Hemidactylus, 441, 482 flaviviridis, 474, 480 persicus, 44-1, 474, 480 turcicus, 474 Hemilepidotus, 580 hemilepidotus, 567, 579 hemilepidotus, Hemilepidotus, 567, 579 herculeus, Cerebratulus, 532 Herpetology of Afghanistan, Third contri- bution to the, by Alan E. Leviton, and Steven C. Anderson, 329-339 Herse cingulata, 258, 299 heterocercus, Cyrtodactylus, 439, 474, 482 Heteronemertini, 516 heteroporus, Plutellus, 188 heufleri, Schizothrix, 267 Hexagrammid fishes, with a synopsis, oste- ological characteristics and affinities of the, by Jay C. Quast, 563-600 Hexagrammidae, 567, 589, 590 Hexagrammos, 565, 570, 575, 580, 581, 582, 583, 589 agrammus, 567, 576, 579 decagrammus, 567, 570, 575, 576, 579 lagocephalus, 565, 567, 570, 571, 575, 576, 579 octogrammus, 567, 583 otakii, 567, 576, 579, 583 stelleri, 567 superciliosus, 565, 567, 576, 579 Heliofugus, INDEX 709 Hinnites multirugosus, 114, 343 Hipponix, 114, 115 antiquatus, 111, 112, 113, 114, 118 Hipponix antiquatus with notes on north Pacific pulmonate limpets, Further ob- servations on, by C. M. Yonge, 111-— 119 hirtella, Euphorbia polycarpa, 15, 24, 33, 46 hirundo, Leicottus, 567 hitchockii teres, Neidium, 352, 354 hofmannii, Scytonema, 265, 302 hohenackeri, Elaphe, 477 hollandi, Calamaria gervisi, 379, 393, 394, 403, 409 Holocentridae, 566 Holocentris suborbitalis, 566, 579 Hologerrhum, 373, 374, 375, 412 philippinum, 374, 390, 391, 403, 404, 408 Hoplichthyidae, 593 Hoplonemertini, 517 hoppi, Dicladocera, 83 horsfieldi, Testudo, 474, 481 hortensis, Eisenia, 224 Howascolex, 194, 213 (Graceevelynia) lasiurus, 215 (Graceevelynia) sauerlandti, 216 (Graceevelynia) tecumumami cus, 216 Hubbs, Carl L., see Radford, Keith W. hubbsi, Anoptichthys, 679 Hugelia virgata, 36 hurdi, Plenoculus, 2, 4, 5, 10, 23 Hurria, 373, 376 Hybomitra, 89 (Tylostypia), 89 Hyla, 430 arborea, 432, 479 arborea savignyi, 424, 430, 432, 473 Hylidae, 430, 473 Hypnea, 303 spinella, 303 Hypomesus olidus, 674 Hypopelma amabilis, 89 quadripuncta, 89 Hypsalonia, 511 Hypsagonus quadricornis, 572 Hypsiglena, 376 vulcani- ibera, Testudo graeca, 474, 481 iberus, Telescopus fallax, 468, 478, 481 710 CALIFORNIA ACADEMY OF SCIENCES Icelidae, 594 Icelus, 594 Ichthyopis, 405 monochrous, 380 ignava, Phyllophaga, 559, 562 imparispinosus, Amphiporus, 517, 518, 542, 546 similis, Amphiporus, 546 impotens, Parapetasia (Loveridgeacris), 229 impressus_ cribricephalus, Heliofugus, 136, 137, 139, 160, 166 Heliofugus, 132, 134, 135, 139, 141 Heliofugus (Heliofugus), 132 Heliofugus impressus, 130, 134, 135, 136, 138, 140, 159, 166 impressus, Heliofugus, 130, 134, 135, 136, 138, 140, 159, 166 punctatus, Heliofugus, 140, 159, 166 Tabanus, 95, 96, 100 inaequalis, Sapindus saponaria, 276 indica, Waltheria, 276 inea, Stypochela, 85, 100 incipiens, Triceratomyia, 79 incola, Paranemertes, 539 inconstans, Neidium, 351, 354 inflatum, Eriogonum, 33 Influence of light of different wave lengths and intensities and total darkness, Behavior and natural reactions of the northern anchovy, Engraulis modrax Girard, under the, by Antole S. Lou- kashkin, and Norman Grant, 361-692 (Inseutoheliofugus) , Heliofugus, 130 kuscheli, Heliofugus, 130 intermedia nigrocellata, Eremias, 476 undulata, Stenopterobia, 356, 365 intermedius, Dictyophorus (Tapesiella) griseus, 229 Interpopulation the colubrid snake Natrix pryeri from the Riukiu 183, 136, 139, variation on Islands, with description of a new subspecies by Edmond VY. Malnate, and Harold E. Munsterman, 51-67 Interpretation of the invertebrate fauna from the Upper Pleistocene Battery Formation near Crescent City, Cali- fornia, by Warren O. Addicott, 341- 347 interruptus, Archoplites, 577, 580 [Proc. 4TH Serr. interscapularis, ?PPhrynocephalus, 475 intestinalis bilineata, Maticora, 374, 385, 401 Maticora, 374 nigrotaeniata, Maticora, 388 phillippina, Maticora, 374, 391, 393, 394, 382, 404 suluensis, Maticora, 374, 387, 388, 410, 411 Ipomoea, 274, 288, 290, 291, 297, 298, 299, 300 pes-caprae, 256, 258, 275, 277, 280, 285, 290, 292, 293, 294, 297, 298 pes-caprae brasiliensis, 276 triloba, 277, 285 tuba, 277 iranica, Eirenis, 477 iridescens, Calamaria gervaisi, 384, 398, 399, 402, 409 iris, Phymateus (Phymateus), 233 irpex, ?Acanthodrilus, 213 Acanthodrilus, 214 Eodrilus, 213 Ramiellona, 213, 218, 219 irus, Macoma, 344 Iscutholieliofugus, 126, 130 ishigakiensis, Natrix pryeri, 61, 62, 65 islandica, Arctica (Cyprina), 550 isolepis, Agama agilis, 445 ivis, Puntius, 386 jaculus, Eryx, 477, 481 Jania, 303 japonica, Engraulis, 633 Malacobdella, 551 Pempheris, 672 japonicus, Pneumatophorus, 633 Trachurus, 633, 672 Jenkinsia lamprotaenia, 633, 672, 679 johni, Eryx, 477, 480 joloensis, Calamaria, 374, 387, 388, 410 jordani, Anoptichthys, 679 Mycteropera, 566 jubata, Eumetopias, 604 jugularis asianus, Coluber, 477 juncea, Brassica, 274, 294 jucunda, Scaptia, 70 juncundus, Scaptia grisea, 70 Juniperus, 559 VoL. XXXI] kachhensis, Cyrtodactylus, 438 kaiseri, Neurergus crocatus, 473 Kaloula conjuncta, 400 karelini, Coluber, 466, 477, 481 Triturus cristatus, 473 kenoyeri, Chara, 268 kessleri, Alosa kessleri, 633 kessleri, Alosa, 633 volgensis, Alosa, 633 Kevan, D. Keith McE., Pyrogomorphidae (Orthoptera: Acridoidea) collected in Africa, by E. S. Ross and R. E. Leech, 1957-1958, with descriptions of new species, 227-248 kineaidii, Navicula, 353, 354 kirmanensis, Agama, 475, 482 Cyrtodactylus, 438, 474 kisutch, Oncorhynchus, 680 kochi, Myrmecodema, 162, 165, 166, 167 kotschyi, Cyrtodactylus, 438, 474, 479 kuscheli, Heliofugus, 126, 158, 166 Heliofugus (Inseutoheliofugus) , 130 Lacerta, 483 brandti, 476, 481 chlorogaster, 476 media, 476 muralis, 476 pipiens, 334, 335, 336 princeps, 476 sanguinolenta, 446 saxicola, 480 saxicola defillipi, 476 strigata, 476, 480 viridis, 476 Lacertidae, 456, 475 lactea, Carinomella, 551 lactescens, Corticium, 260 lacteum, Octolasium, 224 laeta sublaeta, Chrysops, 75 laevis, Alsophylax, 334, 336 lagerheimii, Lyngbya, 266 lagocephalus, Hexagrammos, 565, SA, SIS, SO, Seo) lamellosa, Thais, 343, 344 lamiella, Carcharhinus, 632 lamprotaenia, Jenkinsia, 633, 672, 679 lanatum, Bryum argenteum, 269, 300 lanosum, Penicillium, 263 Larrea divaricata, 25 INDEX (ail lasiura, Ramiellona, 215, 219 lasiurus, Howascolex (Graceevelynia), 215 Ramiellona, 201, 216, 219 laticeps, Leptotyphlops, 477 laticinctus, Dictyophorus (Tapesiella) , 229 laticollis, Euschatia, 137 Heliofugus, 137, 138, 139, 141, 144, 159, 166 Heliofugus (Heliofugus), 137 latifasciata, Boiga dendrophila, 374, 393, 394, 404, 409 latifolia, Najas marina, 271 Zostera marina, 270 latipes, Oryzias, 673 latipinnis, Zaniolepis, 567, 579 latirostris, Pseudocerastes, 472 lebetina, Vipera, 478, 480 Leech, R. E., see Ross, E. S. leechi, Heliofugus, 145, 148, 159, 160 Heliofugus leechi, 146, 160, 166 Heliofugus (Heliofugus), 147 leechi, Heliofugus, 146, 160, 166 maulensis, Heliofugus, 160, 166 maulensis, Heliofugus (Heliofugus), 148 legumen ornata, Pinnularia, 362 Leguminosae, 274, 281 Leiocottus hirundo, 567 Leiolopisma, 482 Lepeta, 104 caeca, 104 concentrica, 103, 105, 106, 107, 109, 344 (Cryptoctenidia) concentrica, 104 Lepomis gibbosus, 677 leporinum leporinum, 392, 399 Oxyrhabdium, 374 Oxyrhabdium leporinum, 391, 392, 399 visayanum, Oxyrhabdium, 384, 398, 399, 409 leprosus, Phymateus (Maphyteus), 234 leptacanthus, Amphiporus, 542 Leptocottus armatus, 567, 579 from Colombia, Oxyrhabdium, 391, Leptodactylid frogs Two new genera, by Coleman J. Goin, and Doris M. Cochran, 499-505 Leptodeira, 376 Leptotyphlopidae, 477 Leptotyphlops hamulirostris, 477 laticeps, 477 712 CALIFORNIA ACADEMY OF SCIENCES macrorhynchus, 477, 479 Lepturus repens, 284 leuciodus, Amphiporus, 542, 543 Leuciscus brandti, 674 leucomaenis, Salvelinus, 674 leucospila, Chrysops, 75 Leucotabanus exaestuans, 91 leucothorax, Diatomineura, 100 Listrapha (Scaptia), 70 leucothorax, Scaptia atra, 70 Leviton, Alan E., Remarks on the zoogeog- raphy of Philippine Terrestrial snakes, 369-416 Leviton, Alan E., see Banta, Benjamin H. Leviton, Alan E., and Steven C. Anderson, Third contribution to the herpetology of Afghanistan, 329-339 libertate, Opisthonema, 633 lichen, Pyrenocarp, 263 lima, Thais, 343 limbithorax, Scaptia, 70 limnocharis, Rana, 54 lineaalba, Tanita, 246 linearus, Cephalothrix, 526 lineata, Natrix, 374, 379, 380, 393, 404, 409 lineatus, Cyclocorus, 374 Cyclocorus lineatus, 384, 390, 392, 394, 395, 398, 403, 407 lineatus, Cyclocorus, 384, 390, 392, 394, 395, 398, 403, 407 nuchalis, Cyclocorus, 379, 394, 404, 409 Lineidae, 516 lineolata, Eremias, 476, 481 lineolatus, Cerebratulus, 532 Psammophis, 478, 480 Lineus, 516, 527, 528 bilineatus, 528 flavescens, 529 pictifrons, 516, 518, 528 pseudo-lacteus, 528 ruber, 516, 518, 527, 528, 529 rubescens, 528 sanguineus, 527, 528 striatus, 532 torquatus, 528 vegetus, 529 viridis, 527, 528 wilsoni, 533 Liopeltis philippina, 374, 382, 385, 411 philippinus, 401 [Proc. 4TH Serr. semicarinata fritzei, 54 tricolor, 374, 378, 382, 385, 401, 411 Liparis dennyi, 567, 579 florae, 567, 579 Listrapha (Scaptia) leucothorax, 70 litigiosus, Phaeotabanus, 81 Loligo opalescens, 674 vulgaris, 674 longicauda, Pelecorrhynchus, 99 longiceps, Cerebratulus, 516, 519, 531 Pseudorabdion, 409 longipes, Crytodactylus, 438, 474, 482 Triturus cristatus, 473 longissimus, Elaphe, 478, 480 loosi, Tanita, 245 Lophotabanus, 95 (Lophotabanus) flavicorpus, Tabanus, 94, 100 lophus, Tabanus, 92 lophus, Tabanus (Lophotabanus), 92 loricatus, Centrotrachelis, 453 Uromastix, 453, 455, 475, 481, 485 Lottia, 103, 108 gigantea, 104, 118 Loukashkin, Anatole S., and Norman Grant, Behavior and natural reactions of the northern anchovy, Engraulis mordax Girard, under the influence of light of different wave lengths and inten- sities and total darkness, 631-692 (Loveridgeacris) impotens, Parapetasia, 229 lucidus, Strongylodon, 282 lucioceps, Snyodus, 632 Lumbricidae, 221, 224 lumbricoideum, Prostoma, 548 Lumbricus rubellus, 224 terrestris, 224 lumsdeni, Stenodactylus, 474 lurida, Maura, 228 luristanicus, Bufo, 430, 473, 481, 482 luteoguttatus, Phrynocephalus, 475, 481 luzonensis, Dendrelaphis caudolineatus, 374, 383, 390, 391, 395, 403, 404, 407 Oryzias, 396 Zaocys, 374, 389, 391, 403 Lycodon, 375, 412 aulicus, 374, 377, 386 aulicus capucinus, 378, 384, 386, 390, 392, 393, 395, 398, 401, 402, 404 dumerili, 374, 375, 379, 393, 394 Vor. XXXI] INDEX 713 miilleri, 374, 375, 380, 391, 395, 403, 408 grossa, 517, 518, 549, 551 striatus bicolor, 478, 482 japonica, 551 subcinctus, 378 mercenaria, 549 subcinctus sealei, 401 minuta, 551 tessellatus, 374, 391 obesa, 549 Lyngbya, 267, 302 Malacobdellidae, 517 aesturaii, 266 Malnate, Edmond V., and Harold E. Mun- confervoides, 266, 302 sterman, Interpopulation variation in guaymasensis, 266 the colubrid snake Natrix pryeri from lagerheimii, 266 the Riukiu Islands, with description of semiplena, 266 a new subspecies, 51—67 versicolor, 266, 268 Malpolon moilensis, 478, 479 Lytorhynchus, 482 monspessulana, 479 diadema, 478, 479 Malvaceae, 276 gaddi, 478 Mantle cavity, habits, and habitat in the ridgewayi, 478 blind limpet, Lepeta concentrica Mid- dendorff, by C. M. Yonge, 103-110 Mabuya, 463, 482 Maphyteus, 233 aurata, 464, 479 (Maphyteus) baccatus, Phymateus, 233 aurata septemtaeniata, 423, 463, 464, leprosus, Phymateus, 234 476, 484 marginata, Euphorbia, 25 macroschisma, Pododesmus, 114 marginatus, Cerebratulus, 531 Macoma irus, 344 marginenevris, Tabanus, 95 nasuta, 342, 344 marina latifolia, Najas, 271 macrocephalus, Gadus morhua, 634 latifolia, Zostera, 270 macroceras, Stenotabanus, 79, 81, 100 Najas, 266, 303 Macrocormus, 92 Zostera, 303, 518 (Macrocormus) sorbillans, Tabanus, 91 marioni, Tetrastemma, 546 macrodon, Rana, 387 maritima pacifica, Ruppia, 270 visayanum, Rana, 387 rostrata, Ruppia, 270 macrophthalma, Sardinella, 633, 672 Ruppia, 270, 302 macropoma, Diplectrum, 566, 579 marmoratus, Cirrhitus, 566, 579 macropterus, Neothunnus, 634, 675 Plutellus, 193, 224 macrorhynchus, Leptotyphlops, 477, 479 Scorpaenichthys, 567, 574, 575, 579, 583 macula, Dicladocera, 83, 84 Sebastiscus, 566, 579 macularius, Cyrtodactylus, 435 marshalli, Chirindites, 241 Eublepharis, 435, 436, 474, 480 Maura, 227 maculatus, Natrix, 374 martini, Tarbophis, 467, 468 Oligodon, 374, 393, 394, 409 Marukawichthys, 594 Phrynocephalus, 475, 480 Mastigocoleus, 302 maculifrons, Stenotabanus, 79 testarum, 265, 302 maetoica, Engraulis encrasicholus, 633 Maticora intestinalis, 374 Rhombus, 673 intestinalis bilineata, 374, 382, 385, 401 magacarpa, Dioclea, 281 intestinalis nigrotaeniata, 388 major, Caesalpinia, 281 intestinalis philippina, 374, 391, 393, 394, Procephalothrix, 527 404 majus, Closterium parvulum, 267 intestinalis suluensis, 374, 387, 388, 410, makahana, Pinnularia, 354, 359 411 Malacobdella, 517, 549, 551 Maura, 228 auriculae, 551 antennata, 228 714 CALIFORNIA ACADEMY OF SCIENCES fitzgeraldi, 228 flavifrons, 228 lurida, 228 marshalli, 227 maulensis, Heliofugus (Heliofugus) leechi, 148 Heliofugus leechi, 160, 166 mcclungi, Calliophis calligaster, 403, 408 mcgregori, Trimeresurus flavomaculatus, 380 mcintyrei, Triceratomyia, 74 mecmahoni, Eristocophis, 472 Pseudocerastes, 478 mcnamarae, Pseudorabdion, 374, 398, 399 mearnsi, Calamaria, 374, 393, 394, 409 meda, Eirenis, 477 media, Lacerta, 476 megalonyx, Agama, 475, 481 Megascolecidae, 219, 224 melanocephala, Boiga trigonata, 477 Melanogrammus aeglefinus, 634 Melanoplus, 503, 506 femurrubrum, 506, 508 militaris, 506, 508 melanops, Sebastodes, 566 melanosticta, Sardinops sagar, 633 melanura, Agama, 475, 481 Elaphe, 411 meleagrinum, Penicillium, 263 Melilotus, 42 membras, Clupea harengus, 633 Menidia starksi, 632 menziesil, Egregia, 518 mercenaria, Malacobdella, 549 merlangus euxinus, Odontogadus, 634 Merremia, 283 tuberosa, 283 391, (Mesalina) watsonana, Eremias, 460 Mesomyia, 71 carbo, 71, 72, 100 (Coracella), 71, 72 (Coracella) carbo, 71 (Coracella) rubricornis, 72 (Coracella) rufopilosus, 72 rubricornis, 72 (Vepriella) frontalis, 72 mexicana, Ramiellona, 209, 214, 218 mexicanus, Plenoculus, 2, 4, 5, 7, 17, 18, 19 meyerinki, Oligodon, 374, 381, 387, 402, 405, 407, 410, 411 [Proc. 4TH Ser. michelbacheri, Myrmocodema, 157, 163, 164, 165, 166, 167 micrantha, Hemicarpha, 255, 273, 300 microcarpus, Sapidus saponaria, 276 Microcystis flos-aquae, 264 Microgecko, 336, 440 helenae, 440, 441, 474, 481 microlepis, Agama, 475 Spalerosophis, 478 Teratoscincus, 474, 482 Uromastix, 475, 481 micropholis, Acanthodactylus, 475, 481 Micropogon altipinnis, 566, 579 Micropterus salmoides, 566, 579 Micrura, 516, 533 alaskensis, 533 nigrirostris, 534 olivaris, 534 pardalis, 534: verrilli, 516, 518, 519, 532, 533 wilsoni, 516, 518, 533 Microscolex dubius, 224 phosphoreus, 224 microtus, Gymnodactylus, 336 microtympanum, Agama, 475 micro-virido-citrinus, Aspergillus, 261 miliaris, Eryx, 477, 480 Ophiomorus, 476 militaris, Melanoplus, 506, 508 mindorensis, Bulbalus, 396 minos, Dasyrhamphis, 97 Stypochela, 97 minuta, Malacobdella, 551 minutum, Pseudorabdion, 399 miolepis, Naja naja, 401 Mirounga angustirostris, 602 missionum, Tabanus, 82 mitchelli, Amphisbaena, 613 mitra, Acmaea, 343 modesta, Eirenis, 477 modestum, Oligodon, 374, 393 Oxyrhabdium, 374, 379, 380, 386, 389, 391, 393, 398, 399, 404, 407, 408 moilensis, Malpolon, 478, 479 mojavensis, Plenoculus davisi, 3, 38, 39, 45 Moniliaceae, 261 monitor, Varanus, 475, 480 monochrous, Ichthyophis, 380 Ochthyophis, 380 VoL. XXXI] INDEX 715 monopterygius, Pleurogrammus, 567, 571, nycterinoides, 161, 162, 166, 167 576, 579, 581 Myrmecosoma, 121, 161 monspessulana, Malpolon, 479 montana, Anacystis, 264, 302 Parasphenula, 240 montanum, Pseudorabdion, 374, 398, 399 montgomeryi, Cerebratulus, 516, 519, 529, 531 mordax, Engraulis, 631, 633, 634, 651, 658, 659, 661, 662, 664, 665, 666, 667, 668, 669, 670, 681 morgani, Naja, 471 morhua, Gadus morhua, 634 macrocephalus, Gadus, 634 morhua, Gadus, 634 Mucoraceae, 260 mucosus, Ptyas, 329, 338, 478 Mucuna, 282, 283 mutisiana, 282 sloanei, 275, 282, 289 urens, 275, 282 Mugil auratus, 673 cephalus, 632, 672 milleri, Lycodon, 374, 375, 380, 391, 403, 408 Stengonotus, 374, 389, 393, 394, 404, 409 multicincta, Boiga dendrophila, 374, 378, 400 multigattatus, Alphestes, 566, 579 multirugosus, Hinnites, 114, 343 munoai, Amphisbaena, 613 Munsterman, Harold E., see Malnate, mond V. muralis, Lacerta, 476 Musci, 268 musculus, Sibbaldus, 169, 178 mutabilis, Carinoma, 516, 519, 526 mutata, Eleocharis, 255, 272, 285, 298, 299, 300 mutisiana, Mucuna, 282 Mycteropera jordani, 566 myerinki, Oligodon, 381 395, Ed- myersi, Trachyphrynus, 502, 503 Myersophis, 373, 374, 392, 412 alpestris, 373, 374, 391 Myripristis berndti, 580 clarionensis, 566, 579 Myrmecodema, 121, 161, 167 kochi, 162, 165, 166, 167 michelbacheri, 157, 163, 164, 165, 166, 167 mystaceus, Phrynocephalus, 475, 480 mystes, Scorpaena, 566, 579 mysticetus, Cetengraulis, 632 Mytilus californianus, 343, 344 Mytis, 161 Myxomycetes, 259 Myxophyceae, 264 Nacella, 103 nainiana, Ramiella, 195 naivashensis, Parasphena, 234 Naja, 482 naja miolepis, Naja, 401 Naja morgani, 471 naja, Naja, 374, 378 Naja naja, 374, 378 naja miolepis, 401 naja oxiana, 478, 480, 482 naja philippina, 374 naja philippinensis, 391, 395, 408 naja samarensis, 374, 380, 389, 393, 394, 404, 409 naja oxiana, Naja, 478, 480, 482 philippina, Naja, 374 philippinensis, Naja, 391, 395, 408 samarensis, Naja, 374, 380, 389, 393, 394, 404, 409 Najadaceae, 271 Najas, 285 marina, 266, 303 marina latifolia, 271 nana, Niceforonia, 499, 500 nancaguensis, Heliofugus ventriosus, 145 nasuta, Macoma, 342, 344 Natrix, 51, 64 auriculata, 374, 379, 380, 389, 393, 394, 404, 409 chrysarga, 374, 378, 382, 385, 401 craspedogaster, 65 dendrophiops, 374, 394 dendrophiops, barbouri, 391, 395, 099, 408 dendrophiops dendrophiops, 379, 93, 394, 399, 409 dendrophiops negrosensis, 384, 392, 398, 399, 402, 409 lineata, 374, 379, 380, 393, 404, 409 maculatus, 374 716 CALIFORNIA ACADEMY OF SCIENCES natrix, 480 natrix, Natrix, 480 Natrix natrix persa, 478 natrix persa, Natrix, 478 Natrix piscator, 64 popel, 65 pryeri, 51, 52, 54, 55, 56, 57, 59, 60, 61, 64, 65, 66 pryeri ishigakiensis, 61, 62, 65 pryeri pryeri, 64 sauteri, 64, 65 spilogaster, 374, 380, 390, 391, 403, 408 tessellata, 479 vibakari, 65 naucrates, Echeneis, 172 Navicula contortula, 352, 353, 354 evexa, 353, 354 exilissima, 353 kineaidii, 353, 354 pseudosilicula olympica, 354, 358 rainierensis, 354, 358 tecta, 352, 353 Neavella, 80, 81 nebulifer, Paralabrax, 566, 579 Nectarges nepenthe, 632 neesii, Emplectonema, 534, 535 negrosensis, Natrix dendrophiops, 384, 392, 398, 399, 402, 409 Neidium hitchcockii teres, 352, 354 inconstans, 351, 354 Nelloscolex, 205 Nemerteans from California and Oregon, by Diva Diniz Corréa, 515-558 Nemertes gracilis, 534, 535, 552 Nemertini, 516 Nemertopsis gracilis bullocki, 552 neominos, Stypochela, 96, 100 Neopilina, 108 neosubmacula, Dicladocera, 83, 84 Neothunnus macropterus, 634, 675 Neotropical Tabanidae (Diptera) mostly from Peru, Further records of, by Cornelius B. Philip, 69-102 nepenthe, Nectarges, 632 neuqueni, Heliofugus, 126, 130, 158, 166 Heliofugus (Rugosiheliofugus) , 129 Neurergus crocatus, 481 crocatus crocatus, 473 crocatus kaiseri, 473 [Proc. 4TH SER. New and rare diatoms from Oregon and Washington, by H. E. Sovereign, 349- 368 Niceforonia, 499, 502, 505 nana, 499, 500 Nicotiana, 285 glauca, 278, 279 niger, Aspergillus, 261 nigriflavus, Phaeotabanus, 81 nigrifrons, Prostoma, 546 Tetrastemma, 517, 518, 546, 547 nigrirostris, Micrura, 534: nigrocellata, Eremias intermedia, 476 nigrocincta, Euborlasia, 552 nigronotata, Esenbeckia, 74 nigropicta, Parasphena, 240 nigropunctatus, Tabanus, 89 nigrotaeniata, Maticora intestinalis, 388 nigrum americanum, Solanum, 278 Solanum, 294, 295 niphobles, Fuga, 672 niphonius, Scomberomorus, 672 niruri, Phyllanthus, 275 Nitelopterus, 46 Nitzschia bella, 356, 364 dissapata undulata, 356, 365 perspicua, 356, 365 Normanichthyidae, 596 Northern elephant seal (Mirounga angus- tirostris) off Central California, Re- establishment of the, by Keith W. Radford, Robert T. Orr, and Carl L. Hubbs, 601-612 Nostocaceae, 265 nostocorum, Plectonema, 302 nothus, Tubulanus, 525 notospilotus, Artedius, 567, 579 notospilus, Oligodon vertebralis, 374, 378, 382, 393, 394, 401, 411 nubila, Pennularia, 354, 360 nubilis, Balanus, 114 Balanus (Balanus), 344 nuchalis, Cyclocorus lineatus, 379, 394, 404, 409 nucifera, Cocos, 273 nupta, Agama, 336, 423, 442, 444, 446, 447, 448, 475, 480, 484 nycterinoides, Myemecodema, 161, 162, 166, 167 VoL. XXXI] obesa, Malacobdella, 549 oblongus, Bufo, 473 obseura, Parasphenula, 235, 236, 238, 239, 240 obscuremarginatus, Stenotabanus, 79 obscuripilus, Phaeotabanus aphanopterus, 81 obscurus, Stenotabanus, 77 obtusa, Pinnularia, 356, 361 ocellatus, Astronotus, 677 Chalcides, 479 Chalcides ocellatus, 476 ocellatus, Chalcides, 476 occidentalis, Cerebratulus, 516, 518,519, 530 Frustulia rhomboides, 351 Ocnerodrilus, 224 Taphronota, 230 Occidentosphena ruandensis, 234 Ochthyophis monochrous, 380 Ocnerodrilidae, 224 Ocnerodrilus occidentalis, 224 octaedra, Dendrobaena, 224 Octoblepharum albidum, 269, 292 Octochaetidae, 194, 224 octogrammus, Hexagrammos, 567, 583 Octolasium lacteum, 224 octolineatus, Oligodon, 387 oculus, Tabanus, 94 odendaali, Chirindites, 241 Odontogadus merlangus euxinus, 634 Oedogoniaceae, 267 Oerstedia dorsalia, 553 ohigginsi, Heliofugus 141 Heliofugus penai, 159, 166 oldendaali, Chirindites, 241 oleracea, Portulaca, 273, 284, 299 olidus, Hypomesus, 674 Oligocara, 124 Oligodon alcalai, 409 ancorus, 374, 391, 392, 394, 395, 404, 405, 408, 409 annulifer, 409 bipartita, 409 maculatus, 374, 393, 394, 409 meyerinki, 374, 381, 387, 402, 405, 407, 410, 411 modestum, 374, 393 octolineatus, 387 perkinsi, 374, 375, 385, 409, 410 (Heliofugus) penai, INDEX Ala purpurascens, 394 typica, 409 vertebralis, 374 vertebralis notospilus, 374, 378, 382, 393, 394, 401, 411 olivaceus, Bufo, 429, 430, 473, 481 olivaris, Micrura, 534 olympica, Navicula pseudosilicula, 354, 358 Ommastrepes sloani pacificus, 674 On Amphisbaena heathi Schmidt and A. car- valhoi, new species, small forms from the northeast of Brazil (Amphisbae- nia: Reptilia), by Carl Gans, 613- 630 Oncorhynchus kisutch, 680 On some earthworms of Eisen’s collection, by G. E. Gates, 185-225 Oocystaceae, 267 Oocystis solitaria, 267 opalescens, Loligo, 674 Opheodrys semicarinata, 54 semicarinata fritzei, 54 Ophiodon, 568, 569, 570, 571, 575, 577, 579, 580, SS, 583, 586m O87, 568, 959) 590, 591, 592, 593, 594, 595, 596 elongatus, 565, 567, 576, 577, 578, 579, 582 Ophiomorus, 464 blanfordi, 476, 481 brevipes, 464, 465, 476, 481 miliaris, 476 persicus, 476 tridactylus, 476, 481 Ophiophagus hannah, 374, 378, 387, 391, 393, 395, 398, 401 Ophisaurus, 483 apodus, 475, 479 Ophisops blanfordi, 476, 481 elegans, 462, 480 elegans ehrenbergi, 476 elegans elegans, 462, 476 Opisthonema libertate, 633 Opisthotropis alcalai, 374, 393, 409 typica, 374, 387, 391, 393, 395, 398, 401 Oplegnathus fasciatus, 672 Opunita, 286 orbicularis, Emys, 473, 480 ornata, Pinnularia legumen, 362 Pinnularia platycephala, 356, 362 ornatus, Phrynocephalus, 475, 481 718 CALIFORNIA ACADEMY OF SCIENCES orqueta, Chloroscombrus, 632 Orr, Robert T., see Radford, Keith W. Oryzias latipes, 673 luzonensis, 396 Oscillatoriaceae, 266 Osteological characteristics and affinities of the hexagrammid fishes, with a synop- sis, by Jay C. Quast, 563-600 otakii, Hexagrammos, 567, 576, 579, 583 Ototyphlonemertes spiralis, 552 oxalicum, Penicillium, 263 oxiana, Naja naja, 478, 480, 482 oxycephala, Gonyosoma, 374, 378, 380, 390, 398, 401 oxycephalum, Pseudorabdion, 374, 398, 409 Oxylebius, 572, 575, 580, 581, 582, 583, 585, 589, 590, 592 pictus, 565, 567, 569, 573, 576, 677, 679 Oxyrhabdion, 374, 375 Oxyrhabdium, 373, 392, 412 leporinum, 374 leporinum leporinum, 391, 392, 399 modestum, 374, 379, 380, 386, 389, 391, 393, 398, 399, 404, 407, 408 leporinum visayanum, 384, 398, 399, 409 pacifica, Ruppia maritima, 270 pacificus, Ommastrepes sloani, 674 padanum, Prostoma, 548 Palaeonemertini, 516 pallasii, Clupea, 633, 675 pallida, Paranemertes, 539 pallipes, Stenotabanus, 77 Palmae, 273, 281 palmarum, Plenoculus, 2, 4, 5, 6, 22, 25, 28, 40 palms, Cocoid, 281 palousiana, Pinnularia, 355, 360, 361 paniculatus, Chrysothamnus, 15 pannonicus, Ablepharus, 329, 377, 380, 476 papillifer, Plutellus, 188, 191, 192, 194, 223, 224 Parabrachirus, 577 Paracirrhites arcatus, 566, 579 paradisi, Chrysopelea, 373, 378, 379, 482, 383, 384, 387, 388, 389, 390, 393, 395, 398, 401, 403, 404, 405, 406 Paralabrax auroguttatus, 566, 579 clathratus, 566 nebulifer, 566, 579 [Proc. 4TH Serr. Paranemertes, 517, 537, 538 biocellata, 538 californica, 539 carnea, 539 incola, 539 pallida, 539 peregrina, 517, 518, 519, 537, 539, 540 plana, 539 Paranthias colonus, 566, 579 Parapetasia (Loveridgeacris) impotens, 229 Parasphena dubia, 242 nigropicta, 240 naivashensis, 234 pulchripes, 234 teitensis, 235 Parasphenella dubia, 242 Parasphenula grandis, 240 montana, 240 obseura, 235, 236, 238, 239, 240 pardalis, Micrura, 534. parietina, Calothrix, 265 parma, Surirella, 356, 366 parryi, Chorizanthe, 36 parva, Euschatia, 152 parvipinnis, Cynoscion, 632 Pavlovskia, 3, 44 tadzhika, 2, 3, 44 parvulum, Closterium, 267 majus, Closterium, 267 parvus, Bimastos, 221, 224 Heliofugus, 151, 152 Plenoculus, 2, 4, 5, 22, 25, 47 pastinaca, Trygon, 673 Pataecidae, 589 Patella, 103, 104, 106, 108 (Patelloida) , Acmaea, 108 tessulata, Acmaea, 103 Patina, 104, 108 (Helcion), 103 paucispinis, Sebastodes, 566, 579 pectinata, Siphonaria, 115 pectinatus, Potamogeton, 269, 270, 303 Pelecorrhynchidae, 99 Pelecorrhynchus elegans, 99 longicauda, 99 vulpes, 99 xanthopleura, 99 pellucida, Chama, 114 pellucidas, Tubulanus, 525 Pelobates fuscus, 473 VoL. XXXI] peltoides, Williamia, 115 Pempheris japonica, 672 penai, Heliofugus, 141, 151, 159 Heliofugus (Heliofugus) penai, 140 Heliofugus penai, 159, 166 ohigginsi, Heliofugus, 159, 166 ohigginsi, Heliofugus (Heliofugus), 141 penai, Heliofugus, 159, 166 penai, Heliofugus (Heliofugus), 140 Penicillium, 263 chrysogenum, 262 citrinum, 262 commune, 262 cyclopium, 262 funiculosum, 262 lanosum, 263 meleagrinum, 263 oxalicum, 263 piscarium, 263 Penoculus davisi davisi, 2, 30, 37, 43, 44, 47 pequinensis, Stenotabanus, 77 Eercan:/O; Sif o79 peregrina, Paranemertes, 517, 518, 519, 537, 539, 540 Peridiniaceae, 268 peringueyi, Plerisca, 243, 244, 245 Peristegas squarrosus, 231 perkinsi, Oligodon, 374, 375, 385, 409, 410 perniger, Plenoculus boregensis, 2, 21, 48 persa, Natrix natrix, 478 persica, Agama, 451, 452, 453, 474, 475, 481 Agamura, 474, 482 Eirenis, 466, 467, 477, 480 Eremias velox, 476 persicus, Ablepharus, 476 Bufo, 430, 473, 481, 482 Bunopus, 439, 474 Cerastes, 472 Cyclophis, 466 Hemidactylus, 441, 474, 480 Ophiomorus, 476 Phrynocephalus, 475 Pseudocerastes, 472, 478, 480 persona, Acmaea, 344 perspicua, Nitzschia, 356, 365 peruviana, Dicladocera, 86 Stypochela, 86 pes-caprae brasiliensis, Ipomoea, 276 Ipomoea, 256,°258, 275,° 277, 280, 285, 290, 292, 293, 294, 297, 298 INDEX 719 pestifer, Solierella, 37 Phaeotabanus, 81 aphanopterus, 81 aphanopterus obscuripilus, 81 litigiosus, 81 nigriflavus, 81 punctatus amabilis, 90 phalericus, Sprattus sprattus, 633 Phanerogamia, 269 Phaseolus, 285, 297 adenanthus, 297 Pheretima, 219 californica, 224 diffringens, 219, 224 hawayana, 224 Philip, Corneilus B., Further records of neo- tropical Tabanidae (Diptera) from Peru, 69—102 philippina, Boiga, 374, 390. 391 Dryophiops, 374, 390, 395, 398, 405, 406 Elaphe erythrura, 378, 381, 382, 385, 401 Liopeltis, 374, 382, 385, 411 Maticora intestinalis, 374, 391, 393, 394, 404 Naja naja, 374 Phillipine terrestrial snakes, Remarks on the zoogeography of, by Alan E. Leviton, 369-416 philippinensis, Naja naja, 391, 395, 408 philippinum, Hologerrhum, 374, 390, 391, 403, 404, 408 philippinus, Liopeltis, 401 mostly philopagus, Acrodectes, 503 Phoca vitulina, 604 phoenicis, Aspergillis, 261 pholis, Blennius, 677 phosphoreus, Microscolex, 224 Phrynocephalus, 482 helioscopus, 475 interscapularis, 475 luteoguttatus, 475, 481 maculatus, 475, 480 mystaceus, 475, 480 ornatus, 475, 481 persicus, 475 scutellatus, 475, 481 phrynoides, Rana, 334 Phycomycetes, 259 Phyiophagus hannah, 378 720 CALIFORNIA ACADEMY OF SCIENCES Phyllanthus, 285, 290, 294 amarus, 275 niruri, 275 Phyllodactylus, 442 elsiae, 44.2, 474, 481 Phyllophaga, 559, 560 galeanea, 559 ignava, 559, 562 pleroma, 562 saylori, 560, 561, 562 torta, 562 Phyllophaga saylori, new species, from Nuevo Leon, Mexico (Coleoptera: Scarabaei- dae), by Milton W. Sanderson, 559-— 562 Phymateus, 230, 243 (Phymateus) aegrotus, Phymateus, 233 Phymateus (Maphyteus) baccatus, 233 (Maphyteus) leprosus, 234: (Phymateus) aegrotus, 233 (Phymateus) iris, Phymateus, 233 Phymateus (Phymateus) iris, 233 (Phymateus) viridipes, 233 (Phymateus) viridipes, Phymateus, 233 Phymaella, 243 capensis, 243 Physeter, 179 piceus, Tabanus, 96, 100 pictifrons, Lineus, 516, 518, 528 picturata, Tanita, 246 pictus, Dendrelaphis, 374, 378 Dendrelaphis pictus, 382, 383, 384, 385, 387, 388; 390; 393; 9395, 398) 401- 402, 403, 404 Oxylebius, 565, 567, 569, 573, 576, 577, 579 pictus, Dendrelaphis, 382, 383, 384, 385, 387, 6885 390) 3935395. 398) 408 402, 403, 404 pilchardus sardina, Sardina, 633 pilsbryi, Zirfaea, 344 Pinnularia, 359, 362 convexa, 354, 359 divergens, 359 divergens elliptica, 359 legumen ornata, 362 makahana, 354, 359 nubila, 354, 360 obtusa, 356, 361 palousiana, 355, 360, 361 [Proc. 4TH SER. platycephala ornata, 356, 362 pluviana, 354, 362 subpalousiana, 354, 361 umbrosa, 356, 363 Pinus, 559 teocote, 560 pipiens, Alsophylax, 329, 334, 336, 474 Ascalabotes, 334, 336 Gymnodactylus, 336 Lacerta, 334, 335, 336 Piptocephalidaceae, 260 piscarium, Penicillium, 263 piscator, Natrix, 64 Pisonia grandis, 284 plagiodon, Stenella, 170, 179 plana, Paranemertes, 539 planatus, Geocarcinus, 258 platycephala ornata, Pinnularia, 356, 362 Platycephalidae, 567, 592, 593 Platycephalus, 568, 571 Plectonema nostocorum, 302 terebrans, 302 Plenoculus, 1, 2, 3, 4, 20, 21, 22, 44, 46, 47, 48 abdominalis, 32, 38, 43, 44 apicalis, 13, 32, 40, 44, 47, 48 atlanticus, 43 beaumonti, 33, 44 boharti, 2, 6, 7, 28, 30, 34, 41 boregensis, 2, 4, 5, 6, 8, 20, 21, 22 boregensis boregensis, 21 boregensis perniger, 2, 21, 48 cockerellii, 2, 4, 5, 7, 12, 15, 17, 18, 21, 28, 29, 40, 45, 46, 47 cuneatus, 2, 4,5, 7, 12, 13, 15, 18, 19 Gknvi A sin Oy My ISH AD, 27, 28, 29) SO) 31, $2). 33, 34; 35, 38.03oueqOuelse 44, 46, 47, 48 davisi atlanticus, 3, 37, 39, 43, 47 davisi gracilis, 3, 29, 39, 42 davisi mojavensis, 3, 38, 39, 45 davisi transversus, 3, 13, 34, 37, 38, 40, 42 deserti, 2, 6, 7, 26, 27, 45 gillaspyi, 2, 4,5, 8, 10, 23, 47 hurdi, 2, 4, 5, 10, 23 mexicanus, 2, 4, 5, 7, 17, 18, 19 palmarum, 2, 4, 5, 6, 22, 25, 28, 40 parvus, 2, 4, 5, 22, 25, 47 Vou. XXXI] propinquus, 2, 4, 6, 7, 9, 25, 26, 27, 28, 34, 41, 47, 48 propinquus rufescens, 25 sinuatus, 2, 4,5, 6, 7, 22, 23 stygius, 2, 6, 7, 12, 31, 32, 34, 45, 46, 48 tadzhika, 2, 3, 44 timberlakei, 2, 4, 5, 6, 9, 11, 12, 13, 15, 17, 18, 19, 22, 28, 29, 40 Plerisca, 244, 245 peringueyi, 243, 244, 245 rubripennulis, 243, 244, 245 senecionicola, 245 pleroma, Phylophaga, 562 pleuroden, Rana, 334 Pleurogrammus, 570, 580, 582, 583, 586, 589, 590, 591, 592 monopterygius, 567, 571, 576, 579, 581 plumbeum, Dictydiaethalium, 259, 260 Plutellus, 187 collinus, 224 fenderi fenderi, 224 heteroporus, 188 marmoratus, 193, 224 papillifer, 188, 191, 192, 194, 223, 224 sierrae, 224 umbellulariae, 223, 224 pluviana, Pinnularia, 354, 362 Pneumatophorus diego, 633, 634 japonicus, 633 Pododesmus macroschisma, 114 Poeciloderos, 89 callipareus, 230 cincta, 230 polifolium, Eriogonus, 34 polillensis, Calamaria gervaisis, 403 Cornufer, 403 polycarpa, Euphorbia, 16, 24, 32 hirtella, Euphorbia, 15, 24, 33, 46 polymorpha, Carinella, 520 polymorphus, Tubulanus, 516, 518, 519, 520, 525 Pomacentridae, 566 pontica, Engraulis encrasicholus, 633, 677 Pontoscolex, 205, 219 corethrurus, 205, 219 popei, Natrix, 65 porcus, Scorpaena, 673 Portulaca, 285 oleracea, 273, 284, 299 Portulacaceae, 273 INDEX 721 Portunus, 552 Potamogeton, 270, 285 pectinatus, 269, 270, 303 Potamogetonaceae, 269 prasina, Ahaetulla, 373 prasina, Ahaetulla, 378, 382, 385, 400, 405 preocularis, Ahaetulla, 379, 380, 383, 384, 387, 388, 389, 390, 393, 394, 398, 402, 404, 410 prava, Achnanthes, 351, 354 preocularis, Ahaetulla prasina, 379, 380, 383, 384, 387, 388, 389, 390, 393, 394, 398, 402, 404, 410 Ahaetulla presiana, 374, 379, 380, 384, 387, 388, 389, 390, 393, 394, 398, 402, 404, 410 presbiter, Veprius, 71, 100 presiana preocularis, Ahateulla, 374, 379, 380, 383, 384, 387; 388, 389, 390; 394, 398, 402, 404, 410 pretrei, Amphisbaena, 615 princeps, Lacerta, 476 Prionotus, 571 albiostris, 567, 579 stephanophrys, 567, 579 Pristurus rupestris, 474, 480 Proboscoides, 73 fairchildi, 74: Procephalothrix, 516, 526, 527 major, 527 spiralis, 516, 519, 526, 527 procumbens, Triumfetta, 284 Propilidium, 104 propinquus, Plenoculus, 2, 4, 6, 7, 9, 25, 26, 27, 28, 34, 41, 47, 48 rufescens, Plenoculus, 25 Tabanus, 91 prosopidis, Solierella, 3 Prosorhochmus albidus, 552 Prostoma, 517, 548, 549 aquarum-dulcium, 549 asensoriatum, 549 eilhardi, 548 graecense, 548, 549 grande, 548 lumbricoideum, 548 nigrifrons, 546 padanum, 548 722 CALIFORNIA ACADEMY OF SCIENCES puteale, 548 rubrum, 517, 54.8, 549 Protococcus grevillei, 268, 302 Protothaca, 344 ruderata, 344 staminea, 343, 344 proxima, Euschatia, 154 proximoides, Heliofugus, 145, 147, 149, 160 Heliofugus (Heliofugus), 146 Heliofugus proximoides, 160, 166 proximoides, Heliofugus, 160, 166 rotundangulus, Heliofugus, 160, 166 rotundangulus, Heliofugus (Heliofugus), 147 proximus, Heliofugus, 132, 141, 146, 152, 156, 160 Heliofugus (Heliofugus), 154 Heliofugus proximus, 146, 147, 160 proximus, Heliofugus, 146, 157, 160 punctatosulcatus, Heliofugus, 156, 158, 167 punctatosulcatus, fugus, 155 pryeri ishigakiensis, Natrix, 61, 62, 65 NaiiaEs, Sil, bP4, 544, 55, SO, 57, 59, GO, wil, 64, 65, 66 Natrix pryeri, 64 pryeri, Natrix, 64 Tropidonotus, 52 Psallus seriatus, 47 Psammodynastes pulverulentus, 379, 380, 381, 382, 386, 398, 401, 402, 403, 404 Psammophis lineolatus, 478, 480 schokari, 423, 470, 478, 480 Pseudoboa carinata, 471 Pseudocerastes, 472, 482 latirostris, 472 mcemahoni, 478 persicus, 472, 478, 480 Pseudochama exogyra, 114 pseudoculus, Tabanus, 94 pseudo-lacteus, Lineus, 528 134n135ee 139, Heliofugus (Helio- 374, 387, 378, 393, Pseudophallus starksii, 632 Pseudorabdion, 394, 409 albonuchalis, 409 ater, 374, 393, 409 longiceps, 409 mcnamarae, 374, 391, 398, 399 minutum, 399 [Proc. 41H SrEr. montanum, 374, 398, 399 oxycephalum, 374, 398, 409 saravacensis, 409 taylori, 374, 393, 394, 409 pseudosilicula olympiea, Navicula, 354, 358 pseudotaeniotes, Stenotabanus, 77 Psychrolutidae, 596 Pterois antennata, 566, 579 Ptyas mucosus, 329, 338, 478 Ptygosphex, 44 Ptyodactylus hasselquisti, 474, 479 pugetensis, Chitonotus, 567 Pugettia, 552 pulcher, Amphiporus, 542 pulchripes, Paraspena, 234 pulverulentus, Psammodynastes, 374, 370, “380, Bi," 3s2yhssonncen 391, 393, 398, 401, 402, 403, 404 punctata, Dermatolepis, 566, 572 Euschatia, 135 Girella, 672 punctatissimus, Chilomeniscus, 309, 313, 315, B72 322 punctatolineata, Eirenis, 477 punctato-striata, Euschatia, 136 punctatostriatus, Heliofugus, 135 Heliofugus (Heliofugus), 136 punctatosulcatus, Heliofugus, 155 Heliofugus (Heliofugus) proximus, 155 Heliofugus proximus, 156, 158, 167 punctatulus, Amphiporus, 546 punctatus amabilis, Phaeotabanus, 90 Heliofugus, 135 Heliofugus impressus, 133, 136, 139, 140, 159, 166 punctipennis, Tabanus, 89, 90 378, 389, pungens, Tabanus, 91 Tabanus (Taeniotabanus), 90 punnetti, Baseodiscus, 552 Puntius, 396, 397 hemictenus, 396, 397 ivis, 386 purpurascens, Oligodon, 394 puteale, Prostoma, 548 Pyrenocarp lichen, 263 Pyrgomorpha, 244 cylindrica, 245 dispar semlikiana, 245 granulata, 245 VoL. XXXI] Pyrgomorphella, 242, 243, 244 albini, 242 arachidis, 241 rugosa, 24.2, 243, 244 senecionicola, 243, 244, 245 serbica, 244 Pyrgomorphidae, 227 Pyrgomorphidae (Orthoptera: Acridoidea) collected in Africa by E. S. Ross and R. E. Leech, 1957-1958, with de- scriptions of new species, by D. Keith McE. Kevan, 227-248 Pyrrophyceae, 268 Python, 482 reticulatus, 374, 378, 379, 380, 384, 387, 389, 391, 393, 395, 398, 401, 402, 403, 404, 407 quadricornis, Hypsagonus, 572 quadrilineatum, Tetrastemma, 547 quadripuncta, Hypopelma, 89 quadripunctatus amabilis, Tabanus, 89, 100 Tabanus, 89, 90 quaturolineata, Elaphe, 478 quillotaensis, Heliofugus, 148, 149, 155, 160, 166 Heliofugus (Heliofugus), 148 racemosum, Syncephalastrum, 260 raddei, Vipera, 478 Radford, Keith W., Robert T. Orr, and Carl L. Hubbs, Reestablishment of the northern Elephant Seal (Mirounga an- gustirostris) off Central California, 601-612 rainierensis, Cymbella, 356, 363 Denticula, 356, 364. Navicula, 354, 358 Ramiella, 194, 195, 217, 218 americana, 215 nainiana, 195 Ramiellona, 192, 194, 195, 218 americana, 215, 218 balantina, 202, 205, 214, 218 eiseni, 209, 213, 216, 218 guatemalana, 195, 205, 209, 212, 213, DiS 216, 217, 218, 219 irpex, 213, 218, 219 lasiura, 215, 219 lasiurus, 201, 216, 219 mexicana, 209, 214, 218 INDEX 723 sauerlandti, 216, 218 stadelmanni, 214, 216, 218 strigosa, 206, 209, 213, 216, 218 tecumumami, 213, 214, 216, 218 vulcanica, 214, 216 Rana, 432 boulengeri, 334 cyanophlyctis, 430, 433, 473, 480 erythraea, 406 limnocharis, 54 macrodon, 387 macrodon visayanum, 387 phrynoides, 334 pleuroden, 334 ridibunda, 424, 432, 434, 479 ridibunda ridibunda, 432, 473 signata, 389 signata similis, 389 sternosignata, 329, 332 tibetanus, 334 Ranidae, 432, 473 ravergieri, Coluber, 477, 480 Reestablishment of the northern Elephant Seal (Mirounga angustirostris) off Central California, by Keith W. Rad- ford, Robert T. Orr, and Carl L. Hubbs, 601-612 reflexa, Dioclea, 282 refracta, Wislezenia, 16 Remarks on the colubrid genus Chilomenis- cus (Serpentes: Colubridae), by Ben- jamin H. Banta, and Alan E. Leviton, 309-327 Remarks on the zoogeography of Philippine terrestrial snakes, by Alan E. Leviton, 369-416 Remilegia, 170, 171, 172 australis, 169, 170, 171, 172, 173, 174, WS, Wii, Lae scutata, 172 Remilegia australis (Bennett) from Califor- nia, with meristic data, First records of the echeneidid fish, by W. I. Fol- lett, and Lillian J. Dempster, 169— 184 Remora, 170 australis, 170, 171 remora, 171, 172 Scuitatase lial 2 remora, Remora, 171, 172 724 CALIFORNIA ACADEMY OF SCIENCES renardi, Vipera, 478, 480 reniforme, Eriogonum, 36 Rentz, David C., see Gurney, Ashley B. repens, Lepturus, 284 Reptilia, 434, 473 Reptiles from Iran, Amphibians and, by Steven C. Anderson, 417-498 rerio, Brachydanio, 679 reticulata, Tetraclita, 114 Trimusculus (Gadinia), 111 reticulatum, Tetrastemma, 54.7 reticulatus, Python, 374, 378, 379, 380, 384, 385 SE, SOI, BOs, BOS, SOS. Aoi, 402, 403, 404, 407 Trimusculus, 114, 119 Revision Der Chilenischen Misolampini: Gat- tungen Heliofugus Guerin Und Myre- mcodema Gebien (= Myremcosma Germain) (Coleoptera: Tenebrioni- dae) by Heinz Freude, 121-168 rhabdotus, Tubulanus, 522, 523 Rhabdotylus venenata, 87 viridiventris, 87 Rhacophorus appendiculatus appendiculatus, 386 hecticus, 405 rhinopoma, Telecopus, 468, 478, 481 Rhizopus arrhizus, 260 rhodensiensis, Afrosphena, 24.1 rhodorachis, Coluber, 329, 338, 423, 465, 466, 477, 480 Zamenis, 465 rhombifolia, Sida, 276, 294 rhomboides oecidentalis, Frustulia, 351 Rhombus maeoticus, 673 Rhynchocalamus satunini, 478, 481 Rhynchocoela, 516 ribulatus, Cirrhitus, 566, 569, 575, 584 Ricardoa, 73 ridgewayi, Lytorhynchus, 478 ridibunda, Rana, 424, 432, 434, 479 Rana ridibunda, 432, 473 ridibunda, Rana, 432, 473 Riopa, 482 Rivulariaceae, 265 rivulatus, Cirrhitus, 569 Roccus, 572 saxatilis, 568, 576, 577, 580 rosaceus, Solen, 519 [Proc. 4TH SER. rosea, Canavalia, 281 Eisenia, 224 rossi, Chrysops, 75, 100 Heliofugus, 154, 156, 160, 167 Heliofugus (Heliofugus), 153 rostrata, Ruppia maritima, 270 rothi, Eirenis, 477, 481 rotundangulus, Heliofugus proximoides, 147 Heliofugus proximoides, 160, 166 ruandensis, Occidentosphena, 234: rubellus, Amphiporus, 546 Lumbricus, 224 rubens, Zygenupolia, 552 ruber, Lineus, 516, 518, 527, 528, 529 rubescens, Dryophiops, 374, 385 Lineus, 528 Tetriclita squamosa, 114 rubida, Dendrobaena, 221, 223, 224 rubiginosus, Scirpus, 272, 273, 300 rubra, Carinella, 520 rubricornis, Mesomyia, 72 Mesomyia (Coracella), 72 rubrigularis, Agama, 475, 481 rubripennulis, Plerisca, 243, 244, 245 (Heliofugus) rubripes, Fuga, 672 Tabanus, 92 rubrofemorata, Bellardia, 92 rubrum, Prostoma, 549 ?Prostoma, 517, 548 ruderata, Agama, 453, 475, 480 Protothaca, 344 rufescens, Plenoculus propinquus, 25 rufitarsis, Heliofugus, 158 rufopilosus, Mesomyia (Coracella), 72 rugosa, Pyrgomorphella, 242, 243, 244 (Rugosiheliofugus) Heliofugus, 128, 130, 166 neuqueni, Heliofugus, 129 suleatulus, Heliofugus, 128 Ruppia, 285 maritima, 270, 302 maritima pacifica, 270 maritima rostrata, 270 rupestris, Pristurus, 474, 480 russowl, Cyrtodactylus, 474 Rutidoderes cinctus, 231 concolor, 231, 232 squarrosus, 231 Vor. XXXI] sachalinensis, Spisula, 551 Sachet, Marie Hélene, Flora and vegetation of Clipperton Island, 249-307 sagar melanosticta, Sarinops, 633 saida, Boreogadus, 634 saira, Cololabis, 674 Salamandridae, 473 salicina, Baccharis, 8 saliens, Dichogaster, 224 Salientia, 428, 473 sallei, Tabanus, 91, 100 salmoides, Micropterus, 566, 579 Salvelinus leucemaenis, 674 samarensis, Naja naja, 374, 380, 389, 393, 394, 404, 409 Sanderson, Milton W., Phyllophaga saylori, new species, from Nuevo Leon, Mex- ico (Coleoptera: Scarabaeidae), 559- 562 sanguineus, Lineus, 527, 528 sanguinolenta, Agama, 445, 446, 451 Agama agilis, 445 Lacerta, 446 Sapindaceae, 275, 282 Sapindus, 282, 283 saponaria, 275, 282 saponaria inaequalis, 276 saponaria microcarpus, 276 saponaria inaequalis, Sapindus, 276 microcarpus, Sapindus, 276 Sapindus, 275, 282 saravacensis, Pseudorabdion, 409 Sardina pilchardus sardina, 633 sardina, Sardina pilchardus, 633 Sardinella aurita, 633 macrophthalma, 633, 672 Sardinops caerulea, 633, 634 sagar melanosticta, 633 Sartorya fumigata, 260 satanica, Dicladocera, 84 satunini, Rhynchocalamus, 478, 481 sauerlandti, Howascolex (Graceevelynia), 216 Ramiellona, 216, 218 Sauria, 435, 474 sauteri, Natrix, 64, 65 savagei, Chilomeniscus, 309, 313, 315, 317, 322 savignyi, Hyla arborea, 424, 430, 432, 473 saxatilis, Roccus, 568, 576, 577, 580 saxicola defillipi, Lacerta, 476 INDEX 72 On Lacerta, 480 Saxidomus giganteus, 344 saylori, Phyllophaga, 560, 561, 562 scaber, Cyrtodactylus, 437, 438, 441, 474, 480 Stenodactylus, 437 scabra, Absidia, 260 Scaevola, 284 sericea, 284 Scaptia, 70 albithorax, 70 atra, 70, 100 atra leucothorax, 70 grisea, 70 grisea jucundus, 70 jucunda, 70 (Scaptia) leucothorax, Listrapha, 70 Scaptia limbithorax, 70 schirazianus, Spalerosophis, 478 Schizothaerus capax, 343, 344 Schizothrix heufleri, 267 schlingeri, Esenbeckia, 72, 100 schmidti, Acanthodactylus cantoris, 457,475 schneideri, Eumeces, 479 variegatus, Eumeces, 476 schokari, Coluber, 470 Psammophis, 423, 470, 478, 480 schultzei, Trimeresurus, 374, 375, 378, 401, 411 Scianenidae, 566 Scincidae, 462, 476 Scincus, 462, 482 conirostris, 462, 463, 476, 481, 485 scincus, Teratoscincus, 474, 481 Scirpus, 285, 300 rubiginosus, 272, 273, 300 Scomber scombrus, 633 Scomberomorus niphonius, 672 scombrus, Scomber, 633 Scorpaena, 572 guttata, 566, 579 mystes, 566, 579 456, porcus, 673 Scorpaenichthys, 580 marmoratus, 567, 574, 575, 579, 583 Scorpaenidae, 566, 567, 587 scripta, Eremias, 476 scutata, Echeneis, 170, 171, 176, 177 Remilegia, 172 726 CALIFORNIA ACADEMY OF SCIENCES Remora, 171, 172 scutellata, Dicladocera, 84: scutellatus, Phrynocephalus, 475, 481 Scutiger, 334 Scytoemataceae, 265 Scytonema, 302 hofmannii, 265, 302 sealei, Lycodon subcinctus, 401 Sebastes, 580 viviparus, 577 Sebastiscus marmoratus, 566, 579, 583 Sebastodes, 572 chlorostictus, 566, 579 melanops, 566 paucispinis, 566, 579 serriceps, 567, 573 Sebastolobus alascanus, 567, 568, 576, 579 altivelis, 567, 579 seclineata, Carinella, 521 semicarinata fritzei, Liopeltis, 54 fritzei, Opheodrys, 54 Opheodrys, 54 seminuda, Esenbeckia, 74 semiplena, Lyngbya, 266 semlikiana, Pyrgomorpha dispar, 245 Semotilus atromaculatus, 677 senecionicola, Plerisca, 245 Pyrgomorphella, 243, 244, 245 senior, Tabanus, 98 septemtaeniata, Euprepis, 463 Mabuya aurata, 423, 463, 464, 476, 484 serbica, Pyrgomorphella, 244 seriatus, Psallus, 47 sericea, Scaevola, 284 Serpentes, 465, 476 Serranidae, 566 serriceps, Sebastodes, 567, 573 Setarches, 574 sexannulatus, Stenotabanus, 79 sexlineata, Carinella, 521 sexlineatus, Tubulanus, 516, 518, 519, 521, 523, 525 Sibbaldus musculus, 169, 187 Sibynophis bivittatus, 374, 378, 385, 401, 410 geminatus, 378, 411 Sida, 274, 285, 294, 295, 296, 301 rhombifolia, 276, 294 sierrae, Plutellus, 224 382, [Proc. 4TH SER. signata, Rana, 389 similis, Rana, 389 signifer, Tetrastemma, 54:7 Silurus glanis, 673 similis, Amphiporus imparispinosus, 546 Rana signata, 389 sinuatus, Plenoculus, 2, 4, 5, 6, 7, 22, 23 Siphonaria, 108, 117, 118 alternata, 111, 115, 116, 118 brannani, 115 pectinata, 115 thersites, 111, 115, 116, 117, 118 Siphonariidae, 115 slevini, Amphisbaena, 613 sloanei, Mucuna, 275, 282, 289 sloani pacificus, Ommastrepes, 674 smithi, Sparganophilus, 224 Solanaceae, 278 Solanum, 258, 285, 290, 294, 295, 296, 301 nigrum, 294, 295 nigrum americanum, 278 Solen rosaceus, 519 Solierella, 2, 3, 4, 46, 47, 48 albipes, 3 pestifer, 37 prosopides, 3 solitaria, Odcystis, 267 sonomae, Sparganophilus, 224 sorbillans, Tabanus, 92 Tabanus (Macrocormus), 91 Sovereign, H. E., New and rare diatoms from Oregon and Washington, 349— 368 Spalerosophis diadema, 480 diadema diadema, 478 microlepis, 478 schirazianus, 478 Sparganophilidae, 220, 224 Sparganophilus, 186, 187, 220 eiseni, 220 smithi, 224 sonomae, 224 speciosa, Carinella, 520 Sphaerellaceae, 268 Sphyrna zygaena, 632 spilogaster, Natrix, 374, 380, 390, 391, 403, 408 spinella, Hypena, 303 spinosa, Dalea, 12, 16 spiralis, Ototyphlonemertes, 552 Vout. XXXI] Procephalothrix, 516, 519, 526, 527 Spisula sachalinensis, 551 sprattus balticus, Sprattus, 633 phalericus, Sprattus, 633 Sprattus sprattus, 633 sprattus, Sprattus, 633 Sprattus sprattus balticus, 633 sprattus phalericus, 633 sprattus sprattus, 633 spumans, Dictyophorus (Dictyophorus) , 228 Squalus acanthias, 673 Squamata, 435, 474 squamosa rubescens, Tetraclita, 114 squarrosus, Peristegas, 231 Rutidoderes, 231 stadelmanni, Ramiellona, 214, 216, 218 stali, Taphronota, 231 staminea, Protothaca, 343, 344 starksi, Menidia, 632 starksii, Pseudophallus, 632 Stauroneis, 358 Stegonotus, 412 miilleri, 374, 389, 393, 394, 404, 409 stellaris, Calothrix, 265 stellatus, Acipenser, 673 stelleri, Hexagrammos, 567 Stenella plagiodon, 170, 179 stenocephalus, Tabanus (Taeniotabanus), 90 Stenodactylus lumsdeni, 474 scaber, 437 Stenopterobia 365 Stenotabanus, 77, 78, 80, 82, 88 albilinearis, 78, 100 flavofemoratus, 77 fulvistriatus, 77 macroceras, 79, 81, 100 maculifrons, 79 obscuremarginatus, 79 obscurus, 77 pallipes, 77 pequinensis, 77 pseudotaeniotes, 77 sexannulatus, 79 taeniotes, 77 xenium, 77 intermedia undulata, 356, Stephanolepis cirrhifer, 672 stephanophrys, Prionotus, 567, 579 Sterculiaceae, 276 Stereolpeis gigas, 566 INDEX 97 sternosignata, Rana, 329, 332 steudneri, Tropiocolotes, 440 Stigonemataceae, 265 stipes, Antherina, 679 stramineus, Chilomeniscus, 315/321, 323 Chilomeniscus stramineus, 310, 317, 323 esterensis, Chilomeniscus, 315, 317, 323 fasciatus, Chilomeniscus, 313, 317, 321, 322 stramineus, Chilomeniscus, 310, 317, 323 strauchi, Eremias velox, 476 striatus bicolor, Lycodon, 478, 482 Heliofugus, 151, 152, 157 Lineus, 532 strigata, Lacerta, 476, 480 strigosa, Ramiellona, 206, 209, 213, 216, 218 Strongylodon lucidus, 282 stygius, Plenoculus, 2, 6, 7, 12, 31, 32, 34, 45, 46, 48 Stypochela, 86 inea, 85, 100 minos, 97 neominos, 96, 100 peruviana, 86 Stypommia, 89 abdominalis, 88, 100 flavescens, 88 Styposelaga, 79 subannulata, Dryocalamus, 374, 401 subcinctus, Lycodon, 378 sealei, Lycodon, 401 subcylindrica, Tanita, 246 sublaeta, Chrysops laeta, 75 309, sublima, Agnostokasia, 504, 505, 506, 507, 508, 509, 511, 512 submacula, Dicladocera, 83, 84 suborbitalis, Holocentris, 566, 579 subpalousiana, Pinnularia, 354, 361 subprotumidum, Cosmarium, 267 subverrucosa, Taphronota, 230 sulcata, Euschatia, 128 sulcatulus, Heliofugus, 125, 126, 130, 132, 158, 166 Heliofugus (Rugosiheliofugus) , 128 sulcatus, Heliofugus, 123, 128, 129, 131, 132, 138, 151, 156, 158, 159, 166 Heliofugus (Heliofugus), 131 sulcipennis brevipennis, Heliofugus, 143, 159, 166 728 CALIFORNIA ACADEMY OF SCIENCES brevipennis, Heliofugus (Heliofugus), 143 Euschatia, 142, 143 Heliofugus, 140, 142, 143, 150, 156, 159 Heliofugus (Heliofugus), 142 Heliofugus sulcipennis, 143, 149, 166 sulcipennis, Heliofugus, 143, 159, 166 suluensis, Calamaria, 383 Maticora intestinalis, 374, 387, 388, 410, 411 superciliosus, Hexagrammos, 565, 567, 576, 579 surdus, Bufo, 430, 473, 481 Surirella beadensis, 356, 365, 366 parma, 356, 366 sydowl1, Aspergillus, 261 symmetricus, Trachurus, 633, 634 Synancejidae, 588, 589 Syncephalastrum racemvsum, 260 Synodus lucioceps, 632 Tabanidae, 99 Tabanus, 71, 77, 80, 81 albibarbis, 98 alboater, 98 albocirculus, 93, 94 albonotatus, 94 albopruinosus, 94 amabilis, 89 ameghinoi, 91 angustifrons, 98 angustivittus, 91, 100 annulicornis, 71 atricornis, 98 basivittus, 95, 98 bipartitus, 94 bitinctus, 95 callosus, 91 carbo, 71 carneus, 90 contirmatus, 83 dasyphrytina, 89 desertus, 91, 100 discus, 99 elegans, 89 flavescens, 88 fumomarginatus, 94, 95 impressus, 95, 96, 100 lophus, 94 (Lophotabanus) flavicorpus, 94, 100 [Proc. 4TH SER. (Lophotabanus) lophus, 92 (Macrocormus) sorbillans, 91 marginenevris, 95 missionum, 82 nigropunctatus, 89 oculus, 94 piceus, 96, 100 propinquus, 91 pseudoculus, 94 punctipennis, 89, 90 pungens, 91 quadripunctatus, 89, 90 quadripunctatus amabilinus, 89, 100 rubripes, 92 sallei, 91, 100 senior, 98 sorbillans, 92 (Taeniotabanus) callosus, 82 (Taeniotabanus) carneus, 90 (Taeniotabanus) claripennis, 91 (Taeniotabanus) pungens, 90 (Taeniotabanus) stenocephalus, 90 testaceus, 92 viduus, 95 xipe, 93 Tachysphex, 46 tadzhika, Pavlovskia, 2, 3, 44 Plenoculus, 2, 3, 44: taeniolatus, Eumeces, 476 Taeniotabanus, 82 (Taeniotabanus) callosus, Tabanus, 82 carneus, Tabanus, 90 claripennis, Tabanus, 91 pungens, Tabanus, 90 stenocephalus, Tabanus, 90 taeniotes, Stenotabanus, 77 Tanita, 245 lineaalba, 246 loosi, 245 picturata, 246 subcylindrica, 246 (Tapeseilla) griseus, Dictyophorus, 229 Dictyophorus, 229 griseus intermedius, Dictyophorus, 229 laticinctus, Dictyophorus, 229 Taphronota, 231 amaranthina, 230 calliparea, 230, 231 corallipes, 229 ferruginea, 229, 230 griseus fuscoroseus, VoL. XXXII] occidentalis, 230 stali, 231 subverrucosa, 230 Tarbophis martini, 467, 468 tessellatus, 467 tataricus, Eryx, 477 taylori, Pseudorabdion, 393, 394, 409 tecta, Navicula, 352, 353 tecummami, Eodrilus, 214 Ramiellona, 213, 214, 216, 218 vulcanicus, Howascolex nia), 216 Tegula brunnea, 118 funebralis, 343 teitensis, Parasphena, 235 Telephoraceae, 260 Telescopus, 467, 468 fallax iberus, 468, 478, 481 rhinopoma, 468, 478, 481 tessellatus, 478, 481, 482 tessellatus tessellatus, 467, 468, 470 Telmatobius, 334 tenipunctatus, Heliofugus, 135, 136 Heliofugus (Heliofugus), 136 teocote, Pinus, 560 tepicana, Esenbeckia, 74 Teratoscincus, 482 bedriangai, 474, 482 microlepis, 474, 482 scincus, 474, 481 terebrans, Plectonema, 302 teres, Neidium hitchcockii, 352, 354 terrestris, Lumbricus, 224 terreus, Aspergillus, 261, 262 terrificus, Dendrelaphis caudolineatus, 374, SSe SO, 380) 382, 388; 389), 393; 394, 398, 399, 403, 404, 406, 411 tessellata, Natrix, 478, 479 tessellatus, Lycodon, 391 Tarbophis, 467 Telescopus, 478, 481, 482 Telescopus tessellatus, 467, 468, 470 tessellatus, Telescopus, 467, 468, 470 tessulata, Acmaea, 107 Acmaea (Patelloida), 103 testaceus, Tabanus, 92 (Graceevely- testarum, Mastigocoleus, 265, 302 Testrastemma candidum, 54.7 Testudinidae, 474 Testudo, 482 INDEX 779 caspica, 434 graeca, 479 graeca ibera, 474, 481 horsfieldi, 474, 481 zarudnyl, 474, 482 Tetraclita reticulata, 114 squamosa rubescens, 114 tetraedra, Eiseniella, 224 Tetrastemma, 517, 546, 547 bilineatum, 547 caecum, 546 hansi, 546 marioni, 546 nigrifrons, 517, 518, 546, 547 quadrilineatum, 547 reticulatum, 54.7 signifer, 547 Tetrastemmatidae, 517 thaelephorus, Gryllus, 230 Thais lamellosa, 343, 344 lima, 343 thalassina, Zygonermertes, 541 thersites, Siphonaria, 111, 115, 116, 117, 118 The wasps of the genus Plenoculus (Hyme- noptera: Sphecidae, Larrinae), by Francis X. Williams, 1-49 Third contribution of the herpetology of Afghanistan, by Alan E. Leviton, and Steven C. Anderson, 329-339 thomasi, Eriogonum, 16 thrissina, Harengula, 632 tibetanus, Rana, 334 tigrinus, Amphiporus, 545 Tiliaceae, 276 timberlakei, Plenoculus, 2, 4, 5, 6, 9, 11, 12, 135 15) 17,918. 1922)078" 2040 Timeresurus, 375 Tinca tinca, 680 tinca, Tinca, 680 tinetipennis, Dicladocera, 84, 100 Tonoscolex, 205 torquatus, Lineus, 528 torta, Phyllophaga, 562 Tournefortia, 284 argentea, 284 Trachurops crumenophthalmus, 632 Trachurus japonicus, 633, 672 symmetricus, 633, 634 trachurus, 633, 673 trachurus, Trachurus, 633, 673 730 CALIFORNIA ACADEMY OF SCIENCES Trachyphrynus, 502 myersi, 502, 503 translucens, Esenbeckia, 73 transversus, Plenoculus davisi, 3, 13, 34, 37, 38, 40, 42 trapezoides, Allolobophora, 224 Triceratomyia incipiens, 79 mcintyrei, 74 Trichoderma viride, 263 trichopes, Eriogonum, 8 trichopodum, Eriogonum, 12, 32 tricolor, Liopeltis, 374, 378, 382, 385, 401, 411 tridactylus, Ophiomorus, 476, 481 Trigaster, 194 Triglidae, 567, 587, 588 trigonata, Boiga, 480, 482 melanocephala, Boiga, 477 triloba, Ipomoea, 277, 285 Trimeresurus flavomaculatus, 374, 376 flavomaculatus flavomaculatus, 380, 383, 387, 388, 389, 391, 393, 394, 395, 398, 410 flavomaculatus halieus, 403 flavomaculatus mcgregori, 380 schultzei, 374, 375, 378, 401, 411 wagleri, 374, 378, 379, 387, 389, 391, 393, 398, 401, 404 Trimusculus, 108, 112, 116 (Gadinia) reticulata, 111 reticulatus, 114, 119 Trionychidae, 474 Trionyx, 483 euphraticus, 474, 481, 483 tripolitanus, Tropicolotes, 440 tristrigatus, Dryocalamus, 374, 378, 401 cristatus karelini, 473 cristatus longipes, 473 Triumfetta procumbens, 284 Tropidonotus pryeri, 52 Tropiocolotes, 336, 440 helenae, 336, 440 steudneri, 440 tripoltanus, 440 Trygon pastinaca, 673 tuba, Ipomoea, 277 Tuberculariaceae, 263 tuberculata, Bunopus, 336 tuberculatus, Bunopus, 439, 474, 480 tuberosa, Merremia, 283 [Proc. 4TH Ser. Tubulanidae, 516 Tubulanus, 516, 519, 522, 525 albocinctus, 525 borealis, 522, 523 capistratus, 525 cingulatus, 516, 519, 524, 525 frenatus, 525 nothus, 525 pellucidas, 525 polymorphus, 516, 518, 519, 520, 525 rhabdotus, 522, 523 sexlineatus, 516, 518, 519, 521, 523, 525 Tupinambis griseus, 455 turcicus, Hemidactylus, 474 turgida, Allolobophora, 224 turgidus, Chroococcus, 264 Two genera of Leptodactylid frogs from Colombia, by Coleman J. Goin, and Doris M. Cochran, 499-505 (Tylostypia), Hybomitra, 89 Typhlopidae, 476 Typhlops, 482 braminus, 476, 480 vermicularis, 477, 480 wilsoni, 477 typica, Oligodon, 409 Ophisthotropis, 374, 387, 391, 393, 395, 398, 401 tyria, Coluber, 477 Ulva, 303 umbellulariae, Plutellus, 223, 224 umbra, Corvina, 579, 673 umbrosa, Pinnularia, 356, 363 undulata, Nitzschia dissapata, 356, Stenopterobia intermedia, 356, 365 unicolor, Dicladocera, 85 Xenopeltis, 374, 378, 381, 387, 388, 401 urens, Mucuna, 275, 282 urmana, Apathya cappadocica, 475, 481 Uromastix, 453, 482 asmussi, 475, 481 loricatus, 453, 455, 475, 481, 485 microlepis, 475, 481 365 vanzolinii, Amphisbaena, 613 Varanidae, 455, 475 Varanus, 455, 482 eriseus, 455, 475, 480 monitor, 475, 480 VoL. XXXII] variegatus, Eumeces schneideri, 476 Zonocerus, 234 vegetus, Lineus, 529 velox persica, Eremias, 476 strauchi, Eremias, 476 venenata, Rhabdotylus, 87 ventriosus, Heliofugus, 160 Heliofugus (Heliofugus), 144 Heliofugus ventriosus, 144, 166 nancaguensis, Heliofugus, 145 ventriosus, Heliofugus, 144, 166 ventromaculatus, Coluber, 466, 477, 480 Vepriella, 72 (Vepriella) frontalis, Mesomyia, 72 Veprius, 71, 72 (Coracella) , 72 presbiter, 71, 100 veriformis grayi, Calamaria, 374, 379, 393, 394, 404, 409 vermicularis, Amphisbaena, 615 Typhlops, 477, 480 vermiformis, Calamaria, 373 vernalis, Williamia, 111, 115, 117, 118, 119 verrilli, Micrura, 516, 518, 519, 532, 533 versicolor, Aspergillus, 216, 262 Calotes, 475, 480, 482 Lyngbya, 266, 268 vertebratis notospilus, Oligodon, 382, 393, 394, 401, 411 Oligodon, 374 vibakari, Natrix, 65 viduus, Tabanus, 95 violacea, Amphithrix, 265, 302 violaceo-fuscus, Asphergillus, 262 Vipera, 483 lebetina, 478, 480 raddei, 478 renardi, 478, 480 Viperidae, 471, 478 virescens, Amphiporus, 541 Zygonemertes, 517, 519, 540, 541 virgata, Hugelia, 36 389, SAR S83 virginea, Acmaea, 103, 107 viride, Emplectonema, 534, 535 Trichoderma, 263 viridipes, Phymateus (Phymateus), 233 viridis arabicus, Bufo, 473 Bufo, 329, 330, 331, 428, 430, 479, 481 Bufo viridis, 473 Lacerta, 476 INDEX 731 Lineus, 527, 528 viridis, Bufo, 473 viridiventris, Rhabdotylus, 87 visayanum, Oxyrhabdium 398, 399, 409 Rana macrodon, 387 vitulina, Phoca, 604 viviparus, Sebastes, 577 volgensis, Alosa kessleri, 633 vulcanica, Ramiellona, 214, 216 vulcanicus, Howascolex (Graceevelynia) tecumumamil, 216 vulgaris Loligo, 674 vulpes, Pelecorrhynchus, 99 leporinum, 384, wagleri, Trimeresurus, 374, 378, 379, 389, 391, 393, 398, 401, 404 Walterinnesia, 470, 482 aegyptia, 470, 471, 478, 481 Waltheria, 285 indica, 276 watsonana, Eremias guttulata, 427, 460, 461, 462, 476, 484 Eremias (Mesalina), 460 Williamia peltoides, 115 vernalis, 111, 115, 117, 118, 119 Williams, Francis X., The wasps of the genus __ Plenoculus (Hymenoptera: Sphecidae, Larrinae), 1-49 wilsoni, Atractaspis, 470 Lineus, 533 Micrura, 516, 518, 533 Typhlops, 477 Wislezenia, 16 refracta, 16 wrighti, Eriogonum, 36 329, 337, xanthopleura, Pelecorrhynchus, 99 Xantocles, 591 xenium, Stenotabanus, 77 Xenopeltis unicolor, 374, 378, 381, 387, 388, 401 xipe, Tabanus, 93 Xylophaga dorsalis, 112 Xylophis, 373, 375, 392 yaito, Euthynnus, 634, 675 Yoldia cooperi, 551 Yonge, C. M., Further observation on Hip- ponix antiquatus with notes on North 732 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Pacific pulmonate limpets, 111-119 zephyreus, Astroscopus, 632 Mantle cavity, habits and habitat in zeylanica, Chara, 268, 303 the blind limpet, Middendorff, Lepeta Zirfaea, 342 concentrica Middendorff, 103-110 pilsbryi, 344 zonifer, Erilepis, 567, 569, 572, 579, 581, 597 Zonocerus elegans, 234: variegatus, 234 Zostera, 270 marina, 303, 518 marina latifolia, 270 zunasi, Harengula, 633 zygaena, Sphyrna, 632 Zygeupolia rubens, 552 Zygnopsis brevipes, 464 Zygonemertes, 517, 540, 541 albida, 541 capensis, 540 Zalophus californianus, 603 zamboangensis, Calamaria, 374, 393, 394, 409 Zamenis rhodorachis, 465 Zaniolepididae, 591, 592, 597 Zaniolepis, 570, 571, 576, 580, 584, 591 frenata, 567, 579 latipinnis, 567, 579 Zaocys, 396 carinatus, 374, 378, 401 luzonensis, 374, 389, 391, 403 zarudnyi, Cyrtodactylus, 438, 474, 482 Diplometopon, 456, 475, 481 glandulosa, 540 Eumeces, 476, 482 thalassina, 541 Testudo, 474 virescens, 517, 519, 54.0, 541 ERRATA Page 13. Line 4 from top: for davis read davisi. Page 115. Line 2 from bottom: for A. thersites read S. thersites. Page 119. Line 8 from top: for Szphonaris read Siphonaria. Page 138. Line 12 from bottom (caption under figure): for barossi read barrosi. Page 144. Line 1 from top (caption under figure): for barossi read barrosi. Page 155. Line 7 from top (caption under figure): for bibioensis read biobioensis. Page 158. Line 4 from bottom: for H. sulcatus read H. sulcatulus. Page 160. Line 6 from top: for barossi read barrosi. Page following 160, after running head, add 161 Page 374. Line 8 from bottom: for bivattatus read bivittatus. Page 391. Line 2 from top: for tesselatus read tessellatus. Line 3 from top: for philippinus read philippina. Page 402. Line 2 from bottom: for grasina read prasina. Page 409. Line 20 from top: for Ntrix read Natrix. Page 438. Lines 22, 23 from top: for C. kermanensis read C. kirmanensis. Page 440. Line 17 from bottom: for Peters (italics) read Peters (roman). Page 450. At top of page add “450” and running head California Academy of Sciences [ Proc. 4th Ser. Page 451. At top of page add Vol. XXXT] running head Anderson: Iranian Herpetology and 451.” Pages 503-505. These pages have been duplicated in nos. 17 and 18. Page 549. Between lines 13, 14 from bottom: insert as a heading, Genus Malacobdella Blainville, 1827. Page 597. After line 1 from top: insert “9, Cyclopteridae—Cont.” PROCEEDINGS OF THB California Academy of Sciences FOURTH SERIES Vol. XXXI, No. 1, pp. 1-49, 90 figs. July 8, 1960 THE WASPS OF THE GENUS PLENOCULUS (HYMENOPTERA: SPHECIDAE, LARRINAE) BY FRANCIS X. WILLIAMS Research Associate California Academy of Sciences SAN FRANCISCO PUBLISHED BY THE ACADEMY 1960 COMMITTEE ON PUBLICATION Dr. Ropert C. Mituer, Chairman Dr. LEo G. HERTLEIN Dr. Epwarp L. Kesseu, Editor Ria fie ng! ms i it =i Py a au haa) a 4 a | di} é sat > BL/WHOL LIBRAR