. F i \ i pe ae a ’ ne ia A A r yey i e Tei tr, SP APAMP HG ys ’ 7 ant Sy ‘pf Tye ea Ae aM a idee (il tid a a SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 O Nee,” whey te S800, wee Ceeeescee® * EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS CES RT VATIONS) RESEARCHES, AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN ”—SMITHSON (PusticaTion 3899) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION 1947 The Lord Baltimore Press BALTIMORE, MD., U, & A. ADVERTISEMENT The Smithsonian Miscellaneous Collections series contains all the publications of the Institution except the Annual Report, and occa- sional publications of a special nature. As the name of the series implies, its scope is not limited, and the volumes thus far issued relate to nearly every branch of science. Papers in the fields of biology, geology, anthropology, and astrophysics have predominated. A. WETMORE, Secretary of the Smithsonian Institution. (iii) an 6. N 10. Lee 13. 14. CONTENTS WermorE, ALEXANDER. The birds of San José and Pedro Gonzalez Islands, Republic of Panama. 60 pp., 4 pls. Aug. 4, 1946. (Publ. 3845.) Ertanson, C. O. The vegetation of San José Island, Republic of Panama. 12 pp., 2 pls., 1 fig. July 18, 1946. (Publ. 3846.) HitpEBRAND, SAMUEL F. A list of fresh-water fishes from San José Island, Pearl Islands, Panama. 3 pp. June 10, 1946. (Publ. 3847.) Cocuran, Doris M. Notes on the herpetology of the Pearl Islands, Panama. 8 pp. June 24, 1946. (Publ. 3848.) Crark, Austin H. Echinoderms from the Pearl Islands, Bay of Panama, with a revision of the Pacific species of the genus Encope. 11 pp., 4 pls. July 18, 1946. (Publ. 3849.) Morrison, J. P. E. The nonmarine mollusks of San José Island, with notes on those of Pedro Gonzalez Island, Pearl Islands, Panama. 49 pp., 3 pls. Sept. 12, 1946. (Publ. 3850.) Ketiocc, Remrncton. Mammals of San José Island, Bay of Panama. 4 pp. July 18, 1946. (Publ. 3851.) Scumipt, Kart PatrEerson. Turtles collected by the Smith- sonian Biological Survey of the Panama Canal Zone. 9 pp., I pl. Aug. 1, 1946. (Publ. 3852.) Witson, Mitprep Stratton. The species of Platycopia Sars (Copepoda, Calanoida). 16 pp., 2 figs. Aug. 23, 1946. (Publ. 3853.) Stewart, T. D. A reexamination of the fossil human skeletal remains from Melbourne, Florida. With further data on the Vero skull. 28 pp., 8 pls., 7 figs. Aug. 9, 1946. (Publ. 3854.) Cuapin, Epwarp A. Review of the New World species of Hippodamia Dejean (Coleoptera: Coccinellidae). 39 pp., 22 pls. June 14, 1946. (Publ. 3855.) . Deicnan, H. G. Descriptions of two new leafbirds from Siam. 3 pp. June 24, 1946. (Publ. 3856.) GILMoRE, CHarLEs W. A new carnivorous dinosaur from the Lance formation of Montana. 19 pp., 4 pls. Sept. 12, 1946. (Publ. 3857.) Rivas, Luis Rene. A new dussumieriid fish of the genus Jenkinsia from Bermuda. 4 pp., 1 pl. 1 fig. Nov. 22, 1946. (Publ. 3859.) (v) V1 Lon to bo SMITHSONIAN MISCELLANEOUS COLLECTIONS Dirxe, G. H. Ladybeetles of the genus Epilachna (sens. lat.) in Asia, Europe, and Australia. 183 pp., 27 pls., 6 figs. Jan. 20, 1947. (Publ. 3860.) WETMORE, ALEXANDER. New birds from Colombia. 14 pp. Dec. 20,1946. ACE ubl.’ 38622) Kwnicut, J. Brookes. Some new Cambrian bellerophont gastro- pods: i pp: .2.pls.! Jan, 3, 1947, (CPubleséo5.) Heperetu, Jorr. W. On the evolutionary significance of the Pycnogonida. 53 pp., I pl., 16 figs. Mar. 24, 1947. (Publ. 3860. ) Reese, Arsert M. The lamina terminalis and preoptic recess in Amphibia. QO pp., 4 pls. Jan. 27, 19047.) (Publ) 386073) 3ARTSCH, PAuL. A monograph of the West Atlantic mollusks of the family Aclididae: 29 pp., 6 pls: Feb. 24, 1647;¢@2nne 3868. ) . WEINTRAUB, Rospert L., AND PRICE, LEONARD. Developmental physiology of the grass seedling. II. Inhibition of mesocotyl elongation in various grasses by red and by violet light. 15 pp., 5 figs. May 8, 1947. (Publ. 3869.) . Crayton, HW. H. Solar cycles. 18 pp., 9 figs. Mar i5aanonee (Publ. 3870.) y 4 aS Len Ty AN oR ity tie Mak SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 1 THE BIRDS OF SAN JOSE AND PEDRO GONZALEZ - ISLANDS, REPUBLIC OF PANAMA (Wore Four PLares) Re ALEXANDER WETMORE Secretary, Smithsonian Institution (PUBLICATION 3845) GITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION - AUGUST 5, 1946 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 1 THE BIRDS OF SAN JOSE AND PEDRO GONZALEZ ISLANDS, REPUBLIC OF PANAMA (WitTH Four PLATES) BY ALEXANDER WETMORE Secretary, Smithsonian Institution (PUBLICATION 3845) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION AUGUST 5, 1946 Bee sy WL ee SS Be Oo gt) ARS ay WAM EK Th eG Pees » te Alene The Lord Baltimore Press BALTIMORE, MD., U. 8. A. THE BIRDS OF SAN JOSE AND PEDRO GONZALEZ ISLANDS, REPUBLIC OF PANAMA By ALEXANDER WETMORE Secretary, Smithsonian Institution WitH Four PLATES Until recently the natural history of the Archipiélago de las Perlas in the Gulf of Panama has been known mainly from investigations on the largest island, Isla El Rey, called San Miguel by early collectors from its principal settlement, and on various others of the northern islands of the group. Isla San José, prior to 1944, had been visited by naturalists casually, so that only parts of its fauna and flora have been known. Its outlying position in the southwest of the group, distant from the other islands, its lack of settlements, and its forests without trails, have made the interior of San José difficult of access until the recent establishment there of a laboratory of the Chemical Warfare Service of the United States Army with roads and trails through which all sections of the island are now accessible. Army engineers came to San José late in January 1944, established a temporary camp back of Ensenada de la Bodega, and began roads and other construction work immediately. Accompanied by Dr. J. P. E. Morrison, of the division of mollusks, United States National Museum, I arrived on San José on February 7, 1944, to make studies of the fauna. I remained until March 14 when I returned to Balboa, C. Z., while Dr. Morrison continued our investigations until October 3. Our work centered on San José, and this report on the birds deals principally with that island. In view of the comparatively small amount of published information on Pedro Gonzalez, which lies im- mediately to the north, I have included also my notes made on that island during the days of March 9g and 11, as well as some data from later visits by Morrison. There are also a few notes on the birds of Moreno Island, lying off the western side of El Rey, which was visited by Morrison on May 109. I have to thank Brig. Gen. E. F. Bullene for his interest and as- sistance in the arrangements for our work, and am deeply indebted to other officers and personnel. Among these I must mention es- pecially Lt. Col. L. W. Glazebrook, who had charge of the arrange- SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, No. 1 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 ments for the installations at the camp in the beginning, Col. H. M. Woodward, Jr., Col. Robert D. McLeod, Jr., and Lt. Col. D. W. Leavens, all of whom were uniformly interested and helpful. Our early studies were aided especially by Russell Foster, of the Engineer Division of the War Department, in charge of construction on San José, and by William Langenbruner, his assistant. Our investigations centered on a survey of the principal land fauna of San José, including the mammals, birds, reptiles, amphibians, fishes, and the land and fresh-water mollusks. Reports on all these groups, as well as on certain echinoderms that we secured along the beaches, will be included in the series now under publication. In addition to the field studies just outlined, the Smithsonian Institution had the cooperation of the Bureau of Plant Industry of the United States Department of Agriculture, through Carl O. Erlan- son who made a series of ecological studies on the plants of San José. Mr. Erlanson reached San José April 6, 1945, and continued his work there until August 27, making detailed observations”on the vegetation. His investigations were forwarded and assisted extensively by the work of Dr. Ivan M. Johnston, of the Arnold Arboretum, Harvard University, who has made systematic studies and collections of the flora during three visits at the end of 1944, for about 6 weeks in March and April, 1945, and for a further period at the end of 1945 and the early part of 1946. Dr. Johnston has supplied names for many species of the plants prior to publication of his own report. The account by Mr. Erlanson published in this same volume of the Smithsonian Miscellaneous Collections gives a general description of the vegetation of the island. His plant specimens have been placed in Dr. Johnston’s hands for his use in the systematic account of the plants to be published elsewhere. NATURAL FEATURES San José, as noted, is the most southwestern of the islands of the Perlas group and is second in size, Isla El Rey being much larger. San José is of irregular shape, being roughly 74 miles long from north to south, and constricted to about a mile wide at the narrowest point near the center, the average width being 23 to 3 miles from east to west, with one extension of a little less than 14 miles on the east terminating in Punta Cruz. The principal bay on the west side is Ensenada de la Bodega (pl. 1, fig. 1) and on the east Ensenada Playa Grande. The shore line is rough for much of the circumference, rising abruptly in rocky bluffs from 20 to 50 feet high (pl. 1, fig. 1; pl. 3, NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 3 fig. 1). At Ensenada Playa Grande there are sandy beaches ex- tending for about 2 miles, with swamps at the mouth of the Rio Marina near the southwestern end. Small sand beaches are numerous at the heads of indentations along the shore, and there are a number of rocky islets that have been cut off by erosion from the main island. The largest of these are found at the southeast. The surface of San José is rolling, rising at the highest point to about 350 feet above sea level. The stream valleys are narrow, and in places steep-sided, with outcroppings of rock that form series of rapids and low falls. There are two streams of fair size, the larger being the Rio Marina (pl. 2, fig. 2), draining a considerable area in the central and northern part of the island. The Rio Mata Puerco, which cuts more than halfway across at the narrow central constriction already noted, is remarkable principally for the extent of its tidal channel, bordered by red mangroves that reach back for half a mile from the shore. There are numerous smaller streams. Water flowed to the sea in the larger of these throughout the dry season without dropping below a certain level, though smaller ones became dry. Almost half of the island is covered with gallery forest, the trees rising to 60 and 8o feet with occasional arboreal giants of greater size (pl. I, fig. 2; pl. 2, fig. 1). While there is considerable under- growth, more or less tangled with vines, travel with a fair amount of cutting was not too difficult. Extensive growths of spiny-trunked black palms in lowland areas offered one of the principal difficulties. These blocked passage in many places, since the tough wood fibers turned and battered the edge on the best of machetes. At the mouth of the Rio Marina there was an extensive swamp, with tidal channels cutting through it, where the trees were tall, and the muddy ground beneath completely open. The red mangrove swamp at the mouth of the Mata Puerco was fairly large, and there was another small swamp where we collected regularly back of the main beach at Ensenada de la Bodega. Near the sea, for the greater part of the circumference of the island, there were extensive areas of brush grown with creepers and vines so closely matted that they were completely impassible except by making trails with much hard labor. Mr. Erlanson, who investigated the plant ecology, is convinced that these represent old clearings that date back to some early period when there was agriculture on the island. One of the curious natural features is Bald Hill, a small savanna area in the north of the island. In several other sections, notably about mid- way between the Rio Marina and East Harbor, and in another section on the East Loop Road near where this joins the East Road, the 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 forest is composed of a few scattered trees with little or no under- growth. Small areas on the summits of some of the headlands were grown with grasses without shrubbery. The surrounding sea was shallow and supported an abundant marine life that, in turn, was attractive to many sea birds. My records for these were made on various launch trips. Pedro Gonzalez Island is third in size of the group, being less than 4 miles long by 1 to less than 2 miles wide. The north shore is very irregular with several indentations. Isla Trapiche (pl. 4, figs. 1 and 2) lies inside the northwestern point, and the village of Cocal (pl. 3, fig. 2) is situated on a low point looking across a bay to Trapiche and the sea beyond. The general conformation is like that of San José, the shore line being rocky and often precipitous, rising 30 to 60 feet or more from the sea. Sandy beaches, broken by ex- posures of rock, line the bays. There is an extensive beach at Cocal, and I noted others near the southern end in passing along the western side of the island. There is a small lagoon back of the seashore in the southwestern part of the island, but I did not have time to visit it. The interior of the island is rolling, with the highest point rising 390 feet above the sea. Pedro Gonzalez is forested (pl. 4, fig. 2), but much of the original growth has been cut to allow planting. These clearings were being extended in the usual wasteful manner as the older fields were invaded by tough grasses and other growth. ORNITHOLOGICAL INVESTIGATIONS The earliest note of ornithological collections on San José that I have found is that of Dr. Carl Bovallius, of the Royal University of Upsala, who, during a 2-month survey of the Pearl Islands in the spring of 1882, visited San José for 2 days or so at the end of April. He came down in a schooner from Pedro Gonzalez, landed in a cove, and followed a fair-sized stream inland. His decription ? was probably written some time later when some of the details of his observations had become obscure, since there is no river of the size he describes on the island. His collection of birds was identified by Hialmar Rendahl,? who lists three birds collected on San José on April 26 and 27, Buteo magnirostris alius, Buteogallus anthracinus subtilis, and Milvago chimachima cordatus. Ludlow Griscom and Maunsell S. Crosby secured 39 birds on San José from February 15 to 17, 1927, during a cruise through the Pearl 1Ymer, 1886, pp. 13-14. 2 Ark. for Zool., vol. 13, No. 4, 1920, p. I-56, I map. NOL I BIRDS OF SAN JOSE ISLAND—WET MORE 5 Islands and to eastern Panama, their specimens being in the Ameri- can Museum of Natural History. This was the first important gather- ing from the island, but was not reported in detail in publication. I have included pertinent records from this material through the courtesy of the American Museum of Natural History. Robert Cush- man Murphy in the launch Wilpet on September 9, 1937, traveled southward through the passage between San José and El Rey, re: cording several species of birds. In 1941, during his investigations on the schooner Askoy, he passed along the east and south coasts of San José on February 13, landing at Playa Grande in the afternoon. On February 16 he made observations along the east and north coasts, and on February 21 passed west and south of the island. Later, on May 23, he sailed along the southern and eastern shores of the island. He returned to the island from November 7 to 28, 1945, and at that time, working from the headquarters of the Chemical Warfare Service, added several migrant species to the previously known list of birds. These data, together with the records from his earlier visit, mainly of aquatic species, he has very liberally placed at my disposal for the present paper so that the list might be complete to date. I am much indebted to him for this assistance. There have been other observations but none of them extensive, and as far as I am aware nothing else has been published relating to the birds. Other naturalists among my friends who have visited San José include Thomas Barbour, James Zetek, and Gerrit S. Miller, Jr., who were interested in the main in other matters than the avifauna. I have not taken time to make an extended search for statements rela- tive to the island in older publications, but may note that I have seen reference to descriptions of scorpions, and mention of a few other natural-history items. Pedro Gonzalez has been a little better known, as a good many have come to the fishing club on Isla Trapiche, an islet cut off from Pedro Gonzalez by a shallow channel so narrow that Trapiche may be said to be almost a part of the adjacent island. Griscom and Crosby, mentioned above, secured 48 skins on Pedro Gonzalez February 18 and 19, 1927, but did not publish on them as a collection. A few of their records are included in the present report. Mrs. Bertha B. Sturgis includes a number of observations from Pedro Gonzalez in her Field Book of Birds of the Panama Canal Zone. My inclusion of this island in the present account has been mainly to record the notes that I made there personally on March 9 and 11, 1944, so that this paper relates almost wholly to San José. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 DISCUSSION OF THE AVIFAUNA The number of species of birds found in the whole of the Pearl Islands is remarkably small in comparison with the adjacent mainland. The resident birds of San José number between 40 and 50 species, uncertainty as to the actual number arising from lack of information as to the breeding of a number of aquatic or other wide-ranging species whose nesting colonies at present are not wholly known. Ac- tually I have listed 46 species as probably breeding, omitting the brown pelican, Colombian booby, Brazilian cormorant, little blue heron, egret, snowy heron, black-crowned night heron, black vulture, and turkey vulture as uncertain. These nine species nest nearby, and some may rear young on San José, but this has yet to be ascertained. It is probable that the gray-breasted martin, Progne chalybea chalybea, may establish colonies now that there is human settlement, but at present this bird is only a visitor. The turkey vultures seem to be casual visitors, as is the blue-headed parrot, Pionus menstruus. There is no form of bird, according to present understanding, that is restricted to San José Island alone. The nearest to this is the wood rail, Aramides cayennensis morrisoni, which is found also on Pedro Gonzalez, the birds from these two islands differing from Aramides c. latens of Isla El Rey and Viveros in being slightly darker colored. In identifying the small resident land birds I have been interested to note in several a tendency toward larger bills that is visible only on close comparison and that is too slight to be evident in series of mea- surements. The tendency, however, is not always peculiar to San José but relates in some cases to the archipelago as a whole. This character is slightly indicated in Formicivora grisea alticincta, M yi- ophobus fasciatus furfurosus, Elaenia chiriquensis chiriquensis, and Coereba flaveola cerinoclunis. It reaches a maximum in Crotophaga ani, in this case to an extent that more specimens should be collected for study. The heavy orange pigmentation of occasional specimens of the warbler Dendroica petechia aequinoctialis is also worthy of notice, occasional males being so deeply colored that the difference is apparent in life, while in the hand they are more richly orange than any individuals of the other races of this bird that I have seen. As San José and Pedro Gonzalez are the outliers of the Pearl Islands, in spite of their relatively considerable size there are a num- ber of species not known from them that are found on other islands, particularly on Isla El Rey (and Viveros, immediately adjacent) to the east and north and nearer the mainland. The more noteworthy of these are the following: a tinamou, Crypturellus soui panamensis ; a woodpecker, Centurus rubricapillus seductus; a black ant-shrike, NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 7 Cercomacra nigricans; and a wren, Thryophilus leucotis conditus. A seed-eater, Sporophila nigricollis nigricollis, is known from Saboga, the Agami heron, Agama agami, has been taken once on El Rey, the caracara, Polyborus cheriway cheriway, has been found on Pacheca, and a small kingfisher, Chloroceryle inda inda, was secured by Boval- lius on Viveros. Three forms that are known from Pedro Gonzalez have not been reported from San José, viz, a chachalaca, Ortalis garrula cinereiceps, a screech owl, Otus choliba luctisonus, and a barred ant-shrike, Thamnophilus doliatus nigricristatus. Only one specimen of the chachalaca is recorded from Pedro Gonzalez, a bird killed by a pearl diver and brought to El Rey to W. W. Brown, and there is only one record to date for the screech owl, which was taken by Bovallius. The native boy who helped me in my two visits to Cocal described a small owl to me, so that it may be more common than the single record indicates. The barred ant-shrike is common. San José Island has no resident land birds that are not known elsewhere in the archipelago. There is thus a regular diminution in _ the universally poor avifauna as regards number of species in pro- ceeding from the islands nearest the mainland on the north and east to San José which is the most distant to the southwest. MIGRANT SPECIES Of the species at present recorded from San José 42 are migrants from the north, assuming that the laughing gull, Larus atricilla, Cabot’s tern, Thalasseus sandvicensis acuflavidus, and gull-billed tern, Gelochelidon nilotica vanrossemi, come from the more northern colonies. Of these migrant species only the laughing gull, the spotted sandpiper, and the two races of the northern water-thrush are at all common, the others being found casually and in small number. The kingbird and dickcissel pass in migration in fair abundance but do not remain in winter. It is in the group of northern migrants that many additions to the list remain to be made, since there are numerous species that come through Panama that have not been reported as yet from San José. Richmond’s swift, Chaetura vauxi richmondi, is recorded at pres- ent from southern México to Costa Rica, the one that we secured being the only report at present south of that area. It must be considered an unusual visitor. The swallow-tailed kite was seen in northward migration in March 1944 but is not known at present to breed here. The turkey vultures offer ground for interesting speculation since the six specimens examined seem to belong to three distinct races. One 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 of these, Cathartes aura teter, is a migrant from western North America; a second, Cathartes aura aura, is the resident form of Panama. These two need no further comment except to state that the latter possibly may be found nesting. The third one, curiously enough, appears to be a representative of the form found in north- western South America which at present is called Cathartes aura ruficollis. This must be considered a wanderer. More material of these birds is necessary to determine their status. The gray-breasted martin, Progne chalybea chalybea, and the blue-headed parrot, or casanga, Pionus menstruus, while present in numbers for a brief season, appear to be wanderers from breeding grounds on the main- land. Such species as the black petrel, Loomelania melania, the flut- tering petrel, Halocyptena microsoma, and the skua, Catharacta skua chilensis, are wanderers over the ocean, to whose number there will be added other species with continued observation. The plumbeous kite, Jctinia plumbea, arrived as a migrant, pre- sumably from South America, on March 11, 1944, and immediately established itself on its nesting grounds. The annual movements of this species at present are not well understood. The yellow-green vireo was present in abundance on San José on our arrival early in February, but probably had not been there for long since this bird withdraws after its breeding season to South America. The period of its migration is not clearly known. ANNOTATED LIST OF*SPECIES In the following list I have given the data that I secured during my stay when San José Island was being developed for the work of the Chemical Warfare Service, with the addition of records made by Dr. Morrison, to present as clear a picture as possible of the kinds of birds found on the island, as well as their abundance, at the time when the island forests were first disturbed and the avifauna was still in its original state. These data will serve for future record in further studies that it is hoped to make from time to time on changes and shifts due to the establishment of clearings, the presence of considerable numbers of men, and also the effect of the experiments of the Chemical Warfare Service. Through the kind assistance of Dr. Robert Cushman Murphy I have been able to add the additional species that he has found, mainly migrants and sea birds, so as to male the list as complete as possible. His field observations in November 1945, which it is hoped he will report in such detail as may be necessary, indicate that some interesting changes had already taken place. 1 NO...2 BIRDS OF SAN JOSE ISLAND—WET MORE 9 Family PROCELLARIIDAE PUFFINUS GRISEUS (Gmelin): Sooty Shearwater Procellaria grisea GMELIN, Systema naturae, vol. I, pt. 2, 1780, p. 564 (New Zealand). Robert Cushman Murphy recorded this shearwater at sea near San José, February 21, 1941. The record is backed by specimens that he secured nearby during the Askoy expedition. Family HypDRroBATIDAE LOOMELANIA MELANIA (Bonaparte): Black Petrel Procellaria melania BONAPARTE, Compt. Rend. Acad. Sci. (Paris), vol. 38, No. 14, Apr. 3, 1854, p. 662 (near San Francisco, Calif.) .3 Dr. Murphy observed these petrels near San José February 13 and 21, and May 23, 1941. As we crossed from San José Island to Balboa March 14, black petrels appeared in midpassage, flying singly or in couples just above the water. Though they passed regularly in front of us they did not follow the wake of the boat. I saw the last one on this day about 10 miles off Taboguilla Island. OCEANODROMA TETHYS KELSALLI (Lowe): Peruvian Storm Petrel Thalassidroma tethys kelsalli Lowe, Bull. Brit. Orn. Club, vol. 46, Nov. 4, 1925, p. 6 (Pescadores Islands off Ancon, Pert). Robert Cushman Murphy recorded this petrel off San José Sep- tember 9, 1937, February 21 and May 23, 1941. Specimens collected nearby belong to this race so that the sight records are assumed to be the same. It is probably this bird that I saw at a distance in crossing to Balboa March 14, 1944. HALOCYPTENA MICROSOMA Coues: Least Petrel Halocyptena microsoma Cours, Proc. Acad. Nat. Sci. Philadelphia, June 30, 1864, p. 79 (San José del Cabo, Baja California). March 14 in midpassage from San José Island to Balboa I recorded half a dozen small petrels, uniformly dark in color with long, cuneate tails, which flew just above the surface of the water. Their wing motion was quicker and more fluttering and they moved more er- ratically than the black petrels about them. I was uncertain of their 3 For the use of the genus Loomelania instead of Oceanodroma for this species see Murphy, Oceanic birds of South America, vol. 2, 1936, pp. 726, 743. Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 identity at the time, but in subsequent investigation Dr. Robert Cushman Murphy showed me an excellent series of the least petrel from this same region, leaving no doubt as to the bird I had seen. He recorded it near San José May 23, 1941. Family PELECANIDAE PELECANUS OCCIDENTALIS CAROLINENSIS Gmelin: Eastern Brown Pelican Pelecanus carolinensis GMELIN, Systema naturae, vol. 1, pt. 2, 1780, p. 571 (Charleston Harbor, S. C.). Brown pelicans are common in the Gulf of Panama between the Pearl Islands and Balboa so that they were seen frequently during the crossing to and from San José on February 7 and March 14. At San José I saw them regularly in the small bays, and on those occa- sions when I was out in launches found them in numbers along the shores of the island. When schools of minnows (pl. 3, fig. 1) were about, sometimes as many as a hundred pelicans gathered, and several times I observed a bird with the pouch distended by small fishes which it gulped down as soon as the water had run off. At Pedro Gonzalez March g and 11, pelicans were even more abundant, so that they were continually passing, their shadows sweep- ing by me as I walked along the shore. Their constant movement remains as one of my clearest memories of my visit to this island. At various places along the steep shore line they had perches in the trees, the ground below being white with excrement. On San José I shot a male in breeding dress on February 26, and Morrison secured two females in nonbreeding plumage September 23 and 27. We also found a skull of a recently dead bird on the beach. Colombian and Venezuelan laborers called them pelicano. These birds of the Pearl Islands show some approach in darker hindneck in breeding dress, and in darker body coloration, to the race murphyi ** of western Colombia and Ecuador to the south, but are decidedly nearer to carolinensis. Family SULIDAE SULA NEBOUXII Milne-Edwards: Blue-footed Booby Sula nebouxit MILNE-Epwarps, Ann. Sci. Nat. Zool., vol. 13, art. 4, 1882, p. 37, pl. 14 (Chile). Robert Cushman Murphy recorded two of these boobies half a mile off the east coast of San José September 9, 1937. This is the 8a See Wetmore, Auk, vol. 62, 1945, p. 583. NOT BIRDS OF SAN JOSE ISLAND—WET MORE [I only record to date, which seems strange as the birds are common on some of the northern islands. SULA LEUCOGASTER ETESIACA Thayer and Bangs: Colombian Booby Sula etesiaca THAYER and Bancs, Bull. Mus. Comp. Zodl., vol. 46, June 1905, p. 92 (Gorgona Island, Colombia). To see the boobies at San José it is necessary to use a boat as they range offshore and only occasionally appear around the rocky headlands. They came especially along the eastern side of the island, flying and soaring steadily in the breeze from 2 to 20 feet above the waves. On March 4, in a run along the eastern side from the south end to East Bay, boobies passed every few minutes singly or in little groups of three or four. Others were observed March g and 11 in crossing to Pedro Gonzalez, and on March 14 I recorded half a dozen in the passage to Balboa, one of them within 8 or 10 miles of Taboguilla Island. Morrison collected one on June 8. The four taken show plumage change from the gray immature to the full adult. Two illustrate complete transition between these two stages, having varying mixtures of gray and white on the feathers of the abdomen. Family PHALACROCORACIDAE PHALACROCORAX OLIVACEUS OLIVACEUS (Humboldt): Brazilian Cormorant Pelecanus olivaceus HuMBotpt, in Humboldt and Bonpland, Recueil d’ Observa- tions Zoologie et d’ Anatomie Comparée, 1805, p. 6 (Magdalena River, lat. 8°55’ N., Colombia). The cormorant, known to the American sailors as the Panama duck, is one of the most prominent species of the sea birds about the islands, whose flocks and flights remain as one of my pleasant and interesting memories of my observations there. Usually I counted on one or two in any walk along the rocky headlands, and regularly I found large flocks whose presence seemed to be governed by the stage of the tide, and by the run of the schools of small fish that provided their food. Flocks, often numbering hundreds, came flying in early morning 300 to 500 feet above the sea, quartering about until fish were sighted, and then dropping precipitately to the surface to swim and dive among the plunging pelicans and boobies and the swooping man-o’-war birds and laughing gulls, while mackerel and amber-jack made the water boil in their surging rushes through the 12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 close-packed masses of small fishes (pl. 3, fig. 1). There was no fighting apparent among the participants in these relentless assaults, though sometimes I wondered if there were not occasional collisions that might result in serious injury, so apparently heedless were the many birds joined in the melee. At a distance the dark forms of cormorants suggested geese by their size, form, and method of flight. When not in search of food they rested on sandy beaches or on rocky headlands, standing close together in rows and columns, sometimes in the edge of the water. The mouths of streams were favorite places. Single birds rested anywhere along the rocks. On Pedro Gonzalez on March 9, and again on March 11, I found hundreds on the shore and in the water at the little village of Cocal. Flocks flew back and forth from these resting bands continually, with no regard for the people,* and in turn completely unnoticed by the inhabitants of the little settlement. Their passing shadows constantly attracted my eye as long as I was near the shore. In many places these birds frequent fresh or slightly brackish waters, so that it is interesting here to find them exclusively, and in large numbers, in the salt waters of the sea. As stated above, these cormorants appear black in the air, though the immature birds are brownish or grayish. An adult male prepared March 12 has a few small filaments of the white of the breeding plumes on the flanks and neck. This specimen has the following measurements, which are usual for the typical form of this bird: Wing 285, tail 168, culmen from base 63.5, tarsus 68 mm. A fully grown female in brownish immature dress collected by Morrison August II measures as follows: Wing 279, tail 180, culmen from base 62.0, tarsus 57.2 mm. Family FREGATIDAE FREGATA MAGNIFICENS Mathews: Man-o’-War bird Fregata magnificens Matuews, Austr. Av. Rec., vol. 2, Dec. 19, 1914, p. 120. (Barrington Island, Galapagos Islands). The angular, long-winged forms of frigate-birds appear constantly in the sky of San José, following the shore line, beating with slowly flapping wings over the sea if fish appear, or soaring, especially in afternoon when steady winds blow, across the rocky headlands. It seemed strange at times in the heavy jungle of the center of the island, with no visible indication that the ocean was near, to look out NOS tT BIRDS OF SAN JOSE ISLAND—_-WET MORE 13 through some opening among the trees to see a frigate-bird or two high overhead in passage across the island. These birds, here, were fishermen in their own right, swooping down over the waves to seize little fishes in their bills without alighting. It was interesting always to watch them descend swiftly from an elevation of 20 or 30 feet, glide forward just above the water, and then, dropping the head, grasp at a fish while the body continued its forward movement, seemingly leaving the head, attached by the long neck, behind to snap forward with a wriggling fish in the bill at the very second when it seemed as though the bird must over- turn. Bill and sometimes head might be buried in the crest of a wave, but neither wings nor body touched the water. When large schools of small fish appeared offshore, frigate-birds joined the peli- cans and cormorants in the swirling cloud of voracious predators that, in and out of the water, made constant attack on their unfor- tunate prey. Even under these circumstances the frigate-birds cap- tured their own supplies, and I saw no instance where they robbed other birds. It must be added, however, that boobies and terns, which I have observed to be the bullied companions who have had to give up their fish to the frigate-birds of another species, Fregata minor, farther west in the Pacific, here were widely scattered. February 13 and 16 we saw males flying with the distended red throat pouches that are developed fully only in the breeding season. All the frigate-birds that I saw closely enough to identify were the present species, marked in the adult male by the plain wing, in the female by the blackish throat, and in the immature by pure white head and neck. There should be careful watch for the related form Fregata minor known from nearby waters both north and south. In this species, regardless of geographic race, the male has a distinct brown band across the middle and lesser coverts of the wing, the female has the throat grayish (not black), and the immature shows a varying amount of light reddish brown on the white of head, neck, and upper breast. Family ARDEIDAE ARDEA HERODIAS Linnaeus: Great Blue Heron Ardea Herodias LinNaEus, Systema naturae, ed. 10, vol. 1, 1758, p. 143 (Hudson Bay). March 1 I saw one on the beach at the mouth of the Rio Marina. Robert Cushman Murphy recorded six or more during November 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106° 1945. These are supposed to be migrant individuals of the typical form from eastern North America. BUTORIDES VIRESCENS MARGARITOPHILUS Oberholser: Pearl Islands Green Heron Butorides virescens margaritophilus OBERHOLSER, Proc. U. S. Nat. Mus., vol. 42, Aug. 29, 1912, p. 553 (Isla El Rey, Archipiélago de las Perlas, Panama). An adult male taken March 1 has the following measurements: Wing 161, tail 51.8, culmen 60, tarsus 46.7 mm. The posterior part of the lower surface is distinctly dark in color, agreeing with a speci- men from Isla El Rey, the type locality. Another male, secured by Morrison April 18, measures as follows: Wing 170, tail 59.5, culmen 61.8, tarsus 48.3 mm.; and a female taken April 18 as follows: Wing 162, tail 57.3, culmen 57, tarsus 45.1 mm. Another female shot July 25 has the following dimensions: Wing 166, tail 62.7, culmen from base 58.5, tarsus 47.7 mm. The second male is especially dark below, being nearly uniform grayish slate, while the females are colored about as in the first male mentioned. These birds were found in small numbers in the mangroves at the mouth of the Rio Marina, and occasionally were seen in the woods along other small streams near where these entered the sea. The notes and mannerisms were those usual to these small herons. FLORIDA CAERULEA (Linnaeus): Little Blue Heron Ardea caerulea LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 143 (South Carolina). March 1 I recorded an adult bird in blue-gray dress and two others in white phase at the mouth of the Rio Marina. I had glimpses of other herons here from time to time that may have been this same species but did-not have opportunity to check their identity. Robert Cushman Murphy secured one in pied plumage in Navy Cove November 23, 1945, and recorded others, particularly along the lower Rio Marina and Playa Grande. LEUCOPHOYX THULA THULA (Molina): Snowy Egret Ardea Thula Mottna, Sagg. Stor. Nat. Chili, 1782, p. 235 (Chile). Robert Cushman Murphy recorded a few during November 1945 on San José, and November 26 collected a male that has the following measurements: Wing 248, tail 82.1, culmen from base 78.7, tarsus go mm. These small dimensions place it with the eastern race. NOL ft BIRDS OF SAN JOSE ISLAND—WET MORE I5 CASMERODIUS ALBUS EGRETTA (Gmelin): American Egret Ardea Egretta GMELIN, Systema naturae, vol. 1, pt. 2, 17890, p. 629 (Cayenne). On both San José and Pedro Gonzalez Islands egrets were fairly common, so that I saw them daily whenever I worked along the shores. They fed along the sandy beaches, in the mangroves at the mouths of streams, and at low tide ranged over the rocky shelves exposed along the headlands. It was common at all times to see them perched on the black rocks of the rougher sections of the shore line, where their white bodies showed clearly at a considerable distance. Morrison shot an immature male on Pedro Gonzalez July 21, and Dr. Murphy collected a male on San José November 21, 1945. NYCTICORAX NYCTICORAX HOACTLI (Gmelin): Black-crowned Night Heron Ardea Hoactli GMELIN, Systema naturae, vol. 1, pt. 2, 1789, p. 630 (Valley of México). An adult that I saw on February 21 at the mouth of the Rio Marina seems to be the first report of this species for the Pearl Islands. NYCTANASSA VIOLACEA CALIGINIS Wetmore: Panama Yellow-crowned Night Heron Nyctanassa violacea caliginis WeEtTMorE, Proc. Biol. Soc. Washington, vol. 59, March 11, 1946, p. 49 (San José Island, Archipiélago de las Perlas, Panama). On San José these herons were common in the mangrove swamps at the mouth of the Rio Marina, and were seen elsewhere along the eastern side of the island, sometimes in woods at the summits of the rocky bluffs. On Pedro Gonzalez they were common about the village of Cocal as there were swampy areas nearby, and here they walked about on the beaches in midmorning like any other herons. Morrison shot an immature bird on Moreno Island May 19. Our five other specimens were taken on San José February 20, March 22, and April 6 and 18, and on Pedro Gonzalez March 11. The allocation of the Pearl Islands birds has led to an extended review of available specimens of the species with results of interest. The characters of the other subspecies are summarized in the follow- ing synopsis. 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Nyctanassa violacea violacea (Linnaeus) : Ardea violacea LINNAEUS, Systema naturae, ed. 10, vol. 1, ‘1758, p. 143, (South Carolina). Characters —Adult, dark gray above and below, with the black central streaks on the dorsal feathers averaging wider; immature averaging dark; bill slender in adult and immature individuals. .Males (30 specimens from the eastern United States), wing 281- 300 (294), tail 102.0-118.7 (109.2), culmen from base 64.5-75.6 (70.9), depth of bill at nostril 19.0-21.9 (20.8), tarsus 93.6-106.2 (99.4) mm. Females (22 specimens from the eastern United States), wing 271- 305 (290), tail Io1.1-115.4 (107.8), culmen from base 64.2-75.3 (69.9), depth of bill at nostril 19.4-21.9 (20.8), tarsus 90.5-105.8 (97.1) mm. Range.—Breeding from southern Texas, southeastern Kansas, southern Illinois and Indiana, New Jersey and Massachusetts (rarely ) to South Carolina and Florida; probably to eastern México. In migration to the West Indies (specimens seen from Cuba, Jamaica, Hispaniola, Puerto Rico, Tortola, Sombrero, Barbuda, Montserrat, Guadeloupe, Mayer Island, Grenadines, Barbados, Swan Island, Great Corn Island), México ( Nayarit, Rio Coahuyana between Colima and Michoacan, Guerrero, Tres Marias Islands, Tamaulipas, Tlascala, Quintana Roo, Chiapas), and Central America (Corinto, Nicaragua ; Guanacaste, Costa Rica; Garray Creek, Permé, Bocas del Toro, Changuinola, and Darién, Panama). Remarks.—The slender bill is an excellent character for distinguish- ing individuals of this race in migrant status south of its breeding range where it enters the territory of the large-billed bancrofti and caligimis. The extent of its breeding range to the southwest is not now certain, but it seems probable that it nests in northeastern México, and that it may breed south through eastern México to the Caribbean coast of Central America. It is possible that one examined from Boca de Paila, Quintana Roo (February 2, 1926), and another from Rio Managua, Guatemala (January 31, 1895), represent resi- dent groups. Specimens seen from the west coast of México are unquestionably migrants as they are found in the same area as typical examples of bancroftt. Nyctanassa violacea bancroftt Huey: Nyctanassa violacea bancrofti Hury, Condor, vol. 29, May 15, 1927, p. 167 (Scammon Lagoon, Baja California). NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 17 Characters —Adult, paler gray above and below, dark dorsal streaks averaging narrower; immature averaging paler; bill heavier and deeper in both adult and immature. Adult males (28 specimens), wing 265-308 (286), tail 100.0-118.8 (110.7), culmen from base 64.6-79.0 (73.4), depth of bill at nostril 22.8-27.9 (24.5), tarsus 86.0-106.1 (92.8) mm. Adult females (16 specimens), wing 260-300 (278), tail 98.4-114.8 (106.3), culmen from base 62.1-77.2 (71.3), depth of bill at nostril 22.3-25.2 (23.5), tarsus 75.9-96.3 (89.0) mm. Range.—Breeding from central Baja California (lat. 28°31’ N.) south along the Pacific coast through western México (Maria Madre, Tres Marias Islands), and from the Bahamas (Stranger Key, Exuma, Andros, Great Abaco) through the Greater and Lesser Antilles to Tobago; specimens examined from Cuba (Preston, Guantanamo), Jamaica (Spanishtown, Trelawney), Hispaniola (Petit Port Il’Ecu and Tle a Vache, Haiti, Cafio Hondo and Samana Peninsula, Dominican Republic) Mona, Puerto Rico (Bayamon), Vieques, St. Thomas, Tortola, Antigua, St. Eustatius, Barbuda, Dominica (Bath, Luzon Park), St. Lucia, St. Vincent, Montserrat (Cass Bay), Grenadines (Carriacou, Mayreau, Little St. Vincent), Grenada (Levera Lake, Beausejour Lake, St. Andrews), and Tobago (Plymouth) ; Panama (Garray Creek, Almirante Bay). Remarks.—tThe resident West Indian group of yellow-crowned night herons was recognized as distinct by Bangs and Penard in 1918,* while Huey in 1927 pointed out that the birds of Baja Cali- fornia were different from those of the southeastern section of the United States. One of the wholly unexpected results of the present survey has been to find that the West Indian and Pacific coast groups are identical as far as I have been able to determine, so that both must bear the same name. Both are marked by heavy, swollen bills, and in the adult by paler gray color. The two populations seemingly are isolated from contact with one another. The name to apply to this race, with the two groups considered identical, has required some study. Bangs and Penard called the West Indian bird Nyctanassa violacea jamaicensis (Gmelin), from Ardea jamaicensis Gmelin, Systema naturae, vol. 1, pt. 2, 1789, p. 625, which is based entirely on Latham’s Jamaica night heron.® In the work indicated Latham treats first (p. 52) what he terms the “Night Heron Male,” which is the adult of Nycticorax nycticorax. This is 4 Bull. Mus. Comp. Zodl., vol. 62, April 1918, p. 31. 5 Latham, General synopsis of birds, vol. 3, pt. 1, 1785, pp. 54-55. 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I06 followed (p. 53) by an account of the “Night Heron Female,” based apparently on a bird in its second year, as the neck is said to be streaked, the under surface gray, and abdomen whitish, while above the markings, except on the neck and wings, seem to be largely absent. Then comes the “Jamaica Night Heron,” basis of Gmelin’s name, which Latham remarks he received from Jamaica “where it goes by the name of Clucking Hen..... Has a great affinity to the female Night Heron, but is larger.” The description is that of an immature night heron, but of which species it is impossible to determine with certainty. It may be noted, however, that the length of the bill is given as 4 inches (probably measured along the gape), which points toward the longer- and more slender-billed Nycticorar. It may be added that in Jamaica the clucking hen of the countrymen is the limpkin and not a night heron. Under these circumstances it seems necessary to discard Gmelin’s name jamaicensis as indeterminate, leaving Nyctanassa violacea bancrofti Huey available for the race here under discussion. Nyctanassa v. bancrofti wanders to some extent, as I have ex- amined one well-marked example with heavy bill (Carnegie Museum No. 103,973) from El Limon, Distrito Federal, Venezuela, taken by E. G. Holt, January 30, 1929, the depth of the bill at the nostril being 23.2 mm. The range south of western Mexico is uncertain. Van Rossem has said that the birds of El Salvador are darker, so that it may prove that these belong with caliginis. In the southern Lesser Antilles there is apparent tendency toward the South American race cayennensis, beginning in St. Vincent and continuing through Grenada and the Grenadines to Tobago, as some specimens show a shortened wing and a more slender bill. Nyctanassa violacea cayennensis (Gmelin) : Ardea cayennensis GMELIN, Systema naturae, vol. 1, pt. 2, 1780, p. 626 (Cayenne). Characters ——Very similar to N. v. violacea, but averaging darker, with dark streaks on dorsal feathers averaging narrower ; wing aver- aging very slightly shorter; bill slender as in violacea. Males (six specimens), wing 271-292 (284), tail 101.5-114.8 (109.7), culmen from base 68.4-73.8 (70.2), depth of bill at nostril 19.9-22.0 (21.0), tarsus 98.3-103.0 (99.8) mm. Females (five specimens), wing 263-288 (279), tail 97.4-107.8 (101.7), culmen 61.8-71.7 (67.2), depth of bill at nostril 20.3-21.0 (20.5), tarsus 92.7-99.2 (96.6) mm. NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 19 Range.—The north coast of Colombia (Bonda and Mamatoca, Depto. Magdalena; Lorica, Depto. Bolivar), Venezuela (Margarita Island, Corosal, Cariquito, Puerto La Cruz, near Mérida), and Trinidad (Caroni Swamp), to Surinam (Braamspunt, coast near Diana Creek) and northern and eastern Brazil. Remarks.—Few South American specimens are available at present in collections in the United States, and from these it appears that the groups of the southeastern United States (typical violacea) and of northern South America are remarkably similar. Both are marked by slender bill, leaving a slightly darker color and an average shorter wing as the only characters for the far distant South American bird. Some specimens cannot be separated on this basis, especially since many museum specimens are discolored by grease which darkens them decidedly. Birds seen from Trinidad seem identical with those of the South American mainland. Nyctanassa violacea gravirostris van Rossem: Nycticorax violacea gravirostris VAN RossEM, Occ. Pap. Mus. Zool. Louisiana State Univ., No. 15, Nov. 22, 1943, p. 266 (Socorro Island, Revilla Gigedo Group, México). Characters —Similar to N. v. violacea, but with brown wash on crown in adult darker and more extensive; bill strong and heavy; tarsus short ; average size smaller. Males (two specimens), wing 255-275 (265), tail 96.5-105.3 (100.9), culmen 64.7-68.8 (66.7), depth of bill at nostril 21.0-23.0 (22.0), tarsus 80.1-85.7 (82.9) mm. One adult, sex not marked, wing 264, tail 100.5, culmen from base 66.7, depth of bill at nostril 22.0, tarsus 82.9 mm. Range.—Socorro Island, Revilla Gigedo group, México. Remarks.—The curious adaptation by which this heron on Socorro Island has come to live in the dense, dry scrub that covers this hilly island, where no lagoons or marshes are available, has been mentioned by various naturalists. Apparently it is a habit of long standing, since it is reflected in the form of the birds, their differences from others being much more definite than can be indicated on paper. The short, heavy bill suggests that of bancroftit, while the darker color points toward pauper. The measurements of the three in the National Museum collection agree with the six examined by van Rossem in preparing his original description. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Nyctanassa violacea caliginis Wetmore : Characters.—Similar to N. v. violacea, but with thicker, heavier bill; adult decidedly darker gray: Similar to N. v. bancrofti in size of bill, but decidedly darker in color. Measurements——Males (12 specimens), wing 282-299 (290.1), tail 101.7-116.9 (108.7), culmen 67.6-81.3 (73.8), tarsus 87.7-101.3 (92.1), depth of bill at nostril 22.2-25.1 (23.4) mm. Females (2 specimens), wing 288-291 (289.5), tail I01.1-109.0 (105.0), culmen from base 73.7-74.7 (74.2), tarsus 96.5-97.4 (97.0), depth of bill at nostril 22.2-23.5 (22.9) mm. Range.—Panama (Balboa, Taboga Island, Obaldia) including the Pearl Islands (San José, El Rey, Saboga) to the Pacific coast of Colombia (Bahia de Cuevita) and Ecuador (Isla de Jambeli and Vaqueria). . Preliminary comparisons of our specimens taken on San José indi- cated that they were darker in color, but I was uncertain as to the stability of this character until I had examined the series in the Museum of Comparative Zodlogy from Isla El Rey (San Miguel) and Saboga, and found that they were likewise darker. It is inter- esting to observe that this tendency toward darker color is in the direction of the very dark N. v. pauper of the Galapagos Islands. Birds from the Pacific side of Panama likewise belong with the new race which extends along the mainland of the Pacific coast from Panama to Colombia and Ecuador. Nyctanassa violacea pauper (Sclater and Salvin) : Nycticorax pauper SCLATER and SAtvINn, Proc. Zool. Soc. London, 1870, p. 327 (Indefatigable Island, Galapagos Archipelago). Characters.—Darker than other races, in both adult and immature plumage, especially on lower surface ; crown in adult washed heavily with dark brown; size small. Males (15 specimens), wing 259-282 (271), tail 98.0-110.6 (103.8), culmen 64.0-73.7 (68.9), depth of bill at nostril 20.2-23.9 (21.8), tarsus 85.9-95.4 (90.3) mm. Females (4 specimens), wing 260-271 (265), tail 99.8-104.0 (101.9), culmen 63.8-70.8 (66.9), depth of bill at nostril 20.3-22.0 (21.1), tarsus 86.0-91.0 (88.2) mm. Range.—Known from all the principal islands of the Galapagos Archipelago except Culpepper and Wenman in the north. Remarks.—The dark color and small size make this the most dis- tinct of the forms of this bird. Like the bird of Socorro Island it NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 21 is often found in the brush inland, though common also about the lagoons. HETEROCNUS MEXICANUS (Swainson): Cabanis’ Tiger Bittern Tigrisoma mexicana SWAINSON, in Murray, Encycl. Geogr., July 1834, p. 1383 (México). The four specimens, three adult and one immature, shot on San José appear to average darker when examined with a series from México and Central America, if comparison is made with birds that are in the same color phase. These herons, however, show great variation. There appear to be two distinct phases in this bird, one that is grayish, with the brown of the under surface considerably lighter and the neck grayer, and one that is more brownish, where the under surface is decidedly more rufous brown and the neck has a heavier brownish wash. In some birds, probably younger indi- viduals, the breast is buffy and brown mixed with broad but in- distinct bars of neutral gray. This is apparently an influence from the heavily barred immature dress that may carry over in some individuals through the first molt toward adult plumage. The single immature specimen from San José, in the barred plumage usual to that stage, has the brown, especially of the dorsal surface, darker and more rufescent than most others seen. In fact in a considerable series I find no exact duplicate. Occasional speci- mens from elsewhere show the darker color in part .but none carry it throughout. A young bird, taken from the nest by Morrison March 24, has the pin feathers of the plumage just appearing. The down is very light grayish white, except that the filaments on the crown, which are longer than elsewhere on the body as is usual in herons, are pure white. Van Rossem ® calls attention to the fact that the earliest specific name for this bird, usually called Heterocnus cabanisi from Heine’s description in the Journal fur Ornithologie, 1859, p. 407, is mexicanus Swainson which dates from 1834. Van Rossem and Hachisuka have described a northern form? on two adult birds that are said to have the barring on the forepart of 8 Auk, 1942, p. 572. 7 Heterocnus cabanist (sic) fremitus van Rossem and Hachisuka, Proc. Biol. Soc. Washington, vol. 50, Sept. 30, 1037, p. 161 (Guiracoba, southern Sonora). 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 the body heavier, and the blackish streaks on the foreneck paler, while the size is said to be slightly larger, the wing being 365 mm. in the male and 375 in the female. These dimensions are definitely larger than those that the authors give for four adults from El Sal- vador. In the National Museum there is, however, an adult male from Los Reyes, Michoacan, with the wing 372 mm., a female from Hacienda Angostura, Rio Verde, San Luis Potosi, with the wing 365 mm., and a female from Chamelicon, Honduras, with the wing 305 mm. The alleged difference in size therefore is not definite. This is the most common heron on San José, a matter of some interest since earlier collectors did not obtain it on Isla El Rey. Mrs. Sturgis speaks of it as “fairly common on the Pearl Islands,” and Bovallius shot one on Bayoneta.® Occasionally I found tiger bitterns in heavy woods along the lower course of the Rio Marina, where they rested along the stream or flew up to perch in the trees 40 feet above the water, often uttering a harsh call. They were more common, however, along the sea, where they rested in low trees along the edge of the low cliffs, or on the rocks in more open locations. In habits they reminded me much of night herons, as they remained motionless for long periods, resting with neck drawn in when at ease, or with the head in the air when watchful. At such times the shortness of the legs in contrast with the length of the neck was very evident. Their appearance on the wing, both in method of flying and in shape, was especially similar to that of the black-crowned night heron. When perched in the par- tial concealment of low trees they often remained while I walked underneath and around them, but on the open rocks they were more wary, though here their colors blended with the background in such a way that often I did not see them until they flew. The native boys from Panama knew them as the grulla. A female taken February 9 was about to lay, and I saw one nest containing a small young on February 22. They appear to be solitary in breeding as the nests seen were placed singly, not associated with others. The usual location was on a tree limb projecting out from a cliff, where it was over water at least during high tide. The nests were flattened platforms of twigs somewhat bulkier than is often the case with herons. ’ 8 Rendahl, Ark. for Zool., vol. 13, No. 4, 1920, p. 17. NO. I BIRDS OF SAN JOSE ISLAND—WETMORE 23 Family THRESKIORNITHIDAE GUARA ALBA (Linnaeus): White Ibis Guara alba LINNAEUS, Systemae naturae, ed. 10, vol. 1, 1758, p. 145 (South Carolina). On February 18 I shot an adult female from a flock of a dozen resting on high limbs under the shade of the tall mangroves at the mouth of the Rio Mata Puerco. White ibises or their tracks were seen here regularly until the end of the month, and on March 1 I saw two at the mouth of the Rio Marina. Morrison shot a male April 30. On Pedro Gonzalez Rendahl ® lists one shot by Bovallius high up in dense forest, and Mrs. Sturgis ?° records a number seen on a wooded cliff. Family CATHARTIDAE CORAGYPS ATRATUS (Meyer): Black Vulture Vultur atratus Meyer, Zool. Annal., vol. 1, 1794, p. 290 (St. Johns River, » UPI). Black vultures were few in number on San José Island and were restricted mainly to the beaches in search for food since these were the only natural open places except for the small savanna on Bald Hill in the north. They were observed regularly but usually not more than two or three together, and seemed rather wild, probably because there have been no houses here. While occasionally one or two passed the camp area they paid no attention to it because of the careful refuse disposal. On Pedro Gonzalez on March g and 11 I recorded several at the village of Cocal. CATHARTES AURA (Linnaeus): Turkey Vulture Vultur Aura LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 86 (Veracruz, México). Turkey vultures, while not abundant in 1944, were recorded on most of the days that I was afield on San José, being seen regularly over the shore where they followed along the beaches, and also ob- served constantly soaring over the forested interior of the island. I found them on Pedro Gonzalez March 11. Because of their rela- tively small number I did not have opportunity to shoot birds for ® Ark. for Zool., vol. 13, No. 4, 1920, p. 18. 10 Field book of birds of the Panama Canal Zone, 1928, p. 79. 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 specimens until February 12. On this day, on the beach at Ensenada de la Bodega, I dropped one at long range. This one I laid on the sand with wings partly spread, when it served immediately to decoy down a companion which I also shot. Through the summer and fall Morrison observed turkey vultures regularly, and on May 12 secured a female that, while fully grown in every way, is shown to be a bird of the year by the dark-colored bill and the considerable amount of down on the back of the head and hindneck. These three proved so interesting that at my suggestion Dr. Murphy collected three more on November 24, 1945, which give much better understanding of the occurrence of forms of this bird on the island. The two that I obtained in February prove to be representative of two races. The first to be noted, a male, has the wing 485 mm., and the tail 245 mm., and is obviously of the typical form Cathartes aura aura (Linnaeus). A male (wing 470 mm.) and a female (wing 480 mm.) obtained by Dr. Murphy belong also to this race. When skinning the two secured February 12 I noticed that one was decidedly heavier and larger in body than the other. And on examination in the Museum it develops that the larger specimen, a female, is the western turkey vulture, Cathartes aura teter Fried- mann.'? It has the wing 492 mm., and the tail 257 mm. A female collected by Dr. Murphy with the wing 511 mm. and the tail 264 mm. is also of this race. The longer tail and the more grayish shade of the brown edgings on the wing coverts are the characters that mark teter from typical aura. It has been known for some time that there is migration among the turkey vultures through southern México and Central America, but because of the failure of most collectors to prepare skins there is not much recorded concerning their races. The western turkey vulture, C. a. teter, has been recorded once before from Panama by Aldrich and Bole, who secured one at Paracoté on the Azuero Peninsula.” The specimen taken by Morrison May 12 is obviously distinct from the other two, and after comparison with available skins it is identi- fied as of the race found in the northern part of the continent of South America, Cathartes aura ruficollis Spix.’* This race is dis- tinctly blacker than the three known from North America, and in addition has the lighter margins on the wing coverts much reduced 11 Cathartes aura teter Friedmann, Proc. Biol. Soc. Washington, vol. 46, Oct. 26, 1933, p. 188 (Riverside, Calif.). 12 Sci. Publ. Cleveland Mus. Nat. Hist., vol. 7, Aug. 31, 1937, pp. 40-41. 13 Cathartes ruficollis Spix, Avium Spec. Nov. Brasiliam, vol. 1, 1824, p. 2 (Interior of Baia and Piauhy, Brazil). NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 25 and very much darker brown. Although immature birds of the North American subspecies are blacker than the adults, none of them ap- proach the South American birds in depth of color, while in addition the edgings of the wing coverts are much lighter. The San José specimen, a female, has the wing 495 mm. and the tail 257 mm. The separation of these few specimens into so many subspecies is somewhat startling, and the identifications have been made only after careful and detailed study. It appears that there is far more move- ment among these vultures than has been known. The extent to which the South American individuals wander will be an interesting matter to ascertain, and there should be check made to determine more clearly their characters, as it seems probable from what I have seen in various visits to the range of what is known as ruficollis that the population now bearing this name will need to be further divided. Family ACCIPITRIDAE ELANOIDES FORFICATUS (Linnaeus): Swallow-tailed Kite Falco forficatus LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 89 (South Carolina). On March 13, over an area of tangled vines and brush, two swallow-tailed kites turned in graceful circles with widely spread forked tails. As they passed on, a flock of six appeared briefly and then continued over the forest. Evidently these were migrants travel- ing northward. As the one that I shot fell at a distance where I could not find it, I was not able to ascertain which subspecies was repre- sented. The species has not been recorded previously from the Pearl Islands. ICTINIA PLUMBEA (Gmelin): Plumbeous Kite Falco plumbeus GMELIN, Systema naturae, vol. I, pt. 1, 1788, p. 283 (Cayenne). The plumbeous kite is evidently a migrant in these islands, as it is elsewhere through much if not all of the northern part of its range outside South America. The first recorded were seen March 11 circling over the forested slopes of Pedro Gonzalez Island. The following day I noted three flying high overhead near East Harbor on San José, and on March 13 I shot a pair near the center of the latter island. These two were mating, so that they had evidently arrived on their nesting grounds in readiness to breed. The female, ready to produce eggs, was extremely fat, the male, with testes fully developed, only moderately so. That these were newly arrived migrants from some other region was easily apparent as they are 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 conspicuous and could not have been overlooked, since I had been in the field daily on San José for several weeks. They soar and circle swiftly and gracefully, spending much time in the air. On the wing the adults appear entirely dark until, when they turn, the sun reflects from the lighter-colored gray head so that it sometimes appears almost white. The wings are long like those of a falcon, and the slight notch in the tail is usually evident. In the hand the dull gray body and the black wings and tail form a pleasing contrast, the wings showing bright brown inner webs on the longer primaries. Three of the outer primaries are notched on the inner web. At rest the birds are partial to the open branches of the guarumo (Cecropia) and similar exposed perches. Morrison reports that this kite remained fairly common through the summer, at least until the latter part of September. He shot females April 23 and June 27, and also prepared one juvenile bird, with wings and tail less than one-third grown, with the light breast heavily streaked. This had been taken from a nest when in white down and kept alive for 3 weeks or so by one of our friends. Robert Cushman Murphy recorded one November 12, 1945. Dr. George M. Sutton ** in an instructive study of these kites has expressed the opinion that the Mississippi kite, Ictinia misisippiensis, and the plumbeous kite, J. plumbea, should be rated as geographic races of a single form. That the two are closely allied is evident on the most casual examination, but after careful study of a consid- erable number of birds I find that I cannot agree with my friend in this matter, since to my understanding the two groups are sufficiently different to be held as separate species. In these interesting birds the criteria for distinction appear mainly in the immature plumage, which should show the greatest resemblance if the two were in reality closely allied. In the adult stage, through the parallelism of the general gray, gray-black, and black color, only slightly relieved by touches of white and of brown, little concrete difference is evident. Turning then to the immature individuals we find that in the Mis- sissippi kite the prevailing streaking of the lower surface is decidedly brown, in the darkest the feathers being only partly edged with Chaetura brown, while the brighest are much lighter. The throat is lightly or not at all marked, and on the grayer dorsal surface there is much concealed white on the inner wing coverts, scapulars, and back. In the plumbeous kite, on the contrary, the streaks are deep, 14 Wilson Bull., vol. 56, 1944, pp. 3-8. NO?! T4108 BIRDS OF SAN JOSE ISLAND—WETMORE 27 dark, or blackish mouse gray, without brown, the throat is heavily streaked, and there is little or no concealed white on the very black dorsal surface. The two are so completely distinct that any specimen may -be identified as to its kind at a glance across the work table. The differences are not those used as modern criteria for subspecies, and the two are seen to be readily and entirely separate. The close alliance between the two kites is shown by their allocation in the same genus in which no others are included. In this genus I con- sider that they should be maintained as distinct though closely allied species. The immature birds of both have the under side of the tail barred with white. In the adult plumbeous kite this mark always remains, while in the full adult of the Mississippi kite it is lacking. In those occasional individuals of the latter in which the white tail bar’ is found, this is due to a juvenile influence that usually is evident else- where in the plumage, often in a mottling of the inner webs of the flight feathers, and in barring or spotting of the under wing-coverts. BUTEO MAGNIROSTRIS ALIUS (Peters and Griscom): Pearl Islands Hawk Rupormis magnirostris alia PETERS and Griscom, Proc. New England Zool. Club, vol. 11, Aug. 30, 1920, p. 48 (San Miguel, Isla El Rey, Archipiélago de las Perlas.) The large-billed hawk, well known in a variety of geographic races throughout much of tropical America, was fairly common on San José Island, where we secured specimens on February 14, March 3 and 6, April 14 and 16, and May 1. March 11 I heard one calling on Pedro Gonzalez but did not collect it. Though these hawks seemed to be fairly uniformly distributed through the forests of San José, both near streams and away from them, it was somewhat difficult to ascertain their true abundance since it was their habit to remain quiet much of the time. As they perched in the upper branches of the trees they were usually hidden by leaves so that I observed them ordinarily only when I heard their querulous, squealing cries, or when the yellow-green vireos were scolding them. Occasionally one of the hawks came when I was squeaking to call other birds. They were sometimes seen resting on open perches around the larger clear- ings. On February 15 I saw one calling near a partly completed nest of sticks placed in a tree 40 feet above the Rio Marina. The specimens from San José agree closely with those of Rey Island. The race, while not strongly characterized, is sufficiently distinct to merit recognition. 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 BUTEOGALLUS ANTHRACINUS SUBTILIS (Thayer and Bangs): Mangrove Black Hawk Urubitinga subtilis THAYER and Banes, Bull. Mus. Comp. Zodl., vol. 46, June 1905, p. 94 (Gorgona Island, Colombia). On San José Island these hawks were fairly common. They were seen mainly along the coast in the borders of small mangrove swamps, or on the rocky headlands above the beaches. Occasionally I found them along the streams farther inland. They were usually unafraid so that undoubtedly I often missed seeing them when they were perched in the heavy forest. A pair came frequently to circle in the rising air currents back of the coastal bluffs at camp, maintain- ing position with skill and certainty. Sometimes I saw them stand- ing on gravel bars along the lower course of the Rio Marina, and on other occasions resting on prominent rocks on the seashore. One hot day when I stopped to rest for a moment beneath a spreading tree at the border of a mangrove swamp I found that I was sharing the cool comfort of this shelter with one of these hawks which remained without fear 50 feet away. f lebruary 11 I shot a female nearly in adult dress that decoyed to me in a growth of white mangroves, and on February 13 secured a male in immature dress in a similar location. I saw several on Pedro Gonzalez Island March 11. The male has the wing 322 mm., the female 352 mm. Dr. Murphy secured a male November 19, 1945, with the wing 342 mm. The present form is certainly only a geographic race of Buteogallus anthracinus, separated mainly by smaller size. As far as I could tell all those recorded were this small race, though the true status of this bird should be checked with more specimens. Thayer and Bangs ** recorded these hawks from Isla El Rey under the name of the larger anthracinus, while Swann later described these specimens under the name Urubitinga anthracina bangsi.1® Griscom 17 detected these errors and reported subtilis from the Pearl Islands. Rendahl ** lists adult females taken by Bovallius on Bayoneta April 19, with the wing 347 mm., and on San José April 26, with the wing 341 mm., under the name anthracina. From his measure- ments these two birds are certainly subtilis. This race therefore is the only one recorded now from the Pearl Islands. 15 Bull. Mus. Comp. Zodl., vol. 46, September 1905, p. 144. 16 Syn. Accipitres, pt. 1, Sept. 28, 1921, p. 08. 17 Bull. Mus. Comp. Zodl., vol. 78, April 1935, p. 300. 18 Ark. for Zool., vol. 13, No. 4, 1920, p. 20. NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 29 The adult in life appears black with a prominent white band across the tail. The immature is streaked irregularly with white and buffy white below. Family PANDIONIDAE PANDION HALIAETUS CAROLINENSIS (Gmelin): Osprey Falco haliaétos carolinensis GMELIN, Systema naturae, vol. 1, pt. 1, 1788, p. 263 (South Carolina). The osprey, a winter migrant from the north, was seen February 21 at the mouth of the Rio Marina, and on February 21 and 26 along” the shore at Ensenada de la Bodega. Dr. Murphy recorded it at intervals during November 1945, once seeing two together. These are the first reports of this bird for the Pearl Islands. Family FALCONIDAE MILVAGO CHIMACHIMA CORDATUS Bangs and Penard: Yellow-headed Caracara Milvago chimachima cordata BANGs and PENARD, Bull. Mus. Comp. Zodl., vol. 62, April 1918, p. 35 (Isla El Rey, Archipiélago de las Perlas, Panama). These small hawks, known to the native boys as aguirre, were only fairly common. On San José we secured specimens February 11, March 3, June 1, and September 23. On Pedro Gonzalez I saw one March g and another on March 11, and Morrison collected two there June 8. I found them sometimes along the beaches half flying, half running actively across the sand to capture crabs. On other occa- sions I noted them inland near mangrove swamps along the roads, or in other places where the forest was not too heavy. Immature birds with streaked breasts were taken June 8 and September 23. From a very limited series these Pearl Islands birds seem to average slightly deeper buff on head and breast than those from northern South America, but this does not appear a wholly definite difference, especially since this color is certainly affected by fading. FILCO PEREGRINUS ANATUM Bonaparte: Duck Hawk Falco Anatum Bonaparte, Geogr. and Comp. List, 1838, p. 4 (Egg Harbor, Brel) Robert Cushman Murphy recorded one resting on a rocky islet near the northern coast of Ensenada de la Bodega on San José on November 20, 1945. 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 FALCO COLUMBARIUS COLUMBARIUS Linnaeus: Eastern Pigeon Hawk Falco columbarius LINNAEUS, Systema naturae, ed. I0, vol. I, 1758, p. 90 (South Carolina). On February 9 on my first visit to the forest along the Rio Marina on San José Island one of these falcons knocked down an adult pale- vented pigeon in flight at my feet, striking its quarry only Io feet from me. Instantly it saw me and rose to a branch where I shot it. The pigeon, actually larger than its attacker, lay bleeding under the wing, with most of its tail feathers torn out from this one blow. As I returned after retrieving the hawk, the other bird recovered and flew away. The pigeon hawk is a winter migrant from the north that has not been recorded previously in the Pearl Islands. FALCO ALBIGULARIS ALBIGULARIS Daudin: Bat Falcon Falco albigularis DAupin, Traité d’ ornithologie, vol. 2, 1800, p. 131 (Cayenne). Though found regularly on the mainland, this is the first report of this small falcon in the Pearl Islands. Morrison secured a pair on May 12, a male on July 6, and saw several others during the summer. The two taken May 12 had a nest in a hole in a tree about 40 feet from the ground with two small young covered with pure white down. After 2 days in captivity they died and were preserved in alcohol. One of them when first taken regurgitated feathers of the yellow honey-creeper. Apparently the bat falcon is fairly common on San José, but is difficult to find because of the heavy forest and the sedentary habits of the bird. FALCO SPARVERIUS Linnaeus: Sparrow Hawk Falco sparverius LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758 p. 90 (South Carolina). One bird was seen over the clearing where we lived February 27, and March 6, 7, and 14, possibly the same individual. There is no question but that this was a migrant from the north. Dr. Murphy recorded the sparrow hawk frequently in November 1945. It has not been recorded previously from these islands. Family RALLIDAE AMAUROLIMNAS CONCOLOR GAUTEMALENSIS Lawrence: Uniform Rail Corethrura Gautemalensis LAWRENCE, Proc. Acad. Nat. Sci. Philadelphia, vol. 15, June 1863, p. 106 (Guatemala). NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 31 The only specimen is one secured by Morrison August 23. This bird resembles skins from México and Central America in color and size. Measurements are as follows: Wing 113.6, tail 45, culmen from base 26.4, tarsus 41.4 mm. Dr. Murphy, in November 1945, recorded them frequently running across the roads and trails. It seems strange that this little-known species should occur when so many of the more usual continental species are missing. ARAMIDES CAJANEA MORRISONI Wetmore: Morrison’s Rail Aramides cajanea morrisoni WETMORE, Proc. Biol. Soc. Washington, vol. 59, March 11, 1946, p. 50 (San José Island, Archipiélago de las Perlas, Panama). Wood rails were common on San José but their presence was known mainly from their calls and from their tracks in the dust of roads and trails. They were found throughout the forested areas of the island, regardless of the presence of water; though they were heard or seen at times near streams, they were noted more often across the drier, upland areas. During February and March they seemed to range in pairs that called in duet, mainly in morning and evening and less frequently during the day. Occasionally they circled around me in the brush, keeping always under cover, and more rarely in early morning they were seen in the roadways cut through the forest. On such encounters they ran instantly into the cover of the brush, and regardless of how long I waited did not return. Though I devoted much time to hunting them, I did not succeed in making a successful shot. I had the impression from the sound of their calls at dusk that at times they went into the trees to roost, but of this I was not certain. As the season advanced, and as they became more accustomed to the presence of men, they were somewhat less shy, so that finally, after my departure, Morrison secured a fine series of 12 adults. These were taken on San José March 31, April 14, 18, and May 21. He also collected a juvenile in down on July 22. On Pedro Gonzalez I was told that they were fairly common, and on July 21 Morrison secured a female there. The downy young several days old, though as yet with no indica- tion of contour feathers, has the pileum, upper parts and sides of head and throat sayal brown; an indefinite line of deep mouse gray extending up upper breast onto the foreneck; rest of body fuscous black. This is the first of the species that I have seen in full down plumage. 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Family HAEMATOPODIDAE HAEMATOPUS PALLIATUS PALLIATUS Temminck: Oyster-catcher Haematopus palliatus TEMMINCK, Manuel d’ ornithologie, ed. 2, vol. 2, 1820, p. 532 (Venezuela). Oyster-catchers were distributed around the rocky headlands of San José, and were seen March g near the village of Cocal on Pedro Gonzalez. In February and March they were in pairs, and in the latter month I heard them calling both by day and by night along the sea below our camp. Occasionally they came out on the sandy beaches, but it was more usual to find them resting on rocks. While not abundant, they were seen frequently and seemed to be widely dis- tributed. The specimens taken are two males February 16 and April 23, a female April 18, and a young bird half grown secured May 10. The stomach of one taken February 16 contained the opercula of the abundant mollusk Nerita and a bit of a barnacle. ‘he fat in these birds is orange in color. While some place this as a race of Haematopus ostralegus of the Old World, the characters of the two seem sufficient to maintain them as separate species. The white back and lower rump of ostralegus are found even in the juvenile, while in young palliatus this area is dark as in the adult. The three adults from San José agree with palliatus from the Atlantic part of the range, showing no points that verge toward the differences under which the more southern form H, p. pitanay has been established. Family CHARADRIIDAE CHARADRIUS WILSONIUS BELDINGI (Ridgway): Belding’s Plover Pagolla wilsonia beldingt RipGway, U. S. Nat. Mus., Bull. 50, pt. 8, June 26, 1919, p. 112 (La Paz, Baja California). Morrison collected a male on a beach on San José, August 2, that is typical of this form in the darker dorsal color. CHARADRIUS HIATICULA SEMIPALMATUS Bonaparte: Semipalmated Plover Charadrius semipalmatus Bonaparte, Journ. Acad. Nat. Sci. Philadelphia, vol. 5, August 1825, p. 98 (coast of New Jersey). On February 18 I shot two females from a flock of a dozen at the mouth of the Rio Mata Puerco, and recorded them again on Febru- ary 21. When the tide was in, they went far back from the open NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 33 beach among the roots of the mangroves. The two taken were begin- ning the molt into breeding dress. Morrison collected an immature male on August 13, an early date for the return of this migrant from the far north. 6 CHARADRIUS VOCIFERUS VOCIFERUS Linnaeus: Killdeer Charadrius vociferus LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 150 (South Carolina). Robert Cushman Murphy recorded one November 14 around the extensive clearing for military headquarters on San José, noting what was probably the same bird frequently on subsequent days. Family SCOLOPACIDAE ACTITIS MACULARIA (Linnaeus): Spotted Sandpiper Tringa macularia LINNAEUS, Systema naturae, ed. 12, vol. I, 1766, p. 249 (Pennsylvania). Spotted sandpipers, as migrants from the north, ranged in small numbers on the rocky areas bordering the headlands along the shores of San José. Seldom were more than two or three seen during a day. Morrison shot a female beginning the molt into the spotted breast of the breeding season on April 6. August 13 he secured another female in worn breeding dress, apparently recently arrived. An adult male, beginning the molt, and an immature male were taken Sep- tember 3. CATOPTROPHORUS SEMIPALMATUS INORNATUS (Brewster): Western Willet Symphemia semipalmata inornata Brewsver, Auk, 1887, p. 145 (Larimer County, Colo. ). Our record for San José is of a female taken by Morrison on Sep- tember 23. This bird has the wing 197 mm. and the culmen 58.7 mm. This is another northern migrant. EROLIA MINUTILLA (Vieillot): Least Sandpiper Tringa minutilla Vie1tLot, Nouv. Dict. Hist. Nat., vol. 34, 1819, p. 466 (Halifax, Nova Scotia). On February 20 I shot three from a dozen found around some rocky headlands bordered by a sandy beach on the eastern side of San José. 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Family STERCORARIIDAE STERCORARUS PARASITICUS (Linnaeus): Parasitic Jaeger Larus parasiticus LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 136 (coast of Sweden). i Robert Cushman Murphy recorded one near the coast of San José February 21, 1941, during his investigations on the Askoy. CATHARACTA SKUA CHILENSIS (Bonaparte): Chilean Skua Stercorarius antarcticus b. chilensis BONAPARTE, Conspectus generum avium, vol. 2, 1857, p. 207 (Chile). On February 7 I saw two skuas between 3 and 4 miles outside the harbor of Balboa, and another 10 miles off San José Island. March 14 I noted one 15 miles out from San José as it crossed the bow of our boat. A short distance away it turned to pursue a laugh- ing gull, buffeting it about without, however, securing any food, and then flew on its way. While this seems to be the first published rec- ord for these waters, the birds should occur regularly in migration. Family LarmAr LARUS ATRICILLA Linnaeus: Laughing Gull Larus atricilla LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 136 (Bahamas). On February 6 and March 14 I recorded laughing gulls in scattered flocks over the Bay of Panama from Balboa across to San José and Pedro Gonzalez Islands. February 24 I shot two females just begin- ning to molt into breeding dress from a flock of 300 off the south- eastern shore of San José. The birds rested in a scattered flock on the water until our launch approached and then flew off in a long line. Later a flock joined pelicans, man-o’-war birds and cormorants in fishing. March 9 in a trip to Pedro Gonzalez several hundred were scattered over the open sea, and I noted that many were acquiring the dark hood of the summer dress. Hundreds were seen March 14 in crossing to Balboa. They seemed at this season to remain in the open sea as it was rare to note one along the beaches. Dr. Murphy recorded them around San José February 13 and May 23, 1941, and November 27, 1945. No. I BIRDS OF SAN JOSE ISLAND—WETMORE 35 THALASSEUS MAXIMUS MAXIMUS (Boddaert): Royal Tern Sterna maxima BoppaErt, Table des planches enluminéez, 1783, p. 58 (Cayenne). Dr. Murphy recorded the royal tern between San José and El Rey September 9, 1937, and reported others around San José February 13 and 21, 1941. He found them frequently during his work in November 1945. THALASSEUS SANDVICENSIS ACUFLAVIDUS (Cabot): Cabot’s Tern Sterna acuflavida Casot, Proc. Boston Soc. Nat. Hist., vol. 2, 1847, p. 257 (Tancah, Yucatan). On March 14 about midway between San José Island and Balboa one passed near at hand, the light tip of the bill shining clearly. GELOCHELIDON NILOTICA VANROSSEMI Bancroft: Western Gull-billed Tern Gelochelidon nilotica vanrossemi BANCROFT, Trans. San Diego Soc. Nat. Hist., vol. 5, No. 19, Dec. 10, 1920, p. 284 (Salton Sea, Imperial County, Calif.). On February 6 several hundred gull-billed terns were feeding over the open sea from 10 to 30 miles offshore from San José. Allocation to subspecies is made on geographical grounds as none were taken. CHLIDONIAS NIGRA SURINAMENSIS (Gmelin): Black Tern Sterna surinamensis GMELIN, Systema naturae, vol. 1, pt. 2, 1789, p. 604 (Surinam). Robert Cushman Murphy recorded this tern at sea near San José on February 21 and May 23, 1941. Family CoLUMBIDAE COLUMBA*CAYENNENSIS PALLIDICRISSA Chubb: Pale-vented Pigeon Columba pallidicrissa CHusB, Ibis, 1910, p. 60 (Costa Rica). This was the most conspicuous pigeon on San José, distributed universally through forested areas, absent only in sections of brush completely overgrown with matted creepers. Many frequented the growths of mangroves where streams entered the sea. In late after- noon and evening birds alone, or in little groups of two to six, flew in swift, direct flight over the forest, often passing the clearing at camp. As I walked quietly through the trees they often flushed over- head with clapping wings. They had considerable curiosity and some- 36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 times swung in on sighting me to alight in the tops of the trees. As they detected any movement on my part, however, they walked or fluttered behind the cover of leaves, or flew precipitately. February and early March marked the breeding season, and I heard their hooting calls, like the notes of some owls, constantly in the forest. Males often soared over the trees or through openings with the wings held stiffly, sometimes elevated above the back, and sometimes extended straight and flat. After the first week in March they were less noisy and many then had nests. February 19 I located a nest 12 feet from the ground below the top of a spiny-trunked black palm. It was a shallow and frail structure of twigs and contained one egg which I broke in attempting to obtain it. March 5 I picked up the shell of an egg that had just hatched, and on March 13 in the northern part of the island I flushed several birds from nests placed in tangles of vines. March 11 on Pedro Gonzalez I saw several and collected a full- grown squab. Our series of five adult males from San José averages definitely darker than specimens from Costa Rica, the type locality of the race pallidicrissa. There is, however, considerable variation in these birds so that it does not seem desirable to attempt separation of another race, occasional continental specimens being as dark as those from the Pearl Islands. COLUMBIGALLINA TALPACOTI NESOPHILA (Todd): Pearl Islands Ruddy Ground-dove Chaemepelia rufipennis nesophila Toop, Ann. Carnegie Mus., vol. 8, May 8, 1913, p. 590 (Isla El Rey, Pearl Islands, Panama). This ground dove, in 1944, was of local distribution on San José, being restricted to fairly open grassy areas on a headland at the East Harbor and the small, open section at Bald Hill in the north of the island. During February and March we did not penetrate into these two places, but as trails opened, Morrison found the birds and col- lected specimens on March 28 and 29, and May 6 and 18. It is probable that they will spread to other sections in recent clearings that have been made. On Pedro Gonzalez these birds were more abundant, ranging around abandoned fields. The natives there told me of them on my two visits, and later on May 16 and 18 Morrison collected two males. . The series from San José includes four females that agree with Todd’s observations that three females he had examined from the NO. OL BIRDS OF SAN JOSE ISLAND—WET MORE a7 Pearl Islands were darker colored, and so substantiate the race that he named tentatively. When compared with a good series from Central America and Panama they are darker and browner both above and below, though more especially so on the lower surface. An occasional bird from the mainland is fairly close, but even these in the main are slightly more grayish in tone. Males, as far as the four obtained show, two from San José and two from Pedro Gonzalez, are identical with those from the mainland. The race in a way thus is not wholly definite, possibly being in what may be considered a formative stage. Todd’s findings have been discounted on the possible ground that his specimens were immature males instead of females as marked by the collector. This supposition cannot apply to the four females in the present collection. As my findings, made inde- pendently, substantiate completely those recorded by Todd, I have no hesitance in recognizing the Pearl Islands birds as distinct. LEPTOTILA VERREAUXI VERREAUXI Bonaparte: Verreaux’s Dove Leptotila verreauxt BONAPARTE, Compt. Rend. Acad. Sci. Paris, vol. 40, 1855, p. 99 (Colombia). This dove was widely distributed through the forests, particularly where tree growth was high and the ground level fairly open below. They ranged alone or in pairs, and usually I saw several on each trip afield. In February and March they were nesting, and I heard their calls regularly. February 14 I flushed one from a compact, cup-shaped nest made of dried vegetation, firmly woven, in the crown of one of the spiny-trunked black palms, and placed about 8 feet from the ground. The nest was empty. The birds were tame and often ex- hibited curiosity ; as I moved quietly through the undergrowth one might fly toward me to alight near at hand and peer at me. Suddenly the tip of the tail would be lowered, then brought quickly up above the back and spread widely, while simultaneously the head was nodded quickly. In early morning, if the air was damp, these doves remained quiet until the sun had warmed the forest, when they became active. When the early morning air was dry they seemed to move around early, and on such occasions I found them sometimes walking about the roads that had been cut through the forest. On Pedro Gonzalez Island also they were fairly common, two being taken. These, with seven others from San José, average a little darker above and below than our series from the mainland, being especially darker than those from western Costa Rica. The differences, however, are bridged by individual specimens. 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 OREOPELEIA MONTANA MONTANA (Linnaeus): Ruddy Quail-dove Columba montana LINNAEUS, Systema naturae, ed. 10, vol. I, 1758, p. 163 (Jamaica). It has been a surprise to see a male of the ruddy quail-dove taken by Morrison on San José March 31. He recorded three others at various times in the forest, but did not have opportunity for another shot. Dr. Murphy recorded one November 10, 1945. These are the first records for the Pearl Islands. Family PsItTACIDAE AMAZONA AUTUMNALIS SALVINI (Salvadori): Salvin’s Parrot Chrysotis salvini SALVApoRI, Catalogue of the birds in the British Museum, vol. 20, 1891, p. 271 (Lion Hill Station, Canal Zone, Panama). Salvin’s parrot, distinguished by the red forehead and the bluish edgings on the feathers of head and nape, was common on San José Island in the more heavily forested areas. I found them especially in the wooded swamp at the mouth of the Rio Marina, and in the region beyond toward the East Harbor. In early morning parrots flew across the sky in pairs, appearing at sunrise and continuing active and noisy until about 10 o’clock. From then until evening I came across them at times feeding on figs and other fruits, and at sunset they began flying again but with less noise. They were always in pairs at this season. They nested in hollow trees in the swamp mentioned above, and here, in the early part of the day, were extremely noisy. There were a few on Pedro Gonzalez, where I shot a female March 9. Eleven were taken on San José February 28 and 29, March 7, June 25, July 16, and August 23. These birds do not differ from those of the mainland. The voice is more varied, and with more chattering calls, than in the other species found here. When Salvin’s parrot is flying, the longer tail is plainly evident. There seemed to be some competition between the two species, as occasionally I saw the present one flying aggressively at the yellowheads when they came too near. AMAZONA OCHROCEPHALA PANAMENSIS (Cabanis): Panama Parrot Chrysotis panamensis CABANIS, Journ. fiir Ornith., 1874, p. 349 (Panama). The Panama parrot, marked by yellow on the crown and by the distinctly shorter tail when on the wing, was less common than Salvin’s parrot, but still was present on San José Island in fair numbers. The two species inhabited the same areas, as their nesting NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 39 places and their food are similar, but they did not mix in friendly fashion. The voice in the Panama parrot is much louder, and though present in smaller number, it was this species that contributed mainly to the noise in early morning in the wooded swamp at the mouth of the Rio Marina, that at times completely covered all other bird sounds. We secured specimens February 28 and 29, May 26, and August 10. On Pedro Gonzalez I saw several on March g and collected a male. There was also a young bird kept as a pet in the village of Cocal. Mrs. Sturgis 1° has reported this species as seen on Pedro Gonzalez, this being the only previous record for the Pearl Islands. Our specimens do not differ from those of the mainland. PIONUS MENSTRUUS (Linnaeus): Blue-headed Parrot Psittacus menstruus LINNAEUS, Systema naturae, ed. 12, vol. 1, 1766, p. 148 (Surinam). The first of these birds was recorded on San José at the beginning of May with the ripening of the fruit of the membrillo near the camp in the southern part of the island. Morrison secured a pair here May 3 and another May 4. At first they were quiet and shy, but in a month a flock of 40 or more was present, including one partial albino, and the birds flew about in a noisy group like other parrots. Others were taken June 3, 4, and 17, and two were secured from high trees back of the settlement of San Sebastian on Pedro Gonzalez Island June 8. These specimens, including immature birds and adults in worn plumage, appear identical with those of the mainland. This parrot, small in size with head and neck completely blue, has not been recorded previously in the Pearl Islands. As we did not find it until 3 months of constant field observation had passed, and then in areas that we had covered repeatedly, it appears that it may be a wanderer from Panama and not resident on the islands. , . Family CUuCULIDAE COCCYZUS ERYTHROPTHALMUS (Wilson): Black-billed Cuckoo Cuculus erythropthalmus Witson, American ornithology, vol. 4, 1811, p. 16, pl. 28, fig. 2 (near Philadelphia, Pa.). The migrant female secured by Morrison on San José April 28 is the first record for the Pearl Islands. 19 Field book of birds of the Panama Canal Zone, 1928, p. 158. 40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 CROTOPHAGA ANI Linnaeus: Ani Crotophaga ani LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 105 (Jamaica). On San José Island I found the ani fairly common in little groups of four to eight that frequented the densest growth of canes and matted vines outside the areas of high forest. I saw them or heard their complaining calls almost daily but did not obtain specimens for some time as it was hard to get near them in the tangles that they frequented. And when I did shoot one it was difficult to find. In the end we secured three males and one female, and later on July 21 Morrison shot two females on Pedro Gonzalez. This ani has been familiar to me for years in many widely separated localities. My first impression of the San José birds was that they seemed large, and on comparing specimens in the Museum it was evident that they were different from most of those of continental areas in the greater development of the keel on the bill. Specimens in the Museum of Comparative Zodlogy from Isla El Rey, however, bridge the gap, and there are some specimens from the mainland of Panama that are quite similar to those of San José. At first I believed that a distinct form was indicated, but it appears on examination of more material that attempt to make a separation is hardly warranted as the differences, while evident, do not seem trenchant. Family TyTONIDAE TYTO ALBA GUATEMALAE (Ridgway): Central American Barn Owl Strix flammea var. guatemalae RipGway, Bull. Essex Inst., vol. 5, December 1873, p. 200 (Chinandega, Nicaragua). The first record for the Pearl Islands is a male shot by Morrison on San José, June 24. The bird rested in broad day in a tree at the edge of the open savanna at Bald Hill in the north end of the island, being visibleefor a long distance. Its stomach was stuffed with young iguanas. Others were seen or heard occasionally at night. The specimen is a male in light phase, quite different in appear- ance from the more common darker examples of this owl. From examination of available specimens the characters of guatemalae, as distinguished from the North American barn owl Tyto alba pratin- cola, are darker color in general, a heavier mottling of gray on the dorsal surface, and more vinaceous on the facial disk. The southern race may average slightly smaller, though this is not wholly certain as most of the skins of guatemalae at hand do not have the sex marked. Examples in light phase are quite similar to darker speci- NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 4I mens of perlata, but differ in average greater extent of gray on the dorsal surface, which also is slightly darker, and in the darker and more extensive vinaceous markings of the facial disk. Recently I have had occasion to examine Tyto alba subandeana Kelso,?° described from a trade skin marked “W. Evans” and en- tered in the National Museum catalog February 17, 1860. In the original description this was compared with twidara. It appears now that it is an example in light phase of guatemalae, as is a specimen received from Dr. Armando Dugand from Los Pendales, Depto. Atlantico, Colombia. This therefore extends the range of guatemalae into Colombia. Family CAPRIMULGIDAE NYCTIDROMUS ALBICOLLIS INTERCEDENS Griscom: Central American Pauraque Nyctidromus albicollis intercedens Griscom, Amer. Mus. Noy., No. 370, Oct. 17, 1929, p. 8 (Tela, Honduras). Our principal observations on these birds were made at night as we traveled the roads in jeeps. From time to time ahead of us we would see what seemed to be an orange-red coal, as the eye of the bird reflected our headlights. Or the rays of a flashlight would pick one up in the brush alongside. The eye color was exactly that of the live coals of the burning logs from the road clearing, so that some- times I had to look carefully to determine which was bird and which the remains of fire. Occasionally I heard one calling. They were reported to me on Pedro Gonzalez. Our series of six males and five females collected from February 10 to August 23 agrees with skins from Panama, as other workers have indicated ; and birds from this area are referred under present under- standing to intercedens of Central America. The San José specimens as a whole average dark with the under surface rather heavily barred, but are matched in this by dark specimens from Costa Rica and elsewhere. CHORDEILES ACUTIPENNIS INFERIOR Oberholser: Central American Nighthawk Chordeiles acutipennis inferior OBERHOLSER, U. S. Nat. Mus. Bull. 86, Apr. 6, 1914, pp. 24, 100, 109 (Triunfo, Baja California). The only record is that of a female taken near Bald Hill on San José by Robert Cushman Murphy November 13, 1945. This bird 20 Tyto alba subandeana Kelso, Biol. Leaflet No. 9, Apr. 21, 1938, p. I. (“Bogota,” Colombia). 42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 has the wing 168 mm., and the color characters of this race. It is a migrant, and one from near the extreme southern limit of this form. Dr. Murphy reports further that on November 22 he heard the booming of the other species of nighthawk, Chordeiles minor, above the clearing at headquarters. . Oberholser, in the reference given above, described the breeding nighthawks of this species from México and Central America as two distinct subspecies, Chordeiles acutipenms inferior from Baja ° California, and C. a. micromeris from the rest of the area. Van Rossem *! has demonstrated that the supposed characters of the bird from Baja California do not hold, a fact that is evident on com- parison of specimens. He unites them under the name muicromeris, the full account of which is found on page 100 of Oberholser’s work while the account of inferior comes on page 109. Peters, however (in litt.), calls my attention to the fact that in a key to the races on page 24 Oberholser gives his two supposed forms with the characters clearly indicated that separate them from the nearest relative. In this key* the name inferior is given first. Under the rules of nomen- clature, therefore, on line anteriority, inferior becomes the name for the Mexican and Central American race instead of micromeris. Family MicroPoDIDAE CHAETURA VAUXI OCHROPYGIA Aldrich: Pale-rumped Swift Chaetura vauxi ochropygia AtpricH, Sci. Publ. Cleveland Mus. Nat. Hist., vol. 7, Aug. 31, 1037, p. 68 (Paracoté, 1 mile south of the mouth of the Rio Angulo, Montijo Bay, Veraguas, Panama). These small swifts were constantly present on San José Island during our work, but were easily overlooked as usually. they flew over the forest where they were hidden from below. I recorded them most often from low hills where roads had been opened, and found them most active in the warmer hours of the day and in early evening. In February and March they were often seen in pairs that at inter- vals set the wings at an angle above the back and so sailed close together for short distances. Usually a dozen or so fed together, with occasional flocks of 50 or more. They circled rapidly with quickly beating wings, moving back and forth above the trees. Their high- pitched, chattering calls were heard infrequently. They were seen all through the summer, but I feel certain that they shifted constantly back and forth from the mainland to the islands, which they could do with ease. 21 Condor, 1942, p. 73. NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 43 Between February 16 and September 10 we secured a long series of specimens, 39 in all, in which the characters of more grayish brown in lower back, rump, and upper tail coverts that Aldrich pointed out in separating this race from richmondi, appear constant. In addition, with this larger series, it is evident also that ochropygia is grayer below, with the extreme upper breast, foreneck, and throat lighter, more whitish. Immature birds, taken July 6, have the rump and upper tail coverts grayer, less brownish. Molt is evident in the wing coverts of occasional birds taken June 27 and July 4, and primaries were being renewed September Io. CHAETURA VAUXI RICHMONDI Ridgway: Richmond’s Swift Chaetura richmondi Ripcway, Proc. Biol. Soc. Washington, vol. 23, Apr. 10, 1910, p. 53 (Guayabo, Costa Rica). The only swift secured on Pedro Gonzalez Island is a female taken by Morrison May 16. This bird is distinctly browner on the back, rump, and under tail coverts than the others collected, and agrees with richmondi of farther north, seeming only slightly intermediate. It is assumed to be a migrant. Family TROCHILIDAE PHAETHORNIS ANTHOPHILUS HYALINUS Bangs: Pearl Islands Hermit Hummingbird Phaéthornis hyalinus Bancs, Auk, vol. 18, January 1901, p. 27 (Isla El Rey, Pearl Islands, Panama). This interesting hummer was found on San José in the more open areas of the forest, often along the banks of the larger streams. It was seen infrequently, so that our eight specimens represent much hunting. The dates are as follows: March 5, 14, 15, April 9, 26, May 14, July 14, and September 13. March 11 I saw two on Pedro Gonzalez along an open, shaded trail adjacent to new clearings. These decoyed to a squeaking sound several times, but on each occasion came flying swiftly to me to hover a few inches from the openings in the barrel of my gun, to examine them curiously, and then, as I moved slightly, darted away out of sight. The birds obtained on San José differ uniformly from typical anthophilus in darker green, less bronzy dorsal surface, with less brownish evident. They are slightly darker on the crown than the only other specimen of hyalinus at hand, a female from Isla El Rey. 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 CHLOROSTILBON ASSIMILIS Lawrence: Allied Emerald Hummingbird Chlorostilbon assimilis Lawrence, Ann. Lyc. Nat. Hist. New York, vol. 7, 1861, p. 292 (Lion Hill Station, Panama Railway, Canal Zone, Panama). This was the most common of the hummingbirds on San José, where it was distributed through the forests, usually near water. I found them especially in mangroves at the mouths of the streams. Occasionally I saw them gleaning insects over the trunks of trees. The call was a low chewp chewp. The males did not seem particu- larly aggressive, so that it was usual to find several near one another. The morning of February 26 the thermometer in my tent dropped to 69.5° F., and we all felt cold in contrast to the warmer tempera- ture to which we had been accustomed. At the mouth of Rio Mata Puerco soon after sunrise I found a male of this hummer fluttering about on the ground unable to rise so that I caught it in my hand. It was in a place exposed to the wind, and I was inclined to believe that it was merely numb from cold as it was in perfect condition, and showed no mark of any injury. On Pedro Gonzalez I saw several, and collected two on March 9 and II. Our series of a dozen specimens, when compared with birds from the mainland, averages darker green, particularly in the males, with- out the brassy sheen that shows in most of the latter. However, two birds from Taboga Island are similar to those from the Pearl Islands. SAUCEROTTIA EDWARD MARGARITARUM Griscom: Pearl Islands Hummingbird Saucerottia edwardi margaritarum Griscom, Amer. Mus. Novy., No. 282, Sept. 12, 1927, p. 4 (Pedro Gonzalez Island, Pearl Islands, Panama). This hummingbird was fairly common in the forests of San José, usually along the streams. I found them in the mangrove swamps, as well as inland. Our eight specimens were taken February 11, 12, 13, and 15, and March 1, 7, and 18. On Pedro Gonzalez, where they were fairly common, two were taken March 9 and 11. The series bears out fully the characters assigned in the original descrip- tion of this well-marked race. Family ALCEDINIDAE MEGACERYLE ALCYON ALCYON (Linnaeus): Belted Kingfisher Alcedo alcyon LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 115 (South Carolina). NO. I BIRDS OF SAN JOSE ISLAND—WETMORE 45 Single belted kingfishers, migrants from the north, were seen on the rocky shores of San José February 25 and 26. Dr. Murphy collected one that he preserved in alcohol. This bird, from the plumage a female, is an individual of the eastern race, which is the only one recorded to date from Panama. MEGACERYLE TORQUATA TORQUATA (Linnaeus): Ringed Kingfisher Alcedo torquata LINNAEUS, Systema naturae, ed. 12, vol. I, 1766, p. 180 (México). A few of these large kingfishers were found at the mouths of the larger streams, along channels bordered by mangroves. And once I saw one fishing in salt water along the open shores of East Harbor, where the habitat was beach and shore with no fresh water near. We collected four on March 1, 10, and 30, and May 28. Family FoRMICARIIDAE THAMNOPHILUS DOLIATUS NIGRICRISTATUS Lawrence: Black-crested Ant-shrike Thamnophilus nigricristatus LAWRENCE, Proc. Acad. Nat. Sci. Philadelphia, 1865, p. 107 (Lion Hill, Panama Railroad, Canal Zone, Panama). March g on Pedro Gonzalez Island as I came up the trail from the beach beyond Cocal into the brushy upland forest I heard the characteristic note of this bird, and presently secured a female. March 11 I made special search for them and found them common, taking six. Morrison secured another May 16. They ranged in thick growth in pairs, moving quietly behind cover, but constantly uttering their peculiar calls. On San José we searched for them without success, and I was certain that they were not found on that island. To date they are recorded from El Rey, Viveros, and Pedro Gonzalez Islands. The series from Pedro Gonzalez has the bill averaging slightly heavier than skins from the mainland, but there is considerable variation. FORMICIVORA GRISEA ALTICINCTA Bangs: Pearl Islands Ant-bird Formicivora alticincta BANcs, Proc. New England Zool. Club, vol. 3, Mar. 31, 1902, p. 71 (Isla El Rey, Pearl Islands, Panama). This small bird, known as the pavita, is the most common of the avian species found on San José Island, being distributed universally through the forest, and from this cover penetrating into the borders of the mangrove swamps, and the matted growths of the more open slopes. In the forested areas they ranged from tangles of vines just 46 — SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 above the ground to the lower and middle branches of the trees. In the dark shadows of the denser growths they were seen with diffi- culty, but they had considerable curiosity and often came near me. Males show flashes of black and white as they fly, and both sexes flit the wings and jerk the tail continually. By March 7 a few males began singing in early morning, uttering a repetition of a single note that suggested the song of the black-crested ant-shrike of Pedro Gonzalez and the mainland. By the middle of the month this singing increased, and the sexual organs of birds taken indicated that the breeding season was beginning. They were as common on Pedro Gonzalez as on San José. Our series of specimens has the bill averaging somewhat heavier than those from El Rey, the type locality, but in some the dimensions are identical. Family TyRANNIDAE TYRANNUS TYRANNUS (Linnaeus): Eastern Kingbird Lamus tyrannus LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 94 (South Carolina). On San José Morrison found the kingbird common in migration at the end of April and early May, and collected a female May 12, the last one for the season. In fall they were also present in fair numbers, and he secured another September Io. TYRANNUS MELANCHOLICUS CHLORONOTUS Berlepsch: Lichtenstein’s Kingbird Tyrannus chloronotus BERLEPSCH, Ornis, vol. 14, 1907, p. 474 (Temax, Yucatan). This tropical kingbird was found in small numbers along the beaches of San José, and in the more open areas inland. It is a conspicuous species that rests on open perches where it is easily seen, so that it was recorded regularly. March 7 two that were evidently a pair, from their difference in size, flew high overhead in the cool air of early morning, finally ending with the male pursuing the female. Several were seen this day, and it may be that there was some migratory movement among them. I secured five on San José in February and early March, one on Pedro Gonzalez March 11, and Morrison collected two on Moreno Island May 19. MYIARCHUS FEROX PANAMENSIS Lawrence: Panamé4 Flycatcher Myiarchus Panamensis Lawrence, Ann. Lyc. Nat. Hist. New York, vol. 7, May 1860, p. 284 (Panama). Like the crested flycatcher of eastern North America this species is an inhabitant of forests, though it frequents the open branches @ NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 47 among and under the canopy tops of the trees, rather than remaining hidden above. It is therefore easily seen. Though resembling the northern bird in color pattern—having a gray throat, yellow lower breast and abdomen, and olive-green upper surface—it is conspicu- ously different in the color of the tail, which is dark without rufous- brown markings. The birds rest quietly to watch for insects, but seem to have much curiosity as frequently when I remained quiet for a few minutes one flew to nearby perches to peer at me with craning neck. They were found also among the more open trees at the forest edge bordering the beaches. March 9 and 11 I recorded this species on Pedro Gonzalez Island, taking three specimens. Morrison shot a female and a juvenile male on Moreno Island. The young bird has the inner primaries, outer secondaries, and the rectrices edged prominently with bright brown. Most of these flycatchers have the bill fuscous or brownish in color but in an occasional one it is black. This seems to be individual variation, though I note that in the juvenile listed above the color is black. On close scrutiny of our series of 12 from San José and 3 from Pedro Gonzalez the bill appears very slightly heavier than in birds from the mainland. The difference is only a tendency and is so slight that it does not stand out in measurements. EMPIDONAX VIRESCENS (Vieillot): Acadian Flycatcher Platyrhynchos virescens Vieittot, Nouv. Dict. Hist. Nat., nouv. ed., vol. 27, 1818, p. 22 (near Philadelphia, Pa.). Robert Cushman Murphy collected a male on San José November 21, 1945. The bird is recorded as a migrant in Panama though this is the first report of it from the Pearl Islands. MYIODYNASTES MACULATUS DIFFICILIS Zimmer: Noble Flycatcher Myiodynastes maculatus difficilis ZimMeR, Amer. Mus. Nov., No. 963, Nov. 18, 1937, p. 91 (Bebedero, Costa Rica). These flycatchers lived in the higher branches of the forest trees, where they were probably more common than the 11 specimens taken on San José and Pedro Gonzalez indicate. At the end of February they seemed to be mated, and Morrison collected a young bird just out of the nest on April 3. I saw them feeding on berries. The series of specimens does not seem to differ from the birds of the mainland. There is much variation in the size of the bill. 48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 MYIOPHOBUS FASCIATUS FURFUROSUS (Bangs): Bran-colored Flycatcher Myiobius naevius furfurosus BANGs, Bull. Mus. Comp. Zodl., vol. 46, September 1905, p. 152 (Saboga Island, Pearl Islands, Panama). The first of these small flycatchers that I saw on San José, on February 27, was in the lower branches of the trees in fairly open forest on the banks of the Rio Marina. It rested quietly watching for insects. On March 8 while looking for land rails, Col. W. H. W. Komp, my companion, located a pair at the edge of an open area on the road to East Harbor. They acted like the small flycatchers of the genus Empidonax, resting quietly until insects passed, and then making short sallies to capture them. Subsequently, as the trails on the island were opened, Morrison found more of them so that in all we secured 17. They are fairly common but because of their quiet habits are easily missed in the dense growths in which they live. A juvenile bird, recently from the nest, is cinnamon brown above, with the under surface colonial buff, washed heavily with honey - yellow across the breast, which is streaked indistinctly as in the adult. In the next plumage the abdomen is white and the breast chamois, while often they are brighter brown above, being tawny. My impression of this series is that the birds of San José average brighter brown above than five seen from the Canal Zone. There is much variation among them, however, some being inseparable. MYIOCHANES RICHARDSONII RICHARDSONII (Swainson): Western Wood Pewee Tyrannula richardson Swainson, Fauna Boreali-Americana, vol. 2, 1831 (1832), p. 146, pl. 46, lower fig. (Cumberland House, Saskatchewan). Among the wood pewees collected by Dr. Murphy there is an adult male, taken November 18, 1945, of this race. The wing mea- sures 88.4 mm. and the tail 66.4 mm. This is the first record for any wood pewee from the Pearl Islands. MYIOCHANES RICHARDSONII SORDIDULUS (Sclater): Central American Wood Pewee Contopus sordidulus ScLAaTER, Proc. Zool. Soc. London, May 1859, p. 43 (Orizaba, Veracruz). Of the four wood pewees collected by Dr. Murphy 3 males taken November 10 and 21, 1945, belong to the present race, which breeds from southern México to Guatemala. They are marked by shorter tails, measurements being as follows: wing 76.5, 79.7, 81.8 mm., tail 59.2, 59.4, 59.8 mm. Two are immature as is shown by the NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 49 definite whitish bars on greater and middle coverts. The adult is slightly darker above than the adult richardsonu secured. The two immature birds are browner above, and have the dark breast band more or less interrupted in the middle. One is much yellower below and browner above than the other, the color of the underparts in fact suggesting Myiochanes virens. On careful comparison, however, it is seen to be darker above, and is also separated by the short tail. PIPROMORPHA OLEAGINEA PARCA (Bangs): Bangs’ Pipromorpha Mionectes oleagineus parcus BANGs, Proc. New England Zodél. Club, vol. 2, Sept. 20, 1900, p. 20 (Loma del Leon, Panama). On San José this bird was found resting on high perches in heavy forest along the Rio Marina. It was quiet, and because of its colors was difficult to find. Our seven specimens were taken February 10, April 11, 14, 21, May 3, 27, and August 8. The last one listed is an immature male, barely grown. It is like the adults but with more extensive buffy bars on the wing coverts. @ The series is similar to specimens of parca from the Canal Zone. MYIOPAGIS VIRIDICATA ACCOLA Bangs: Panama Placid Flycatcher Myiopagis placens accola Bancs, Proc. New England Zool. Club, vol. 3, Jan. 30, 1902, p. 35 (Boquete, Chiriqui, Panama). This small flycatcher, identified by the partly concealed yellow in the crown, on San José is found in the denser forest near the streams. It ranged in the lower limbs and was difficult to find. We secured adults February 17, March 1, April 14 and 26, and August 8. Morri- son shot an immature male April 11. This differs from the older speci- mens in having an indefinite brownish-gray band across the chest, and in being darker above. In addition the yellow in the crown is restricted to a small, concealed, central spot. ELAENIA FLAVOGASTER SILVICULTRIX Wetmore: Pearl Islands Elainea Elaenia flavogaster silvicultrix WWETMorE, Proc. Biol. Soc. Washington, vol. 59, March 11, 1946, p. 51 (San José Island, Archipiélago de las Perlas, Panama). The Pearl Islands elainea was found near the streams, where it ranged from the lower branches of the trees to their summits. Febru- ary seemed to be the beginning of the mating season, and I heard their loud, wheezy notes constantly. The birds were quite active, and often posed with raised crests. A female taken February 12 would have laid within a week, and another secured March 5 was about 50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 to lay. There seems to be variation in their breeding, however, as I shot a young bird in juvenal dress March 3, and by the end of the first week in March they were less noisy than during February. I saw them eating berries of various kinds. We secured 11 specimens on San José, and 3 more on Pedro Gonzalez, where I found them in fair numbers on March g and II. ELAENIA CHIRIQUENSIS CHIRIQUENSIS Lawrence: Lawrence’s Elainea Elainea Chiriquensis LAWRENCE, Ann. Lyc. Nat. Hist. New York, vol. 8, 1867, p. 176 (David, Chiriqui, Panama). These elaineas, marked by smaller size, were less common on San José than the larger species, but still were not rare as we secured 13 specimens. They ranged in the forests but also appeared in more open sections, and in early morning as the sun came over the island were prone to come out into the branches where its rays penetrated. In February and early March they were nesting, and on March 6 a mating pair flew through camp several times, once coming into my tent. The call is a clear whees. This series is like a similar set of specimens from the Canal Zone. Morrison shot an immature male August 6 in which the wing bars are grayish brown, with the lighter adult feathers in these bands just appearing. A juvenile only recently from the nest has the abdomen dull white, the breast grayish brown, and the dorsal surface grayish brown without the greenish-olive cast that comes later. Though Griscom ** records this bird “throughout the arid tropics of Pacific slope” it ranges also into the more humid areas, as we have one specimen taken by Goldman at Majagual, near Colon, March 13, IQII. SUBLEGATUS GLABER ARENARUM (Salvin): Smooth Flycatcher Elainea arenarum SAtvin, Proc. Zool. Soc. London, August 1863, p. 190 (near Puntarenas, Costa Rica). These small flycatchers, much like elaineas in form and habits, were fairly common on San José, usually frequenting the border of the higher forest, where light and air enter freely. I found them also along the course of the Rio Marina, where similar conditions prevail, and occasionally at the border of mangrove swamps. The note is a low swees. I shot two and saw several others on Pedro Gonzalez March 11, and Morrison secured a juvenile male on Moreno Island May 19. 22 Bull. Mus. Comp. Zodl, vol. 78, 1935, p. 353- NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 51 The latter is cross-barred with grayish white on head, wing-coverts, and back, like a young elainea. The identification of these birds as arenarum is tentative and sub- ject to later revision. Zimmer, in his paper in American Museum Novitates (No. 1109), has assigned birds from the Pearl Islands to atrirostris (type locality Cartagena), but the series of 12 skins that I have from San José and Pedro Gonzalez is definitely duller colored than one from Cartagena. There is considerable variation in these birds and the whole group is subject to further revision. CAMPTOSTOMA OBSOLETUM MAJOR Griscom: Pearl Islands Camptostoma Camptostoma pusillum major Griscom, Bull. Mus. Comp. Zool., vol. 72, January 1932, Pp. 353 (San Miguel, Isla El Rey, Pearl Islands, Panama). The only specimen available from San José is one secured by Robert Cushman Murphy November 17, 1945, at Ensenada de la Bodega. This bird is slightly grayer above than a series from the Canal Zone, as are two from Isla Fl Rey (San Miguel) in the National Museum. The difference is very slight but seems sufficient to characterize the island race as distinct. The alleged difference in size, the Pearl Islands birds having been said to be larger, seems to me very slight, and I doubt that it will hold. The sex of the specimen from San José could not be ascertained. On two occasions in early March, 1944, I saw one of these tiny flycatchers flying near the road to East Harbor, but did not succeed in collecting one. Griscom and Crosby shot one on Pedro Gonzalez February 18, 1927. Family H1RUNDINIDAE PROGNE CHALYBEA CHALYBEA (Gmelin): Gray-breasted Martin Hirundo chalybea GMELIN, Systema naturae, vol. 1, pt. 2, 1789, p. 1026 (Cayenne). On San José Island, about 9 in the morning on March 7, six or eight martins flew overhead near East Harbor, these being the first I had seen. They crossed high overhead, traveling directly without circling to feed. On March 14, when crossing to Balboa, I saw one flying north across the gulf. Martins did not nest on San José, but appeared again early in July, being noted through September. On July 20 Morrison collected 10 from a flock of 40 or 50, all being immature birds. He took another immature bird September 1, and two adults September Io. 52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106: HIRUNDO RUSTICA ERYTHROGASTER Boddaert: Barn Swallow Hirundo erythrogaster BoppArrt, Table des planches enluminéez, 1783, p. 45 (Cayenne). On March 11 I recorded one at sea midway between San José and Pedro Gonzalez Islands, flying to the northwest toward distant Panama. On San José, Morrison shot an adult male April 18, and in the fall secured an immature female September 10, and adult and immature females September 24. Robert Cushman Murphy recorded one between San José and El Rey September 9, 1937, and collected one at Bald Hill November 12, 1945. He found them com- mon around open areas during November. RIPARIA RIPARIA RIPARIA (Linnaeus): Bank Swallow Hirundo riparia LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 192 (Sweden). March 14, midway between San José Island and Balboa, I recorded several flying north across the sea. Morrison collected three on San José September 10. PETROCHELIDON PYRRHONOTA (Vieillot): Cliff Swallow Hirundo pyrrhonota Vieritot, Nouv. Dict. Hist. Nat., nouv. ed., vol. 14, 1817, p. 519 (Paraguay). While we were crossing the Gulf of Panama on March 14, en route between San José and Balboa, several passed our boat flying north. Family TROGLODYTIDAE TROGLODYTES MUSCULUS INQUIETUS Baird: Panama House Wren Troglodytes inquietus Batrp, Review of American birds, vol. 1, September 1864, pp. 138, 143 (Panama Railroad). Though the house wren has been reported only twice from the Pearl Islands, once from Isla El Rey and once from Isla Trapiche, I found it fairly common. It inhabited tangles of vines in forest, and was so shy that it was difficult to secure specimens. Our series of 10 from San José and 1 from Pedro Gonzalez was obtained only by much hunting. Males were singing in February and March, and it was through their songs that I usually found them. But when I attempted to approach they slipped away into heavy cover. They were not nesting at that season. Morrison secured young birds not fully grown July 14, August 8, and September 12. NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 53 Compared with specimens from Panama, these from the Pearl Islands average very slightly brighter brown on the flanks, and some- what browner above. In addition the bill is slightly heavier, though this does not appear in measurements. The variation appears to be slight, and as yet only a tendency that from this material does not warrant a name. ° Family BoMByCILLIDAE BOMBYCILLA CEDRORUM Vieillot: Cedar Waxwing Bombycilla cedrorum ViEtLLot, Oiseaux de |’ Amérique septentrionale, vol. 1, 1808, p. 88, pl. 57 (eastern North America). On February 29 on San José I had a clear view of a flock of about 25 flying over the large open area near East Harbor. The birds were traveling north. While this is a sight record, I have no hesitance in including it here as a valid occurrence, as there was no question as to the identity of the birds. Family VIREONIDAE VIREO FLAVOVIRIDIS FLAVOVIRIDIS (Cassin): Yellow-green Vireo Vireosylvia flavoviridis Cassin, Proc. Acad. Nat. Sci. Philadelphia, vol. 5, 1851, p. 152 (San Juan de Nicaragua, Nicaragua). These vireos were distributed throughout the forests of San José on my arrival the first week in February, and remained among the most common of the birds of the island throughout my stay. By the first week in March they were mating, and by the middle of the month a good many seemed located on their nesting grounds, as at camp at this time I noted two that sang daily from two differ- ent clumps of trees. In display males posed with spread tail and crest feathers raised. All through this period it appeared that mi- grants were passing also. The song in general is like that of the red-eyed vireo (Vireo olivaceus) of eastern North America, but differs so definitely as to be characteristic. The sound, in part, is the same in both birds, but in the yellow-green vireo the utterances are less regular, the spacing in time between the different phrases varying widely and irregularly, so that the song does not proceed with the steady, almost mechanical insistence of that of the northern bird. Some of the sounds, too, are more slurred. These vireos ranged in the middle and upper branches of the trees, searching for their food in the usual leisurely vireo fashion. On March 9 and 11 I found them on Pedro Gonzalez Island and collected specimens. 54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 As these birds were common we secured a series of 21 skins to check on the question of the supposed form insulanus, described by Bangs ** from Rey Island and later considered to be the bird of Panama also. There is variation in depth of color among these, some being decidedly darker, but this seems to be entirely individual. As far as this material goes, insulanus seems identieal with flavoviridis. r) VIREO FLAVIFRONS Vieillot: Yellow-throated Vireo Vireo flavifrons ViEILLoT, Oiseaux de |’ Amérique septentrionale, vol. 1, 1808, p. 85, pl. 54 (eastern United States). March 1 I collected a female in the mangrove swamp at the mouth of the Rio Marina on San José Island. Family CoEREBIDAE COEREBA FLAVEOLA CERINOCLUNIS Bangs: Pearl Islands Bananaquit Coereba cerinoclunis BANGS, Proc. New England Zool. Club, vol. 2, Feb. 8, 1901, p. 52 (San Miguel —Isla El Rey, Archipiélago de las Perlas, Panama). This honey-creeper was common through the forest on San José, but could be easily overlooked because of its small size and weak notes. Our series of specimens includes a juvenile taken February 13, while on February 21 I noted a nest partly constructed. On Pedro Gonzalez these birds were decidedly common on March 9g and 11. CYANERPES CYANEA CARNEIPES (Sclater): Blue Honey-creeper Coereba carneipes ScLater, Proc. Zool. Soc. London, 1859 (February 1860), p. 376 (Playa Vicente, Oaxaca). This honey-creeper was common in the forests of San José, where it ranged usually in the tops of the taller trees, often in little flocks of half a dozen. In flight, even when near at hand, the males appeared blackish, only occasionally displaying a flash of color. In the tree- tops they were difficult to see because of their small size. In the mangrove swamp at the mouth of the Rio Marina, where fresh water entered, I found them coming down to drink, and here secured speci- mens with ease. Once, while I was watching here, two females came tumbling through the branches together to rest a few feet away, posing with bills at a 45° angle, threatening one another with flitting wings while they called softly. Presently they were followed by a 23 Vireo insulanus Bangs, Proc. New England Zodl. Club, vol. 3, Mar. 31, 1902, p. 73 (San Miguel = Isla El Rey). NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 55 male that assumed the same attitude and pecked insistently at one female until she left. The six males and six females taken appear identical with birds from the mainland. Griscom and Crosby collected a male on Pedro Gonzalez Feb- ruary 18, 1927. Family MNIoTILTIDAE MNIOTILTA VARIA (Linnaeus): Black-and-white Warbler Motacilla varia LINNAEUS, Systema naturae, ed. 12, vol. 1, 1766, p. 333 (Hispaniola). This northern migrant was seen on three occasions on San José, February 12 when I shot a female, February 17, and March 12. Its habits in the Tropics are identical with those in the north. VERMIVORA PEREGRINA (Wilson): Tennessee Warbler Sylvia peregrina Witson, American ornithology, vol. 3, 1811, p. 83, pl. 25, fig. 2, (banks of the Cumberland River, Tenn.). On March 1 there was a small migration of these birds in man- groves near the mouth of the Rio Marina. The three that I shot were females. Griscom and Crosby secured a female on San José February 16, 1927. PROTONOTARIA CITREA (Boddaert): Prothonotary Warbler Motacilla citrea BoppaErt, Table des planches enluminéez, 1783, p. 44 (Loui- siana). This is a common winter resident. On February 12 we secured one on the Rio Marina on San José, and February 20 saw another in an extensive mangrove swamp. Dr. Murphy obtained specimens November 10, 17, 26, and 27, 1945. DENDROICA PETECHIA AEQUATORIALIS Sundevall: Panama Golden Warbler Dendroeca petechia h) aequatorialis SUNDEVALL, Ofvers. Kongl. Vetensk.-Akad. Forhandl., vol. 26, 1869 (1870), p. 609 (Panama City, Panama). In the excellent series obtained on San José Island there is evident a tendency toward heavier pigmentation when these birds are com- pared with those from El Rey and from the mainland. This is apparent in the deeper golden color of the lower surface, in the somewhat darker chestnut of the crown, and in the tendency toward heavier and more abundant streaking on the sides and lower breast. 56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Birds from Pedro Gonzalez agree with those from San José in these characters. One male from the latter locality shows this trend toward deeper color to an abnormal degree as the entire plumage is suffused with orange that covers the yellow normal in the lighter markings, and that shows even in the more greenish shades of the dorsal surface. While an average difference for specimens from the two outer islands may be shown, there are birds from the other localities, particularly one (U.S.N.M. No. 41534) from near Panama City, so closely simi- lar that it does not seem valuable to indicate this difference by name, at least not until more specimens from the whole coastal area of Central America are at hand. This was one of the common birds of both islands that was re- corded regularly, and of which we secured a good series of speci- mens illustrative of the considerable variation in plumage. My first encounters with them were in or near mangroves near the mouths of the streams, but as I became more familiar with the area I found that they were distributed widely through the higher forest growths. On San José I recorded them away from the shore mainly in the taller trees near or over water, but on Pedro Gonzalez March g and It they were common over the forested hills far from any such habitat. Because of the heavy vegetation they were more evident along the shores, as elsewhere they remained concealed in the leaves 60 to 75 feet from the ground. In various places small trees with thin, light green leaves grew along the shore, often projecting out from rocky ledges or sandy banks over the tide line. These were favorite haunts, as were open-branched trees growing at the tops of higher, steeper bluffs. Many times these warblers were shy so that they were seen with difficulty, and usually I located them by their chipping notes or songs. It was especially hard to find them where the wind was agitating the leaves, as then they slipped away unnoticed. February 19 in the mangrove swamp at the mouth of the Rio Marina I shot a fine male nearly in breeding stage, and a young female molting into adult dress came down to it immediately. It appeared that the two were mated. DENDROICA FUSCA (Miiller): Blackburnian Warbler Motacilla fusca MULLER, Natursyst. Suppl., 1776, p. 175 (French Guiana). The only present record is one collected on San José by Robert Cushman Murphy November 17, 1945. NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 57 DENDROICA CASTANEA (Wilson): Bay-breasted Warbler Sylvia castanea Witson, American ornithology, vol. 2, 1810, p. 97, pl. 14, fig. 4 (Pennsylvania). This warbler was present in small numbers as a winter resident in the heavy forest along the Rio Marina. February 1o I secured a male that was beginning to molt into spring plumage. The flanks are chestnut, and brown feathers are appearing on breast and crown. Two females, shot February 27 and March 6, are in winter dress, and one taken April 14 is in spring plumage. In addition to these I recorded two March 12. SEIURUS NOVEBORACENSIS NOVEBORACENSIS (Gmelin): Northern Water-thrush Motacilla noveboracensis GMELIN, Systema naturae, vol. 1, pt. 2, 1780, p. 958 (New York). These water-thrushes were fairly common on San José in Febru- ary and March, some being apparently in northward migration as early as February 11. I assumed this, since I found them frequently in trails across dry hillsides far from water, where no bird of this group would normally range. It was more usual to encounter them among the jumbled boulders behind the sandy beaches and rocky points, which is a normal habitat, so that probably some of these latter were of the winter resident group. Though they were found in the open, usually they flew to cover immediately when I appeared. On the beaches this meant retreat behind the huge boulders where endless shaded passages offered a safe hunting ground, completely screened from observation. On the open trails they had only to fly into the adjacent coverts to be hidden from sight in an instant. They were silent at this time, seldom calling. It is impossible to distinguish the races of this bird in life, and I was at some trouble to secure the 10 skins found in our collection. Four birds of the typical form were obtained on February 11, March 5 and 8, and Pepi D1: One recorded March 9 on Pedro Gonzalez was not collected and is therefore of unknown race. SEIURUS NOVEBORACENSIS NOTABILIS Ridgway: Grinnell’s Water-thrush Seiurus naevius notabilis Ripaway, Proc. U. S. Nat. Mus., vol. 3, 1880, p. 12 (Black Hills, Wyo.). ° The six specimens from San José attributed to this form were taken February 20 and 23, March 1, 3, 5, and 12. These are more 58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 slaty, less brownish above, and usually, but not always, less yellow below. Dr. Murphy secured one at Bald Hill November 9g, 1945. SEIURUS MOTACILLA (Vieillot): Louisiana Water-thrush Turdus motacilla ViEtLLot, Oiseaux de |’ Amérique septentrionale, vol. 2, 1807, (1808?), p. 9, pl. 65 (Kentucky). These birds appeared early in fall migration, as Morrison secured a male on August 26. He shot a female on September 12. SETOPHAGA RUTICILLA (Linnaeus): American Redstart Motacilla Ruticilla LinNaEus, Systema naturae, ed. 10, vol. 1, 1758, p. 186 (Virginia). On Pedro Gonzalez Island I collected an adult male March 11. Morrison secured a female on San José September 7. Family IcTERIDAE CASSIDIX MAJOR PERUVIANUS (Swainson): Great-tailed Grackle Quiscalus Peruvianus Swainson, Animals in menageries, Dec. 31, 1837, p. 354 (Pert). The grackle was common along the shores of San José and Pedro Gonzalez, the natives at Cocal on the latter island calling it changamé. The birds ranged along sandy beaches and rocky areas, feeding on the ground and perching in the trees above. In these forested islands they did not go inland, except into the mangrove swamps at the mouths of the streams. They were especially in evidence when low tide exposed extensive feeding grounds. A female was observed carrying nesting material on February 16. They were often found in pairs. Swainson’s description of his Quiscalus Peruvianus from “Mr. W. Hooker’s Collection Mus. Nost.,” taken from a bird from Pert, applies in every detail to a male of this grackle as pointed out by Hellmayr.** Family THRAUPIDAE PIRANGA RUBRA RUBRA (Linnaeus): Summer Tanager Fringilla rubra LINNAEUS, Systema naturae, ed. 10, vol. 1, 1758, p. 181 (South Carolina). ‘ Morrison secured males of this northern’ migrant April 2 and 16. 24 Cat. Birds Amer., Publ. Field Mus. Nat. Hist., vol. 13, pt. 10, 1937, p. 92. NO. I BIRDS OF SAN JOSE ISLAND—WET MORE 59 THRAUPIS EPISCOPUS DIACONUS (Lesson): Blue Tanager Tanagra (Aglaia) diaconus Lesson, Rev. Zool., June 1842, p. 175 (Realejo, Nicaragua). The blue tanager was fairly common on San José so that we obtained a series of eight specimens in February and March. These birds delight in long, undulating flights over the forests, dropping down into the tops of the tallest trees. At such times they seem very small, so that they give the impression of being some other bird. Because of their strong flight it seems probable that they may move from island to island, or even cross to the mainland. Griscom and Crosby secured one on Pedro Gonzalez February 18, 1927. RAMPHOCELUS DIMIDIATUS LIMATUS Bangs: Pearl Islands Crimson-backed Tanager Rhamphocelus limatus Bancs, Auk, January 1901, p. 31 (San Miguel = Isla El Rey, Pearl Islands, Panama). This beautiful tanager, in the adult male crimson with black wings and tail, was common on San José and Pedro Gonzalez though the heavy forests made it difficult to judge the actual number present. The birds were found frequently in small parties of half a dozen or more, with a predominance of those in immature plumage. They were at- tracted readily, and frequently uttered a low chewp chewp. At the end of February they were mating. A male in immature dress taken February 12 had the testes half developed. Our excellent series agrees with birds from Isla El Rey, the type locality. Family FRINGILLIDAE SPIZA AMERICANA (Gmelin): Dickcissel Emberiza americana GMELIN, Systema naturae, vol. 1, pt. 2, 1789, p. 872 (New York). Morrison found these birds in the open area at Bald Hill and secured five specimens on April 29 and 30. Robert Cushman Murphy collected a female November 12, 1945, noting several small flocks during the month. SALTATOR ALBICOLLIS SPERATUS Bangs and Penard: Pearl Islands Streaked Saltator Saltator striatipictus speratus BANGS and PENARD, Bull. Mus. Comp. Zodl., vol. 63, June 1919, p. 33 (Saboga Island, Pearl Islands, Panama). These streaked finches were among the commoner birds on San José and Pedro Gonzalez Islands, ranging in the undergrowth and 60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 through the lower branches of the trees, more rarely going up into the forest crown. Their sweet, cardinal-like songs made them the principal song birds of the island forests, and in February and March I heard them constantly. A young bird under care of its parents was taken February 26. Morrison secured a male on Moreno Island May 19. This race is rather poorly differentiated from S. a. isthmicus of the mainland, but our series of 17 skins indicates that it is separable. VOLATINIA JACARINA ATRONITENS Todd: Blue-black Grassquit Volatinia jacarini atronitens Topp, Proc. Biol. Soc. Washington, vol. 33, Dec. 30, 1920, p. 72 (Campeche, Campeche). On Pedro Gonzalez Island the blue-black grassquit was commen around the little open gardens of the natives. I shot an adult male here March 11, and Morrison secured a small series on May 16 and 18. Morrison also found a few in the opening at Bald Hill on San José, and secured specimens there April 29 and May 6. ORYZOBORUS FUNEREWUS Sclater: Lesser Rice Grosbeak Oryzoborus funereus P. L. Scrater, Proc. Zool. Soc. London, 1859 (February 1860), p. 378 (Suchapam, Oaxaca). February 18 I collected a male still in brown juvenile dress back of the sandy beach at the mouth of the Rio Mata Puerco. It rested in a bush 6 feet from the ground, and sang a sweet but indefinite warbling song. Morrison secured a small series April 30, August I1 and 24, and September 1. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 1, PL. 1 1, ENSENADA DE LA BODEGA, SAN JOSE ISLAND, MARCH 6, 1944 2, LOOKING ACROSS THE FORESTED SURFACE OF SAN JOSE ISLAND, BACK OF ENSENADA DE LA BODEGA, MARCH 6, 1944 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES LOG) NOS i Plea 1. ROAD CUT THROUGH THE FOREST, NEAR THE RIO MARINA, SAN JOSE ISLAND, FEBRUARY 13, 1944 2, A FOREST-BORDERED POOL IN THE LOWER COURSE OF THE RIO MARINA, FEBRUARY 13, 1944 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 106; NOS 2, PEs 3 1. PELICANS AND CORMORANTS OVER A SCHOOL OF FISH ON THE EAST SIDE OF SAN JOSE ISLAND, FEBRUARY 24, 1944 2. VILLAGE OF COCAL, WITH A NATIVE-MADE CAYUCO, PEDRO GONZALEZ ISLAND, MARCH 9, 1944 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 1, PL. 4 1. ISLA TRAPICHE, SENORA. AND SENORITA (LEFT TO RIGHT) AS SEEN FROM COCAL ON PEDRO GONZALEZ ISLAND, MARCH 9, 1944 2. HARBOR AT COCAL, PEDRO GONZALEZ ISLAND, ISLA TRAPICHE AT RIGHT, MARCH 9, 1944 ig Pe it eneet . a ap P Wy , us "SMITHSONIAN MISCELLANEOUS COLLECTIONS iy VOLUME 106, NUMBER 2 A THE VEGETATION OF SAN JOSE ISLAND, ao. _ REPUBLIC OF PANAMA (WiTH Two ‘Ptares) | BY 4 CG. O. ERLANSON U. S. Department of Agriculture ha SOX 2a AHS, Lies (PUBLICATION 3846) _ CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION _ JULY 18, 1946 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106 NUMBER 2 THE VEGETATION OF SAN JOSE ISLAND, REPUBLIC OF PANAMA (WitTH Two PLATEs) BY GC. O. ERLANSON U. S. Department of Agriculture ey WEN Ss, GE'A ad io 70, On Ga (PUBLICATION 3846) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JULY 18, 1946 Cha Cah cok aim The Lord Baltimore Press BALTIMORE, MD., U. 8 As Fa “es se ty : he oe oe on ! q Lh y A ; a. A gy { ws > wt "y ‘ 5 ¥ wit ‘ ay : a ae : ‘ ee Pe a, ‘ yh * . #; i + é ; pat MN q a , A, A © ; r a we \ 3 ate. ay " ay oa Pe SG ee 7 . ) t Al. ae eal - MEE OTL Mite Rote al dpae JOU ee ' SA hate ee | Ge THE VEGETATION OF SAN JOSE ISLAND, REPUBLIC OF PANAMA By C. O. ERLANSON U. S. Department of Agriculture (WitrH Two PLATEs) San José is the farthest west of a group of small islands lying in the Bay of Panama which have been called the Islas del Rey (King Islands) or Islas de las Perlas (Pearl Islands) by mariners who have visited them during the past several centuries. In all, they are said to cover an area of approximately 400 square miles. The three largest islands are Isla del Rey (Rey Island), 17 miles long by 10 wide; San José, less than half as large; and Pedro Gonzalez, still smaller. The rest are small islets, some of them hardly more than protruding rocks. All are volcanic in origin, heavily eroded and rounded to low-lying, rolling tableland. Probably the highest points above sea level are on Rey Island, where elevations between 600 and 700 feet are reported. Apparently the islands have always been sparsely populated. Rey Island at present has the largest population, and on its northern side is situated a small village, San Miguel, which has been in existence since Spanish Colonial days and is the seat of the local government of the islands. It was the center of what was once a flourishing pearl-fishing industry, now all but discontinued. The livelihood of the people, living on hardly more than a subsistence basis, depends largely on fishing during the dry season and on migra- tory agriculture during the rainy season. Inquiries made at San Miguel indicate that some 300 hectares of land are at present cleared for crops, which consist mostly of manioc, maize, rice, bananas, and plantains. The agriculture practiced is of the crudest type. Small patches of forest land are cleared by cutting and burning, and crops are planted without further cultivation. After two or three crops the land is abandoned to the inevitable scrub jungle that encroaches upon it, and new land is cleared. Since access to new land is much easier by water than through the forest, the coastal areas are largely denuded of virgin growth no matter how steep the terrain. The island of Pedro Gonzalez has probably only a handful of people SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 2 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 residing there permanently. The villages observed consisted of only 5 to 10 crude huts built of the native cane (Gynerium) and thatched with coconut-palm leaves. Around the huts were a few trees of mango, calabash gourd, bananas, and plantains, as well as chickens and pigs. Fishing appeared to be the principal means of livelihood. The island of San José has not been occupied for about 80 years, although there is evidence that much of the coastal region has been denuded of its virgin growth by migratory agriculture in past centuries. San José lies between longitudes 79°4’ and 79°9’ and latitudes 8°12’ to 8°19’ N. It is approximately 7 miles long and 3 miles wide, being constricted in the center to a neck hardly more than a mile in width. The coast line is very irregular for the most part, rising steeply from the sea as rocky bluffs 30 to 50 feet high. The largest sand beach is Playa Grande, bordering a large bay and 2 miles long, on the eastern side of the island. Other sandy beaches are very small, usually marking a valley through which a stream flows to the sea. The interior is gently rolling with an average elevation between 100 and 200 feet, the highest hills being around 350 feet. There are no lakes or ponds, and the very few streams which flow throughout the year arise as springs. The largest of the perennial streams is the Rio Marina, which arises in the northern half of the island and enters the sea in the Ensenada Playa Grande on the eastern side of the island. Another stream, the Rio Mata Puerco, flows northward from the southern half of the island and empties into the sea on the western side of the island. None of the streams can be described as anything more than a brook. There are many ravines and small gullies which carry water only during the rainy season, and then only after heavy rains. There are insufficient data to draw any reliable conclusions as to the average annual rainfall, but it probably approximates that of the region about Panama City, about 71 inches a year. Judging from the island’s location and low elevation it will probably prove to be less than on the immediately adjacent mainland. The effect of the precipitation on the plant life cannot be judged by the total amount of rainfall per year, but depends rather on its seasonal distribution and the manner in which it falls. The rainy season begins in April, apparently becoming heaviest 1n July and August and diminishing in October. The dry season extends from the latter part of Novem- ber through March, and during this period the relative humidity is high. Much of the rain that falls during the wet season is torrential in nature and the run-off is severe. NO. 2 VEGETATION OF SAN JOSE ISLAND—ERLANSON 3) Around the first of April, at the end of the dry season, the de- ciduous trees are bare of leaves, and even some of the evergreen types have withered foliage owing to the dryness. The grasses and other herbaceous vegetation have either disappeared or dried to brittle stiffness. During April, even before the rains begin, there is a marked change in the growth rhythm of the vegetation. Trees and shrubs begin to send out new shoots of growth, and the bare deciduous trees start putting out new leaves, anticipating the rains to come. In 1945 the rainy season apparently was late in arriving, and much of the new growth again withered and died for lack of water. With the first rains a great surge of flowering took place lasting through the early part of May, followed by lush growth. Ferns, liverworts, mosses, grasses, and other herbaceous material had largely matured new growth and begun to fruit by the middle of July. The island is volcanic in origin, being composed of dark igneous rock and tufa, with occasional dikes of basalt protruding through it. The soils are reddish-brown, here and there taking on a purplish tinge, and in the richer forested valleys becoming a dark chocolate in color. They are very friable, absorb moisture rapidly, and tend to fissure quickly on drying. Over much of the island the soils are thin, and outcrops of the basic rock or large angular boulders are frequent. Stream beds are usually free of soil or sand and generally have a pavement of stone on which are scattered erratic boulders worn and pocked by the action of water. The plant life of the Pearl Islands has been practically unknown. Aside from a few collections made by naturalists aboard various ships which have stopped at the islands for water, and collections recently made from short visits by Panamanian botanists to Rey and Pedro Gonzalez Islands, no material was available for study. The main difficulties have been the lack of transportation to the islands and subsistence problems which arise in an extended stay. Early in 1944 the United States Army established an outpost on San José, built roads and trails to all parts of the island, and thus made available for study an area uninhabited for at least 80 years, and probably never occupied by more than a handful of people at any time. This we may construe from the general absence of the common tropical weeds which follow man wherever he settles for any length of time. In one location on the western side of the island potsherds have been found which the Smithsonian Institution has reported as belonging to the Spanish Colonial period. In other spots on both eastern and western sides of the island, a few sour oranges, mangos, and plantains have been found. The considerable areas of 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 scrub jungle which occur near the coast, especially in the northern half of the island, are probably at least in part the result of migratory agriculture. Here and there straight lines of demarcation between scrub and forest can be explained only as the result of clearing by man. But probably the whole interior of the island had remained, until a year and a half ago when the Army moved in, completely unaltered by man. Dr. Ivan Johnston, of the Arnold Arboretum of Harvard Uni- versity, made extensive plant collections on the island during 1944 and 1945. From April through August of 1945, while on assignment to the War Department through the Smithsonian Institution, the writer was based on San José and engaged in making certain plant investigations for the United States Army. During this time he was able to make an additional series of collections which have been identified for the most part by Dr. Johnston. The writer is indebted to him for such technical names as appear in this paper. The map (fig. 1) shows the distribution of the canopied forest in relation to open scrub. Although the forest is now generally confined to the interior depressions and shallow valleys, there is evidence that it once had a wider spread over most of the island. Shrubs and small trees normally found only in the shade of the forest canopy still survive in what is now low scrub jungle exposed to the sun. The forests of the island belong to the type which may be called semi- evergreen seasonal forest. Many of the trees normally drop their leaves during the long dry season, and others which usually remain evergreen are capable of dropping their leaves if the drought is exceptionally severe. At the end of the dry season they flower and send out new shoots and leaves with the rains, much’ as in early spring in temperate regions. The forest canopy is on the average between 35 and 60 feet above the ground, depending on the dominant species which form it. It is sufficiently dense to allow little or no sunlight to reach the ground and, where it is best developed, the undershrubs and small trees are sparse, allowing visibility at the ground level for 75 to 100 feet. Except in ravines or other protected areas where conditions of mois- ture and humidity are higher, there are few mosses or other epiphytes on the tree trunks, which are mostly smooth and gray in color. Trunk diameters of trees forming the canopy are from 6 to 20 inches, but the majority are 10 or 12 inches. Rising above the canopy are occa- sional giant trees reaching about 100 feet in height with trunk diame- ters up to 3 feet. The three commonest are Bombax ceiba, Bombax sessilis, and Didymopanax morototoni; others less frequent are Ana- = AD , 2 é =, Fic. 1—Map of San José Island showing distribution of canopied forest in relation to scrub. Prepared July 1945, by C. O. Erlanson. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 cardium excelsum, Manilkara chicle, and Ficus. lf the rainfall were more evenly distributed throughout the year and more abundant, these giant trees might become frequent enough to form a canopy, and the forest would then approach the structure of a tropical rain forest such as is found on the Atlantic side of Panama. In regard to the dominant trees that form the canopy there are three types of forest on the island. One kind covers the entire southern half of the island up to the valley of the Rio Mata Puerco at the narrowest width of the island. Another type covers the northern half of the island, reaching its greatest development in the valley of the Rio Marina, and the third and smallest lies immedi- ately below the main camp in the small valley leading to Main Beach. Although the majority of species are found well distributed over the island, there are enough differences between the vegetation of the southern half and that of the northern half to give rise to the belief that earlier in geologic time, as the land rose from the sea, the two parts of the island were separated by a narrow channel of water somewhere in the vicinity of the Rio Mata Puerco, causing the two land masses to develop a vegetation with characteristic differences. The dominant trees of the forests in the southern half of the island are Cassipourea podantha, Ternstroemia seemannii, Pera ar- borea, and Ilex guianensis, which form a canopy with an average height of hardly more than 4o feet above the ground. Trunk diameters are usually about 10 inches. The bark is smooth, gray in color, and rarely with mosses or other epiphytes. Associated trees are Eugenia banghami and Xylopia aromatica, which form part of the canopy, and Hirtella racemosa, Ouratea wrightii, and Amaioua corymbosa, which commonly grow in the shade under the canopy as shrubs or small trees. The common vines hanging from the canopy as lianas are Connarus panamensis and Cnestidium rufes- cens. The ground is too shaded to allow much growth of herbs, but several ferns belonging to the genera Adiantum, Dryopteris, and Trichomanes are rather common, as also is the sedge, Rhynchospora cephalotes. In the deeper gullies where there is more moisture the black palm, Bactris balanoides, may appear together with other moisture-loving shrubs such as Inga and Pithecellobium. Where trees have fallen, leaving holes in the canopy for sunlight to reach the ground, a tangle of scrub growth soon forms an impenetrable thicket. Often the dominating plant in such openings is the climbing fern, Dicranopteris pectinata, or the vicious saw grass, Scleria secans. a NO. 2 VEGETATION OF SAN JOSE ISLAND—ERLANSON 7 The forests in the northern half of the island are made up largely of one species, Tetragastris panamensis, which forms a uniform canopy 60 to 70 feet in height with trunk diameters running from 10 to 20 inches. With this tree are associated a number of others, but seldom in quantities to form associations of their own. In the deeper valleys the wild cashew, Anacardium excelsum, becomes rather common along with the two species of Bombax. The giant tree Zanthoxylum sp. is sometimes seen with trunk diameters up to 4 feet. Other common trees forming part of the canopy are Cordia bicolor, Zue- lania guidonia, Ternstroemia seemannii, Protium sp., Didymopanax morototom, and Manilkara chicle. Under the shade of the canopy are various evergreen trees and shrubs, the commonest of which are Eugenia, Xylopia, Calycolpus, Hirtella, Amaioua, and Miconia. The black palm is plentiful along streams and dry gullies. In addition to the two common lianas found in the southern half of the island, there appears here as the most common vine a species of Bauhinia, locally called monkey-ladder. The shade of the canopy is sufficiently dense to discourage all but the most shade-loving plants, and much of the area can be traversed without the use of a machete. The third and most limited forest type, lying in the valley above Main Beach, has been separated from the other two extensive types because it contains several species found nowhere else on the island. Dominating a part of the forest are-two species of Guarea, G. parva and G. culebrana. Common here also is Gustavia superba, a relative of the brazil nut, Protium sp., and several other species rather remark- able for the island. In places the canopy reaches a height of 60 feet, and the forest here more closely approaches the rain forests of the mainland than anywhere else on the island. Throughout most of the area the black palm forms an understory layer 10 to 15 feet high, in places so thick that one can hardly squeeze between the slender thorny trunks. The ground is too shaded to support any grasses or other thin-leaved herbs. Only occasionally occurs an evergreen fern of the maidenhair type, Adiantum, or some species of Dryopteris. On the older tree trunks epiphytes are fairly common, including the orchid Mazillaria, the fern Lygodium, and various mosses and liverworts. On the rocky bluffs of the coast and less frequently inland wherever rock outcrops occur, a special association of trees replaces the regular types. The two characteristics that these trees have in common is a very shallow horizontal root system which allows them to take advantage of the thin layer of soil, and the ability to drop their leaves during the dry season when the soil can obtain practically no ® 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 subsoil moisture because of the impervious rock. The largest in this series of trees is the cigar-box cedar, Cedrela fissilis, which some- times forms dense stands with a canopy about 75 feet in height. Associated with the cedar, but very rarely in homogeneous stands, is the smooth, brown, curly-barked gumbolimbo, Bursera simaruba, the nance, Byrsonima crassifolia, and two species of Luehea, L. seemannii and L. speciosa. Almost invariably on the ground beneath these trees may be found the spiny, narrow-leaved wild pineapple, Ananas magdalenae. The vegetation of sandy beaches is very characteristic and contains species seldom found elsewhere. The beach at Playa Grande is the best developed on the island and may be used as an example of the other smaller strips found scattered along the island’s generally rocky coast line. Just above high tide level on the bare sand may be found clumps of the beach morning-glory, /pomoea pes-caprae, common on most tropical beaches, along with the seagrape, Coccoloba uvifera. Farther up on the beach occur colonies of the ‘white spiderlily, Hymenocallis americana, and the sand-binding grass, Uniola pit- tiert, which has the power of growing to any length necessary to keep itself from being buried by the shifting sand. Above these pioneer plants usually occurs a low stabilized dune covered by low thicket containing a number of typical beach shrubs and vines. Commonest among the shrubs is the clambering Dalbergia browni, whose branches twist like tendrils to help it climb, and the ubiquitous yellow-flowered wild hibiscus, Hibiscus tiliaceus. Other woody plants found in the thicket are Rourea glabra, Caesalpimia crista, Dioclea, Elaeodendron, Cissus, Davilla, Terminalia, and Arrabidaea. Here and there is found a tree of the poisonous manchineel, Hippomane mancinella, with its glossy leaves and green, applelike fruits. Usually coming down to sandy beaches from inland is a small stream, and, if the valley is wide and the gradient gradual, the salt water of high tide passes up into the valley, making estuary swamps. Dominating such swamps is the mangrove, Rhizophora mangle, with its yellow-green leaves and twisted, interlaced trunks growing out of the mud. Perched on these trunks are the bromeliad, Vriesia sanguinolenta, tillandsia, and one or two species of orchids. Also growing out of the mud are the long-stalked leaves of the dumb cane, Dieffenbachia, and the large fern, Acrostichum aureum. Near the mouth of the estuary and spreading out on the sand of the beach are white mangrove, Laguncularia, the button-mangrove, Conocarpus, palo del sal, Pelliciera, and usually also a grove of coconut palms, Cocos nucifera. The four most extensive mangrove swamps occur NO. 2 VEGETATION OF SAN JOSE ISLAND—-ERLANSON 9 at the mouths of the Rio Mata Puerco, Rio Marina, at Main Beach, and in the Naval Cove. Occasionally plants usually found only on the beach invade the slopes of coastal valleys and form definite associations for some dis- tance inland. One such formation is common around the Main Camp area, consisting of Hibiscus tiliaceus and the giant cane, Gynerium sagittatum, whose plumy inflorescences may be seen for considerable distances waving above the scrub during the rainy season. On the cliffs of the coast, hanging to the bare rocks and exposed to the salt spray, are three species of cactus, one with 5-angled stems, Acanthocereus pentagonus, one with 3-angled stems, Hylocereus monacanthus, and the prickly-pear, Opuntia elatior. With the cacti there usually may be found an orchid with white and very fragrant flowers, Brassavola nodosa. On top of these cliffs where the soil is thin and dry for half the year, the frangipani, Plumeria acutifolia, with its white fragrant flower, Jatropha urens, with its stinging hairs, and Apeiba tibourbou, a small tree with sea-urchin-like fruits, are common. On these cliffs, with the beginning of the rains a number of very small, mostly annual, herbaceous plants belonging to the genera Polygala, Phyllanthus, Marsypianthes, Borreria, and others, shoot up rapidly. Probably the most extensive type of vegetation on the island is scrub jungle, covering most of the slopes that face the sea and occur- ring inland wherever the canopied forest has disappeared for one reason or another. Once the scrub jungle invades and dominates an area, it may take several hundred years for the forest again to gain a foothold. Undoubtedly large areas of the scrub now found on the island are the direct result of migratory agriculture of the past century. As mentioned above, places can be found where the bound- aries between forest and scrub are so artificial and clear-cut that only such an explanation will cover the distribution of the two types of vegetation. No particular species dominate the scrub jungle, but rather it is a conglomeration of all available shrubs and vines that thrive in open sunlight, the whole matted down into an impenetrable thicket by the preponderance of woody vines which grow so much more rapidly than the shrubs or trees, and are represented by so many different species, that other types of vegetation cannot compete against them. The scrub varies in height from 4 to 20 feet, depending upon the shrubs and small trees that act as foundational support for the vines. Shooting up through this jungle are scattered trees about 40 feet in height, soft-wooded and short-lived, which have managed 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 to grow fast enough to escape being swamped by the vines. The commonest of these are the guarumo, Cecropia arachnoidea, Cordia alliodora, Cordia bicolor, and Zuelania guidonia. In depressions the black palm is often found with the shrubs, and associated with it is Costus villosissimus, a member of the ginger family growing to a height of more than 9 feet. The commonest of the shrubs are Cal- licarpa acuminata, Solanum extensum, Miconia, and Hirtella ameri- cana. Of the vines, those belonging to the bignonia family are by far the most frequent and make a wonderful display of bloom with their large trumpet-shaped flowers, white, yellow, pink, or purple. Some of the genera of this large family commonly occurring here are Phryganocydia, Anemopaegma, Amphilophium, Arrabidaea, Ad- enocalymna, and Cydista. Other common vines are Gouania polygama, Tetracera oblongata, Davilla lucida, Bauhinia, Connarus, Rhodospathe, Serjania, Caesalpinia, Dioclea, and Cnestidium. ' A study of the invading plants on areas bulldozed clear of all vegetation and along the edges of roads made during the past 18 months gives some idea as to the species which first pioneer the area and lead to the ultimate formation of scrub thickets. The first plants to come in are seedling trees—not trees which ultimately make up the canopied forest, but weedy, short-lived species. The two com- monest are Cecropia arachnoidea and Trema micrantha, which grow so rapidly and thickly that at first nothing else gets a chance to survive in many places. Of these two, Cecropia is the faster grower, becoming 12 to 15 feet in height during one rainy season, as com- pared with Trema, which attains about 8 feet. Here and there along the roads the very fast-growing balsa, Ochroma lagopus, has come in, being seeded from two old trees growing in the valley of the Rio Mata Puerco. Balsas measured in April have grown as much as 10 feet by the end of June. Other slower-growing seedling trees coming in under the shade of Cecropia, Trema, and Ochroma are Zanthox- ylum, Byrsonima, Eugenia, and Pithecellobium. As these trees get a slow start, vines begin to appear between the seedlings, the com- monest being Phryganocydia, Gouania, Tetracera, Davilla, Anemo- paegma, and Arrabidaea, as well as the shrubs Callicarpa, Hirtella, and Solanum extensum. In a year’s time the vines have begun to get the upper hand, climbing up into the branches of the young trees which have slowed down their growth owing to competition with one another, and by the end of the second rainy season the trees are heavily matted down, only an occasional Cecropia showing its head above the tangled thicket with vines dangling from its trunk. Ap- parently the trees such as Trema and Ochroma eventually disappear NO. 2 VEGETATION OF SAN JOSE ISLAND—-ERLANSON If entirely and are replaced by other kinds such as Cordia alliodora, Cordia bicolor, and Zuelania guidoma, some of which manage to ' break through the scrub to grow into isolated trees 30 to 50 feet high, but I have seen little evidence of the scrub allowing trees of the canopied forest to gain a foothold. In the north-central part of the island, at Bald Hill, is a small area of perhaps Io or 15 acres in extent which is devoid of either forest or scrub but is entirely covered by erect bunch grasses, mostly Trachypogon. A similar area was seen by the writer on adjacent Rey Island near the village of San Miguel. During the dry season the grasses were crisp and withered and the soil between the clumps was bare. With the rains the clumps soon became green, and between the clumps appeared herbs such as S piranthes orchioides, Habenaria pauciflora, Hyptis lantanifolia, Curculigo scorzoneraefolia, Crotalaria pilosa, sedges, and grasses not seen elsewhere on the island. Around the edges of the bald occur the usual trees and shrubs of sunny places, with certain rarities such as Clidemia rubra and Cornutia grandifolia. This area of dominant grassland may be due to the soil, which appears to be different in color and texture from that found elsewhere. It has been interesting to note to what extent the common cos- mopolitan weeds have been able to invade the island during the 2-year period that it has been occupied. Weeds usually invade new territory by means of seed that arrive as contaminations in the seed of agricul- tural crops brought in for planting, but since the island’s occupation was not for the purpose of agriculture and little seed of any kind has been deliberately introduced, it is to be expected that the weed flora would be relatively small. Most weeds are annuals producing seed abundantly and, given a cleared area in which to establish themselves without competition, will spread very rapidly. Only about 35 species of weeds have been noted as probably introduced during the past 2 years, and the greater proportion of these have as their focal point the naval station on the southern shore of the island. Since few sup- plies are brought in at that point, the presence of so many weeds there would be surprising except that the Navy made an attempt to beautify the grounds about the buildings by sowing imported grass and other seed, which is never free from weed-seed contamination. The second greatest concentration of weeds is around the main beach, which was to be expected, the seeds probably arriving on the clothing of incoming personnel. On the bluffs at East Harbor occur a few weeds not seen elsewhere, and because they are so well established, I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 it is probable that they are relics of the weed flora brought in at a much earlier period by settlers. Only two of the introduced weeds are likely to become a nuisance to the personnel of the island. Solanum scabrum, with its curved thorns and clambering growth, will tear clothing and rip the skin of anyone trying to pass through it. Cenchrus viridis is a grass pro- ducing very sharp-thorned burrs which, when ripe, fall on the ground and become a hazard to bare feet. The affinity of this grass for sandy beaches makes it all the more hazardous because of the presence there of bathers. It would not be difficult to eradicate these plants now, but given a few years to spread, they will constitute a much greater problem. The native sedge, called cut-grass or cortadera, Scleria secans, finds disturbed sections along roads and trails ideal for its growth and in some places covers considerable areas, growing to a height of I5 feet in roadside thickets. It is a very objectionable weed because its rough three-cornered’ stem easily cuts the skin, making painful sores. Probably the two weed trees Ochroma lagopus (balsa) and Trema nucrantha, so common along the roads, were brought in, but at some time previous to the Army’s occupation. One cannot help but speculate as to what prospects the island would have agriculturally if the land were properly handled. Under the economy practiced on Rey Island, a mixed fishing and migratory agriculture economy, perhaps as many as 100 families could main- tain themselves on a subsistence level. Any attempt to grow produce beyond the subsistence level brings in the problem of transportation to markets on the mainland, a considerable obstacle for an island in San José’s position. The product would have to obtain a high price to cover the cost of its transportation. The friable nature of the soil and its tendency to erode badly would necessitate extensive ter- racing to make much of the rolling land available to intensive agri- culture. The writer can think of no crop in the ordinary sense of the word that could be expected to bring in an adequate income under the circumstances. What could bring in a fairly comfortable in- come, if not to the present owners then to their descendants, would be the growing of tree crops—in other words, a well-balanced forestry program. It has been demonstrated that teak, mahogany, and African mahogany, to mention only three valuable woods, will thrive under conditions such as occur here. Aside from keeping the scrub growth from strangling the trees during their seedling stage, such plantations require a minimum of care and would eventually supply a steady annual income in valuable timber. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES 106; NO: 2; PEs 1 vex - 1. A VIEW OF THE GRASS-DOMINATED BALD HILL AREA 2. A COLONY OF SPIDERLILY, HYMENOCALLIS AMERICANA, GROWING UNDER COCONUT PALMS ON A SMALL SANDY BEACH SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 106; >NOS%2;5Pie2 1. SEEDLINGS OF BALSA, OCHROMA LAGOPUS, COMING IN ALONG ROAD BUILT ABOUT 6 MONTHS PREVIOUSLY 2. GRASSY COASTAL BLUFFS WITH FRANGIPANI, PLUMERIA ACUTIFOLIA, IN THE BACKGROUND ray es MOS. aye ie Pu UN ag ‘ 4k Ny Mi | \ _ SMITHSONIAN MISCELLANEOUS COLLECTIONS : VOLUME 106 NUMBER 3 ae | ALIST OF PRESH. WATER FISHES FROM SAN JOSE Fy ISLAND, PEARL ISLANDS, PANAMA BY ~* SAMUEL F. HILDEBRAND Ichthyologist, U. S. Fish and Wildlife Service ; (PUBLICATION 3847) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JUNE 10, 1946 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106 NUMBER 3 A LIST OF FRESH-WATER FISHES FROM SAN JOSE ISLAND, PEARL ISLANDS, PANAMA BY SAMUEL F. HILDEBRAND Ichthyologist, U. S. Fish and Wildlife Service (PUBLICATION 3847) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JUNE 10, 1946 The Bord Baltimore ress BALTIMORE, MD., U. & As A LIST OF FRESH-WATER FISHES FROM SAN JOSE ISLAND, PEARL ISLANDS, PANAMA By SAMUEL F. HILDEBRAND Ichthyologist, U. S. Fish and Wildlife Service No fresh-water fishes seem to have been recorded heretofore from the Pearl Islands. The species listed below were collected on San José Island in 1944 by Dr. Alexander Wetmore and Dr. Joseph P. E. Morrison. All the species taken occur also on the mainland of Panama. References are given for each species to the two general works on the classification of the fresh-water fishes of Panama listed at the end of this article. The earlier publication (1916) contains rather complete synonymies, keys, and descriptions ; and the later one (1938) brings the synonymies up to date and includes descriptions of species discovered in Panama since the publication of the earlier work. Family CHARACINIDAE GEPHYROCHARAX ATRICAUDATA (Meek and Hildebrand) Gephyrocharax atricaudata MEEK and HILDEBRAND, 1916, p. 277; Hildebrand, 1938, p. 253. Seventy-five specimens, 13 to 51 mm. long (U.S.N.M. No. 128473), pools near head of tributary of Rio Marina; II specimens, 33 to 50 mm. long (U.S.N.M. No. 128474), upper Rio Marina; and 28 specimens 31 to 51 mm. long (U.S.N.M. No. 128475), head- waters of Rio Marina. Common in the Chagres Basin and in the Pacific drainage from the Canal Zone to the Tuyra Basin. ASTYANAX RUBERRIMUS Eigenmann Astyanax ruberrimus MEEK and HILDEBRAND, 1916, p. 281; Hildebrand, 1938, p. 258. Two specimens, 55 and 68 mm. long (U.S.N.M. No. 128486), pools in upper Rio Mata Puerco. Both slopes of central Panama, ranging into Colombia on Pacific slope ; extremely abundant in vicinity of Canal Zone. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 3 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 tbo Family SYNBRANCHIDAE SYNBRANCHUS MARMORATUS Bloch Synbranchus marmoratus 1 HILDEBRAND, 1938, p. 202. Two specimens, 165 and 230 mm. long (U.S.N.M. No. 128476), from small stream at southwest part of island; 5 specimens, 82 to 430 mm. long (U.S.N.M. No. 128477), head of small stream in west drainage; I specimen, 245 mm. long (U.S.N.M. No. 128478), small stream in southwest drainage; 5 specimens, 35 to 77 mm. long (U.S.N.M. No. 128479), exact place of capture missing ; 2 specimens, 225 and 305 mm. long (U.S.N.M. No. 128480), south side of Hog Creek valley, above falls; 29 specimens, 100 to 410 mm. long (U.S.N.M. No. 128481), upper reaches of Hog Creek; and 65 speci- mens, 75 to 525 mm. long (U.S.N.M. No. 128514), middle reaches of Rio Marina. A widely distributed nocturnal fresh-water eel, ranging from México to, and probably beyond, the Amazon; occurring on both slopes of Panama. Family POECILIIDAE DARIENICHTHYS DARIENSIS (Meek and Hildebrand) Priapichthys dariensis MEEK and HILDEBRAND, I916, p. 321, fig. 7. Dariemchthys dariensis HILDEBRAND, 1938, p. 305. Forty-two specimens, 12 to 30 mm. long (U.S.N.M. No. 128482), small stream near north end of east beach; 3 specimens, 20, 24, and 25 mm. long (U.S.N.M. No. 128483), upper reaches of small stream flowing into Ensenada de la Bodega. Known heretofore only from the Pacific slope of Panama from La Venta to the Tuyra Basin. Family ELEOTRIDAE GOBIOMORUS MACULATUS (Giinther) Philypnus maculatus MEEK and HILDEBRAND, I916, p. 352. Gobiomorus maculatus HILDEBRAND, 1938, p. 340. Five specimens, 113 to 220 mm. long (U.S.N.M. No. 128463), pools in upper reaches of stream flowing into Ensenada de la Bodega: 1 Known from Panama since 1904, though not reported by Meek and Hilde- brand (1916) ; included, however, in their work, “The Marine Fishes of Panama,” Field Mus. Nat. Hist., Zool. Ser., vol. 15, pt. I, p. 131, 1923. NO. 3 FISHES FROM SAN JOSE ISLAND—-HILDEBRAND 3 I specimen, 170 mm. long (U.S.N.M. No. 128468), lower section of stream on southwestern tip of Island; 5 specimens, 112 to 184 mm. long (U.S.N.M. No. 128469), Rio Marina; 1 specimen, 108 mm. long (U.S.N.M. No. 128470), stream flowing into Ensenada de la Bodega, taken under a light at night. Pacific slope from Baja California to Ecuador; very common in Panama. ELEOTRIS PICTA Kner and Steindachner Eleotris picta MEEK and HILDEBRAND, I9QI6, p. 357; Hildebrand, 1938, p. 344. Eleven specimens, 97 to 325 mm. long (U.S.N.M. No. 128471), mid-reaches of Rio Marina ; and.1 specimen, 350 mm. long (U.S.N.M. No. 128472), lower section of stream flowing into Ensenada de la Bodega. Pacific slope from Baja California to Ecuador ; common in Panama. REFERENCES MEEK, SetH E., and H1ILpEBRAND, SAMUEL F. 1916. The fishes of the fresh waters of Panama. Field Mus. Nat. Hist., Zool. Ser., vol. 10, pp. 217-374, pls. 6-32, 10 text figs. HILDEBRAND, SAMUEL F, 1938. A new catalogue of the fresh-water fishes of Panama. Field Mus. Nat. Hist., Zool. Ser., vol. 22, pp. 217-359, figs. 2-13. 7 . he >a 1 tiie “ff . ‘ ul “ . " fai) , ; ri , we u . 4 * - a ‘ ‘ r, - , 7 * w~ ‘ ‘ . y i . i, ¥ ’ “7 w : "y - ’ ’ cw ie _ THE NONMARINE MOLLUSKS OF SAN JOSE ISLAND, WITH NOTES ON THOSE OF PEDRO GONZALEZ ISLAND, PEARL ISLANDS, PANAMA By J. P. E. MORRISON Assistant Curator, Division of Mollusks U. S. National Museum (WitH THREE PLATES) INTRODUCTION Under the joint auspices of the United States Army Chemical Warfare Service and the Smithsonian Institution, the writer spent the months of February to September, 1944, inclusive, on San José Island, Archipiélago de las Perlas, Republic of Panama. During this time all types of animals were collected for the United States National Museum, but particular emphasis on the molluscan fauna was the natural outcome of the writer’s primary interest. San José Island is the farthest outlying, as well as one of the largest of the Pearl Islands, lying approximately 60 miles south of Panama City, nearly in the middle of the Bay of Panama. Up to the time of these collections, the island had been uninhabited for some 70-odd years, although at various times earlier it had been cleared for agriculture over extensive areas. As a direct result of this earlier land use, the virgin jungle covered about half the island’s area, and the remainder was secondary in nature, with sour canebrakes and a matted ‘“‘vine” or “brush” jungle almost impossible to penetrate without cutting one’s way foot by foot. There is only a thin layer of leaf mold present anywhere on the island. The soil is a red clay, residual from the weathering of the volcanic breccia which forms the underlying rock of all the island surface. With abundant rainfall, and a maximum elevation of about 350 feet, the numerous, mostly small streams are for the most part rapidly flowing, ecologically young. Since there is no limestone on the island, the land and fresh-water snails must depend for lime on the small amount leached out of the volcanic rock present. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 6 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Pedro Gonzalez Island was visited very briefly on five different occasions, and mollusks were collected there also. It lies directly north of San José at about 5 miles distance, is at present inhabited, and does not exhibit the impenetrable secondary ‘‘vine” jungle habitat. On account of the limited time spent thereon, the mollusks of Pedro Gonzalez Island were very incompletely collected, but are included in this report as far as known. Three new genera, 27 new species, and one new subspecies, de- scribed herein, were found during the course of this study. This rela- tively high number of new forms represents a lack of previous knowl- edge of the fauna, rather than a high degree of endemism on the Pearl Islands. Further information on the general biology and character- istics of the Pearl Islands and San José Island in particular may be found in the other reports of this series.* The writer wishes to express his appreciation to the authorities of the United States Army and of the Smithsonian Institution, who made possible such an extended and intensive study of this insular fauna. In particular, thanks are due to Dr. Alexander Wetmore, whom I had the honor to assist in the field for the first 6 weeks of these studies, and to Dr. Paul Bartsch and Dr. Harald A. Rehder, also of the United States National Museum, for their most helpful advice and guidance in the preparation of this report. MOLLUSCA The molluscan fauna of the Pearl Islands is the fauna of the Panamic region in general. As far as known, the marine fauna is identical with that found on the Pacific shore of the Isthmus of Panama, with the possible addition of a very few endemic species. The nonmarine fauna, the restricted subject of this report, is a true insular fauna, showing clearly its direct relationship and deriva- tion from that of the Panama Isthmus. Its poverty in species may be considered likewise a direct result of the island’s geographic isolation from that isthmus. For the purposes of this report, all the land, fresh-water, and brackish-water mollusks are included. Included also are certain forms such as the littorinid snails which occupy habitats that are transitional to a true land habitat. 1 Smithsonian Misc, Coll., vol. 106, Nos. 1 to 7. NO. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 3 GASTROPODA PROSOBRANCHIA ° Family NERITIDAE Genus NERITA Linné, 1758 NERITA SCABRICOSTA Lamarck 1822. Nerita scabricosta LAMARCK, Hist. Nat. Anim. sans Vert., vol. 6, No. 2, sLOA. 1823. N ee cet, Sowersy, Gen. Shells, Nerita, fig. 4. Ubiquitous, on every rocky portion of the shores of San José and Pedro Gonzalez Islands, often in countless numbers. They occur on rocks in the intertidal zone at the mouths of certain streams, even on the same rocks with Neritina latissima, where salt concentration must vary from fresh to salt with every change of the tide. This change may not alter their activity much, however, since so much of the time they are not actually immersed in water, either salt in their normal habitat, or brackish to fresh at the transition area around the mouth of streams. NERITA BERNHARDI Recluz 1850. Nerita bernhardi Rectuz, Journ. de Conch., vol. 1, p. 285. 1851. Nerita funiculata MENKE, Zeitschr. Malak., vol. 7, p. 160. Not numerous, but found on both San José and Pedro Gonzalez Islands. Seen on rocky shores, principally in the intertidal zone, and along the sea-water side of some of the mangrove swamps. This species may be distinguished from the more abundant N. scabricosta by the lower spire and proportionately more effuse aperture. Genus THEODOXUS Montfort, 1810 THEODOXUS LUTEOFASCIATUS (Miller) 1833. Neritina picta SowerBy, Proc. Zool. Soc. London, 1832, p. 201 (not of Férussac, 1825). 1879. Neritina picta luteofasciata MILtER, Malak. Blatt., n.s., vol. 1, p. 168. 1923. Theodoxus luteofasciatus H. B. BAKer, Proc. Acad. Nat. Sci. Phila- delphia, vol. 75, p. 158. The individuals of this species collected on San José Island have the same obscurely reticulated dark greenish color as the Neritina latissima from the island. The similarity to some young individuals of N. 1. globosa is confusing. It may be differentiated by habitat, as far as personal experience goes; luteofasciatus is to be found on the Ay SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 margin of mangrove swamps, on the muddy margin of lagoons or seepages of those swamps, but not in running fresh water, which is ‘the habitat of NV. latissima. The proof of distinction lies in the slight difference in proportions of shell, prominence of opercular hypophyses, and the distinct micro- scopic beading. In contrast, the fresh epidermis of N. latissima shows almost regular threaded spiral sculpture, but it is not beaded, and usually shows a more velvety surface. Genus NERITINA Lamarck, 1816 NERITINA LATISSIMA Sowerby 1833. Neritina latissima SowerBy, Proc. Zool. Soc. London, 1832, p. 200. 1836. Neritina latissima Sowersy, Conch. Ill., Neritina sp. No. 20, figs. 3, 16. The typical. (extreme) alate form of adults was abundant in several streams on the Island of San José. On the other hand, local geography or stream size or elevation above sea level seemed to have little apparent effect on the variations of this enormously widespread species. The phase of the species (shell form and average adult size) seemed to be set for each stream in which they were found. For example, of the six streams, stream courses, or seepages in which this species was collected on Pedro Gonzalez Island, only two showed any trace of the alate condition, one north- and one south-flowing stream, both in the eastern part of the island. Six stream channels examined on the north side, west of the village of Cocal, ranging from a fresh- water seepage at high-tide line containing Littorina conspersa also, to a permanent stream up to 50 feet elevation and up to 200 yards inland, all showed the globosa form exclusively. Likewise, the globose, intermediate, and alate forms were all found in dominance on the west side of San José, but in different streams. In the Rio Marina on the east side, and in the small southwest streams it was evident that the alate maximum adults were to be found at the upstream limit of their zonal occurrence in the fresh water. In only one medium small stream on the west side did globosa go all the way up to the headwater pools as it did in one stream on the north side of Pedro Gonzalez Island. This difference is probably due to some unmeasured local chemical or physical difference of the stream habitat. Neritina latissima latissima, N. 1. intermedia, and N. l. globosa were all found on the Island of San José; Jatissima and globosa forms are both found on Pedro Gonzalez Island. NO. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 5 The globosa form was taken at the mouth of at least one small stream on San José, on the rocks in company with Nerita scabricosta; its habitat must here alternate between salt and fresh water with every tide! This species as seen in the Rio Juan Diaz, a little east of Panama City, Panama, was predominantly of the intermedia form. Genus NERITILIA Martens, 1879 Genotype.—N. rubida Pease. The distribution of this genus is most interesting. The genotype is widespread in the West Pacific; N. succinea and guatemalense are known from the West Indies and Central America in the West Atlantic. The following new species from the (Panamic) East Pacific region closes one gap in the tropical circle of its distribution. NERITILIA PANAMEWNSIS, new species Plate 1, fig. 2 Shell small, obliquely neritiform, corneous (covered with a blackish deposit in the types seen), the spire hardly projecting above the volu- minous body whorl. Nuclear whorls minute, smooth; postnuclear whorls 14, rapidly expanding, regularly ornamented with fine lines of growth. The aperture is large, suborbicular, with a columellar callus as wide as the operculum ; the columellar margin is not straight but sinuous (pinched-in medially), producing a subelliptic opening to be closed by the thin, translucent, pinkish horn operculum. The operculum is subelliptical, irregularly convex externally with a well- defined, humped-up ridge just at the columellar edge of the opercular nucleus. The nucleus is placed two-fifths of the distance from the base, and exactly opposite the slight ridge which continues obliquely downward on the internal face of the operculum to form the single, well-defined peg. The operculum is strengthened within with a transparent shell layer which continues at an angle as the raised “margin” of the baso-columellar region, and is glossy within except for the two distant, subtriangular, impressed muscle scars. The type, U.S.N.M. No. 542133, has 1.5 whorls and measures: Height, 2.9 mm.; diameter, 4.4 mm.; aperture height, 2.9 mm.; aperture diameter, 3.5 mm. It and 62 paratypes, U.S.N.M. No. 542134, were taken in company with Neritina latissima intermedia and thousands of Zetekella frenata, Cochliopa. diazensis, and Cochliopina zeteki along the east bank of the Rio Juan Diaz near Las Sabanas Road east of Panama City. This is in the zone of rapid water just a little above tidewater. One 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I06 large lot of the amnicolids was screened principally from rocks in the edge of the current. A second large lot screened from roots and plants along the sandy and muddy margin of the river contained only five additional specimens, U.S.N.M. No. 542135. These speci- mens from the muddy margin were covered with a blackish (iron?) deposit, as were most of the amnicolids from this spot. No special search was made here for Neritidae; this new species is apparently present in moderate numbers on the rocks along with the Neritina. This species may be easily distinguished from rubida Pease by the lower spire, the plane or concave, not convex, columellar callus, and the more widely effuse aperture. It differs from all other known species of Neritilia in the position of the opercular nucleus which is subcentral, near the columellar margin, and not subbasal. Family PILIDAE Genus POMACEA Perry, 1811 : POMACEA CUMINGII (King) 1831. Ampullaria cumingti Kinc, Zool. Journ., vol. 5, p. 344. 1856. Ampullaria cumingii Reeve, Conch. Icon., vol. 10, Ampullaria No. 81, DL aIG A sakes tei The Island of Taboga is given as the type locality. The National Museum collection contains one badly worn or acid-cleaned topotype, U.S.N.M. No. 4673, which measures: Height, 34 mm.; diameter, 32.5 mm.; aperture height, 24 mm.; aperture diameter, 17.5 mm.; umbilicus diameter, 3.5 mm. A small form of this species is known from Pacheca Island, the northernmost of the Pearl Islands. A representative individual, U.S.N.M. No. 432879, one of several recently received from James Zetek, has the following measurements: Height (apex eroded), 27.5 mm.; diameter, 27 mm.; aperture height, 20 mm.; aperture diameter, 15 mm.; and umbilicus diameter, 2.7 mm. This species is not found on the Isthmus of Panama as far as known to the writer. All specimens in the National Museum from Panama in the Atlantic drainage, as well as Pacific (Isthmus) streams, belong to the following species (zetek7). POMACEA CUMINGII SANJOSENSIS, new subspecies Plate 1, fig. 1 Shell of medium size, globose, openly umbilicated, solid. The surface dull, ornamented with numerous fine lines of growth, and No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON g minute lines of beaded microscopic sculpture, greenish to dark greenish or blackish horn colored, with numerous (8 to 10) subequi- distant spiral bands of a mahogany reddish brown color. Whorls remaining 4, well rounded, separated by moderately deep sutures, spire usually eroded, variable in height, usually comprising about one-third the shell height. Aperture large, subauriform, almost en- tire; the lip heavy, somewhat flaring, but not reflected. The outer lip is evenly rounded, the base well rounded; the columellar lip is rounded below, straight above across the parietal callus. The aperture is slightly flared, the lip all in one plane except for the heavy parietal callus which is carried forward slightly on the parietal wall. The aperture is hardly insected by the penultimate whorl. The type, U.S.N.M. No. 542136, has 5 whorls remaining, and measures: Height, 44.0 mm.; diameter, 46.5 mm.; aperture height, 33.5 mm.; aperture diameter, 24.5 mm.; and umbilicus diameter, 6.0 mm. This large, heavy subspecies was found in only three small streams (not of contiguous drainage) on the west side of San José Island. It differs from cumingu by its heavier, broader shell, more com- pletely rotund in adult examples, and by its much wider umbilicus. It is faintly remindful of the group of glauca L., from northern South America (Atlantic drainage), but its closest relatives are probably in the region of Rio Tuyra near the Colombian border of Panama, if one specimen of doubtful identification from the Rio Sambt is any indication. Its occurrence in these three smaller streams only, out of the dozen or more permanent streams of suitable habitat personally examined on San José Island, is entirely fortuitous, and could not have been brought about by stream capture. The reproduction of this species is possibly somewhat continuous, as young of various sizes were seen on several occasions. Egg masses in various stages of hatching were seen along with individuals in coitus on March 5, 1944, during the dry season. It is much larger in size than any typical cumingii seen and has a larger umbilicus in proportion. The spire is higher although usually eroded in adults, and the fresh young shells of about 25 mm. are covered with numerous spiral rows of epidermal tufts, giving the shell a velvety appearance. The largest of many specimens seen measures: Height (apex eroded), 52 mm.; diameter, 53.5 mm.; umbilicus diameter, 8 mm. The smallest adult shell measures: Height (apex eroded), 37.5 mm.; diameter, 40.5 mm.; and umbilicus diameter, 4.8 mm. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 POMACEA ZETEKI, new species Plate 1, fig. 3 1899. Ampullaria cumingi MartENs, Biol. Centr.-Amer., Mollusca, p. 422, in part. Shell of medium size, globose, narrowly umbilicate, moderately solid. The surface is dull to shining with fine lines of growth and minute lines of beaded microscopic sculpture, corneous, with nu- merous subequal and subequidistant spiral bands of mahogany color within. Whorls well rounded, separated by a moderate suture. Body whorl a little more abruptly rounded above the periphery, less so below. Spire moderate, about one-third the shell height. Aperture large, ovate-semicircular, somewhat effuse below, lip heavy, thickened but not reflected. The outer lip is evenly rounded in an almost even curve from suture to base; columellar lip well rounded into. the basal curve. The aperture is slightly insected by the penultimate whorl in the region of the moderately thin parietal callus. The holotype, U.S.N.M. No. 542137, has 43 whorls remaining and measures: Height, 40 mm.; diameter, 40 mm.; aperture height, 30.5 mm.; aperture diameter, 23 mm.; umbilical diameter, 3 mm. It and numerous paratypes, U.S.N.M. No. 542138, were person- ally collected from the shallow margin of the Chagres River near Gatuncilla, Republic of Panama, on October 6, 1944. It is named in honor of James Zetek, who has done so much to promote the scientific knowledge of the animals of Panama. Von Martens did not differentiate between the isthmian form as found in the Chagres River and the true cumingii from Taboga Island in the Bay of Panama. P. zeteki (cumingu of authors generally) is easily distinguished by the slightly shouldered appearance of the whorls and the narrow umbilicus. The numerous bright reddish-brown color bands show through the greenish horn epidermis and are a rich mahogany color within the aperture. In clean specimens the microscopic sculpture is seen as spiral bands of scars of the epidermal tufts which produce the velvety appearance of the young shells. Family LITTORINIDAE The littorinid snails are usually considered as marine in habitat, while actually many of the species occupy transitional habitats. It has been actually proved by experiment that certain species of Littorina have an optimum environment of an occasional immersion in salt water (or regular immersion at each high tide) with long No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 9 periods out of water in the air. The consideration of the littorinid snails as occupying a marginal land habitat is further strengthened by their close morphological relationship to the members of the family Cyclophoridae, which are exclusively terrestrial in habit. Genus LITTORINA Férussac, 1822 On San José Island the genus Littorina is well represented, both in species and number of individuals. In the mangrove swamps these snails are usually seen high up on the aerial mangrove roots or the tree trunks, often out of the reach of high tide. Here two larger species about an inch in length were found. LITTORINA ZEBRA (Donovan) 1825. Turbo zebra Donovan, Nat. Repos., vol. 4, pl. 131. 1832. Littorina pulchra SoweErsy, Gen. Shells, pl. 37, figs. 2, 3. This large reddish species has a shouldered body whorl and a striking design of narrow diagonal black stripes, for which it is named. LITTORINA FASCIATA Gray 1839. Littorina fasciata Gray, Zool. Capt. Beechey’s Voyage, p. 139. This large species possesses a more rounded body whorl, a little angulate at the base, and is spirally more roughly ridged than L. zebra. Also, the brown and gray mottling presents a much more subdued color pattern. ?LITTORINA GLABRATA Philippi A third species, much more strikingly conical in shape and inter- mediate in size, was found only in small numbers, and mostly in the young state, on the mangrove roots. It is tentatively identified as Littorina glabrata Philippi (Proc. Zool. Soc. London, 1845, p. 140). The few specimens seen from San José Island and elsewhere in Panama vary in color from light lemon yellow to brown mottled characteristically with darker brown. LITTORINA DEBILIS Philippi 1846. Littorina debilis Puttipri1, Proc. Zool. Soc. London, 1845, p. 140. 1847. Litorina debilis Puttipr1, Abbild. Conchylien, vol. 3, p. 11, Litorina, piyO; tip. 7. This small species, about half an inch long, was found in extreme abundance on and under the drift in the Rio Marina swamp, behind IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 the sand barrier at its mouth, especially on the rotting fronds of the coconut palm, along with great numbers of Detracia snails. It is long, conical, prettily mottled with brown, and varies considerably in the strength of color. An undetermined species of trematode worm was found in some of these snails. On the date of collection, August 2, 1944, these parasites were in the sporocyst stage in the bodies of the snails. On the rocks, near, at, or even above the high-tide lines, there are two species in great abundance: LITTORINA CONSPERSA Philippi 1847. Litorina conspersa Puttiprt1, Abbild. Conchylien, vol. 2, p. 200, Litorina, pl. 4, fig. 14. This species is usually almost entirely whitish in color, with a minute blackish apex. LITTORINA ASPERSA Philippi 1847. Litorina aspersa Puttipr1, Abbild. Conchylien, vol. 2, p. 200, Litorina, pl. 4, fig. 13. Littorina aspersa, in contrast to conspersa, is much darker in color, mottled with black and may have the tip of the shell eroded and white. Family FOSSARIDAE Genus FOSSARUS Philippi, 1841 Along with the species of Littorina on the rocks near or at the high-tide line are two species of Fossarus. Their habitat may like- wise be considered transitional to land or “marginal” land. FOSSARUS ANGIOSTOMUS (C. B. Adams) © 1852. Littorina (?) angiostoma C. B. ApAms, Cat. Shells Panama, p. 170. Fossarus angiostomus (C. B. Adams) is ovate-conic, about a quarter of an inch long, smoothish, and almost entirely white in color. FOSSARUS ATRATUS (C. B. Adams) 1852. Littorina atrata C. B. Apams, Cat. Shells Panama, p. 171. Fossarus atratus (C. B. Adams) is strongly spirally ridged and variable in color from white to heavily spotted with black. These No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON II members of the genus Fossarus may be distinguished from the littorinid snails, with which they occur, by their small size and the presence of a small umbilicus or central opening on the base of the shell. Family AMNICOLIDAE Subfamily HYDROBIINAE Genus ZETEKELLA, new genus Genotype.—Littoridina frenata Pilsbry=Zetekella frenata (Pils- bry). Shell imperforate, elongate-conic, smooth, with shallow sutures separating the appressed whorls. Spire moderate, acute. Aperture long-elliptical, acutely angled above. Columellar lip somewhat cal- loused. Operculum thin, corneous, paucispiral. Verge a little flattened, functionally simple, with a rather blunt tip, and furnished with two small appendages on the left side. The genus Zetekella is a well-marked group of Central American snails, known at present to be distributed from Nicaragua to Panama and the Pearl Islands in the Bay of Panama. As far as known, this genus is confined to streams of the Pacific drainage, above tidal influence. In addition to the genotype, Z. frenata (Pilsbry), from the Rio Juan Diaz, Panama, it is known to include: Z. martensi (Pilsbry), from the Rio Fula, Nicaragua, Proc. Acad. Nat. Sci. Philadelphia, vol. 87, p. 5, fig. 2, 1935; Z. melanoides (Martens), from the Rio de los Platanales, Costa Rica, Biol. Centr.-Amer., Mollusca, p. 436, pl. 22, fig. 8, 1899; Z. panamensis (Bartsch), from the Rio Matasnillo, Panama, Proc. U.S. Nat. Mus., vol. 58, p. 164, pl. 12, fig. 8, 1920; Z. veraguasensis and Z, kompi, new species, described herewith, from Sona, Veraguas, Panama, and San José Island, Pearl Islands, Gulf of Panama. Unfortunately, Z. panamensis (Bartsch) is apparently extinct. Its exact type locality is believed to be the Rio Matasnillo, on the eastern edge of Panama City. A search of this stream above tidewater on October 7, 1944, revealed no amnicolids; the stream’s condition on this date indicated heavy sewage pollution from the encroaching city. It may be noted here that members of the genus Zetekella and of the Cochliopa group are the closest morphological relatives, in the Panam4 region, of the Asiatic genera Oncomelania and Katayama. I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 The extinction of some of these species of the family Amnicolidae by sewage pollution in the Panama region is probably a condition of great medical importance. If we analyze this condition carefully, we find here biological proof that these species do not have the physi- ological or ecological ability to carry the trematode parasites of Asiatic human schistosomiasis (Schistosoma japonicum Katsurada). These facts will also explain why the Asiatic human schistosomiasis has never been endemic in Panama, in spite of the probability that infected human carriers have been present in the region of Panama City for the past three centuries. ZETEKELLA FRENATA (Pilsbry) 1935. Littoridina frenata Pirsspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 87, p. 5, figs. I, Ia. Z. frenata (Pilsbry) was taken in considerable numbers from the Rio Juan Diaz (type locality) east of Panama City, in company with Neritina latissima intermedia, Neritilia panamensis, Cochliopina seteki, and Cochliopa diagensis. It was especially abundant on the matted tree rootlets hanging in the water along the bank of the river. ZETEKELLA VERAGUASENSIS, new species Plate 2, fig. 1; plate 3, fig. 1 Shell small, elongate-conic, imperforate; the spire regularly conic (apex acute), of six flat-sided whorls. Suture linear, shallow, each whorl regularly increasing. Apex minute, apical whorl smooth, closely appressed to the preceding. The suture appears banded be- cause of a subsutural callus filling the appressed summit of the whorl. Postnuclear whorls smoothish, ornamented with minute growth lines and microscopic spirals. Aperture two-fifths of the shell, oval, acutely angled above, narrowly rounded below. Parietal margin al- most straight with a moderate callus on the lower part. Operculum thin, corneous, paucispiral, of about 3 turns; nucleus subbasal. The verge is as described for the genus, attached a little behind the right tentacle. This species may be distinguished by the large size, more obese body whorl, and almost straight parietal lip. The type, U.S.N.M. No. 542139, has 7 whorls and measures: Height, 6.2 mm.; diameter, 3.1 mm.; aperture height, 2.8 mm.; aperture diameter, 1.7 mm. It and several paratypes, U.S.N.M. Nos. 218175 and 516967, were collected by James Zetek from the Rio Tribiqué at Sona, Veraguas Province, Panama. No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 13 ZETEKELLA KOMPI, new species Plate 2, fig. 2; plate 3, fig. 2 Shell small, imperforate, elongate-conic; spire terete-conic (apex acute), of 6 almost flat-sided whorls; suture shallow, linear, between the faintly subangulate, regularly increasing whorls. Apex smooth (eroded ) ; postnuclear whorls, beginning with the second, ornamented with fine growth lines and prominent microscopic spiral lines. Aperture ovate, narrowed posteriorly, evenly rounded anteriorly. The parietal lip is arched somewhat to present a curve almost equal to that of the periphery. The lip is strong but thin, dark-edged, connected in adults by a moderate callus on the parietal wall. The operculum is thin, corneous, paucispiral, of about 2 turns, with subterminal nucleus. The animal is as described for the genus, the verge with two small appendages on the left (forward) side. ’ The type, U.S.N.M. No. 542140, has 6 whorls remaining, and measures: Height, 5.8 mm.; diameter, 2.6 mm.; aperture height, 2.3 mm.; aperture diameter, 1.66 mm. It and numerous paratypes were personally collected from the Rio Mata Puerco and its branches on the west side of San José Island, Archipiélago de las Perlas, Panama. This species may be distinguished by its large size and faintly shouldered whorls. Named for Col. W. H. W. Komp, of the United States Army Sanitary Corps, whose malarial research led to its discovery. Genus AROAPYRGUS H. B. Baker, 1931 Shell amnicoloid, with minute spiral striae on a smooth shell. Operculum thin, corneous, paucispiral. Verge: a large simple penis, unbranched, its terminal half smaller in diameter and pigmented. Genotype.—Aroa ernesti vivens H. B. Baker=Aroapyrgus ernesti vivens (H. B. Baker). The reproductive anatomy of all Panamanian species of “Ammnicola” available has been examined, and found to be that of this genus, now known definitely to occur from Panama to Cayenne. Determination as to which of the species of “Amnicola’ known from Tampico, Mexico, to Costa Rica, belong here also must await a check of their animal characters. A. panamensis Tryon agrees in shell characters; its male repro- ductive anatomy has not been examined. It is believed to belong here, nevertheless. In addition, there are the following new species in the Panama region. ¢ I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 106 AROAPYRGUS ALLEEI, new species Plate 2, fig. 4; plate 3, fig. 3 Shell small, ovate-conic, narrowly perforate, the spire almost regularly conic, of well-rounded, cbscurely shouldered whorls, sepa- rated by a deep suture. Aperture 0.4 of the shell height, narrowed and obtusely angled above, rather evenly rounded below. Parietal lip straight in contact with the penultimate whorl, then evenly rounded to the columellar region. Umbilicus narrow, behind the narrowly thickened lip. Sculpture of microscopic spirals and more prominent, irregular, widely spaced, raised growth lines. Plane of the aperture sinuous, produced a little peripherally. Operculum thin, corneous, paucispiral, of about 3 turns, with subbasal nucleus. Animal as described for the genus, with a large simple verge in the male, attached a little distance behind the right tentacle. The holotype, U.S.N.M. No. 542142, has 44 whorls and measures: Height, 3.1 mm.; diameter, 2.3 mm.; aperture height, 1.7 mm.; aper- ture diameter, 1.1 mm. It and numerous paratypes, U.S.N.M. No. 542143, were personally collected from dead leaves, sticks, etc., in a pool in the Allee Stream on Barro Colorado Island, Gatun Lake, Canal Zone, Panama, on October 8, 1944. It is named for Dr. W. C. Allee, my former professor at the University of Chicago, in whose honor this stream is named. U.S.N.M. ‘No. 432880 includes many additional paratypes col- lected by James Zetek. U.S.N.M. No. 474053 contains additional paratypes collected by Prof. H. J. Van Cleave, of the University of Illinois. This species may be distinguished by its obscurely shouldered whorls. It is larger than the specimen of panamensis Tryon (Rowell’s No. 20) in the National Museum collections. +] AROAPYRGUS CHAGRESENSIS, new species Plate 2, fig. 3; plate 3, fig. 4 Shell small, narrowly perforate, ovate-conic, with the spire conic, of moderately well-rounded whorls, separated by a deep suture. Aper- ture less than 0.4 the shell height, ovate, narrowed a little above, evenly rounded below, made entire by continuation of the lip across the parietal wall. The aperture is outwardly slightly effuse. Plane of the aperture a little sinuous, the thickened lip carried forward a little peripherally and on the parietal wall. Sculpture of microscopic spiral lines and more conspicuous, distant, irregular growth lines. The lip NO. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 15 callus is thickest in the upper parietal region. Operculum thin, cor- neous, paucispiral, of about 2 turns. Animal as described for the génus with a large, simple verge in the male, attached to the neck a little behind the right tentacle. The holotype, U.S.N.M. No. 542144, has 5.5 whorls and measures: Height, 3.7 mm.; diameter, 2.5 mm.; aperture height, 1.6 mm.; aperture diameter, 1.3 mm. The type and many paratypes, U.S.N.M. No. 542145, were screened from small beds of Chara and other plants in shallow water near the southeastern side of the Rio Chagres, near Gatuncilla, Re- public of Panama Another lot of specimens, U.S.N.M. No. 542146, was collected from plants in the rapids of the Rio Chagres, near Madronal, Re- public of Panama. This species is easily distinguished by its roundly ovate, slightly effuse aperture of adult specimens. The whorls are evenly rounded from suture to suture, not faintly shouldered as in A. alleei. AROAPYRGUS JOSEANA, new species Plate 2, fig. 5; plate 3, fig. 8 Shell small, narrowly perforate, ovate-conic to conic, the spire composed of very well-rounded whorls, separated by a deep suture. Apex smooth. Postnuclear sculpture of fine growth lines and faint microscopic spiral lines. The base is unusually well rounded, pro- duced by the marked rotundity of the axially short whorls, subor- bicular in section. Umbilicus distinct, narrow behind the unreflected columellar lip. Aperture ovate-suborbicular, about one-third the shell height, almost entire, ovate, narrowed above, almost evenly rounded below. Plane of the aperture a little sinuous, projecting slightly forward in the columellar region. Lip thickened somewhat, unreflected. The moderate lip callus is heaviest at the baso-columellar region. The operculum is thin, corneous, paucispiral, of about 2 turns. Animal as described for the genus. The males with a prominent simple verge attached on the right of the neck some distance behind the right tentacle. Its outer half (functional tip) is darkly pigmented and much smaller in diameter than the basal portion. It may be variously coiled in the retracted condition seen in preserved specimens. The holotype, U.S.N.M. No. 542147, has 5 whorls and measures ; Height, 3.1 mm.; diameter, 2.1 mm.; aperture height, 1.2 mm.; aperture diameter, 1.1 mm. 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 The type and 12 paratypes, U.S.N.M. No. 542148, were collected from a northern tributary of the Rio Mata Puerco estuary, San José Island, Archipiélago de las Perlas, Panama, about a quarter of a mile above tidewater, and near the upper limits of Amnicolidae in this small stream. In addition, this species was found in small numbers in two streams on the northwest part of San José Island, and in the Rio Marina on the east side. -Aroapyrgus joseana may be easily distinguished from the other Panamanian species by the axially shorter whorls and the rotundity of the base. In this last character it approaches A. ernesti (Martens) from Venezuela, but it may be distinguished from that species by its smaller aperture and narrower conical outline. Genus LYRODES Doering, 1884 Genotype.—Lyrodes guaranitica Doering, 1884. The genus Lyrodes is widespread from southern United States (Texas) to Central America, the West Indies, and south to the Argentine in Latin America. The absence of this genus from San José Island is noteworthy, in view of its widespread occurrence on islands of the West Indies. It may be that there are no sufficiently favorable habitats on the Island. The following new species from the Isthmus of Panama were located in the search for Panamanian species related to the island fauna : LYRODES CHAGRESENSIS, new species Plate 2, fig. 6; plate 3, fig. 6 Shell small, turreted-conic, narrowly perforate, translucent; apex acute. Nuclear whorls smooth, postnuclear whorls spirally lirate; the cord at the shoulder a little above the periphery becoming prominent to form a carina on the third whorl, often prominently nodose to form short “spines” on the fourth and fifth whorls, usually less prominent on the body whorl. The whorls are rather well rounded, separated by a deep suture and furnished with numerous equidistant spiral lirae above and below the shoulder. In the aspinose phase, the shoulder carina is discernible, but little larger than the other spiral lines, and the whorls are more evenly rounded from suture to suture. Aperture small, one-third the shell height, regularly ovate, narrowed very little above. The columellar lip is evenly arched, regularly rounded into the base. Operculum thin, corneous, paucispiral. No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 17 Animal as described for the genus; in the male with a prominent appendiculate verge on the right side of the neck. The verge has a tripartite appendage at the left side (forward) pigmented near the functional tip, and basally on the right a single row of fine short papillae ; female ovoviviparous. The holotype, U.S.N.M. No. 542149, has 6 whorls and measures: Height, 4.2 mm.; diameter, 2.1 mm.; aperture height, 1.4 mm.; aperture diameter, 1.1 mm. It and several paratypes, U.S.N.M. No. 542150, were personally collected from Chara beds, etc., in the shallow water near the margin of the Chagres River, near Gatuncilla, Republic of Panama, in company with Aroapyrgus and Pisidium. This species may be distinguished by its slender form and small aperture, with the characteristic sculpture most prominent on the fourth and fifth whorls, and reduced on the body whorl. The male examined and drawn (Jan. 16, 1946) proved this to be a typical member of the genus, variable as the shell is, on account of the pattern of the verge. This species has the narrow (lirate) form of the genotype variable to spinose (as in Pyrgophorus) and prob- ably proves that these two are just shell phases of one generic group. LYRODES ZETEKI, new species Plate 2, fig. 10 Shell small, elongate-conic, narrowly perforate, translucent, apex smooth, acute; postnuclear whorls spirally lirate and shouldered. The shoulder carina is most prominent on the fourth and fifth whorls, and often nodose to form the characteristic short “spines.” Whorls well rounded, shouldered, separated by a deep suture. In the aspinose form the whorls are narrower, more flatly rounded, the sculpture is much reduced, and may be without a discernible shoulder or spiral lirations on the body whorl. Aperture moderate, about three-eighths the shell height, ovate, narrowed above. The lip is thin, simple, in some individuals with a callus across the parietal region to produce an entire aperture. Operculum thin, corneous, paucispiral. Females ovoviviparous, the embryonic young in the uterus with short amnicoloid shells of 15 whorls. The holotype, U.S.N.M. No. 542151, has.53 whorls and measures: Height, 4.8 mm.; diameter, 2.6. mm.; aperture height, 2 mm.; aper- ture diameter, I.4 mm. U.S.N.M. No. 432871 includes many paratypes from the same lot collected by James Zetek at Pedro Miguel, Canal Zone, Panama. 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 This species from the Pacific drainage may be distinguished from L. chagresensis by its large aperture and less slender shape. The nodose spines (when present) are larger and less numerous than in that species. A critical examination of the species hitherto referred to the genus Cochliopa has shown that they fall into three groups, distinct from each other on the bases of shell and of anatomy. In fact all the species previously referred to Cochliopa, except the genotype, are proved by the present anatomical studies to differ generi- cally from that species. The species involved comprise three natural groups which are here separated. Genus COCHLIOPINA, new genus Shell conic-helicoid to planorboid, without umbilical keeling. The whorls are separated by moderate to deep sutures, not closely ap- pressed ; the sculpture is often composed of subequal spiral lirations, which may be distinctly colored. The aperture is oblique, subcircular to subtrapezoidal, usually almost or quite entire, sometimes a little solute in mature shells. Operculum thin, corneous, paucispiral, but almost multispiral in appearance, of 3 to 5 whorls, and has a sub- central nucleus. The radula has the formula: 47174 253 3e 13: «© (in C. minor). The verge is long, entirely simple, without appendages (pl. 3, fig. 7). Genotype.—Cochliopa riograndensis Pilsbry and Ferriss, 1906= Cochliopina riograndensis (Pilsbry and Ferriss). The distribution of known members of the genus Cochliopina is very interesting. All the northern species, ranging from Texas through Mexico to eastern Guatemala as far as Lake Izabal and the Rio Dulce, are found only in river systems draining into the Gulf of Mexico and the Caribbean. In contrast, all the species known from the southeast corner of Guatemala, southward through Nica- ragua, Costa Rica, Panama, and the Pearl Islands in the Gulf of Panama, occur only in streams of the Pacific drainage. Undoubtedly this interesting and apparently complete switch from one side of the Continental Divide to the other has its source in the biological history of this region in past geological time. It is extremely interesting to note that in the two known streams in which species of Cochliopina and Cochliopa are found living together, the Cochliopina species is much the smaller of the two. No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 19 An even more striking difference in ecological habit was observed on San José Island. The only Cochliopa found on the island was present from tidewater almost to the sources of the Rio Mata Puerco system. On the other hand, the Cochliopina found in that stream, and the ones found in five other streams on the island also, were markedly limited to a narrow habitat zone of the stream a little above tidewater. This genus probably includes, besides the genotype, the following: C. picta (Pilsbry), Rio Choy, San Luis Potosi, Mexico, Nautilus, wo 23, NOY 6,.p.,100,:pl., 9, .figs:, 1,2, IQ10; C. compacta (Pilsbry), Rio Choy, San Luis Potosi, Mexico, Nautilus, vol. 23, No. 8, p. 99, pl. 9, figs. 4, 5, 1910; C. infundibulum (Martens), Petén, Guatemala, Biol. Centr.-Amer., Mollusca, p. 429, pl. 23, fig. 3, 1899; C. francesae (Goodrich and Van der Schalie), Rio Pasion, Guate- mala, Univ. Michigan Mus. Zool., Misc. Publ. No. 34, p. 38, pl. I, fig 3, 1937; C. hinkleyi, C. izgabal, and C. izgabal mut. perstriata (Pilsbry), all from Lake Izabal, Guatemala, Proc. Acad. Nat. Sci. Philadelphia, vol. 72, pp. 198, 200-201, figs. 4, 6, 7, 1920; C. dulcensis (Marshall), from the Rio Dulce, Guatemala, Proc. U. S. Nat. Mus., vol. 58, p. 302, pl. 17, figs. 1-3, 1920; C. guatemalensis (Morelet), Rio Michatoya, near Istapa, Guatemala, Test. Noviss., pt. 2, No. 138, p. 22, 1851; Fischer and Crosse, Miss. Sci. Mex. Amér. Centr., pt. 7, vol. 2, Mollusques, p. 302, pl. 48, fig. 2, and pl. 50, fig. 1, Igo00. C. tryoniana (Pilsbry), western Nicaragua and southwestern Costa Rica, Nautilus, vol. 4, No. 5, p. 52, 1890; Proc. Acad. Nat. Sci. Philadelphia, 1891, p. 331, pl. 15, fig. 12, 1892; C. minor (Pilsbry), Polvon, Nicaragua, Proc. Acad. Nat. Sci. Philadelphia, vol. 72, p. 199, fig. 5, 1920; and the following new species: COCHLIOPINA ZETEKI, new species Plate 2, fig. 7; plate 3, figs. 5, 9 Shell small, helicoid, narrowly umbilicate, peripherally subangular, of 5 regularly increasing whorls. Apex low; nuclear whorls about 14; postnuclear whorls spirally striate or lirate; suture shallow. Sculpture variable, in most regularly spirally striate with subangular periphery, others peripherally carinate and striate above and below, in a few multicarinate, due to almost equal prominence of a few strongly developed spiral lirae. In all the base is well rounded. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Aperture subtrapezoidal, oblique, produced on the parietal wall. Lip simple, thickened within, the callus most prominent on the columella. Operculum thin, corneous, of about 5 narrow turns. Animal as de- scribed for the genus; male with simple verge on the back of the neck behind the right tentacle. The snout has two black bands across it, and the tentacles show two bands of black pigment, one at the middle and the second near the tip. The holotype, U.S.N.M. No. 542152, has 5 whorls and measures: Height, 3.1 mm.; diameter, 3.9 mm.; aperture height, 2.1 mm.; aperture diameter, 2.1 mm. The type was personally collected from the leaves and roots along the eastern margin of the Rio Juan Diaz, just below the bridge of Las Sabanas Road, east of Panama City, on October 7, 1944. U.S.N.M. No. 542153 includes several hundred paratypes from the same source. James Zetek had previously collected this species here on February 20, 1932 (U.S.N.M. Nos. 432877 and 432878), and it is named in honor of its discoverer. Formerly considered C. tryoniana Pilsbry, this species differs from that by having the shell less high in adult specimens. The noncarinate phase of this species is markedly lower than the tryoniana seen. COCHLIOPINA JURADOI, new species Plate 2, fig. 8; plate 3, fig. 10 Shell small, low, conic, narrowly umbilicate, peripherally rounded to subangular, of 54 regularly increasing whorls. Spire conic; nuclear whorls minute, 14, smooth; postnuclear whorls finely spirally lirate, sometimes with one subcarinate lira at the periphery. Whorls moder- ately flattened above the periphery, well rounded below. Suture moderately deep. Base well rounded with narrow funiculate um- bilicus. Aperture subtrapezoidal, oblique, produced at the posterior angle; base line of aperture evenly rounded from columella to pe- riphery. Lip thin, simple, markedly calloused on the columella. Operculum thin, corneous, of about 5 closely wound turns and sub- central nucleus. Animal with a prominent pigment band across the tentacles at their distal third, with simple verge attached far over on the right side of the neck a little behind the right tentacle. The verge is as described for the genus, the distal half more slender, and lightly pigmented. The holotype, U.S.N.M. No. 542154, has 54 whorls and measures: Height, 4 mm.; diameter, 4.6 mm.; aperture height, 2.1 mm.; aper- ture diameter, 2.3 mm. No. 6 MOLLUSKS OF SAN JOSE ISLAND—-MORRISON 21 U.S.N.M. No. 542156 includes several paratypes collected July 25, 1044. U.S.N.M. No. 542155 includes many more paratypes collected at the same spot on August 20, 1944. The type was personally collected from rocks in the rapids of the stream leading to the northwest mangrove swamp on San José Island, Archipiélago de las Perlas, Republic of Panama. The species is named for R. B. Jurado, civilian surveyor for the United States Army Engineers, whose joint resurvey of this stream with the writer led to its discovery, and whose exploratory “survey trails” opened many spots on San José Island for their first scientific scrutiny. C. juradoi may be distinguished by its regular conic spire and well- rounded whorls and base with narrow umbilicus. COCHLIOPINA FRATERNULA, new species Plate 2, fig. 9; plate 3, fig. 11 Shell minute, helicoid, openly umbilicated, of 43 whorls (in the type) ; apex low; nuclear whorls smooth; postnuclear whorls finely regularly spirally lirate, well rounded and separated by a moderately deep suture. Base rounded; umbilicus proportionately large. Aper- ture roundly trapezoidal, plane of aperture oblique, somewhat sinuous ; lip thin, thickened within, the callus heaviest at the base. Operculum thin, corneous, of closely wound turns with subcentral nucleus. Animal (male examined) with a black pigment band at the middle and a round black spot at the tip of the tentacles. Verge whitish, the slender distal half lightly pigmented, attached a little behind the right tentacle and a little to the right of the midline, on the back of the neck. The holotype, U.S.N.M. No. 542157, has 4 whorls and measures: Height, 1.5 mm.; diameter, 2 mm.; aperture height, 0.8 mm.; aper- ture diameter, 0.9 mm. It and 4 paratypes, U.S.N.M. No. 542158, and 1 specimen, U.S.N.M. No. 542159, were personally collected in the reaches just above tidewater of the Rio Mata Puerco, San José Island, Archipiélago de las Perlas, Republic of Panama. It is named “‘little brother” on account of its much smaller size than Cochliopa joseana in whose company it was found. Unlike that species (genus) it does not occur throughout the greatest part of the stream, but only in a definite zone a short distance upstream from the mangrove swamp. In this ecological character it agrees with all the other Cochliopina species known from San José Island. 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 It is easily distinguished by its minute size and well-rounded base with proportionately large umbilicus. It is the smallest Cochliopina known from the Panamanian region. A paratype of U.S.N.M. No. 542158 was dissected to check the male genitalia and furnished the sketch. COCHLIOPINA NAVALIS, new species Plate 2, fig. 12; plate 3, fig. 12 Shell small, helicoid, umbilicate, of 5 to 54 well-rounded, regularly increasing whorls. Spire subconic; nuclear whorls minute, 14, smooth; postnuclear whorls regularly spirally lirate, the lirae closely spaced and equally prominent. Suture near the apex well marked, deep, increasingly less prominent between the later whorls, sometimes very shallow on the body whorl. Base well rounded, with moderately open, funicuilate umbilicus. Aperture rounded, subtrapezoidal, the plane of aperture oblique; lip simple, much thickened within; the callus subequal all around. Operculum thin, corneous, of about 5 turns and with subcentral nucleus. Animal with the distal half of the tentacles darkly pigmented. The prominent verge with pigmented distal half is attached a little to the right of the midline, a short dis- tance behind the right tentacle. The type, U.S.N.M. No. 542160, has 54 whorls and measures: Height, 3.7 mm.; diameter, 4.6 mm.; aperture height, I.9 mm.; aper- ture diameter, 2.2 mm. It and numerous paratypes, U.S.N.M. Nos. 542161 and 542162, were collected from the stream flowing into the small bay on the southwest part of San José Island, at which is located the landing for the United States Naval lighthouse maintained on the southwestern part of the island. C. navalis may be distinguished by its larger size, lower conic spire, and slightly shouldered whorls. The base of the body whorl may be somewhat flattened. COCHLIOPINA EXTREMIS, new species Plate 2, fig. 11: plate ¢ ne. 13 Shell small, umbilicate, low-helicoid, of 4$ whorls. Apex low; nuclear whorls 15, smooth; postnuclear whorls finely but regularly strongly spirally lirate. The whorls are apparently a little wider than high, well rounded above, apparently somewhat flattened below, separated by a well-marked deep suture that often becomes shallower on the body whorl. The periphery is well rounded, not angulate. No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 23 Aperture roundly trapezoidal, very oblique, markedly sinuous, most recessive on the outer basal margin. Lip thin, heavily thickened within, the callus heaviest on the columellar side. Operculum thin, corneous, of about 5 turns as in the genus. Animal with the snout darkly pigmented, with two black pigment bands across it, the rear one a little obscure. The tentacles light, with a black pigment band across their middle and a black pigmented spot at their tip. The verge is proportionately large, with pigmented distal half, and attached far over on the right side, at the neck, close behind the right tentacle. The type, U.S.N.M. No. 542163, has 44 whorls and measures: Height, 2.5 mm.; diameter, 3.5 m.; aperture height, 1.5 mm.; aper- ture diameter, 1.6 mm. It and numerous paratypes, U.S.N.M. No. 542164, were personally collected from the small stream with a 100-yard lagoon just east of the southern tip of San José Island, Archipiélago de las Perlas, Panama. The habitat of this species is extremely limited in geographic space. The habitat zone for this species in this stream consists of only the last rapids of the stream just above the lagoon. Actually the species was found living in only 25 yards or less of the stream, at this point not even 25 feet wide. | C. extremis may be distinguished by the small size and the See essed appearance of whorls and shell. COCHLIOPINA AUSTRALIS, new species Plate 2, fig. 13; plate 3, fig. 14 Shell small, umbilicate, helicoid, of 54 whorls. Apex low; nuclear whorls minute, 13, smooth; postnuclear whorls subangular, strongly spirally lirate; usually the lirae are of unequal strength and strong lirae are separated by several fine ones, so as to produce a sort of multicarinate appearance. Whorls of the low spire are faintly shouldered, separated by a moderate suture which becomes shallow near the aperture where the shoulder angle disappears on the body whorl of adults. Base widely reamed around the moderate umbilicus, somewhat flattened near the aperture. Aperture large, subtrapezoidal, very oblique; lip simple, thickened within with a moderate callus. Operculum as in the genus, of about 5 turns with subcentral nucleus. Animal lightly pigmented, with one prominent black band and a second fainter one across the snout. Tentacles with one prominent black pigment band across their middle. Verge whitish, lightly pig- mented distally, and attached far to the right side a little distance behind and lower than the right tentacle. 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 The type, U.S.N.M. No. 542165, has 54 whorls and measures: Height, 3.6 mm.; diameter, 5 mm.; aperture height, 2.2 mm.; aper- ture diameter, 2.3 mm. It and many paratypes, U.S.N.M. No. 542166, were collected in the lowermost reaches of the Musselshell Creek, just above the lagoon. This Musselshell Creek opens on a small steep sand beach near the south (east) end of the island. C. australis may be known by its depressed conic shell of sub- angular whorls, large aperture, and lightly pigmented animal. The body whorl is apparently of much less axial height proportionately than in C. wetmoret. COCHLIOPINA WETMOREI, new species Plate 2, fig. 16; plate 3, fig. 15 Shell large (for the genus), umbilicate, regularly conic, of about 54 whorls. Nuclear whorls minute, 14, smooth; postnuclear whorls somewhat subangular, spirally lirate, narrowly shouldered below a distinct but shallow suture. The body whorl appears higher than wide and is depressed toward the aperture, so as to touch the penul- timate whorl only below the periphery over the last half whorl. The base of the body whorl is markedly flattened just in back of the aper- ture. The aperture is large, roundly subtrapezoidal, very oblique, usually made entire by the heavy callus within the lip which is heavy all around but thickest in the parietal region. Some adults seen have an almost solute aperture. Operculum thin, corneous, of about 5 turns as in the genus. Animal lightly pigmented, with two prominent black pigment bands across both the muzzle and the tentacles. One pigment band is about at the distal third of the tentacle, the other is almost terminal, a small white spot showing beyond at the tentacular tip. Verge large, simple and distally pigmented as in the genus, attached at a little distance directly behind the right tentacle. The holotype, U.S.N.M. No. 542167, has 4 whorls remaining and measures: Height, 4.4 mm.; diameter, 5.3 mm.; aperture height, 2.2 mm.; aperture diameter, 2.5 mm. One of the largest paratypes seen measures: Height, 5.8 mm.; diameter, 6.1 mm.; aperture height, 2.6 mm.; aperture diameter, 2.9 mm. It and thousands of paratypes were collected from the habitat zone of the species in the Rio Marina, the largest stream on San José Island. This habitat zone begins a little above the lowermost rapids of the stream and extends only about a quarter of a mile upstream, through a series of three sets of rapids and the intervening pools. This Rio Marina is the large stream that opens at the middle of the long sand beach on the east side of San No. 6 MOLLUSKS OF SAN JOSE ISLAND—-MORRISON 25 José Island, Archipiélago de las Perlas, Panama. Although the stream was personally traced to almost every branch of its head- waters, there are no Cochliopinas living beyond this short zone above the tidewater lagoon. Genus SUBCOCHLIOPA, new genus Shell conic to helicoid, usually peripherally keeled, and with spiral lirations which may be obsolete on the base which is somewhat flat- tened below the peripheral keel. Aperture oblique, subtrapezoid, produced at upper parietal region. The operculum is thin, corneous, almost multispiral in appearance, of about 4 turns, with a subcentral nucleus. Animal in the male with a simple verge, the functional tip (distal two-fifths) heavily pigmented, and at base of the pigmented area a very small appendage on the left (forward) side. Genotype.—Subcochliopa trochus, new species, described herewith. This new genus possesses spiral lirations similar to those of the Cochliopina group, accentuated in the direction of the peripheral keel and somewhat flattened base of the body whorl. On the other hand, the verge resembles that known for Cochliopa. While there is one small appendage in each case, the general appearance is different enough to indicate they are not identical (congeneric). The striking shell differences will make it easy to separate them from Cochliopa. While it is not yet proved by anatomical study, it is thought that Cochliopa trochulus (Martens)? from southwestern Costa Rica belongs here. SUBCOCHLIOPA TROCHUS, new species Plate 2, fig. 17; plate 3, fig. 20 Shell large (for the genus), regularly conic, narrowly umbilicate, peripherally carinate, of about 5 regularly increasing, more or less flat-sided, whorls; apex obtuse. Nuclear whorls 15, smooth (eroded in the type) ; postnuclear whorls finely and regularly spirally striate, peripherally carinate, and wound so that the suture is at the carina of the preceding whorl. Base somewhat flattened. Aperture sub- trapezoidal, oblique, carried far forward on the parietal wall, entire, the lip margins joined by a moderate callus on the parietal wall. Lip simple, thickened within, the callus heaviest on the columellar margin. Operculum thin, corneous, paucispiral, but apparently arcti- spiral, of 5 narrow turns, and with the nucleus subcentral. Animal in the male with a simple verge, the functional tip (distal two-fifths) 2 Biol. Centr.-Amer., Mollusca, p. 429, pl. 23, fig. 2, 1899. 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 heavily pigmented, and with a very small appendage on the left (forward) side at the base of the pigmented area. The type, U.S.N.M. No. 542168, and numerous paratypes, U.S.N.M. Nos. 217341 and 432875, were collected by James Zetek from the Rio Tribique, at Sona, Veraguas Province, Panama. The type has 54 whorls and measures: Height, 5.9 mm.; diameter, 6.7 mm.; aperture height, 3.3 mm.; aperture diameter, 3.4 mm. The largest paratype seen has 64 whorls and measures: Height, 7.5 mm.; diameter, 7.9 mm.; aperture height, 3.2 mm.; aperture diameter, 3.8 mm. This species is easily distinguished by its greater size and acute peripheral keel, constantly carinate periphery and regular conic shape. The base is more flattened in this species than in C. trochulus (Martens). SUBCOCHLIOPA COLABRENSIS, new species Plate 2, fig. 14; plate 3, fig. 16 Shell smail, helicoid, narrowly umbilicate, peripherally subangular, of 42 regularly increasing whorls. Apex low. Nuclear whorls minute, 14, smooth; postnuclear whorls spirally lirate. Suture well marked, shallow between the moderately rounded spire whorls. Body whorl finely regularly lirate above the prominent peripheral subcarinate liration, flatter below the peripheral angle, where medially the lira- tions become much finer, almost obsolete, and subangulate around the narrow umbilicus. The lirations near the umbilicus are again promi- nent. Aperture ovately subtrapezoid, oblique, much produced on the parietal wall. Lip thickened within, most prominently on parietal and columellar margins. Operculum very thin, corneous, of about 4 turns. Animal with muzzle generally dark, base of tentacles dark, obscuring the eyes. Tentacles light, with an interrupted black pigment band at the basal third and a black spot at the tip. The verge, large, simple, light, with one minute light appendage and the functional tip pigmented beyond this point, is attached to the right of the midline, a little distance behind the right tentacle. The type, U.S.N.M. No. 542169, has 5 whorls and measures: Height, 3.5 mm.; diameter, 4.6 mm.; aperture height, 2.1 mm.; aper- ture diameter, 2.3 mm. It and several paratypes, U.S.N.M. No. 218177, were collected by Meek and Hildebrand from the Rio Colabre, of the Bayano River drainage, Panama. This species may be easily distinguished by the marked flattening of the base of the body whorl below the subcarinate periphery. No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 27 Genus COCHLIOPA Stimpson, 1865 The small (typical) group of species comprises the genus Cochliopa. Genotype-—Amnicola rowelli Tryon=Cochliopa rowelli (Tryon). Unfortunately the genotype is now extinct, apparently having suc- cumbed to sewage pollution of its type locality, the Rio Matasnillo immediately east of Panama City. At least careful search of this stream October 7, 1944, revealed no amnicolids whatsoever living in the heavily polluted stream. Reexamination of type material of rowelli (U.S.N.M. No. 47855) has shown that the “bifid” condition is not a flagellate one as in the Bythinellinae, but is due to the possession of a simple verge with one small appendage, as described by Stimpson. A peculiarity of this species is the pigmentation of the tentacles, to give the appearance of false eye spots, a little distal of the midlength of the tentacles (pl. 3, fig. 17). pe Shell helicoid, rimate to narrowly umbilicate, smooth to spirally lirate ; aperture oblique, subrotund, angled or obtusely rounded above; whorls, especially the body whorl, closely appressed to the penul- timate and depressed below the suture. Operculum thin, corneous, paucispiral, of 2 to 3 turns, with the nucleus near the columella-basal margin. Radula with the formula 5—*—3 ; 3—1—4:18:24, in the genotype. Verge “rather elongated, compressed, geniculated, and bifid, the inner branch being very small, less than one-forth the size of the outer one and arising at the inner angle of the geniculation” (Stimpson). The genus Cochliopa, as here restricted to species having the shell characters and male reproductive anatomy of the genotype, comprises a group of species known only from the Republic of Panama. In addition to the genotype, only the following two new species have been definitely proved to belong to the group. COCHLIOPA DIAZENSIS, new species Plate 2, fig. 15; plate 3, fig. 18 Shell small, conic-helicoid, narrowly umbilicate, smoothish, of a few rapidly increasing whorls. Nuclear whorls minute, 14, smooth; early postnuclear whorls microscopically spirally striate, the lirae be- coming obsolete on the last 2 whorls, except in the umbilical region. Suture shallow, linear at the body whorl, which is narrowly concave below the suture. Aperture subpyriform, entire; plane of aperture 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 oblique, a little sinuous. Lip thickened within; callus heaviest along parietal and columellar region, subangular at base of columella. Umbilicus narrow, steep-sided, margined with a blunt angularity on base of body whorl. Operculum thin, corneous, paucispiral, of about 3 turns; nucleus subcentral. Animal lightly pigmented, except for the snout which is more or less uniformly black. Tentacles light, with a black band across the middle and an irregular black spot at the tip. Verge light-colored, very little pigmented, with a small branch on the left (forward) side near the midlength (in contracted spe- cimens). The holotype, U.S.N.M. No. 542170, has 44 whorls and measures: Height, 3.5 mm.; diameter, 4.1 mm.; aperture height, 2.3 mm.; aperture diameter, 2.1 mm. The type and hundreds of paratypes, U.S.N.M. Nos. 542171 and 542172, were personally collected from leaves, roots, etc., along the east bank of the Rio Juan Diaz just below Las Sabanas Road bridge, east of Panama City. James Zetek had previously collected this species here, but it has been hitherto considered as rowelli. It differs from rowelli by the higher, more regularly conic spire, and an aperture that is narrower above (posteriorly). COCHLIOPA JOSEANA, new species Plate 2; fig. 18;~plate 3) fig: 10 Shell large (for the genus), conic-helicoid, narrowly umbilicate, smoothish, of a few rapidly increasing whorls. Nuclear whorls minute, 13, ‘smooth; postnuclear whorls microscopically spirally striate, the striae obscured on the body whorl by irregular growth lines. Suture shallow, becoming linear along the body whorl which is narrowly concave below the suture. Aperture subpyriform, entire, plane of aperture oblique, somewhat sinuous at the subangular base of the columella. Lip thickened within, most prominently along parietal and columellar region. Umbilicus narrow, flat-sided, mar- gined with a blunt angularity and often irregular because of rough irregular growth lines. Base of body whorl swollen, narrowly rimate at the umbilicus. Operculum thin, corneous, paucispiral, of about 3 turns; nucleus hardly subcentral. Animal generally light-colored, with the snout black, and a black band across the middle of the tentacles and a black spot at their tips. Verge light-colored, with a pigment area along its middle on the left or incurved side in the region of the single small appendage, and with the functional tip un- pigmented. No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 29 The type, U.S.N.M. No. 542173, has 54 whorls and measures: Height, 5.1 mm.; diameter, 5.4 mm.; aperture height, 2.7 mm.; aperture diameter, 2.7 mm. It and numerous paratypes were collected from the Rio Mata Puerco, which opens out into the Ensenada de Bodega on the southwestern part of San José Island, Archipiélago de las Perlas, Panama. Unlike the members of the genus Cochliopina found on San José, this species is not restricted to a narrow zone of the stream above the mangroves, but lives almost up to the headwaters in all the branches of the stream. It was collected in the smaller northeastern tributary of the Rio Mata Puerco mangrove swamp, also indicating that it was probably living in the Rio Mata Puerco system before the main southern stream and this northern tributary were isolated at their mouths by the continued erosion and encroachment of the mangrove swamp. C. joseana may be distinguished by its large size, the greater height of the spire, and the narrow umbilicus. Family POTAMIDIDAE Genus CERITHIDEA Swainson, 1840 This group of snails occupies an ecological transition zone in the mangrove swamp between brackish-water and land habitats. In the swamps studied on San José Island they occur abundantly but only in certain habitats, not too close to fresh water. Under certain con- ditions, as in the estuary of the Rio Mata Puerco, two of the species are found on the sandy mud, out of water a greater part of the time, between times of high tide. On the other hand, the group was represented by all four species known from the Panamanian region in the Rio Marina swamp. Here they may be out of contact with salt water for months at a time, when the sand barrier cuts off the lagoon from the tide during the dry season, and they may be only occasionally in the brackish water. It is interesting to note that three of the species, C, hegewischu, C. valida, and C. pulchra, are found largely on the mud or sandy mud. On the other hand, C. montagne is found consistently a foot or more off the mud, on the roots of the mangroves, and roots and trunks of other trees, in the inactive state sealed to the trunks by mucus threads. Their shells were clean, in con- trast with the other species usually confined to the sandy mud or mud of the swamp floor. There is a marked difference in size between 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 individuals of each species from different mangrove swamps, as much as 100 percent difference in some cases. This difference in growth is apparently the result of some unknown food differences from swamp to swamp. . CERITHIDEA HEGEWISCHII (Philippi) 1848. Cerithium (Potamides) Hegewischii Puiciprt, Zeitschr. Malak., vol. 5, p. I9. 1849. Cerithiwm Hegewischiit Puiipri, Abbild. Conchylien, vol. 3, p. 15 (bis) Cerithium, pl. 1, fig. 6. This species may be distinguished by its slender shell with several varices or former lips, and a network sculpture of subequal vertical and spiral ribs, raised into rounded knobs where they cross. The lip of the adults found on San José Island is whitish as is general in those from the Panama region. CERITHIDEA VALIDA (C. B. Adams) » 1852. Cerithium validum C. B. ApaMs, Cat. Panama Shells, p. 157. This is the earliest valid name for the species often called varicosa. It was found living with C. hegewischii in the Rio Marina swamp without intergrades, hence is specifically distinct. C. valida is more massive, with a proportionately broader shell than hegewischii, of the same type of sculpture (knobbed, reticulate), and also with white lip in mature specimens. The largest valida collected on San José Island exceeds the published records for size, the largest one seen measuring 2 inches long and 1 inch in diameter at the aperture. CERITHIDEA PULCHRA (C. B. Adams) 1852. Cerithium pulchrum C. B. Apams, Cat. Panama Shells, p. 156. This interesting form is specifically different from any other Cerithidea seen. The shell is relatively wider, composed of more finely ribbed, well-rounded whorls, not as coarsely sculptured as in the other species of the region. In addition, the epidermis of this ,shell is markedly different, being very prominent in life, and con- sisting of many spiral rows of cuticular folds, giving the fresh shells a very velvety appearance when cleaned. The lip of these shells is a rich chocolate color. No. 6 MOLLUSKS OF SAN JOSE ISLAND—-MORRISON 31 CERITHIDEA MONTAGNEI (Orbigny) 1837. Cerithium Montagnei Orpicny, Voy. Amér. Meérid., vol. 5, pt. 3, Mollusques, p. 443, pl. 63, figs. 3-4. This species, as mentioned above, is normally found upon the mangrove roots and is clean-shelled. It may be recognized by the bare, coarse, vertical ribs on the turns, not especially marked by spiral sculpture. There is no duplication of former lips on the spire of these shells. The well-expanded aperture is smoky brown or dark chocolate in color within the lip of the same color. Family CERITHIIDAE Genus THERICIUM Monterosato, 1890 The genus Thericium is represented on San José by two species whose inclusion in this list is justified by their apparent ecological preference for situations of transition from salt to fresh water inter- mittently, such as the gravelly runs at the mouth of estuary streams freely running out of mangrove swamps, and at low tide pouring out almost fresh water, and tide pools filled at low tide with fresh water from intertidal seepage springs. THERICIUM ADUSTUM (Kiener) 1841. Cerithiwm adustum KIENER, Spéc. Gén. et Iconogr. Coq. Vivantes, Cerithium, p. 37, pl. 13, fig. 2. One such intertidal spring, brackish and almost fresh enough to the taste to drink, on the east side of San José, was the habitat of one colony of T. adustum extremely variable in size. The majority of individuals collected on April 9, 1944, were not quite mature, but the adults varied in size from a length of 37 mm. and a diameter of 21 mm. to the smallest, a length of 24 mm. and a width of 13 mm. It would be interesting to determine if the extreme change from full salt water to almost fresh at each tide has any causative effect on such extreme variation in size within the species. THERICIUM STERCUS-MUSCARUM (Kiener) 1841. Cerithium stercus-muscarum KiENER, Spéc. Gén. et Iconogr. Coq. Vivantes, Cerithium, p. 47, pl. 10, fig. I. Under estuary conditions this species was abundant. It was found on sandy or gravelly bottom in or near the channel pouring water out of the mangrove swamp with each tide at the mouths of the Rio Mata Puerco, and the stream at the head of the Navy landing station bay, both on the west side of the island. 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 PULMONATA BASOMMATOPHORA Family ELLOBIIDAE (salt-marsh snails) These characteristic salt-marsh snails form a conspicuous group of the molluscan fauna of San José, being well represented in the salt or brackish water to land transition zones. The small number of specimens of this family taken from Pedro Gonzalez Island was due to the lack of time available for examination of the salt-marsh areas. The total of 20 hours spent personally on Pedro Gonzalez was far too little to discover all the species of mollusks living there. Genus PHYTIA Gray, 1821 PHYTIA CONCINNA (C. B. Adams) 1852. Auricula concinna C. B. Apams, Cat. Panama Shells, p. 208. (24 miles east of Panama City.) 1920. Phytia brevispira PItsBry, Nattilus, vol. 33, p. 77, fig. Five specimens, U.S.N.M. No. 542174, were found in company with Ellobium stagnalis, Melampus piriformis, Detracia geteki, D. joseana, and D. graminea, under leaves on the flood plain of the Rio Marina, just above the mangrove swamp. The description of Adams’ concinna calls for the presence of a callus above the most prominent parietal lamella. Some few of the adults seen from near Panama City and from San José Island show a thin white callus in this position, irregularly ridged. The presence of intergrades, together with the known variability of strength in apertural characters in species of the Ellobiidae, leads the writer to regard Adams’ concinna as the heavily calloused form of the species Pilsbry has described as brevispira. Most of the young and adult shells seen totally lack this upper parietal callus. Genus TRALIA Gray, 1840 TRALIA PANAMENSIS (C. B. Adams) 1852. Auricula panamensis C. B. ApAMs, Cat. Panama Shells, p. 210. 1854. Melampus (Tralia) panamensis H. and A. ApAms, Proc. Zool. Soc. London, 1854, p. I0. 1900. Melampus panamensis Martens, Biol. Centr.-Amer., Mollusca, p. 561, pl. 43, figs. 10, 10a. C. B. Adams collected the species abundantly ‘“‘under stones, at high water mark, or crawling over wet stones,’ at Panama City, and also on Taboga Island. No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 33 It was found twice (in numbers only once) on San José Island, in the same ecological habitat. It was taken in numbers, although not as abundantly as M. tabogensis, in whose company it was, on and under cobbles and drift at high-tide line along one of the small bays on the east side of the island. It was active in the early morning out of the sun, crawling between cobbles and taking cover when the sun came up to dry off the cobbles of the beach, wet with dew. Genus DETRACIA Gray, 1840 DETRACIA ZETEKI Pilsbry 1920. Detracia zeteki Pitspry, Nautilus, vol. 33, p. 76, figs. a, b, c. Type locality—Panama City and Paitilla nearby. Collected in small numbers in mangrove swamp margins on both east and west sides of San José, near the mouth of the Rio Mata Puerco and the Rio Marina. One collection of several hundred in- dividuals was taken near the mouth of the Rio Marina, where they were abundant in and under drift, chiefly decaying coconut fronds, in company, with Littorina debilis in great numbers, and Detracia graminea. This species is very close to D. globulus (Orbigny) from Guayaquil, Ecuador, also reported from Tumacao Island, Colombia, but differs by the possession of the heavy parietal callosity and teeth. The species varies considerably in the largest lot collected (several hundred specimens); the variation is principally in the degree of development of the columellar fold and occlusion of the aperture, and in shape from regularly ellipsoidal to subglobular. DETRACIA GLOBULUS (Orbigny) 1837. Melampus margarita Becx, Index Moll., p. 107, No. 10 (nomen nudum). 1837. Auricula globulus OrpicNy, Voy. Amér. Mérid., vol. 5, pt. 3, Mollusques, Pin 327 1854. Melampus (Tralia) globulus H. and A. ApAms, Proc. Zool. Soc. London, 1854, p. II. 1854. Melampus globulus Preirrer, Novit. Conch., vol. 1, p. 23, pl. 6, figs. 23-25. 1901. Melampus globulus Koxsett, Martini and Chemnitz Conch.-Cab., vol. 1, pt. 16, p. 195, pl. 22, fig. 14. Type locality—Guayaquil, Ecuador. Close to zeteki Pilsbry, 1920, but without any parietal callosity of prominence. 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 DETRACIA STRIGOSA (Martens) 1900. Melampus strigosus MARTENS, Biol. Centr.-Amer., Mollusca, p. 560, pl. 43, fig. 9. (Tributary of Rio Coto, Golfo Dulce, Costa Rica.) Represented by topotypes, specimens from the original lot, U.S.N.M. No. 190285. Easily distinguished by the longer stripes, light color, and larger size of the extremely globular adults. DETRACIA JOSEANA, new species Plate 1, fig. 16 Shell small, obese-ovate, reddish brown, with spiral bands of lighter color. Spire banded, darkest brown, with a light band revolving a little below the upper limit of the aperture, body whorl indistinctly banded with lighter reddish brown. Spire angle of diver- gence about 90°, of closely wound whorls, separated by a linear suture. The aperture is narrow, almost three-quarters the length of the shell, sublinear above and contracted below by the moderately developed lamellae. Columellar fold is strong, regular, entering horizontally. There is a prominent callus on the columella below the lower of the parietal folds. The upper parietal fold is smaller, little developed in some of the young shells seen. The lirae within the outer lip are strong, more or less uniformly decreasing in promi- nence upward. The type, U.S.N.M. No. 542175, has 74 whorls and measures: Length, 6.2 mm.; width, 4 mm.; aperture height, 5.1 mm. It was collected from under fallen leaves on the flood plain of the Rio Marina at the upper limits of the tidal mangrove swamp at its mouth on the east side of San José Island, Archipiélago de las Perlas, Re- public of Panama. This species is very close to D. strigosa (Martens) from the Rio Coto, Costa Rica, but differs in being smaller, spirally banded with a reddish brown color over all, and with heavier apertural lamellae. Specifically, the columellar callus is higher; there are 2 lamellae present on the midparietal wall, and the stronger marginal lirae regularly decrease from below upward. In D. strigosa, the lira just above the single parietal lamella is more prominently de- veloped than those above and below. NO. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 35 This species was not found in great numbers, but was present in mangrove swamps on both east and west sides of San José Island, apparently preferring the inner landward or fresh-water edge of the swamps. It was also taken in company with D. zeteki and Phytia concinna at the margin of the Rio Matasnillo mangrove swamp, just east of Panama City, Republic of Panama. DETRACIA GRAMINEA, new species Plate 1, fig. 18 Shell small, regularly ellipsoidal, light horn color, banded with dark reddish brown. There is one broader, very dark band below the suture, a light band at the periphery, and three narrower dark bands almost equidistant below. The 10 closely wound whorls are separated by a sinuous, linear suture; the systrophic nuclear whorls are very small, projecting at the mucronate apex. Aperture narrow, approximately equaling two-thirds the shell height, linear above, wider below. The outer lip is thin, sometimes with 1 to 4 minute lirae below the periphery. The columellar plica strong, subhorizontal, with a narrow sinus above it. The parietal wall immediately above the sinus is furnished with two approximated, moderately strong lamellae with a third small lamella above. The type, U.S.N.M. No. 542177, has 9 whorls: and measures: Height, 5.3 mm.; diameter, 3.3 mm.; aperture height, 3.8 mm. The type and 7 additional specimens, U.S.N.M. No. 542178, were per- sonally collected August 2, 1944, in and under drift (chiefly decay- ing coconut fronds) on the mangrove-swamp side of the sand barrier near the mouth of the Rio Marina, Isla de San José, Archipiélago de las Perlas, Republic of Panama. They were taken in company with innumerable specimens of D. seteki and Littorina debilis. This species had been found previously under fallen leaves at the inner or landward edge of this mangrove swamp on the east side of the island, and of another smaller mangrove swamp on the west side. This species has also been collected on the Isthmus of Panama, U.S.N.M. No. 216282, in a tidal swamp near Corozal, Canal Zone, Panama, by James Zetek. This species is the smallest Detracia encountered on San José; it is about the height of D. joseana but much more slender. The colora- tion resembles that of light-colored young individuals of D. zeteki, but the apertural appearance is distinct. 36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Genus MELAMPUS Montfort, 1810 MELAMPUS PIRIFORMIS (Petit) 1843. Auricula piriformis PETIT, Proc. Zool. Soc. London, 1842, p. 202. (Tumaco Island, western Colombia.) 1852. Auricula trilineata C. B. ApAMs, Cat. Panama Shells, p. 212. (Panama City.) 1900. Melampus bocoronicus Martens, Biol. Centr.-Amer., Mollusca, p. 560 (in part) (not Morch, 1860). (Punta Arenas, Costa Rica.) This species is extremely variable as to size from swamp to swamp. It was taken in moderate numbers from mangrove swamps on both east and west sides of San José Island. MELAMPUS TABOGENSIS C. B. Adams 1852. Melampus tabogensis C. B. ApAMs, Cat. Panama Shells, p. 211. (Taboga Island and Panama City.) 1856. Melampus bridgesii CARPENTER, Proc. Zool. Soc. London, 1856, p. 161. (At mouth of the Bay of Panama.) 1896. Melampus panamensis Datu, Proc. Acad. Nat. Sci. Philadelphia, vol. 48, p. 452 (not C. B. Adams, 1892). (Cocos Island.) Seen from: San Lucas Island, Costa Rica (Valerio) (U.S.N.M. No. 381416); Taboga Island, Panama (Zetek) (U.S.N.M. No. 331820) ; Panama (U.S.N.M. No. 3807) ; Galapagos Islands (Dall) (U.S.N.M. No. 122016); Cocos Island (U. S. Fish Comm.) (U.S.N.M. No 122853). The Melampus panamensis of Dall reported from Galapagos Islands, U.S.N.M. No. 122016, and from Cocos Island, U.S.N.M. No. 122853, are not Tralia panamensis (C. B. Adams), but the large form of Melampus tabogensis (C. B. Adams) reported by Hanna and Hertlein.® Their remarks * on the comparative size of tabogensis and trilineatus refer only to the small form of trilineatus. The large trilineatus seen from Punta Paitilla and Panama City (collected by Zetek) and from San José Island, reach a maximum length of 20+-mm.(?). As they imply, Tralia has not been seen from Cocos or the Galapagos Islands. M. tabogensis was most abundant on cobble beaches and was seen (and taken) in great numbers on and under the cobbles and drift at high-tide line. It was raked by the handfuls off the bottom of drift logs on such a cobble beach on the west side of San José. Like Tralia 3 Allan Hancock Pacific Exp., vol. 2, No. 8, p. 133, 1938. 4Tdem, p. 134. No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 37 panamensis, which was found present less numerously, it was active in the early morning out of the sun; in early afternoon it was under the drift logs and cobbles. MELAMPUS BOCORONICUS Morch 1860. Melampus bocoronicus Morcu, Malak. Blatt., vol. 6, p. 118. Originally described from Los Bocorones near Punta Arenas, Costa Rica, “under stones near the beach.” The National Museum col- lections contain specimens (U.S.N.M. No. 380678) from Flamenco Island, Panama, collected by C. D. Alleman. Very close to M. tabogensis in general appearance, the shells of this species may be distinguished by the heavy white callus next above the columellar fold. Genus ELLOBIUM Roeding, 1798 ELLOBIUM STAGNALIS (Orbigny) 1835. Auricula stagnalis OrpicNy, Guérin’s Mag. Zool., vol. 5, cl. 5, No. 62, B23 1837. Auricula stagnalis OrBIGNy, Voy. Amér. Mérid., vol. 5, pt. 3, Mollusques, p. 325, pl. 42, figs. 7-8. (Guayaquil, Ecuador.) 1844. Auricula papillifera Ktster, Martini and Chemnitz Conch.-Cab., ed. 2, vol. 1, pt. 16, p. 25, pl. 3, figs. 9-10. (Western Colombia.) 1852. Auricula stagnalis C. B. Apams, Cat. Panama Shells, p. 210. (Panama City.) 1860. Ellobium papilliferum Morcu, Malak. Blatt., vol. 6, p. 116. (Sonsonate, El Salvador.) The National Museum collections include numerous specimens from the Pacific coast of Salvador (Biolley) (U.S.N.M. No. 163063), the Republic of Panama, Cocos, and Galapagos Islands, and Tumaco Island, Colombia (Cuming) (U.S.N.M. No. 119545). This species’ optimum habitat on San José Island seems to be in the decaying wood of logs and stumps about at the high-tide line in mangrove swamps. They were seen in greatest abundance on San José in a small mangrove swamp on the west side of the island, in rotting logs subject to partial inundation at the highest tides. More than a hundred specimens were collected here in about an hour, mostly from one large log. They were found burrowing in the rotten “wood floor” in almost every gallery or crevice in the more solid wood. Variation in size is very great in this species, as is the degree of surface sculpturing of the body whorl. Variation in size apparently 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 is controlled at least in part by food conditions affecting growth in general in the locale. Surface ornamentation varies from almost smooth to a completely decussated body whorl in different individuals from the same colony. Variation in comparative height of the spire is likewise individual. The size of adult individuals within a popula- tion does not vary nearly as much as their average size may vary from the average for another colony living less than a mile distant. It would be interesting to determine the effect on size of these snails of feeding on the rotting wood of different species of trees. This species was found drifted on the rocky shore near a large mangrove swamp on the north side of Pedro Gonzalez Island, indicating its presence there also. Family PLANORBIDAE Certain members of the genus Australorbis Pilsbry (1934, Proc. Acad. Nat. Sci. Philadelphia, vol. 86, p. 55) are known to be inter- mediate hosts of the parasite of African schistosomiasis (Schistosoma mansoni Sambon). In none of the collections made to date nor in any published record is there any indication that this genus of snails occurs in, or has been introduced in, the Panama region. Just as in the case of Asiatic human schistosomiasis, there has been an un- scheduled but natural and large-scale human experiment in the Canal Zone for a number of years. The fact is unquestioned that among United States troops of Puerto Rican origin, stationed in the Panama region during both World Wars I and II, there were numbers of human carriers of the parasite Schistosoma mansoni. In the absence of suitable intermediate host species of planorbid snails, this disease has simply not been able to become endemic either on the Isthmus of Panama or on San José Island. Although certain members of the genus Tropicorbis Brown and Pilsbry (1914, Proc. Acad. Nat. Sci. Philadelphia, vol. 66, p. 212) are under suspicion as possible intermediate hosts of S. mansoni, this same natural experiment offers conclusive proof that the native Panama species Tropicorbis isthmicus Pilsbry is not an effective intermediate host. The Island of San José is particularly free at the present time from such disease. In all the search for fresh-water mollusks in every body of water examined, only one specimen of planorbid snail was discovered. Apparently the ecology of stream habitats on the island is not favorable for the existence of this group of snails. No. 6 MOLLUSKS OF SAN JOSE ISLAND—-MORRISON 39 Genus DREPANOTREMA Crosse and Fischer, 1880 DREPANOTREMA ANATINUM (Orbigny) 1835. Planorbis anatinus OrpicNy, Guérin’s Mag. Zool., vol. 5, cl. 5, No. 62, p. 28. 1933. Drepanotrema anatinum Acuayo, Nautilus, vol. 47, pp. 64-68. One specimen only of this species was found in a small stream in the southeastern part of San José Island. Family ANCYLIDAE Genus FERRISSIA Walker, 1903 Subgenus LAEVAPEX Walker, 1903 FERRISSIA (LAEVAPEX) JOSEANA, new species Plate 1, figs. 5, 6 Shell small, elliptic, about three-fourths as wide as long, low, about one-fourth as high as long, pale corneous, thin, transparent. Nucleus large, without sculpture, remainder of shell finely radially striate. The anterior and posterior slopes are subequal in angle, the anterior rounding into the blunt apical curve, the posterior distinctly recti- linear. The apex is situated at the posterior third of the length, and approximately one-third of the width from the right margin. Lines of growth fine, inconspicuous. The. type, U.S.N.M. No. 542179, measures: Length, 4.2 mm.; width, 3.2 mm.; height, 1.2 mm. It and several paratypes, U.S.N.M. No. 542180, were collected from flood-plain pools along the middle reaches of the stream opening into the northwest mangrove swamp on San José Island, Archipiélago de las Perlas, Republic of Panama. This species may be easily distinguished from the widespread F. excentrica by the distinct radial sculpture. It does not have the sharp hooked beak of the described species of Uncancylus. The species is rather widespread on the island, being found also in the Rio Mata Puerco and Rio Marina drainage systems. It is the only fresh-water snail except Pomacea that is not restricted td a narrow supra-tidewater zone, but occurs in the middle and upper reaches of the streams, wherever the habitat is suitable. 40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 STYLOMMATOPHORA Family VERONICELLIDAE Genus VAGINULUS Férussac, 1822 VAGINULUS (LATIPES) OCCIDENTALIS (Guilding) 1925. Vaginulus (Latipes) occidentalis H. B. Baxer, Proc. Acad. Nat. Sci. Philadelphia, vol. 77, p. 174, pl. 5, figs. 18-20. This moderately large land slug (2 inches long) is of uncommon occurrence in the virgin jungle areas on San José Island. In the rainy season it may be found active in the daytime on overcast days or after a rain; otherwise, it is active only at night. Family COCHLICOPIDAE Genus CECILIOIDES Férussac, 1814 CECILIOIDES CONSOBRINA PRIMA (de Folin) 1930. Cecilioides consobrina prima (de Folin), Pirspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 82, p. 351. Only a single specimen was found on San José Island. The species may be present in greater abundance, but no special search was made for this minute subterranean form. Family SUBULINIDAE Genus SUBULINA Beck, 1837 SUBULINA OCTONA (Bruguiére) 1926. Subulina octona Pirspry, Proc. Acad. Nat. Sci. Philadelphia, tail 78, p. 89. This rather widespread land snail, which has evidently been carried by man’s agency over most of the American Tropics, was found in small numbers on San José Island. It was usually to be found in spots of second-growth “scrub” jungle, representing the formerly extensive cultivated areas of the island. In all probability it was introduced during some earlier period of occupancy of the island. Genus LEPTINARIA Beck, 1837 LEPTINARIA LAMELLATA CONCENTRICA (Reeve) 1926. Leptitaria lamellata concentrica Pirspry, Proc. Acad. Nat. Sci. Phila- delphia, vol. 78, p. 89, text figs. This, the largest (and widest) species of Leptinaria living on San José, may be easily distinguished by the presence of the revolving lamella upon the parietal wall. NQ. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 41 LEPTINARIA PANAMENSIS Pilsbry 1910. Leptinaria panamensis Picspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 62, p. 508, fig. 4. 1926. Leptinaria panamensis Pitspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 78, p. OI, text figs. This species is narrower and relatively smooth, without distinct riblets on the shell. On San José Island it seems to be more abundant than L. 1. concentrica. LEPTINARIA FILICOSTATA (Strebel) 1882. Lamellaxis filicostata STREBEL, Beitr. Kenntn. Fauna Mex. Land- und Stissw.-Conch., vol. 5, p. 113, pl. 17, fig. Io. 1926. Leptinaria filicostata Prtspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 78, p. 90, text figs. This species is about the size of L. panamensis but is easily dis- tinguished by the regularly spaced vertical riblets upon all the whorls. It seems to be moderately abundant on San José Island. Genus OPEAS Albers, 1850 OPEAS PUMILUM (Pfeiffer) 1906. Opeas pumilum Pitspry, Man. Conch., ser. 2, vol. 18, p. 200. 1926. Opeas pumilum Pitspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 78, p. 92, text figs. Found only once on San José island. OPEAS GRACILE (Hutton) 1906. Opeas gracile PirsBry, Man. Conch., ser. 2, vol. 18, p. 108. 1926. Opeas gracile Pirspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 78, p. 92, text figs. Found in moderate abundance, mostly in newly cleared areas on San José Island. Family OLEACINIDAE Genus EUGLANDINA Crosse and Fischer, 1870 EUGLANDINA CUMINGI (Beck) 1926. Euglandina cwmingi Prtspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 78, p. 95, pl. 10, figs. 3, 4. This fine, large carnivorous snail was found only in small numbers. Its more common occurrence in the areas of secondary jungle or brushy areas on San José points to its possible accidental introduction on the island. 42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Family ENDODONTIDAE Genus PUNCTUM Morse, 1864 PUNCTUM (?) COLOBA (Pilsbry) 1893. Thysanophora coloba Pitspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 45, p. 403. 1930. Punctum (?) coloba Pirssry, Proc. Acad. Nat. Sci. Philadelphia, vol. 82, p. 346, pl. 29, figs. 2-2b. This exceedingly minute land snail was found only once on San José Island, from the deeper leaf mold around the base of a large tree in a virgin jungle area near the eastern side of the island. Family ZONITIDAE Genus HABROCONUS Fischer and Crosse, 1872 Subgenus ERNSTIA Jousseaume, 1889 HABROCONUS (ERNSTIA) ZETEKI Pilsbry 1930. Habroconus (Ernstia) zeteki Prtspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 82, p. 350, pl. 20, figs. 5, 6. Found in small numbers in second-growth jungle, in places where there was sufficient moisture, such as in deeper ravines. Family BULIMULIDAE Genus DRYMAEUS Albers, 1850 DRYMAEUS TRANSLUCENS (Sowerby) 1926. Drymaeus translucens Picssry, Proc. Acad. Nat. Sci. Philadelphia, vol. 78, p. 83, text figs. 1930. Drymaeus translucens PitsBry, Proc. Acad. Nat. Sci. Philadelphia, vol. 82, p. 340. These smaller tree snails may be recognized by their pale yellow color and extremely thin shells. It is almost impossible to collect the San José individuals without crushing the shells between one’s fingers. They are found in the jungle areas of San José that have not been too disturbed by former plantation clearings, and overrun by vine brush. Unseen for the greater part of the year, they may be easily collected in April and May, when they are coming out of hibernation at the end of the dry season. The adults or nearly adult individuals pass the dry season in the leaf mold of the jungle floor, and a few No. 6 MOLLUSKS OF SAN JOSE ISLAND—-MORRISON 43 weeks after the onset of the rains, start climbing the tree trunks. At this time they are easily discovered, and easily collected in numbers from 2 to 20 feet off the ground on the trunks of certain small smooth- barked trees, which they seem to prefer. None of these arboreal snails were found on trees whose bark was excessively rough in character. In the series taken on San José Island there was con- siderable variation in the height of the spire. DRYMAEUS SEMIMACULATUS Pilsbry 1808. Drymaeus semimaculatus Pttspry, Man. Conch., ser. 2, vol. 11, p. 297. 1899. Drymaeus semimaculatus Pitspry, Man. Conch., ser. 2, vol. 12, pl. 5, figs. 8, 9. One specimen (a broken shell of an adult) was collected on San José Island. More’material is needed to determine its exact habitat on the island. Genus ORTHALICUS Beck, 1836 (not of Thiele) (= OXYSTYLA auct.) ORTHALICUS PRINCEPS DECEPTOR (Pilsbry) 1899. Oxystyla princeps deceptor Pttspry, Man. Conch., ser. 2, vol. 12, p. 116, pl. 24, figs. 19-25. This large tree snail may be easily distinguished by its size and prominent zigzag color pattern. Dead and bleached shells may be found on the jungle floor at any time of year, but collection of living individuals is much more difficult on San José Island. The eggs are laid in the leaf mold at the bases of trees, apparently at the beginning of the dry season (November—December). They are spherical, about 3 mm. in diameter, and covered with a thin white calcareous shell. At the beginning of the rainy season (April and May) they contain young with shells of 2 whorls, ready to hatch. Upon the onset of the rains in April and May they hatch, and the young start their arboreal existence. Half-grown young may be found singly upon the lower part of the tree trunks or upon the lower leaves of trees in midsummer (July and August). Unfortunately, no living adults were personally collected on San José, so it is not certain whether one or two seasons are required to reach maturity. It is also not known whether adults reascend the trees after laying their eggs in the leaf mold. 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Family FRUTICICOLIDAE Subfamily EPIPHRAGMOPHORINAE Genus AVERELLIA Ancey, 1887 . Subgenus TRICHODISCINA Martens, 1892 AVERELLIA (TRICHODISCINA) COACTILIATA (Deshayes) 1838. Helix coactiliata DESHAYES, Hist. Nat. Gén. et Part. Moll. Terr. et Fluv., vol. 1, p. 18, pl. 72, figs 1-5. This small discotdal species, neatly marked with brownish spiral stripes of color, was found in areas of virgin jungle on San José Island, but nowhere in abundance. Perhaps the presence of feral pigs on the island makes existence hazardous for these snails which live in the thin layer of leaf mold or under fallen logs. PREBCY PODA Family CORBICULIDAE Genus POLYMESODA Rafinesque, 1820 POLYMESODA JOSEANA, new species Plate 1, figs. 12-14. Shell large, equilateral, subtriangular, inflated, the ventral margin evenly rounded, anteriorly light colored, posteriorly violaceous under a moderately thick pale greenish-horn epidermis. Lines of growth fine, somewhat irregular, and interspersed by several ill-defined rest marks. Umbones uneroded, high, prominent, pale violaceous. Pos- terior ridge low, ending at the posterior angulation a little above the base. Secondary posterior ridges above this inconspicuous. Lunule broad, shallow, indistinctly defined. Ligament short, corneous. Car- dinal teeth well developed, laterals high, thin in proportion to the light shell. The unique holotype, U.S.N.M. No. 542181, measures: Length, 42 mm.; height, 38.5 mm.; diameter, 30.5 mm. It was found in the drift of the Rio Marina mangrove swamp on the east side of San José Island, Archipiélago de las Perlas, Republic of Panama. This large, thin species has no close relatives except P. maritima (C. B. Adams) described from Panama. It may be distinguished from that species by being smaller and proportionately somewhat higher ; although possessing the approximate size and shape of species from the Atlantic drainage in Nicaragua and Lake Maracaibo, it is at once recognized by its very thin shell. No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 45 Family CYRENOIDIDAE Genus CYRENOIDA Joannis, 1835 CYRENOIDA INSULA, new species Plate 1, figs. 8-11 Shell small, lenticular, suborbicular, vitreous, blue-white, under a pale corneous epidermis. Umbones smooth, little prominent, scarcely projecting above the general outline. Sculpture of minute, crowded concentric striae. Hinge well developed, lightly arcuate, wider an- teriorly. Anterior and ventral margins evenly rounded as one curve. Posterior margin almost evenly rounded, a trifle more abruptly rounded into the dorsal and ventral margins. The holotype, U.S.N.M. No. 542182, measures: Length, 6.7 mm. ; height, 6.2 mm.; diameter, 3.8 mm. It was collected from pools in the mud of a small mangrove swamp on the west side of San José Island. This species was found only in small numbers and did not seem to be abundant in any of the mangrove swamps on the island. It greatly resembles C. guatemalensis Pilsbry, 1920, from the Atlantic drainage in Guatemala, but may be easily distinguished by its much straighter hinge line. Family SPHAERIIDAE Subfamily PISIDIINAE F. C. Baker, emend. The principal anatomical character upon which the late F. C. Baker ° based this subfamily of “‘pill-clams’’ has up to this time not been correctly stated. Odhner, in 1921, briefly reported on the anatomy of Pisidium, stating that the genotype, P. amnicum (Muller) has two siphons, branchial and anal. He erected the subgenus Neopisidium with P. torquatum Stelfox as subgenotype, for those species possessing only the anal siphon. Examination of the animal of P. dubium (Say) 1816° from the Rappahannock River, Virginia, shows that it groups with the type of Pisidium, possessing both siphons, although the branchial siphon appears much smaller than the anal. Most of our American species undoubtedly belong to the Neopisidium group; all of several species personally examined alive or in preserved condition, except dubium, 5 Amer. Midl. Nat., vol. 10, p. 220, 1927, and Fresh Water Moll. Wisconsin, vol. 2, p. 363, 1928. 6 Pilsbry, Nautilus, vol. 59, p. 86, 1946. 46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 and obviously all species examined by Baker possess only the anal siphon. Therefore, the subfamily Pisidiinae must be amended to in- clude Pisidium “with an obsolete or without any branchial siphon.” We may thus avoid the absurdity of splitting the subfamilies in the middle of the genus. Genus PISIDIUM Pfeiffer, 1821 , Subgenus NEOPISIDIUM Odhner, 1921 PISIDIUM (NEOPISIDUM) sp.? A species of Neopisidium was found in most of the streams visited on San José Island, but only in small numbers. The animal of this Neopisidium possesses only the anal siphon, and only one pair of gills. The marsupium is on the inner face of the outer lamina of the gill, in those animals examined, which were collected in a tributary of the Rio Mata Puerco. On account of the lack of adequate material for specific comparison with others from the Panama region, it is thought best not to describe it at this time. In a group difficult to identify specifically, even under the best of conditions, another obscure species name would not be particularly advantageous. The study of complete seasonal series from different localities is desirable in this genus, in order fully to recognize the true specific characters. This is especially true in the tropical regions, where under accelerated life- history conditions adults may perhaps be found only at certain seasons of the year. Family DREISSENSIIDAE Genus MYTILOPSIS Conrad, 1837 MYTILOPSIS ADAMSI, new species Plate 1, figs. 4, 7 Shell small, mytiliform, dorsal and posterior margins rounded into one regular curve; ventral margin straight to incurved, abruptly rounded into the posterior margin at the posterior base. Ridge de- limiting the ventral flattening of the shell well rounded. Umbones smooth, terminal, usually eroded except in very young individuals. Septum narrow, furnished with a small, sharp triangular process in each valve. Substance of shell whitish to clear, vitreous, under a thin corneous epidermis which is irregularly pinched up into cuticular fringing folds along the lines of growth. These cuticular folds in some individuals are accentuated into triangular processes along two No. 6 MOLLUSKS: OF SAN JOSE ISLAND—MORRISON 47 distinct rows, one a little above the ventral ridge, the second midway between that and the dorsal margin. Animal as described for the genus, with a well-developed byssus, rufous in color. The holotype, U.S.N.M. No. 542183, measures: Length, 12.5 mm.; height, 6 mm.; diameter, 5.6 mm. It and numerous paratypes of all ages, U.S.N.M. No. 542185, were personally collected in the upper end of the lagoon at the mouth of Musselshell Creek, which is the largest of the streams in the southeastern part of San José Island. They were principally found attached by the byssus to the underside of rocks in the uppermost end of this (fresh-water) lagoon and in the lowermost part of the stream proper, in situations where there was plenty of stream current remaining. Reminiscent of M. leucophaeatus (Conrad) of eastern North America, this species can be distinguished by its greater proportionate diameter and regularly rounded posterio-dorsal margin. Some in- dividuals, heavily eroded and malformed in the umbonal region, are truly boxlike in appearance. This species is named in honor of Prof. C. B. Adams, who did so much to make known the Panamic molluscan fauna. Family CORBULIDAE Genus PANAMICORBULA Pilsbry, 1932 1852. Potamomya C. B. Apams, Cat. Shells Panama, p. 296 (not of Sowerby, 1839). 1932. Panamicorbula Pitspry, Nautilus, vol. 45, p. 105. 1945. Panamicorbula Voxes, Bull. Amer. Mus. Nat. Hist., vol. 86, pp. 11-12, pl. 2, figs. 1-4. Genotype-—Potamomya inflata C. B. Adams=Panamicorbula in- flata (C. B. Adams). ® It is very interesting to find this group of brackish-water corbulid clams living in the mangrove swamps of San José Island. The genus, according to Pilsbry, is confined to the Panamic Province and consti- tutes another proof of the true picture of the San José Island fauna as a reduced Panamic fauna, changed somewhat by its isolation. PANAMICORBULA CYLINDRICA, new species Plate 1, figs 15, 17 Shell ovate-cylindric; the umbones small, projecting little above the general shell outline, subequilateral; the umbones flattened with 48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 a low but well-defined posterior ridge ending in the subangular mid- posterior region. Shell sculptured with fine concentric ridges which are somewhat irregular over the seasonal rest marks, and especially prominent above the posterior ridge on young shells. Hinge as de- scribed by Vokes for the genus. The right valve with one small cardinal tooth immediately under the umbo, and anterior to the well- defined resilial pit on the ventral face of the hinge; the left valve with a small, broad chondrophore, apparently tripartite on its dorsal face. The anterior part is a rounded radial rib, margined with a groove anteriorly and posteriorly; the central and posterior parts are both radially concave. Muscle scars moderate; the pallial line simple, the pallial sinus represented by a broadening of the pallial scar. The holotype, U.S.N.M. No. 542186, measures: Length, 13.3 mm.; height, 9.3 mm.; diameter, 6.3 mm.; right valve, 3.2 mm.; left valve, 3.4 mm. It and a few broken valves of older individuals, were found in the drift of the Rio Marina mangrove swamp on February 19, 1944. Its exact habitat in the mangrove swamps was never determined. The largest paratype (adult) valve seen, U.S.N.M. No. 542187, measures: Length, 22 mm.; height, 16.3 mm.; diameter, 8 mm. (1.e., 16 mm. for both valves). This spécies may be distinguished by the extreme inflation of adults, and the scarcely projecting umbones. With the extreme in- flation of imjlata, this island species has the prominence of the beaks much reduced to produce an apparent cylindric shape. EXPLANATION OF PLATES (All enlargements are given in approximate figures. ) PLATE I Fic” 1. Pomacea cumingtt sanjosensis, holotype, X #. . Neritilia panamensis, holotype, X 8. . Pomacea zeteki, holotype, < 3. . Mytilopsis adamsi, interior of right valve of holotype, « 23. . Ferrissia (Laevapex) joseana, right profile of paratype, x 8. . Ferrissia (Laevapex) joseana, holotype, X 8. . Mytilopsis adamsi, exterior of left valve of holotype, x 2h. 8-11. Cyrenoida insula, holotype, < 24. 12-14. Polymesoda joseana, holotype, X 3. 15. Panamicorbula cylindrica, holotype, interior of right valve, & 24. 16. Detracia joseana, holotype, X 8. 17. Panamicorbula cylindrica, holotype, exterior of left valve, 24. 18. Detracia graminea, holotype, 8. EN Cn 10S" NWP es N No. 6 MOLLUSKS OF SAN JOSE ISLAND—MORRISON 49 PLATE 2 (All figures X approximately 63.) Fic. 1. Zetekella veraguasensis, holotype. % kompi, holotype. . Aroapyrgus chagresensis, holotype. os alleei, holotype. joseana, holotype. . Lyrodes chagresensis, holotype. Cochliopina zeteki, holotype. ae juradot, holotype. fraternula, holotype. . Lyrodes seteki, holotype. . Cochliopina extremis, holotype. s navalis, holotype. australis, holotype. 14. Subcochliopa colabrensis, holotype. 15. Cochliopa diazensis, holotype. 16. Cochliopina wetmorei, holotype. 17. Subcochliopa trochus, holotype. 18. Cochliopa joseana, holotype. “ OC PIAUE SD y “cc soe =“ Oo PLATE 3 (Diagrammatic sketches of the external male reproductive characters greatly enlarged, but not to an exact scale.) Fic. 1. Zetekella veraguasensis, paratype. Px, i kompi, paratype. 3. Aroapyrgus alleei, topotype. 4. ‘s chagresensis, paratype. 5. Cochliopina setekt, paratype. 6. Lyrodes chagresensis, paratype. 7. Cochliopina riograndensis. 8. Aroapyrgus joseana, paratype. 9. Cochliopina setcki, paratype. 10. + juradoi, paratype. wale e fraternula, paratype. ios . navalis, paratype. na € extremis, paratype. 14. * australis, paratype. 15. 2 qwetmorei, paratype. 16. Subcochliopa colabrensis, paratype. 17. Cochliopa rowelli. 18. : diazensis, paratype. ‘ joseana, paratype. 20. Subcochliopa trochus, paratype. ; f { a oe 5 BTA NEMS tir ; Ee ie ais th } t R al i ¢ Wisk meh LAIST) SOOT AS ; isi) Tole, meta ota ‘ a yr Rtg Ais a La . ed ge hope cry Sete aa m ‘ Y al 1 x7 a Vs 4h . at ayeied hexyl oe y y 5 : 7 ‘ia 7 i : | ae . hi ey =. fy f ele W ‘ } 7 .* it af il BS i ty } i Pa ew Fo eee a F . ; Sea te 7 . vig tist SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 6, PL. MOLLUSKS OF SAN JOSE AND PEDRO GONZALEZ ISLANDs, PEARL ISLANDS, PANAMA (For explanation, see p. 48.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 6, PL. 2 MOLLUSKS OF SAN JOSE AND PEDRO GONZALEZ ISLANDS, PEARL ISLANDS, PANAMA (For explanation, see p. 49.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 6, PL. 3 17 is | 19 20 MOLLUSKS OF SAN JOSE AND PEDRO GONZALEZ ISLANDS, PEARL ISLANDS, PANAMA (For explanation, see p. 49.) ee Riz Pa «Yio i is iJ ert) : ; \ % des eee | + Nr |S 4 ‘ aS ane x $ S j Rt Pay 7? x ae ve SMITHSONIAN MISCELLANEOUS’ ‘COLLECTIONS va eee ; er al VOLUME 106, NUMBER 7 MAMMALS OF SAN JOSE. ISLAND, BAY ae PANAMA ¥* 2 Tl is | icc BY a || : REMINGTON KELLOGG | Sees Curator, Division of Mammals _ q my 2 sg ON Meh 4 U.S, National Museum — a iE i$ 3 5 } We al \ Nee, Seenncee® (PUBLICATION 3851) CITY OF WASHINGTON. PUBLISHED BY THE SMITHSONIAN INSTITUTION JULY 18, 1946 b>. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 7 MAMMALS OF SAN JOSE ISLAND, BAY OF PANAMA BY REMINGTON KELLOGG Curator, Division of Mammals U. S. National Museum (PUBLICATION 3851) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JULY 18, 1946 ; ine Shae ete tee nt rae | i i e | ’ a I ‘ ¢: ap tere oe Se SS ee \ _| | ; { The Lord Baltimore Preas BALTIMORE, MD., U. 8. A. Le ; « MAMMALS OF SAN JOSE ISLAND, BAY OF PANAMA By REMINGTON KELLOGG Curator, Division of Mammals, U. S. National Museum Included in the collections made by Dr. A. Wetmore, Secretary of the Smithsonian Institution, and by Dr. J. P. E. Morrison, of the Division of Mollusks, United States National Museum, on San José Island, Archipiélago de las Perlas, in the Golfo de Panama, are 76 specimens of mammals, representing 9 species and subspecies. Bangs (Amer. Nat., vol. 35, No. 416, pp. 631-644, August Igor; Bull. Mus. Comp. Zool. Harvard College, vol. 46, No. 8, pp. 139- 140, September 1905) and Goldman (Smithsonian Misc. Coll., vol. 69, No. 5, 309 pp., 39 pls., 1920) have recorded the mammals known to occur on Isla del Rey (San Miguel Island), and among them the following eight kinds have not as yet been taken on the nearby San José Island: Marmosa fulviventer Bangs, Didelphis marsupialis par- ticeps Goldman, Sylvilagus gabbi incitatus (Bangs), Diplomys labialis (Bangs), Mus musculus Linné, Rattus rattus rattus (Linné), Rattus rattus alexandrinus (Geoffroy), and:Carollia perspicillata aztecum (Saussure). Goldman (1920, p. 199) has pointed out that the bat erroneously identified by Bangs (1901, p. 644) as Vampyrops helleri is actually a specimen of Uroderma bilobatum. Order/ CHIROPT ERA: Family EMBALLONURIDAE SACCOPTERYX BILINEATA BILINEATA (Temminck): Greater White-lined Bat Ten specimens of this small blackish-brown bat, which is character- ized in part by a pair of whitish longitudinal dorsal stripes, were taken by Morrison. One was collected on March 1, eight on May 28, and one on July 2, 1944. All the specimens are females and are presumably adult, with the exception of one half-grown young. These specimens range in total length from 62.5 to 78 mm.; forearms from 44 to 49 mm. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 7 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Family PHYLLOSTOMIDAE MICRONYCTERIS MICROTIS Miller: Nicaraguan Small-eared Bat Of the 22 specimens of this small rusty-brown bat collected by Morrison, 4 were taken on May 8, 2 on May 13, 12 on May 21, 1 on May 28, 2 on July 26, and 1 on August 1, 1944; 12 are females and 10 are males. The individuals in this series vary in total length from 55 to 67 mm.; forearms from 33.5 to 37 mm. ARTIBEUS JAMAICENSIS JAMAICENSIS Leach: Jamaican Bat Four specimens of this large bat are contained in the collection made by Morrison, and of these only one (U.S.N.M. No. 277176) has a readily discernible pair of whitish stripes on the top of the head between the nose leaf and the ears. On the other three bats these stripes are either lacking or rather indistinct. These four females vary in total length from 80.5 to 85.5 mm.; forearms from 59.5 to 61.5 mm. Three were taken on August 22 and one on August 23, 1944. URODERMA BILOBATUM Peters: Yellow-eared Bat The yellowish ear margins of the four bats collected by Morrison were very noticeable when the collection arrived in Washington, but faded to whitish after the bats were transferred to 75 percent alcohol. All four specimens have the characteristic pair of white stripes extending from the nose leaf to the inside of the ear on the crown, and a single white stripe on the side of the face running from below the eye to the tragus. The two females have dark wood-brown upper- parts, with a longitudinal white stripe extending from the nape to the rump. The two males, however, have fuscous-brown upperparts, with the longitudinal white stripe commencing behind the shoulders and ending at the rump. One specimen was collected on February 20, the others on September 12, 1944. They range in total length from 55.5 to 61 mm.; forearms from 40.5 to 42.5 mm. Order EDENTATA Family MYRMECOPHAGIDAE TAMANDUA TETRADACTYLA CHIRIQUENSIS J. A. Allen: Three-toed Anteater The immature male was shot by Wetmore in March, and the two adult females were obtained by Morrison in August 1944. The two NON 7 MAMMALS OF SAN JOSE ISLAND—KELLOGG 3 females measured, respectively: Total length, 995, 990 mm.; tail, 495, 498; hind foot, 90, 88. The measurements for the skulls of the two females are, respectively: Occipito-nasal length, 117.5, 113.8; tip of maxillary to hinder face of occipital condyle, 113, 108; antor- bital width, 31.5, 30.5; length of nasals, 43.4, 41.8. Ikeeve (Proc: Zool. Soc. London for 1942; vol. 111, ser. A, pp. 297, 301) has questioned the validity of this geographic race and has pre- sented statistical data to support his conclusion that the Panamanian specimens on the basis of skull proportions cannot with certainty be distinguished from British Guiana tetradactyla. For these San José females, the ratio to occipito-nasal length of nasal length is, respec- tivelV, 36.7 and 36.9. For specimens of chiriquensis from the main- land of Panama, Reeve gives ratios varying from 37.8 to 41.5. The ratio to occipito-nasal length of antorbital breadth for the San José females is, respectively, 26.8 and 26.7, and the corresponding ratios cited by Reeve for mainland specimens range from 26.0 to 29.1. The corresponding ratios for specimens from Gatun (¢, U.S.N.M. No. 171077) and Porto Bello (9, U.S.N.M. No. 171482) are only slightly greater than those computed for the San José females. Speci- mens of tetradactyla from British Guiana were not available for comparison. Family ECHIMYIDAE PROECHIMYS SEMISPINOSUS IGNOTUS Kellogg: San José Island Spiny Rat Since San José Island is only about 7 miles distant from the Isla del Rey, it would be expected that the same general climatic condi- tions should prevail on both islands, but nevertheless the series of spiny rats taken on San José is noticeably darker than the race on the larger island. Skins and skulls for two adult females, four adult males, and two juvenile males, as well as three young in alcohol and three odd skulls, were collected by Wetmore and Morrison during March, May, and June, 1944. Order ARTIODACTYLA Family CERVIDAE MAZAMA PERMIRA Kellogg: San José Island Brocket Several officers and men of an engineer detachment, when off duty, hunted deer for sport, and through their cooperation five speci- mens were obtained. Kay L. Thurman presented an adult female and a male fawn which he had killed on August 15, 1944. The skin and 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 skull of an adult male, which had been killed on September 24, 1944, by a party consisting of C. F. Jones, W. N. Collins, and Robert Carder, were preserved by Morrison. A nearly complete skeleton of a male that had been shot and lost was picked up by Morrison, who also received a female fawn from another hunting party. Be "SMITHSONIAN MISCELLANEOUS COLLECTIONS Me et ke VOLUME 106, NUMBER Bi xs TURTLES COLLECTED BY THE " SMITHSONIAN BIOLOGICAL SURVEY OF THE PANAMA CANAL ZONE (Wis ONE eee BY KARL PATTERSON SCHMIDT Chief Curator of Zoology Chicago Natural History Museum : (PUBLICATION 3852) cIry OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION AUGUST 1, 1946 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 8 Pie eho COLLECTED BY THE SMITHSONIAN BIOLOGICAL SURVEY OF THE PANAMA CANAL ZONE (WiTH ONE PLATE) BY KARL PATTERSON SCHMIDT Chief Curator of Zoology Chicago Natural History Museum °, Ss ca . rap 1), \F ° CAAA YAingtow’ (PUBLICATION 3852) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION AUGUST 1, 1946 eee Pads a 7 ha bead at at VS fad Pe Loe Py De pS iM ries ‘ bp pal & “gl tr) idee gate Reentry uv: , The Lord Baltimore Press y BALTIMORE, MD., U. 8 As — > . ’ »% | on, nd 4 a i oe 4 ly ——- = i - ; ‘>, > Va-& -- Phe ae as 5 ‘ <. rts ee = oes TURTLES COLLECTED BY THE SMITHSONIAN BIOLOGICAL SURVEY OF THE PANAMA CANAL ZONE By KARL PATTERSON SCHMIDT Chief Curator of Zoology, Chicago Natural History Museum (WitH ONE PLATE) When my report on the collections of amphibians and reptiles of the Smithsonian Biological Survey of the Canal Zone was prepared in 1933,' the turtles of that collection were reserved by the late Dr. Leonhard Stejneger for study in connection with his long- continued project for a comprehensive review of the turtles of the North American continent. The 13 specimens of turtles in the col- lection are of interest in view of the importance of Panama as the crossroads of turtle emigration from the north and of resurgence from the south; and a formal report upon them is required by the original contract between the Chicago Natural History Museum (at that time the Field Museum of Natural History) and the Smithsonian Institu- tion. Panama turtles available for comparison have been lent by the Museum of Comparative Zoology, the United States National Museum, the Carnegie Museum, and the Museum of Zoology of the University of Michigan. I am indebted to Arthur Loveridge, Dr. Doris M. Cochran, M. Graham Netting, and Norman Hartweg, respectively of these institutions, for their generous aid in this connection. I am especially indebted to Dr. E. R. Dunn, of Haverford College, for unpublished information made available for this report. As far as can be discovered, the land and fresh-water turtles of Panama include eight species : Chelydra acutirostris Geoemyda annulata Kinosternon postinguinale Geoemyda funerea Kinosternon panamensis Geoemyda melanosterna Pseudemys ornata Testudo denticulata It is disappointing to find that actual information as to the distribu- tion of these species in Panama is still extremely scanty. The snapping turtle (Chelydra) and the land turtle (Testudo) are known from Panama only from single specimens. It is probable that representatives of all the genera of sea turtles occur on both the Pacific and Atlantic coasts of Panama, but no 1 Smithsonian Misc. Coll., vol. 89, No. 1, 1933. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 8 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 actual records of sea turtles from that country are to be found in the literature. It is customary to distinguish the Atlantic sea turtles from those of the Pacific and Indian Oceans as follows: ? Atlantic sea turtles Pacific sea turtles Dermochelys coriacea Dermochelys schlegelit Chelonia mydas Chelonia agassizti Eretmochelys imbricata Eretmochelys squamata Caretta caretta caretta Caretta caretta gigas Lepidochelys kempii Lepidochelys olivacea It must be pointed out that this partition is subject to a basic criticism, since the great majority of the littoral marine animals of the Pacific coast of Central America are more closely allied to Caribbean, and hence Atlantic, forms than to the more wide-spread representa- tives of the western Pacific and Indian Oceans.’ In the above list only Chelonia agassizii is based on specimens from the Central Ameri- can Pacific coast. Thus the actual identification of the turtles of the American Pacific coast remains a problem of some interest.* The land and fresh-water turtles of Panama clearly represent a part of what Dunn * refers to as the “Old Northern” element of the tropical American fauna. Even Testudo denticulata, though now a South American species reaching its northern limit in Panama, is indirectly derived from the North American Testudo, though the genus appears as early as the Miocene in South America.* With respect to the definition of a minor faunal province, the two species Chelydra acutirostris and Geoemyda annulata range to western Ecuador, and” thus compare with a number of frogs, lizards, and snakes that have a similar range and a similar northern limit in Panama. Testudo denticulata is a species of the Amazonian forest region that similarly reaches its northern limit in Panama; Kin- osternon panamensis represents the Amazonian scorpioides in Pan- ama; and Pseudemys ornata ranges from southern Mexico south- ward to Panama and perhaps into western Colombia (Dunn, E. R., 2 See Stejneger, Leonhard, and Barbour, Thomas, Check-list of North Ameri- can amphibians and reptiles, 5th ed. Bull. Mus. Comp. Zool., vol. 43, No. 1, 1043. 3 Ekman, Sven, Die Tiergeographie des Meeres. Leipzig, 1936. 4 Schmidt, K. P., Problems in the distribution of marine turtles. Marine Life Occ. Pap., vol. 1, No. 3, pp. 7-10, 1945. 5 Dunn, E. R., The herpetological fauna of the Americas. Copeia, 1931, pp. 106-119. 6 Simpson, G. G., A Miocene tortoise from Patagonia. Amer. Mus. Novit., No. 1200, pp. 1-6, 1942; Turtles and the origin of the fauna of Latin America. Amer. Journ. Sci., vol. 241, pp. 413-420, 1043. No. 8 TURTLES OF THE PANAMA CANAL ZONE—SCHMIDT 3 in litt.). The forested region from Panama to western Ecuador is sharply characterized by such remarkable forms as the tree frog Agalychnis calcarifer and the coral snake Micrurus ancoralis; and while its boundary is not yet adequately defined, this faunal province does not appear to entend beyond the mountain massif of Costa Rica. Micrurus ancoralis falls into two well-marked sub- species within this area. Differentiation of the turtles that characterize this Ecuador—Panama fauna, such as Chelydra acutirostris and Geoemyda annulata, into corresponding subspecies is not demonstrated. CLASS | REP TILIA Order CHELONIA CHELYDRA ACUTIROSTRIS Peters Chelydra serpentina var. acutirostris Peters, Monatsber. Akad. Wiss. Berlin, 1862, p. 627 (Guayaquil, Ecuador). Chelydra rossignonit (nec Bocourt) BouLencer, Ann. Mag. Nat. Hist., ser. 7, vol. 9, p. 49, 1902; Vaillant, Miss. Mes. Arc. Mer. Equatorial, vol. 9 (Zool.), fasc. 2, p. 48, pls. 1-3, 1909; Boulenger, Proc. Zool. Soc. London, 1914, p. 814. Chelydra serpentina (part) Corr, Proc. Acad. Nat. Sci. Philadelphia, vol. 24, iy Ap itey en This species is not included in the Smithsonian Survey collection but is represented in the Museum of Comparative Zoology by a single large specimen from Panama. Comparison material Mus. No. Locality Date Collector M€.Z. 216069...,.-+ Coca Solo, Colon, C. Z: 1925 Thomas Barbour As may be seen from the above synonymy, which omits references not based on specimens, the taxonomic status of the Ecuadorian snapping turtle has been in a state of considerable confusion. Cope, Gunther, and Boulenger at first agreed to the distinction of rossignonii from Central America while referring Ecuadorian specimens to the North American serpentina. When Boulenger became convinced that the Ecuadorian snapping turtle should be distinguished, he applied the name rossignonu, rejecting Peters’ earlier name acutirostris as “not accompanied by an adequate description.” A single large speci- men from Panama in the collection of the Museum of Comparative Zoology and five specimens of rossignonii in the Chicago Natural History Museum, collected by myself in Honduras, enable me to re- examine the problem as to the distinction of the several supposed forms and to form an opinion as to the identity of the Panama species. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 The distinctness of the Central American snapping turtle from the North American form may be regarded as established. The adult specimens from Honduras before me all have a shorter and more angularly pointed head, four barbels on the chin instead of two, a slightly wider plastral bridge, and broader vertebral shields. In the skull the height at the quadrate is greater, .44 to .48, as compared with the length of the skull to the condyle, while in serpentina the same ratio is .38 to .43. In three skulls of rossignonii the width of the palatine ranges from .28 to .30 of the length of the skull, while in serpentina this ratio varies from .19 to .25. The question as to the identity of the single Panama specimen may be examined by means of the literature, since Boulenger (1902, loc. cit.) and Vaillant (1909, loc. cit.) give measurements of Ecuadorian specimens. Comparing the ratios of length of plastron, A, width of bridge, B, and width of second vertebral shield, C, all to the total length of carapace, the result is as follows: A B Cc Ecuadorian specimens (6).......... 71-77 .07—.12 .24—.27 Panama specimen (1). feces ese: Giyl .09 24 Honduras specimens (5)........... 77-81 .06—.08 ~ 32-.34 Thus, as far as may be determined from a single specimen, the Panama snapping turtle is most closely allied to the Ecuadorian form, and should bear the name acutirostris. The specimen examined ap- parently has only one pair of short barbels; the head is acute, like that of rossignonii; and the growth rings are more evident than in the Honduran specimens (though these are smaller). There are five inframarginals on the right and four on the left side. The measure- ments of this specimen are as follows: mm TenatholicakapaCe rc cctrsece ry tel asiecctories 340.0 Wadthitotvcaranace ys as cemiseectee preci seers 286.0 Weneth' ior plastromman: aan. . ve aetsies oe ete ee tie cles 260.0 Width of plastral bridge...... Es SO S93 30.0 Width of 2d vertebral shield................. 82.0 Length of shielded top of head................ 97.0 Width of shielded top of head................ 79.0 KINOSTERNON POSTINGUINALE Cope Cinosternum leucostomum Corr, Proc. Acad. Nat. Sci. Philadelphia, vol. 17, p. 189, 1865; Boulenger, Proc. Zool. Soc. London, 1898, p. 108; Peracca, Bol. Mus. Torino, vol. 19, No. 465, p. 2, 1904. No. 8 TURTLES OF THE PANAMA CANAL ZONE—SCH MIDT 5 Cinosternum brevigulare CopE (nec Giinther), Proc. Amer. Philos. Soc., vol. 2, p. 389, 1885 (type locality, Sipurio, Costa Rica). Cinosternum postinguinale Corr, U. S. Nat. Mus. Bull. 32, p. 23, 1887 (substi- tute name for brevigulare Cope). Three specimens, prepared as skin and skull, collected as follows: U.S.N.M. No. Locality Date | Collector GEBUO Tiie. apap: Gatun March IoII Meek and Hildebrand EOS yaaa Ee Frijoles Feb. 13, IQII e “s or Io wig I tel SE lie Panama sek E. A. Goldman Comparison Material Mus. No. Locality Date Collector ©.M:6878 >. 22:5 Madden Dam, C. Z. Aug. 16,1933 A. M. Greenhall MM IZZOO 4, «0 Kicty Bop Se C.Z. Mar. 21,1934 M. Graham Netting Goa 7701 ees it “cc “ “cc “c “cc “ 73 C.M. 7702 Fe “ if “ ce “ce “ “cc “ “ “ C.M.N.H. 13410... - i I ia sshc 1928 K. P. Schmidt U.M.M.Z. 63732.. “ He SE sues H. T. Gaige U.M.M.Z. 63734. ; “ “ “ “ ry. bs Os eG “ U.M.M.Z, 63735.. H - Cy a ee CORES All three of the Survey specimens are males; they exhibit the well- defined opposing patches of horny tubercles on the femoral and tibial joints of the leg. This character, with various others, amply dis- tinguishes postinguinale from the forms allied to scorpioides. The related species of the Choco Region of Colombia, Kinosternon Spurrelli Boulenger, in view of other faunal relations, might be thought to be referable to the Panama form. Dr. Norman Hartweg (in litt.) regards it as at least subspecifically distinct, and suggests that leuco- stomum, postinguinale, and spurrelli may be regarded as a series of subspecies. The further examination of this question must be based on a much larger amount of material than is at present available. KINOSTERNON PANAMENSIS, sp. nov. Diagnosis —A species of Kinosternon allied to scorpioides, with an elongate shell, keels present but less distinct than in scorpioides, the posterior end of the plastron only very narrowly notched, and the head of the adult male less enlarged. No postfemoral tubercles in the male. Distinguished from integrum by its larger and longer gular shield. Range.—Panama. Type——Cat. No. 117369, U.S.N.M. ¢; Panama Railroad, C. Z., Panama, June 10, 1932; Dr. Thomas Barbour, collector. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Description of type-—Carapace elongate, the sides parallel, with three barely distinguishable keels making the dorsal area flat ; carapace descending abruptly behind with little outward flare of its posterior border; head large, upper jaw strongly hooked; outer four digits connected by web on both front and hind feet; plastron large, the posterior lobe feebly nicked; gular shield large, its length much more than half that of the anterior lobe; inguinal shield large, broadly in contact with the axillary; tail long, ending in a hooked horny nail; no trace of femoro-tibial horny tubercles; carapace dull brown, with- out markings; plastron yellow, unmarked; skin of soft parts ap- parently yellowish without definite markings, as is the case also with the darker horny shields of the jaws. Measurements of type-——Length of carapace 150, width 93, depth 52.3; length of anterior lobe of plastron 42.0, width 68.0, length of gular shield 25; length of middle segment of plastron 39.0; length of posterior lobe 46.5, width 60; width of head 33.3; length of tail about 42. Variation.—The nine paratypes include a series of dry shells of juvenile specimens, the smallest only 50 mm. in length, with very sharp dorsal keels, and with the costal fenestrae 60 percent of the costal length; in a specimen 93 mm. in length, with the keels still sharp, the fenestration is reduced to Io percent of the costal length; while in a specimen measuring 124 mm. the keels are rounded. In the largest specimen, M.C.Z. No. 24957, a male, the length of carapace is 161 mm. The tail is very short ig female specimens. The list of the paratypes is as follows: Mus. No. Sex Locality Date Collector U.S:NEM 7864.5 ented Panama sete ose es Tie Fete se .... | Thomas Barbour MiG.Z 18630. 5.45 fc) Ancon, C. Z. aie . . ST AOS A cenion 3 Panama City eavets € bi ca 27 OLO mace: (shell) Panama City tees UR Re Dn $ . 35486, ceria 1" Rio Coclé, Penomoné .... F-. Jackson AO STAR ee eee . RioGrande, “ Pyare ewe “6 sO) ASG meets % * Sater ew SST ASO, Wah. . y “ «fer ss se Remarks.—This species of Kinosternon, not represented in the Survey collections, proves to be quite as abundant in Panama as is postinguinale. Inspection of the series available convinces me that the Panama form is more closely allied to the Guianan and Ama- zonian scorpioides than to the Mexican integrum, which is not other- wise known from Lower Central America, or even from Guatemala. No. 8 TURTLES OF THE PANAMA CANAL ZONE—SCHMIDT 4 Typical scorpioides is widespread in South America, and there have been some attempts to partition it into subspecies. Miller and Hellmich? conclude that the subspecific arrangement proposed by Siebenrock ® is untenable; Siebenrock gives the range of scorpioides scorpioides as the Guianas, entirely enclosed by that of scorpioides integrum, which is thought to extend from western Mexico to Para and the Madeira River. PSEUDEMYS ORNATA Gray Emys ornata Gray, Syn. Rept., p. 30, 1831. Pseudemys ornata Corr, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, p. 153, 1876. Chrysemys ornata BouLENGER, Cat. Chelon. Brit. Mus., p. 80, 1889. Six specimens, cataloged in part as Pseudemys grayi, offer no distinctive differences from a series from British Honduras. U.S.N. M. No. Locality Date Collector Pango oe Culebra (Rio Cu) Mar. 10, IOTI Meek and Eiidepeana Sapege ss Alhojulla (Limon Creek) Feb. 26, 1911 a BAOB Rei. ts Gatun May 15, I9QII Sle ff 54086...... Panama IQII wa: - [+ 6c alaerar/ Panama IQII laa s 54088...... Mindi Swamp IQII NERS s Comparison material M.C.Z. No. Locality Date Collector BOB A celts be San Pablo 1874 Allan Lesley 25080 uses a8 Barro Colorado : 1927 Thomas Barbour ZSZ00s sae as « & Ke 1929 E. R. Dunn AGG Mere acre Penomoné, Coclé Prov. 1931 F. Jackson BOOBS. eee Panama 1909 Thomas Barbour sroedcr. (ict Margarita Swamp near Colén 1925 % sy Within the great recorded range of ornata, from Sinaloa to Panama (and perhaps to the Chocé in Colombia, per E. R. Dunn, in litt.), geographic variation is to be expected. The range of the species does, however, appear to be continuous—it is the common fresh-water turtle in Guatemala, Honduras, and apparently in Costa Rica and Panama. I have examined four additional specimens in the Museum of Comparative Zodlogy, from San Pablo, Penomoné, and Coldn, and it is recorded also from Barro Colorado Island. 7 Miiller, Lorenz, and Hellmich, Walter, Wiss. Ergeb. Deutsch. Gran Chaco- Expedition. Amph. Rept., pt. I, p. 96, 1936. 8 Siebenrock, Franz, Zool. Jahrb., Suppl., vol. 10, p. 445, 1909. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 GEOEMYDA ANNULATA Gray Geoclemmys annulata Gray, Proc. Zool. Soc. London, 1860, p. 231, pl. 29. [Geoemyda] annulata Dunn, Proc. New England Zool. Club, vol. 12, p. 32, 1930. Rhinoclemmys gabbit ALLEE, Ecology, vol. 7, p. 451, 1926. Two specimens in the Canal Zone Survey collection, with other material examined from Panama as follows: U.S.N.M. No. Locality Date Collector SOAR Oe eee pense aL, oo Panama February 1911 E. A. Goldman 54009 ‘cc 6c “cc 6c Comparison material Mus. No. Locality Date Collector OLSEN UGA A Ry amene oe Panama B BOS77, bee re setae : ¥ sisters Sayan IMEC 272 2220 rae ceva c ere Barro Colorado I. 1926 E. R. Dunn Re PORT OM Laete wie oisisiels - i ‘ 1929 ake mo COMEN TE IGE 24.04). t. 2e8 i) rs . 1924 W. C. Allee e TASB rectrstieicae i * . 1928 K. P. Schmidt The remarkable infestation of Geoemyda annulata by a tick that attacks the bony carapace at the sutures between the horny plates is reported by Allee and Allee.® Every shell examined exhibits traces of infestation by such ticks, which leaves characteristic and un- mistakable scars (pl. 1). The head and neck of Kinosternon postin- guinale also tend to be attacked by ticks, presumably of the same species, to judge from the specimens in the Carnegie Museum. The type locality of Chelopus gabbu Cope being Costa Rica, this name would be available should a northern race of annulata be dis- tinguishable. I have no Ecuadorian specimens at hand, and follow Dunn in referring Panama specimens to the Ecuadorian species. GEOEMYDA FUNEREA Cope Chelopus funereus Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, p. 154, 1876. [Geoemyda] funerea Dunn, Proc. New England Zool. Club, vol. 12, p. 31, 1930. This species is added to the Panama fauna on the authority of Dr. E. R. Dunn (in litt.). I have not seen specimens. ® Allee, W. C., and Allee, Marjorie H., Jungle islands, p. 1o1, fig. 43. Rand McNally, Chicago, 1925. No. 8 TURTLES OF THE PANAMA CANAL ZONE—SCH MIDT 9 GEOEMYDA MELANOSTERNA Gray Geoclemys melanosterna Gray, Proc. Zool. Soc. London, 1861, p. 205. This species was described from “Cherunha, Gulf of Darien” (then Colombia), and Dunn (in litt.) informs me that he has seen a more recent specimen from the Rio Chucuhaque, Panama. TESTUDO DENTICULATA Linnaeus Testudo denticulata LINNAEUS, Systema naturae, ed. 12, p. 352, 1758. The single male specimen, U.S.N.M. No. 50251, is without data other than Panama. I find no other record from Panama for this species; it is abundant in Venezuela and in the Santa Marta region of Colombia, and may be supposed to have a continuous distribution along the Colombian Caribbean coast. CHELONIA AGASSIZII Duméril and Bocourt A single juvenile specimen, U.S.N.M. No. 51500, from Chame Point, collected by Robert Tweedlie represents the green turtle in the Smithsonian Survey collection. This locality is in the Gulf of Panama. A single juvenile specimen in the collections of the Museum of Com- parative Zoology, from the Hassler Expedition, is also presumably from the Pacific side. There is obviously no opportunity to open the question as to the distinctions between Chelonia agassizii and Chelonia mydas on the basis of available collections. CARETTA CARETTA Linnaeus A single juvenile specimen, U.S.N.M. No. 46390, was collected from the coasts of Panama by J. M. Dow. Without information as to which coast it came from, and no comparative material, this specimen obviously affords no information as to the differences between the loggerheads of the two sides of the Isthmus. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 8, PL. 1 ANTERIOR PART OF CARAPACE OF GEOEMYDA ANNULATA, SHOWING TYPICAL INFESTATION BY A TURTLESHELL TICK (X 11f) ay ee Ue WS Lain genet K: LP ae ee a ee Se A ST ars oe ; aaa oo qe is 3 ee "SMITHSONIAN MISCELLANEOUS COLLECTIONS ) _ VOLUME 106, NUMBER 9 | 2 a THE SPECIES OF PLATYCOPIA SARS | (COPEPODA. CALANOIDA) Rey BY ~ MILDRED STRATTON WILSON Paap te" Curator, Division of Marine Invertebrates U.S. National Afcgernn (PuBLICATION 3853) _GITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION AUGUST 23, 1946 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 9 ihe SPECIES OF PLATYCOPIA SARS (COPEPODA, CALANOIDA) BY MILDRED STRATTON WILSON Assistant Curator, Division of Marine Invertebrates U. S. National Museum (PUBLICATION 3853) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION AUGUST 23, 1946 The Lord Baltimore (Pres BALTIMORE, MD., U. S. A. Steen Ra Yee Spe ps 7 * 4 de Pw be iit \ tet a . 7 w - 1 ‘ K t ‘ ' | ‘ 7 ha nt , Pa i C's arte etl AJ % 1 vet y + .Y it i , a , / H ve » THE SPECIES OF PLATYCOPIA SARS (COPEPODA, CALANOIDA) By MILDRED STRATTON WILSON Assistant Curator, Division of Marine Invertebrates U. S. National Musewm In a publication + discussing the sand and mud of beaches as a copepod habitat, C. B. Wilson included a drawing of a first antenna labeled “Arenocalanus tumidus, female, a new genus of sand-dwelling calanids, showing an exceptional increase in the number of aesthe- tasks.” In the text the only reference made to this drawing was' in part of a sentence: “in the female either the number of aesthetasks is multiplied as in figure 4, or..... ” No other figure or description of either genus or species was ever published. Recently, while checking the collections that came to the National Museum after Wilson’s death in 1941, I found a large, partially identified collection from Mount Desert Island, Maine, in which was included a vial labeled “Arenocalanus tumidus.” An examination of the specimens in the vial revealed that they did not represent a new genus but belonged to the genus Platycopia. A study of all the individuals showed that not one, but two species were present, neither of which can be assigned to the species tumidus as characterized by Wilson’s drawing. The specimen dissected for this illustration is apparently no longer extant, for it was not found among the few remaining temporary glycerin mounts that Dr. Wilson made of all his dissections, and which are now in the National Museum. However, some incomplete notes and drawings relating to this species were found among Wilson’s papers. Although they disagree in some details with his published figure of the first antenna, there are enough clues to enable me with some degree of assurance to assign the specific name he used to one of the two species found in his Mount Desert material (see under Remarks, p. 13). The antenna as figured by Wilson in no wise constitutes a description by which the species can be recognized; nevertheless I am describing it as Platycopia tumida (C. B. Wilson) in order to avoid any confusion that may result from bibliographic reference to this hitherto in- adequately characterized species. 1 Wilson, C. B., Smithsonian Misc. Coll., vol. 94, No. 7, p. 7, fig. 4, 1935. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 9 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Because of the similarity in size and outward appearance of the two species it was thought advisable to dissect, at least partially, all the specimens in the collection. The appendages have been mounted in glycerin between cover glasses of unequal width, which have then been inverted on a slide in damar or balsam. They have been studied not only with a high dry but with a 1.8 mm. (95x) oil immersion objective, with which both 10x and 15x oculars were used. This use of oil immersion gave much greater resolution than is possible with high dry objectives. The study of detail in this genus proved easier than it is in many calanoids, because of the unusual flatness of most of the appendages. All drawings have been made to the same scale. I am indebted to Dr. Waldo L. Schmitt, head curator of biology, U. S. National Museum, for encouragement during the progress of the work and suggestions in preparing the final draft of the paper. a SYSTEMATIC DISCUSSION The genus Platycopia hitherto has included but two species, both described by G. O. Sars from the southern coast of Norway, and not since reported. The two additional species described in this paper were collected by C. B. Wilson at Mount Desert Island, Maine (northeastern coast of United States), probably in 1930. Both of the Norwegian species were taken with bottom samples of coarse, muddy sand, but because of the depth from which they were dredged it cannot be stated exactly whether these species lived in the mud, among the grains of sand, or hovered near the bottom. From observations of bottom-living calanoids Sars? concluded that they keep themselves “constantly close to the ground over which they move in a peculiar gyrating manner.” The Mount Desert Island specimens came from a large tidal pool which was described by Wilson * as being about the size of a small room, and containing sand “made up very largely of rounded grains of quartz.” Apparently he obtained these specimens of Platycopia from the washings of the sand from this tidal pool, and believed that they inhabited the sand. This is borne out by the caption under the figure mentioned above which reads “new genus of sand-dwelling calanids.” If that is the case, it may well be that many of the calanoids obtained in bottom samples, even at considerable depths, actually dwell in sand or mud. However, it is possible that these specimens 2 Sars, G. O., Archiv Math. Naturv., vol. 31, No. 7, p. 13, Io1I. 3 Wilson, C. B., Smithsonian Misc. Coll., vol. 94, No. 7, p. 3, 1935. NO. Q SPECIES, OF PLATYCOPIA SARS—-WILSON 3 of Platycopia may have been hovering very close to the bottom of the pool, in the manner that Sars described, and they could thus have been swept up with sand dipped from the bottom. The Norwegian species of Platycopia were both dredged from the same type of bottom at Korshavn, but at different stations. They varied from one another in the presence or absence of inside setae or spines on segments I and 2 of the exopods of legs 3 to 5; in a re- duction of segments in the endopods of legs 2 to 5; and in the shape of the male fifth leg. As the specimens of Platycopia in the Mount Desert collection were placed by the collector in one vial, it is not known whether they came from the same spot in the tidal pool or were sorted out from collections made in two or more places. Finding them so closely associated, however, is very interesting, because they vary from one another in much the same way as do the species from Korshavn. Family PLATYCOPIIDAE Sars, 1911 Genus PLATYCOPIA Sars, 1911 Arenocalanus C. B. WILson, 1935, Smithsonian Misc. Coll., vol. 94, No. 7, p. 7, fig. 4. Diagnosis ——Antenna 1 shorter than first metasome segment, bear- ing on segment 1 a long spine ending in a thin filament; neither antenna geniculate in male. Leg 1 much reduced in size, both rami 2-segmented. Exopods of legs 2 to 5 with 2 outer spines on segment 1; endopods 2- or 3-segmented. Fifth legs in female resembling legs 2 to 4; in male, symmetrical, exopods slightly transformed. Description (emended).—Length between 0.50 and 0.95 mm.; male slightly smaller than female of species. Body robust, somewhat cyclopoid in form; rostrum stout, triangular from above; eye spot distinct or wanting. Metasome consisting of 5 segments; cephalic segment greatly elongated, with anterior portion strongly vaulted above. Urosome of 4 segments in both sexes, narrow; a pair of anal lappets, conspicuous in dorsal profile, extending from third segment ; caudal setae 4, jointed near base. First antennae considerably shorter than first metasome segment, consisting in female of 22 or 23 seg- ments, in male of 15 or 16, both alike, nongeniculate; last segment of female and last two of male elongated; first segment in both sexes much broadened and elongated, bearing beyond the middle a long spine which ends in a thin filament, this spine almost half the length of the antenna; some segments bearing aesthetasks, more numerous and highly developed in the male than in the female. Second antennae 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 compact, outer ramus larger than the inner, 5-segmented, the first 2 segments dilated. Mandibles not particularly stout, with few denticles and slender palp. First maxillae distinctive; masticatory lobe large, ovoid, armed with short, stout, clawlike spines; palp of very short, slender lobes with comparatively few setae. Second maxillae some- what cyclopoid in structure, armed with stout spines; maxillipeds of the usual calanoid type, slender and not conspicuously elongated. Legs flattened, with basal segments greatly broadened. Rami of first legs differing from others, much abbreviated, 2-segmented, somewhat lamellar in shape, bearing plumose setae. Distal basal segment of legs 2 to 5 with outer, distally-placed spine. Exopod of legs 2 to 4 with 3 segments, the first bearing 2 spines outside, distal edge of segments I and 2 on anterior face extended beyond the middle as a hyaline membrane which partly overlies the succeeding segments, this edge serrate on legs 3 and 4; armature of endopod varying from one leg to another. Setae of both rami of legs 2 to 5 mostly trans- formed to flattened spines, many bearing marginal hyaline flanges. Female fifth legs like legs 2 to 4, slightly smaller; species differing in segmentation of endopods, and in armature of both rami. Right and left fifth legs of male alike, exopod slightly transformed, 2- or 3- segmented, bearing terminally thin, variable hyaline lappets, flanked internally by a modified spine, differing specifically in shape and size; endopod variable. . Genotype.—Platycopia perplexa Sars. KEY TO THE SPECIES OF PLATYCOPIA a.4 Endopods of legs 2 to 5 3-segmented; first or second segment of exopods 3 to 5 bearing inner setae or spines (except leg 5 3); male fifth exopod elongated, terminal inner spine styliform. 1. Male fifth exopod bearing I spine at proximal constriction of second segment, distal segment of endopod with 5 spines. P. perplexa Sars, 1911 2. Male fifth exopod bearing 2 spines at proximal constriction of second segment, distal segment of endopod with 4 spines. { P. sarsi, new species b. Endopods of legs 2 to 5 varying in segmentation; first and second segments of exopods 3 to 5 lacking inner setae or spines; male fifth exopod compact, terminal inner spine saber-shaped. 1. Second endopod 2-segmented; female fifth endopod incompletely 3- segmented, the whole bearing 4 spines; male fifth exopod 2-seg- mented, distal segment of endopod with 2 spines. P. pygmaea Sars, 1919 * Because only the male of P. sarsi is known, the female sex is omitted from section a of the key. The female of P. perplexa can, of course, be distinguished from those of section b by the major characters given under a. NO. 9 SPECIES OF PLATYCOPIA SARS—-WILSON 5 2. Second endopod 3-segmented; female fifth endopod 2-segmented, the whole bearing 6 spines; male fifth exopod 3-segmented, distal segment of endopod with 5 spines... P. tumida (C. B. Wilson), 1935 PLATYCOPIA PERPLEXA Sars _ Platycopia perplexa Sars, 1911, Archiv Math. Naturv., vol. 31, No. 7, p. 4, Is. 1-2. te perplexa Sars, 1919, Crustacea of Norway, vol. 7, p. 11, pl. 6, pl. 7, nee Ts E Description (after Sars) —Female. Length about 0.95 mm. Eye wanting. Urosome about 4 the length of the metasome, caudal rami short, slightly longer than broad. Antenna 1 with 23 segments, aesthetasks short, apparently 5 or 6 in number. Masticatory lobe of first maxilla with 6 clawlike spines and 6 setae, most proximally located spine unbranched, bearing spinules on distal margin; rudi- mentary epipodal lobe present on palp. Leg I with seta and a curved spinous process on distal inner edge of basal segment 2. Spines of exopod 2 having margins of hyaline flanges smooth, those of exopods 3 to 5 with serrate margins. Third exopod with inner seta on segment 1, fourth exopod with seta on I and spine on 2. Endopods 2 to 4 3- segmented ; segment I bearing 1 seta; segment 2 unarmed in endopod 2, with I spine in 3 and 4; distal segment of endopod 2 with 6 setae and spines; that of endopod 3 with 7 spines; that of endopod 4 with 6 spines. Exopod of leg 5 with inner seta on segment 1 (this shown only in 1919 reference), inner spine on segment 2; segment 3 bearing 6 spines, afl well developed and having serrate hyaline flanges. Endopod 5 3-segmented, segment I unarmed, segment 2 with 1 spine, segment 3 with 5 small, unequal spines. Male. Length about 0.83 mm. Antenna 1 with 16 segments, aesthetasks much longer than in female, numbering 10 or 12 (number based on illustrations). Legs I to 4 exactly like female. Leg 5 with 2-segmented exopod, distal segment somewhat elongated, constricted near the proximal end and bearing a stout, hyaline flanged spine at this point. End of segment transversely truncated and carrying a thin partly ciliated lamella, flanked internally by a long styliform spine and on the outer edge by 2 unequal spines. Endopod 3-segmented, first segment unarmed, second bearing a long slender spine, third with 5 short spines. Occurrence.—Korshavn, southern coast of Norway; dredged from 60 fathoms, coarse, muddy sand. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 PLATYCOPIA SARSI, new species Figure 1, A-G Specimens examined.—t male, 2 immatures, collected from tidal pool at Sea Wall, Mount Desert Island, Maine, by Charles Branch Wilson. Holotype male (1 slide) U.S.N.M. No. 79917. Diagnosis ——Male. First antenna with 16 segments. Endopods of legs 2 to 5 3-segmented ; exopod 5 with 2 spines at proximal constric- tion of second segment, distal spine greatly enlarged; third segment of endopod with 4 spines. Mature female unknown. Description—Male. Length about 0.70 mm. Urosome like that of P. perplexa, anal lappets somewhat smaller. First pair of antennae (fig. 1, 4) with 16 segments, bearing 12 aesthetasks, most of which are jointed near the middle (the shortness of some of the aesthetasks in this unique specimen is due to loss of the distal part, as indicated by broken lines in the illustration); 4 setae on segment I; 3 on segments 4, 15, and 16; 2 on 2, 6, 9, 10, and 12; 1 on 7, 8, II, 13, and 14; short spines on 3, 5, and 8. Second antennae and mouth parts like those of the genotype as described by Sars, except for slight differences in the spines of the masticatory lobe of the first maxilla (fig. 1, B). This lobe with 6 large clawlike spines and 6 setae; 2 of the setae placed together near center of the anterior face; most proximally located spine with distal margin branched (fig. 1, C); rudimentary epipodal lobe of palp not found in this specimen. Leg I (fig. 1, D) very similar in shape to that of P. perplexa, distal segment of exopod with 6 setae, endopod with 3. Inner seta of endopod spiniform (i.e., stiff and straight), set into a stout, well-defined ridge which appears to be more or less demarcated from the surface. A slender seta with its basal part widened arising from the inner surface of second basal segment and overlying a small ovoid ridge near the inner edge of first segment of endopod. Legs 2 to 4 apparently exactly like those of P. perplexa, all endo- pods 3-segmented, varying from one another in the number of setae and spines. Hyaline flanges of spines of exopod 2 smooth, except that.on large terminal spine, mostly serrate in succeeding pairs. Exopod of leg 2 bearing 4 spines and 2 setae on distal segment; endopod (fig. 1, £) carrying an inner seta on segment I, none on segment 2, and 4 spines and 2 setae on 3. Setae converted to spines in the following pairs of legs, except for seta on segment 1 of endopod. Leg 3 bearing an inner seta on first segment of exopod, distal segment of endopod with 7 spines; leg 4 carrying an inner seta on first segment of exopod and a spine on second; distal segment of endopod with 6 spines. NO. Q SPECIES OF PLATYCOPIA SARS—WILSON 75 Fic. 1.—Platycopia sarsi, new species. Male, A, antenna 1; B, masticatory lobe of maxilla 1; C, proximal spine of B, greatly enlarged; D, leg 1; E, endopod of leg 2; F, leg 5. Female, G, leg 5, late copepodid stage, prior to molting. Platycopia tumida (C. B. Wilson). Female, H, leg 2; I, endoped of leg 3. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Leg 5 (fig. 1, F) resembling P. perplexa in shape and size, and in having a 2-segmented exopod with the inner terminal spine styliform. Differing from perplexa in presence of a second spine at proximal constriction of second segment of exopod; this spine almost twice the length of the first with very broad marginal flanges. A thin, serrate, cuneiform, hyaline structure lying on the proximal surface of the segment near outer edge; a heavier, narrow, crescent-shaped ridge near inner edge. Terminal portion of segment irregularly shaped with inner edge extended into a sharp spinous process. Three small, oval- shaped, very thin hyaline lappets hanging between this process and the outer edge. A spatulate-shaped process with rolled hyaline margin, hanging on the outer edge nearly opposite the point of insertion of the inner spine. Endopod 3-segmented, first segment unarmed, second bearing a long inner spine and having a hairy pad across the surface ; distal segment carrying 2 short spines inside and 2 terminally, outer and terminal edges set thickly with slender spines. Immature forms.—Two immature forms, referable to P. sarsi be- cause of the presence of inner setae and spines on exopods 3 and 4, were present in the collection. Both are copepodid stages; one much less developed than the other may be a male, the other represents a late stage of the female prior to molting, and is briefly described. Female, late copepodid stage. Body more slender, cephalic segment shorter and succeeding segment longer than in mature male. Penulti- mate segment of urosome longer than in mature male. First antenna with 22 segments, segment I somewhat shorter than in mature male, with 4 setae; aesthetasks 5 in number, very slender, segmented near middle. Leg 1 showing same general form as in male, though endopod not completely segmented, ridge and accompanying spiniform seta of endopod well developed, other setae shorter than in mature form. Legs 2 to 4 with segmentation of both rami complete, setae and spines developed, but somewhat smaller than in mature male. Leg 5 (fig. 1, G) not well enough developed to find points to distinguish it from P. perplexa. Segmentation of exopod not com- plete; only 1 spine of first segment and 5 of third segment completely developed; inside spine of second segment partially developed, no evidence of inside seta on segment I as in perplexa. (This seta in perplexa is very slender and was not shown in Sars’ 1911 description. Its absence in this immature specimen may or may not indicate its presence in the adult.) Endopod 2-segmented, but showing evidence of further separation; inside spine of segment 2 present; terminal segment awvith 5 spines. As perplexa carries 5 spines on the end seg- NO. 9 SPECIES OF PLATYCOPIA SARS—WILSON 9 ment there is nothing to suggest what is the differentiation of the females of the two species in the adult form. PLATYCOPIA PYGMAEA Sars Platycopia pygmaea Sars, 1919, Crustacea of Norway, vol. 7, p. 13, pl. 7, fig. 2. Description (after Sars) —Female. Length about 0.60 mm. Eye wanting. Urosome very narrow, nearly half the length of the meta- some ; caudal rami elongated, more than 3 times as long as they are broad. Antenna I with 23 segments, number of aesthetasks not known. Mouth parts and leg 1 said to be very similar to P. perplexa. Legs 2 to 5 lacking inner setae on exopods. Leg 2 with a 2-segmented endopod, distal segment with 6 spines. Endopods of legs 3 and 4 with division of two distal segments incomplete; spinal formula not given for leg 3; endopod 4 having I spine on segment 2, and 5 on segment 3. Leg 5 with third segment of exopod having the 2 inner spines very much shortened and their margins smooth. Endopod incompletely 3- segmented, as in legs 3 and 4, segment 2 with a small inner seta, distal segment with 3 unequal spines. Male. Length about 0.52 mm. Antenna I with 16 segments, num- ber of aesthetasks not known. Leg 5 of a compact structure; exopod 2-segmented, the distal segment bearing proximally 2 large spines, truncated terminally as in perplexa and carrying a thin hyaline lamella flanked internally by a large saber-shaped spine, and externally by a somewhat similarly shaped spine with the inner edge dentate. Endo- pod 2-segmented, segment I bearing a stout spine, segment 2 somewhat lamellar in shape and bearing 2 short spines. Occurrence.—Korshavn, southern coast of Norway; dredged from 30 fathoms, muddy bottom. PLATYCOPIA TUMIDA (C. B. Wilson) Figure 1, H-J, Figure 2 Arenocalanus twnidus C. B. Witson, 1935, Smithsonian Misc. Coll., vol. 94, No. 7, p. 7, fig. 4. Specimens examined.—4 females, 1 male. Collected from tidal pool at Sea Wall, Mount Desert Island, Maine, by Charles Branch Wilson. Lectotype female (slides) U.S.N.M. No. 79920; allotype male (slides) U.S.N.M. No. 79921. Diagnosis.—First antenna of female with 22 segments, that of male with 15; first segment of both sexes with a narrow ridge running lengthwise of the middle of the segment, bearing aesthetasks in male. Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Endopod of leg 2 3-segmented, of legs 3 to 5 2-segmented. Distal segment of female fifth endopod with 6 spines. Male fifth leg with 3-segmented exopod, distal basal segment bearing a large spine in- side; endopod with 5 distal spines. Description Female. Length about 0.72 mm. Eye spot present, relatively small. Anal lappets of urosome conspicuous in profile; caudal rami more elongate than in P. sarsi, apparently not so slender and elongate as in pygmaea. First antenna (fig. 2, 4) with 22 seg-. ments; a long curved ridge running lengthwise of the first segment; a stout spine pointing somewhat obliquely across the segment near distal base of ridge. Aesthetasks 8 in number, al! jointed near the middle; I each on segments I, 3 to 5, 10, 15, 19, and 22. Six setae and I small spine in addition to the larger spine on segment I; 3 setae on segment 22; 2 on segments 3, 5, 8, and 21; 1 each on seg- ments 6, 9 to 15, and 17 to 20; short spines on segments 2, 4, and 7. Seta of segment 16 not found. Second antennae and mouth parts very similar to those of type male of sarsi. Proximal spine (fig. 2, C) of masticatory lobe of first maxilla (fig. 2, B) unbranched, bearing instead 3 spinules on distal side; palp like that of sarsi except for the presence of a small distally placed knob, which may represent the - rudimentary epipodal lobe mentioned by Sars. Leg 1 (fig. 2, D) with same number of setae as sarsi, inside seta of endopod stiff and spiniform, and set in a similar well-defined ridge. Second basal segment bearing inside a short, broad, curved process in addition to the long seta overlying the endopod. Exopod much like that of sarsi, second segment slightly broader with outer margin less elongate and inner more rounded; second segment of endopod much less lamellar, with distal half elongated and narrowed terminally so as to form a distinctly truncated end. Exopods of legs 2 to 4 like those of sarsi, but lacking inner setae or spines, as in pygmaea. Leg 2 (fig. 1, H) with a 3-segmented endopod, inner edge of first segment more bulging and rounded than in sarsi, first spine of inner edge of third segment short and smooth. (In sarsi (fig. 1, E) this spine is much more elongate and bears an inner serrated flange.) Endopods of legs 3 and 4 (figs. 1, J and 2, E) 2-segmented, each carrying a long inner seta on first segment. (This seta may be lacking in pygmaea. This seems-at least to be true of endopod 4.) Distal seg- ment of endopod 3 carrying 6 setae and spines, that of 4 with 7 spines. Leg 5 (fig. 2, F) differing from that of pygmaea in having the two inner spines of the third segment of exopod larger and with strongly serrate edges; endopod distinctly 2-segmented, first segment 1; B, maxilla nlarged; D, leg 1; , greatly e 1; H, endopod of leg 4; I, leg 5. tory lobe of maxilla 1 of mastica 2.—Platycopia tumida (C. B. Wilson). Female, A, antenna 1; C, pro spine i E, leg 4; F, leg 5. Male, G, antenna ximal Fic. (e LZ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 very narrow, second lamellar in shape, bearing 6 spines which de- crease in size from the outer to the inner edge. Male. Slightly smaller than female. The unique specimen examined showing in profile a small distally projected hook in front of the middle of the cephalic segment. Eye spot present, large. Urosome and cephalic appendages, except first antennae, like those of the fe- male. First antenna (fig. 2, G) with 15 segments, bearing 16 aesthetasks, all jointed near the middle, those on segments I and 5 exceptionally elongated ; many attached in an irregular fashion, giving the antenna the appearance of being almost covered by them. First segment with the ridge arising near its center, curved somewhat differently from that of the female, and bearing 3 very long aes- thetasks; the long slender spine set at the distal base of the ridge points diagonally across the segment. This segment carrying a fourth aesthetask near its distal end, 6 setae and 1 small spine along its length; the long modified spine not differing from that of other species. Other aesthetasks borne singly on segments 2-to 9, II, 12, 14, and 15. Four setae each on segments 14 and 15; 2 on segments 3, 5, 8, and I1; I on segments 2, 9, 12, and 13; short spines on segments 4, 6, 7, and 10. Legs 1 to 4 like those of female, except that endopod 4 (fig. 2, H) has the distal segment narrowed and the upper half of the inner edge invaginated. Fifth leg (fig. 2, 7) built on the plan of pygmaea, being broad and compact. Exopod with 3 distinct segments; first segment with the usual 2 outer spines, second bearing a long, stout spine with a broadened hyaline flange. Second segment constricted basally and attached to the first near its inner edge, broad distally and not set off sharply from third segment at inner margin; an elongate club-shaped hyaline process, arising in distal half near inner margin, overlies the surface of the succeeding segment for most of its length. Third segment with an irregular shape, inner margin straight and shorter than outer which is indented near the middle; distal part somewhat bulging and slightly irregular marginally. Outer margin bearing proximally a small spine with a hyaline rim. Distal inner spine saber- shaped like that of pygmaea, very thin and almost completely hyaline. Terminally there is another stout curved spine near the center of the segment, and toward the outer margin a thin hyaline spinelike process somewhat like the inner spine in shape. Overlying these are 3 very thin, ovoid hyaline lappets. These lappets are much larger than those in similar position in sarsi, with a more or less prominent riblike thickening in the center. Endopod 2-segmented, set somewhat deeply into the center of the basipod, lamellar in shape, with a large spine NO. Q SPECIES OF PLATYCOPIA SARS—WILSON 13 inside on the first segment, and 5 somewhat small, subequal spines on second. A long curving spine, about twice the length of the inner margin of first segment, set in basal segment near the inner juncture with endopod. Remarks—The notes found among C. B. Wilson’s papers and labeled Arenocalanus tumidus, n. sp., contained the following state- ments which indicate rather clearly that the female specimen he had dissected was the one described above: First antenna made up of 22 segments. .. . There is a large asethetask at the inner distal corner of the basal segment, another at the tip end of the segment and 7 others at intervals along the inner margin of the segment. ... The second endopod [referring to legs] is 3-segmented. In the third, fourth and fifth endopods the two distal segments are fused. This description disagrees with Dr. Wilson’s published drawing of the antenna which has 23 segments and bears 11 aesthetasks. How- ever, the original of this drawing was labeled and included among the notes with other inadequate sketches of the species, and un- doubtedly belongs with it. It seems likely that the whole represents only a preliminary description and that the inaccurate drawing was published before a complete study of the species had been made. The notes are on file at the National Museum with Dr. Wilson’s papers in the Division of Marine Invertebrates. DISCUSSION The genus Platycopia has always been considered an anomalous one. There are a few other bottom-living calanoids, among which are Pseudocyclops, Pseudocyclopia and Paramisophria, that have the same compact, somewhat cyclopoid body form and short first antennae, but otherwise are very different. The genus Pseudocyclops shows the closest affinities to Platycopia in that the fifth leg of the female is not transformed or reduced, but otherwise the relationship seems remote. The two genera differ markedly from one another in the structure of the cephalic appendages, especially the first antennae and the first maxillae. In Pseudocyclops the male right antenna is distinctly ge- niculate, and neither sex has aesthetasks similar to those of Platycopia. Though both genera have the legs somewhat flattened and carry broad spines with hyaline flanges, there is still considerable difference be- tween them. Leg 1 of Platycopia is very unlike the other pairs of legs, in Pseudocyclops leg 1 is not differentiated; in Platycopia the setae of legs 2 to 5 are almost wholly transformed to spines, in Pseudocyclops the setae are not transformed; and the peculiarities I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 of structure of the male fifth legs are not at all comparable in the two genera. Furthermore, Platycopia has several characters that are unique for the genus and set it apart from all other calanoids. In addition to the distinctive characters of the first maxillae and the transformation of setae to spines on both rami of legs 2 to 5, there are the presence of two spines on the first segment of the exopods of legs 2 to 5, the exact likeness of the right and left fifth legs in the male, and the similarity of the urosome in the two sexes. As no genus has yet been found to which it is very closely related, it is therefore best to retain the separate family Platycopiidae erected for it by Sars. Morphologically, the four known species of Platycopia fall into two groups and it will be interesting to note if future species will also tend toward one or the other of the two species originally described by Sars. Of these two groups, the perplexa—sarsi is the least variable, only the first antennae and fifth legs showing any marked differences. The pygmaea—tumida group, however, exhibits differences not only in the first antennae and fifth legs, but also in the endopods of legs 2 to 4. P. tumida is certainly the most anomalous of the species, having characters not only different from pygmaea, but exhibiting some that are unique in the genus, such as the presence of an eye spot, the unusual ridge of the first segment of antenna 1, the segmentation of the male fifth exopod and the slight sexual difference in the fourth endopod. There is some question as to just how much tumida differs from pygmaea as Sars did not describe and figure that species, especially the male, as completely as he did perplexa. Probably the species did not show any unusual characters in the first antennae; though not figured, they are described as “being composed of 16 joints” and if there had been unusual features they would certainly have been noticed by so careful a worker as Sars. It is not possible to compare exactly legs 3 and 4 as they are undescribed beyond mention of the incomplete division of the last 2 endopod segments, and only the female endopod of leg 4 is figured. The number of spines on the distal segment of endopod 3 differs from that of endopod 4 in the perplexa-sarsi group. This is also true in twmida, but the variation is not the same as that of the other group. Sars shows 5 spines on the distal segment of the fourth endopod of pygmaea, but one can only guess that this number is more or less than that on leg 3. Whether or not pygmaea shows a slight sexual variation in endopod 4 is also open to question, and it must be assumed for the present that it is a character belonging only to tumida. P. perplexa, sarsi, and NO. 9 SPECIES OF PLATYCOPIA SARS—WILSON I5 tunuda, though having very similar first legs, show some slight specific differences. Leg 1 of pygmaea is not illustrated so it cannot be de- termined how much, if any, it differs from the others. Certain things are, however, definitely known. The pygmaea- tumida group differs from the perplexa-sarsi in having no inside setae or spines on exopods 3 and 4, in the variability of the segmentation of endopods 2 to 5, and in the more compact structure of the male fifth leg. P. tumida differs from pygmaea in having 15 rather than 16 segments in the male first antenna, and 22 rather than 23 in that of the female. The second endopod of pygmaea is 2-segmented, that of tumuda is 3-segmented, indeed it is very similar to that of sarsi and perplexa. The endopods of legs 3 and 4 and the female fifth endopod are incompletely segmented in pygmaea; in tumida this incomplete- ness of division is so slight that I have considered them as being visibly 2-segmented. Furthermore, the number of spines on the endopods of leg 4, and possibly of leg 3, is different in the two species. In view of the very apparent natural relationships of the American to the Norwegian species, it is strange to find differences that seem to have no systematic explanation. There is, for instance, no mention in the descriptions of the Norwegian species of the unusual heavy “ridge” of the endopod of the first leg to which is attached the long, spiniform inner seta. This ridge stands out sharply in all the mounted specimens, and is present even in the most immature specimen of P. sarsi in which the endopod is still 1-segmented. I find it hard to believe that a character present in two such unlike forms as the American species should be absent in the equally unlike Norwegian forms. The presence of such a ridge is suggested in the original drawing of perplexa® although the seta is not so stout as it is in sarsi and tumida. There is little indication of it, however, in the subsequent illustration,* but the base of the inner seta is shown as larger than those of the other setae. It is possible, therefore, that this character is common to all representatives of the genus and was overlooked by Sars in describing the Norwegian forms. Likewise, the aesthetasks on the first antennae of the American species are jointed near the middle, but Sars does not mention such a condition or indicate it in his illustrations. The aesthetasks of Platy- copia are true aesthetasks and not modified setae, as they accompany rather than transplant the setae on every segment. They are very slender bodies, and their interior has a mottled appearance in glycerin 5 Sars, G. O., Archiv Math. Naturv., vol. 31, No. 7, pl. 2, fig. 1, 1911. 6 Sars, G. O., Crustacea of Norway, vol. 7, pl. 7, fig. 1, 1910. 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 mounts. They are constricted basally, also slightly at the joint, and usually the distal portion is somewhat more slender than the proximal. The line of division is a “joint” insofar as it represents a real joining of two portions; these lines can be seen to be somewhat thickened and circular in form when the aesthetask is turned at a certain angle. The American species further differ from the Norwegian in lacking the lamella shown by: Sars on the terminal portion of the male fifth exopod. Again it seems strange that these very closely related species should vary so much in what is usually a basic pattern, and in this instance, at least, it may be due to a difference in interpretation. From its position it seems likely that the irregular terminal portion of the segment in sarsi corresponds to the “lamella’’ pictured for perplexa, and that in sarsi it is a stouter structure and not so sharply demarcated from the rest of the segment. No such extension of the terminal portion is present in twmida and I am at a loss in visualizing the structure which Sars described as a “lamella” in pygmaea. Both sarsi and tumida have a few very sheer, hyaline lappets hanging from the terminal portion of the segment. Those in tumida are excep- tionally large. In both species it is necessary to focus carefully on these lappets, else only the edges are seen, giving the appearance of several cilia extending from the terminal portion. It could be that the cilia shown by Sars on the end of the “lamella” of perplexa represent the edges of comparable hyaline lappets. In the case of pygmaea the “lamella” may or may not be homologous to the lappets. I am inclined to consider, therefore, that the terminal portion of the exopod in the perplexa—sarsi group is somewhat narrowed and irregularly extended beyond the point of insertion of the inner modi- fied spine, and has hanging from it a few very thin, hyaline lappets, probably rather small. In the pygmaea—tumida group the terminal part of the segment is not so extended and the hyaline lappets are attached closer to the point of insertion of the inner spine. The “lamella” of Sars is considered in this view to represent in perplexa the extension of the segment, and in pygmaea a lappet. I find no other way to interpret the latter when it is compared to the very similar twmida, which has comparatively conspicuous lappets. Only reexamination of the Norwegian species can, of course, decide these points. “SMITHSONIAN MISCELLANEOUS . COLLECTIONS _ VOLUME 106, NUMBER 10 , @ --\ REEXAMINATION OF THE _ FOSSIL HUMAN SKELETAL REMAINS FROM MELBOURNE, FLORIDA with, FURTHER DATA ON THE VERO SKULL a | (Wire E1cHt Sik EN y BY. T. D. STEWA RT _ Curator, Division of Physical Anthropology EUS Ss National Museum | (PUBLICATION 3854) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION — AUGUST 9, 1946. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 10 A REEXAMINATION OF THE FOSSIL HUMAN SKELETAL REMAINS FROM MELBOURNE, FLORIDA WITH FURTHER DATA ON THE VERO SKULL (With EicHt PLaTEs) BY T. D. STEWART Curator, Division of Physical Anthropology U. S. National Museum (PUBLICATION 3854) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION AUGUST 9, 1946 The Lord Baltimore ress BALTIMORE, MD., U. 8. As A REEXAMINATION OF THE FOSSIL HUMAN SKELETAL REMAINS FROM MELBOURNE, FLORIDA WITH FURTHER DATA ON THE VERO SKULL By T. D. STEWART Curator, Division of Physical Anthropology U. S. National Musewm (WirH 8 PLATEs) INTRODUCTION The Christmas scientific meetings held in New Haven in 1925 were enlivened by a session of the Paleontological Society at which J. W. Gidley exhibited a crushed human skull from Melbourne, Fla., which, he suggested, had been stepped on by a mastodon or mammoth. Ordinarily the paleontologists might not have been excited by Gidley’s offering and its implication of human antiquity on this continent. However, news of his scheduled paper had reached the American Anthropological Association, meeting several blocks away, and this body, led by that veteran antagonist of the antiquity of man in America, Ales Hrdlicka, came en masse to hear the debate. This increased attendance, besides showing the general interest in the subject, also gave emphasis to its controversial nature. The anthropologists, or at least the more vocal of them, were in- tent on defending the generalization, more widely held at that time than now, that man first came to this continent in recent times. The paleontologists, on the other hand, were defending basic concepts of their science; namely, that the faunal inclusions in a stratum when found undisturbed and not redeposited are indicative of the relative age of the stratum, and also that such faunal associations represent in themselves. contemporaneity of existence. In the tense atmosphere engendered by these different viewpoints, the session proceeded with rather more dignity than some had antici- pated. Gidley somewhat reduced the controversial issue by sug- 1 The secretary of the Paleontological Society reported the event thus: “At 2 p.m. the Society convened and had as its guests visiting archeologists and ethnologists to hear the geological evidence for fossil man in Florida. “A spirited discussion followed Dr. Gidley’s presentation, which was illustrated by some of the human remains discovered and by lantern slides. Among the speakers was Dr. Ales Hrdlicka, who made an appeal for closer affiliation be- tween the ethnologists and geologists in studies of ancient man.” [P. 239.] SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 10 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 gesting “. ..a relatively recent date, either late Pleistocene or even post-Pleistocene, for the extinction of the last survivors of the Pleistocene fauna in the south” (Gidley, 1926a). In leading the discussion, Hrdlicka refused to accept this dating and reasserted some of the arguments he had used against the antiquity of the similar finds of E. H. Sellards at Vero, Fla. (Hrdlicka, 1918, pp. 34-38). The essence of these arguments is: (1) that the chances for the accidental inclusion of human remains, especially of more than one individual, in a geological deposit are infinitesimal, and (2) that, judging from Old World finds, “As late as the Aurignacian culture period, approximately 15,000 to 25,000 years ago, man had not yet fully reached modern standards in physical development . . . and cannot possibly be conceived of as having been numerous enough to reach the northeasternmost limits of Asia, from which alone there was a practical way open to the American continent” (p. 37). Since Gidley denied the existence of any evidence pointing to intrusive burial, whereas the anthropologists felt that such evidence could have been overlooked, the debate, needless to say, ended in a draw. e The next year Gidley and Loomis, as co-excavators at Melbourne, both together and separately restated their conclusions: Gidley: .... there was no difficulty in recognizing over a wide area at Melbourne the three principal geologic horizons designated by Sellards at Vero as Nos. 1, 2 and 3. This made the correlation of the beds of the two localities comparatively easy. It was found that at both localities, all the fossil bones taken from Sellards “No. 2” layer were primarily deposited and were definitely of Pleistocene age. Many of the bones of the lower part of No. 3 were also of this age but were often mixed with bones of more modern species. Also that “No. 3” layer usually lies unconformably upon the somewhat unevenly eroded surface of “No. 2.” “No. 3” layer throughout contained numerous evidences of man, apparently of no great antiquity, while no remains of this character were found, in the lower portion at least of “No. 2.” However, at Melbourne there were found, at three relatively widely separated areas, human bones or artifacts associated with un- disturbed, and not redeposited, fossil bones of the Pleistocene fauna. These finds were all near the top of “No. 2” level, just below the contact plane [see plate 1, A and B]. As no human remains or artifacts were found below the top layer of “No. 2” it is assumed that man arrived in Florida about the close of the time marked by the finished deposition of “No. 2” or during the erosion interval between it and “No. 3” and that he seems to have found there a late survival of the Pleistocene fauna, certain species of which may have persisted in the south later than did their relatives in the north country. [1926b, p. 26.] Looms: While this No. 3 bed was forming, the bones of mastodons and mammoths were - included in the material along with both human bones and tools. The whole NO. 10 SKELETAL REMAINS FROM MELBOURNE—STEWART a3 question now hangs on when this last filling in took place. The only data which are now available are the bones of the mastodon and mammoth. At first thought one would say it must have been either in late Ice Age or very shortly after the Ice Age. The end of the Ice period is put at 25,000 years ago. The only possibility of a later date than 15,000 to 20,000 years ago as a mini- mum, lies in the possibility of these elephants surviving in Florida to a later date than elsewhere. In these beds we are not dealing with a remnant of the elephant fauna. They occur in great numbers. . . . The elephants were flourishing during the time that No. 3 bed was laid down. Much the same is true in other parts of the country. Just before they disappeared they were very abundant, then they were all gone, as though swept off by a pestilence. There is no reason to think they lasted longer in Florida than elsewhere. All this, then, would indicate that man of the Neolithic type of culture was in Florida 20,000 years ago at least. [1926, p. 262.] Gidley and Loonus: . it may safely be assumed that man, although of modern type, reached Florida before the total extinction of the mammoths and mastodons. But that this arrival occurred far within Pleistocene time does not seem probable. Mam- moths and mastodons are known to have survived in the Great Lakes re- gions up to the time of the formation of the swamp deposits which followed the last retreat of the glaciers in North America, and it seems not improbable that a considerable remnant of the Pleistocene fauna survived the glacial periods in Florida and other localities of the southern and southwestern United States, and remained there after the great ice sheets had disappeared in the North. However this may be finally concluded, the present evidence as here interpreted seems to indicate quite clearly that the time of man’s first appearance in Florida should be placed at a time much earlier than has hitherto been supposed, possibly early post-Pleistocene, and that these people seem to have preceded the Indians who built the mounds characteristic of the east coast of Florida. [1926, pp. 263- 264. | Gidley continued his search for elephants in Florida each year until his death in 1931. In the course of this work, although he found no more ancient human bones, he seems to have fortified his original convictions regarding the natural association of the human and elephant remains and hence their relative antiquity. (See Ex- plorations and Field Work of the Smithsonian Institution for the respective years. ) FIRST RECONSTRUCTION About 1937 the Melbourne skull and left clavicle were transferred from the Division of Vertebrate Paleontology to the Division of Physical Anthropology in the United States National Museum, where 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 they bear catalog No. 331,402.? At that time, as when exhibited in New Haven, the specimens were still encased in plaster of paris (pl. 2). However, in order to demonstrate the type of the skull, Hrdlicka had one of the minor museum preparators undertake its reconstructicn. In this process, since many parts were missing, the pieces were applied for stability to a mass of modeling clay (plasti- cine). The result (pl. 3) was discussed by Hrdli¢cka at the Inter- national Symposium on Early Man held in Philadelphia in 1937 and exhibited during the same year at the meeting of the American As- sociation of Physical Anthropologists in Cambridge (Hrdli¢ka, 1937a and b). On both occasions he characterized it as the usual type of the Indian crania found in the mounds of this part of Florida. Although this about covers the history of the Melbourne skull, note should be made of an important coincidence relating to the year 1925, when this skull was found. This was the year in which the Folsom fossil pit—across the continent in New Mexico—was first reported. Here in 1926 Figgins was to find the first Folsom point associated with an extinct bison. Since then many other chipped blades of this peculiar type have been found in widely scattered localities, especially in the Southwest, and usually under conditions suggesting a relative antiquity (Roberts, 1939). Unfortunately, no human skeletal remains have been found associated with these artifacts. However, the cir- cumstances surrounding their recovery, both geological and paleonto- logical, constitute the best evidence thus far in favor of ancient man in America. As such it gives renewed significance, as J. C. Merriam has pointed out (1935), to the Florida finds of Gidley and Loomis. It may be noted also in this connection that Sellards, who is well acquainted with these developments in the West, reaffirmed in 1937 his views regarding his Vero finds of 1915-16. As he views the situation after an interval of 20 years, The conclusion that is in accord with the observations at Vero, and at many other localities in the western hemisphere, is that man reached this continent before the close of the Pleistocene . . . and participated in the great drama otf the extinction of the magnificent mammalian fauna of that period. [P. 210.] 2 Gidley and Loomis (1926, p. 259) speak of pieces of finger, arm, and leg bones found near the skull. When the present study was almost completed (1946) these parts, along with some other pieces of the skull, were found in Gidley’s collection in the Division of Vertebrate Paleontology. These parts also have been given the number 331,402 in the Division of Physical Anthropology. Since these extra parts do not change the conclusions already reached, it has been decided not to recast the report, but rather to add a section at the end (see pp. 24-27). Throughout the text, wherever statements require modification on account of this new material, proper footnotes have been added. NO. I10 SKELETAL REMAINS FROM MELBOURNE—STEWART 5 From the above summary it appears that the Melbourne skull, although one of the very few thus far found in America for which such an impressive age is claimed, itself has received only minor attention. Aside from considerations as to its position in the deposits and its flattened condition when found, the sole description by an anthropologist (Hrdlicka, 1937a) is contained in one sentence supple- mented by three small illustrations. This description reads as follows: It is the skull of an Indian male, of advanced adult age, undeformed, brachy- cranic, high, and in general presenting the ordinary Florida mound Indian type and characters. [P. 08.] The mound crania from this part of Florida are indeed relatively broad- and high-headed (Hrdlicka 1940a; see also below). Distri- butional studies (Stewart, 1940), on the other hand, support the belief that the earliest cranial type on the continent was long- and high-headed. What then is the explanation of the rounded form reported by Hrdlicka for the Melbourne skull? Is it, in spite of the paleontological evidence, actually as recent as Hrdli¢ka con- tended? Or were there more than one early type, both long- and round-headed? Or is the reconstruction at fault and the specimen really long-headed ? . SECOND RECONSTRUCTION I have long felt that the crude method used in the original recon- struction of the Melbourne skull was reason enough for reexamining its form. In this version, owing to the solid mass of clay within, only the outer surface of the skull is visible. This makes it difficult to judge whether all fragments are in their proper positions and, where fragments have been united, whether they are tightly joined. The proper union of the fragments is especially important because this mainly determines the form of the total reconstruction. If, as seems to have been the case, the fragments were visually oriented on the surface of the ball of clay, then their relationships depend more upon the size and shape of this ball than upon their proper union. Besides, there are certain morphological details, clearly visible in the illustra- tions, that cast doubt upon this reconstruction. We shall return to these details later. I should point out also that thus far the Vero and Melbourne skulls have been studied only by Hrdliéka, who definitely was biased on the subject of man’s antiquity on this continent. Now that there is more reason to accept the paleontological evidence of man’s presence in America during late Pleistocene or early post-Pleistocene times, it seems high time to examine the skulls from this different point of view. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Recently I have detached the fragments of the Melbourne skull from the ball of clay, cleaned them and made a new reconstruction. In this second version the shape of the whole has been determined primarily from the union of the parts. Temporary support for the assembled fragments was gained by the use of wire braces held in place by ambroid. At a later stage when the result was deemed satisfactory, the gaps caused by missing parts were filled in with a commercial “crack filler.’ Usually this gave sufficient strength so that the braces were no longer needed. Finally, when all the parts that could be joined were in place, certain distinctive parts, such as the temporal bones and zygomatic processes of the frontal bone, which lacked actual contacts, were approximately located. Here also the gaps in the temporal and supraorbital regions were filled in for support. Plate 4 shows the result of this second reconstruction. It is important to indicate the limitations imposed upon the restorer by the imperfections of the fragments. The better to illustrate these details, stereographic drawings have been made of the new version with the aid of the Schwarz stereograph (fig. 1, A-D). Inspection of figure 1, A, makes it clear that only in the midparietal region is there continuity of bone between the right and left sides. The total length of the preserved part of the sagittal suture, which provides this contact, amounts to 3 cm. Furthermore, along the edges of the suture, only the inner table of the bone is preserved, the outer table being irregularly broken away for distances up to 7 mm. laterally (this detail is also shown in pl. 6). The recovered portions of the two parietals thus join along a thin edge. Accordingly, this joint can be moved like a hinge, narrowing or broadening the skull as a whole. In joining these parts I was guided simply by the fit of the edges and my original joint has not been (although perhaps it should be slightly) modified. Aside from the sagittal suture the only indication of the midline is preserved in the form of the frontal crest on the inner side of the frontal bone. This crest is a part of the set of fragments making up much of the left side of this bone. Since damage makes the exact alignment of the frontal and parietal bones along the left coronal suture somewhat uncertain, the only solution was to align the frontal crest with the sagittal suture. Along the right side of the coronal suture, where likewise there was considerable damage, the alignment of the fragments could be judged by the pattern of the vascular impressions on the inner surface (pl. 5). The only other questionable joints occur on the left parietal just above the superior limit of the squamous suture (fig. 1, C). It is SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 067) NO to; REST A, view of a drainage canal at Melbourne, Fla., and the excavation near it, in which was found the human skull. B, the human skull just before it was bandaged. The arrow points to the skull. Above it is No. 3 layer, below is No. 2 layer. (Captions as given by Loomis, 1926, p. 261.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 10, PL. 2 Spi ct os The Melbourne skull and left clavicle mounted in plaster of paris as exhib‘ted in 1925. The box seen here as a frame is recalled as being only about 3 inches deep. This indicates the flatness of the crushed specimen. Also, owing to the paleontological technique of remov- ing such specimens, the parts here exposed were originally on the under side as the specimen was discovered. In witness of this fact, note the crepe paper passing under the clavicle and under the skull. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 10, PL. 3 Four views of the Melbourne skull as reconstructed under Hrdliéka’s direction. Note the roundness of the specimen and the abnormal distance between ear opening and lateral border of the eye. X indicates a misplaced piece of frontal bone. Compare details with the writer’s reconstruction shown in plate 4. A cast of the original is shown here oriented as nearly as possible on the Frankfort horizontal. Approximately 4%; natural size. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 10, PL. 4 Four views of the writer’s reconstruction of the Melbourne skull oriented on the Frank- fort horizontal. Note the elongated form and compare with the earlier reconstruction shown in plate 3. These views are interpreted in stereographic drawings (fig. 1, A-D). The posterior part of the left parietal was found and placed in position after these views were made (fig. 6). Approximately '% natural size. Fic. 1.—Stereogrz (C) norma lateralis : stipple = natural bon endocranial surface; re ee Ww ie Z oy PU OC wu ir y 1 ee re 2 uy hy Fic. 1.—Stereographic drawings of the writer’s reconstruction of the Melbourne skull in (A) norma verticalis, (B) norma facialis, (C) norma lateralis sin., and (D) norma lateralis dex. X indicates cut surface. Compare with photographs in plates 4 and 6. (Coarse ‘tipple = natural bone surface; fine stipple = broken bone surface; included white areas = supporting “filler”; heavy dots = natural tdocranial surface; crosshatch = openings.) %%4 natural size. » NO. 10 SKELETAL REMAINS FROM MELBOURNE—STEWART 7 believed that imperfections in this area account for the flatness of this side as contrasted with the right (fig. 1, B). RECONSTRUCTIONS COMPARED Let us now compare the two versions of the reconstructed Mel- bourne skull. Examination of the photographs (pls. 3 and 4) will show that more parts have been included in the first than in the second version. This is because the second version depends, with few exceptions, upon proven contacts between fragments, and no such contacts can be found for some of the smaller fragments. Indeed, some of the smaller fragments seen in the frontal region of the first version seem to have been placed in position without any justification. The same can be said about the assemblage of fragments occupying the midline near the occiput in the first version (indicated by the X in pl. 3). This assemblage is quite definitely a part of the frontal bone (right side) and has been located accordingly in the second version. In proof of this, as already indicated, I call attention to plate 5, which shows a positive impression of the vascular markings on the inner surface of the right fronto-parietal region. The symbols Fy, F2, and F, denote parts of the frontal bone; P,, Pz, and Ps, parts of the parietal. The assemblage P,, F;, F2, and F; forms a unit with tight joints. Between P, and P, there is a defect in the bone; between P, and P, the joint surface is very thin, being damaged externally and narrowed internally by a vascular impression; and between F, and P; there is likewise contact of the inner table alone. The correct relationship of these fragments is indicated by the continua- tion of the vascular impressions as indicated by the arrows. Also, it is evident from figure 1, A, that the included portion of the coronal suture is symmetrical with that of the other side. In figure 1, A and C, attention is called to a feature of the second re- construction that seems to prove the correctness of the assemblage of fragments composing its left parietal. This feature, which is shown in more detail in plate 6, A and B, consists of an artificially cut edge, beginning at the sagittal suture about 7 cm. back of bregma and running laterally and somewhat diagonally backward. This edge is interrupted by a circular break in the bone, and hence consists of a medial portion 4.5 cm. long and a lateral portion 1.5 cm. long, separated by a gap 4 cm. wide. Nevertheless, both portions of this edge appear to be in the same plane (pl. 6, B), both being beveled so as to face upward as well as backward.* In Hrdlicka’s version 3 This reconstruction was definitely verified by the finding of additional pieces of this parietal (see pp. 24-26). 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 these two portions of the cut edge are not in alignment, and, more- over, in contact with the medial portion is the fragment of right frontal discussed in the foregoing paragraph. The further significance of this cut edge will be discussed later. The method used in reconstructing the second version of the Mel- bourne skull made it possible to obtain an endocranial cast. Besides further verifying the reasonably correct relationships of the cranial fragments, this cast has served to demonstrate two cranial features, the existence of which is completely hidden in the first version. Both of the features are internal surface configurations reflecting, un- doubtedly, brain shape. The first of these features is an asymmetry of the frontal area. Here the right side is flatter than the left. The result is a rather pronounced crest along the midline extending as far back as bregma. There is no evidence of this shape externally ; in- deed, the forehead is nicely rounded. The second feature of the endocranial cast is represented externally by the gap in the artificially cut posterior parietal edge described in the preceding paragraph (pl. 6, A and B). At this point the cast shows a marked prominence. Between the surviving portion of this prominence and the midline is a depression about 5 mm. deep. On the other sides there is a more gradual slope. And, of course, the posterior part, as well as the eminence is missing. The bone sur- rounding the eminence is reduced in thickness as can be seen in plate 6, A, but still amounts to 5 mm. This structure certainly repre- sents a brain tumor of some sort. The occurrence of two such unusual features in the same area, namely, a brain tumor and an artificial opening in the bone, may be a coincidence, because the nature of the cut is not clearly related to the shape of the tumor. Let us turn now to the measurements, which show perhaps the most striking differences between the two versions. Table 1 shows that the first version is hyperbrachycranic, whereas the second version is dolichocranic. For further comparison with the first version I have used a hyperbrachycranic skull from Tampa Bay (U.S.N.M. No. 243,689). No skull with a breadth of 162 mm. is listed among the 700-odd specimens from Florida in the Catalog of Crania (Hrdlicka, 1940a) ; indeed, less than 11 percent of those judged to be undeformed have a cranial index exceeding 85. For comparison with the second version I have used a skull of about the same size and shape from Pensacola Bay (U.S.N.M. No. 242,665). Dolichocrany (70.0-74.9) 4 The subsequent finding of the posterior part of this parietal shows that the swelling in the endocast slopes off gradually in this direction. NO. 10 SKELETAL REMAINS FROM MELBOURNE—STEWART 9 is rare (4 percent) among the Florida skulls listed in the Catalog of Crania. © Comparing now the first version with the round-headed Tampa Bay specimen a great discrepancy will be seen between all measure- ments involving the landmark on the upper border of each ear open- ing known as porion (measurements Nos. 3, 5, 6, and 7). The TABLE I.—Comparative measurements of the two versions of the reconstructed Melbourne skull and of two undamaged skulls representing extreme recent Florida types U.S.N.M. U.S.N.M. 1st No. 2d No. Measurement (mm.) or index version 243,680 version 242,665 Tem Vicrmtiatita len oth. . << a ciewe cite oo cee 3 (182) 1 180 (186) 184 Deed LEA CEL joc. jacocccsseteiem «,acyoie 162 154 136 136 Cranial index (22222), Pi tputets d ans 89.0 85.0 73.1 73.9 I 3. Biporion-bregmatic height ......... 134 124 122 124 Upper mean height index Ox) We yeni td te AR 779 74.2 75.8 76.9 mean of I & 2 4. Minimum frontal diameter......... 102 100 94 97 Fronto-parietal index (422) 68% 64.9 60.1 71.3 2 5. Ectoconchion-porionic length, right. 110 (72) 72 (74) 6 ‘ ae left.. 105 (77) 66 76 Porion position index ( et ofS EEX 100 ) ee Ore 50.1 (41.4) 37.1 (40.8) 7abiporionic breadth <.V.00002%6 Js... 148 124 109 113 Biporion-parietal index (o'") 1.4 80.5 80.1 83.1 2 1 Measurements and indices in parentheses are based upon estimates. distance between the two ears (measurement No. 7: biporionic breadth) in the first version is 24 mm. greater than in the Tampa Bay specimen. Similarly, in measurement No. 3, the biporionic line is Io mm. farther from bregma; and in measurements Nos. 5 and 6 as averaged, the distance from the lateral border of the orbit (ecto- conchion) to porion is 33 mm. greater. Without attempting to furnish more proof, it can be said categorically that the position of the ears in the first version of the Melbourne skull is freakish. No such discrepancies exist between the measurements of the second version and the long-headed skull from Pensacola Bay. Slight distortion of the former probably is indicated by its lesser biporionic [oO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 breadth, minimum frontal diameter, and left ectoconchion-porion length; but these measurements probably are still within normal limits. Summarizing these findings, the two reconstructions of the Mel- bourne skull represent extremes of skull shape; hyperbrachycrany (89.0) in the first version and dolichocrany (73.1) in the second. Only about 15 percent of undeformed Florida Indian skulls fall into these combined classes; the great majority (85 percent) fall into the intermediate classes of mesocrany and brachycrany. It seems certain that the width of the vault arrived at in the first reconstruction (162 mm.) is excessive. Also, the positions of the temporal bones and some of the frontal and parietal fragments in this version are definitely incorrect. In the main, however, the tremendous difference in shape represented in the two versions depends upon the imperfect joint involving about an inch and a half of the sagittal suture in the mid- parietal region. Since this joint, in addition to being damaged ex- ternally, is the sole contact between the two sides, it is capable of a hinge action. As this hinge is opened up, the vault becomes broader, and as it is closed, the vault becomes narrower. The accuracy of the reconstruction depends to a considerable extent, therefore, upon this joint. Although the facts thus far presented probably will convince most readers that the second version of the Melbourne skull is the better of the two, doubtless many are still uncertain as to how nearly the new reconstruction approaches the original form. Owing to the pattern of the missing parts, no one could hope to reach perfection in reestablishing the relationships of the remaining parts.’ Therefore, the best that can be done is to aim for a reasonable reconstruction. This, I believe, has been achieved. THE, VERO. SKULL The question that arises at this point is: How does this new version of the Melbourne skull compare with the Vero skull, for which an equal age is claimed by the paleontologists and which also comes from Florida? Before answering this question it is necessary to discuss the Vero skull. 5]T spent considerable time trying to show that the curvature of the parietal bone in transverse section is closely related to the horizontal shape of the vault. If such a relationship could be found, it could be used in the present case, because the original transverse curvature of the parietal is preserved. However, I found the same kind of parietal curvature in both dolichocranic and brachycranic skulls from Florida. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 10, PL. 5 Plaster cast of endocranial surface in the region of the right half of the coronal suture. F,_, indicate three portions of the frontal bone. P,, indicate three portions of the right parietal. Breaks between these parts have been inked in on the photograph. Arrows show the course of vascular impressions crossing the breaks and thus verifying the correct restora tion of the parts. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 10, PL. 6 Two views of the left parietal showing cut surface. A, the arrows indicate the extent of the two preserved portions of this surface. The circular break between the pairs of arrows perhaps has resulted from a thinning of the bone in this region, which in turn may have been caused by a brain tumor. Note also that the contact between the two parietal bones at the sagittal suture is limited to the inner table. B, the arrows and line of dashes indicate that the cut surface is a plane. : s ; * Ur ae ae a _ wr ‘) norma lateralis dex., and ( e stipple = broken bone surfa ce of supporting “filler.”) ™% nz SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 10, PL. 7 Four views of the Vero skull oriented on the Frankfort horizontal. Note the slender con- nection between the frontal and right parietal. These views are interpreted in stereographic drawings (fig. 2, A-D). Approximately 1% natural size. ‘a . . e as 3 . d D) Fic. 2.—Stereographic drawings of the Vero skull in (A) norma verticalis, (B) norma facialis, (C) aap 2 ae norma occipitalis. Compare with photographs in plate 7. (Coarse stipple = natural bone surface; fine stipple = ro “EI r.”) 4 natu- included white areas — supporting “filler”; heavy dots = natural endocranial surface; circles = inner surface of supporting er. ral size. er ie SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 10, PL. 8 cm. Fragments of bones of the extremities belonging to the Melbourne skeleton. A, left humerus; B, right humerus; C, two parts of the left tibia; D, right tibia: E, left ulna: F, metacarpal (probably right fourth); G, left fibula. The line through the right tibia (D) is at the level of the cross section shown in figure 7. NO. 10 SKELETAL REMAINS FROM MELBOURNE—STEWART II According to Hrdlicka’s study of this skull (1918, pp. 55-57) the vault is incomplete, being made up of 16 pieces mostly from the right side. However, all these pieces join firmly and are sufficiently ex- tensive so that the shape can be determined with some exactitude. Although Hrdli¢ka’s illustration (his pl. 10) shows a dolichocranic form, he fails to give measurements or to mention this fact. His description is limited to the following statements: The skull is large, finely shaped, of thoroughly modern features, and unusually thin. It shows a normal nasal process of moderate breadth (about 2.1 cm.) ; orbits with fairly sharp borders; glabella and supraorbital ridges subdued; forehead not high but well built; the sagittal region oval from side to side; the temporal ridges only mildly marked and at their nearest approach running at a distance of 7 cm. from the median line; parietal eminences gently developed; the occiput smooth and blunt; and the outline of the vault as viewed from above forming a fine ellipse. The thickness of the parietals ranges from 3 to 4 mm., which for a strong Indian is decidedly exceptional. The mastoid process shows normal masculine development; the auditory meatus is rather large and in cross section nearly circular; the zygomatic processes were of moderate masculine strength. [P. 56.] Elsewhere (p. 55) he states that, The skull and bones are Indian, but they seem to belong to a type such as can occasionally be found among the Eastern Algonquian, or among the Sioux, rather than in Florida. The Vero skull was reconstructed by Dr. Riley D. Moore, who was at that time Scientific Aid in the Division of Physical Anthropol- ogy. After the skull had been studied, it was returned to the Florida Geological Survey, but a cast, also made by Dr. Moore, was retained in the Museum (No. 299,245). This cast is unsatisfactory because the position of bregma is not indicated. This landmark is important in the orientation of a skull lacking one of the temporal bones. In order to approximate the pdsition of the conventional Frankfort plane, which is determined by the two ear openings and the lower border of the left orbit, it is necessary to judge the location of the missing ear opening from the relationship between the other ear and the median sagittal plane. In the Vero skull the median sagittal plane can be determined from the three preserved landmarks: nasion, bregma, and lambda. Thus, in order to make certain of the location of bregma, I found it necessary to consult the original.’ 6 Since the sagittal and coronal sutures are almost entirely closed ectocranially, their juncture is determined in the original more by color than by relief. The photographs do not help in locating this point. 7I am indebted to Dr. E. H. Sellards for his help in arranging for the loan of thé specimen and to Herman Gunter, State Geologist of Florida, for granting the loan. IZ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 When the original Vero skull was sent to me, | discovered to my surprise that, like the Melbourne skull, it too was filled with modeling clay (plasticine). Moreover, a block of plaster of paris had been attached to the clay to form a base, and where the clay showed in the skull vault shellac had been applied. Obviously, this was almost as unsatisfactory a specimen for study as had been the first version of the Melbourne skull. Also, because of its weight, it could not be set up in the Schwarz stereograph. And to make matters worse, the oil in the clay had kept the shellac soft and sticky. Owing to these circumstances, I requested and was given permission by Mr. Gunter to remove the plasticine and to reconstruct the parts as in the case of the Melbourne skull. The result is shown in plate 7 and interpreted in the series of stereographic drawings shown in figure 2, A-D. An examination of these illustrations will convince the reader that the surviving portion of the frontal depends for its orientation upon a short (less than 3 cm.) connection with the parietal on the right side. Since the skull as a whole is remarkably thin, and at this point does not exceed 3 mm. in thickness, it should be apparent that con- siderable care is required in reestablishing this joint. Depending upon the nicety of this articulation, then, the glabellar region will vary in position in an anterior-posterior direction, and consequently the maximum length of the skull will be affected. The drawings represent, I believe, the best reconstruction possible and certainly do not exaggerate the length.® MELBOURNE AND VERO SKULLS COMPARED It should be apparent also from the illustrations of the Vero skull that it is similar in shape to the Melbourne skull as newly re- constructed. In order to show this more clearly I shall compare the two specimens both metrically and graphically. Also, because of the metric similarity shown in table 1, I shall include in the comparison the recent dolichocranic Florida skull from Pensacola Bay (U.S.N.M. No. 242,665). The measurements of these three specimens are shown in table 2. According to these figures all three skulls are remarkably similar in shape; all have about the same degree of dolichocrany, and about the same fronto-parietal relationships. However, the Vero skull 8 After the drawings and photographs were made, the frontal became detached in the course of making an endocranial cast. As finally restored by one of the museum preparators, the glabellar region has a more anterior position than in my restoration. As a result, the maximum length is several millimeters greater. NO. IO SKELETAL REMAINS FROM MELBOURNE—STEWART 13 differs from the other two in its relatively low vault (upper mean height index) and thin wall. Graphically these relationships are brought out well in figure 3, A-C. In superimposing the outlines of these three skulls the following parts were made to coincide: 1, the midline; 2, the most anterior projecting part of the supraorbital prominence; and 3, the Frankfort plane. This TABLE 2—Comparative measurements of the Vero, Melbourne (2d version), and Pensacola Bay skulls Melbourne U.S.N.M. 2d No Measurement (mm.) or index Vero version 242,665 Meeiatmitm length: joc aastees sos snot cee wee EOS) (186) 1 184 Pee Marnier VDLEACEH: 6 56.6 d:eidie bis Oy et ee We ks 5 4 fs test: Peli? py SO Lae PY Pt | a he ie EP pepe OSes. thy t fH rs Za UE. Thomas Lincoln Casep Fund ReVIeW OF THE NEW WORLD SPECIES: OF “HIPPO- DAMIA DEJEAN (COLEOPTERA: COCCINELLIDAE) * By EDWARD A. CHAPIN Curator, Division of Insects, U. S. National Museum (WitH 22 PLATES) INTRODUCTION The present paper is offered as a point of departure for an adequate study of the many intricate and fascinating problems in the taxonomy of this division of the family Coccinellidae. Much more collecting and study will have to be devoted to the species of the quinquesignata, sinuata, and convergens groups before a true picture of these species and ali their subspecific segregates is revealed. It is hoped that the data assembled here will be of use in that future study. ACKNOWLEDGMENTS Without the aid of many individuals, always generously given, this paper could not have been written. The writer thanks them all gratefully: Th. Dobzhansky for the use of his collection of more than 2,000 specimens and for his stimulating advice and encourage- ment; John E. Blum, O. L. Cartwright, J. C. Chamberlin, Wm. L. Jellison, G. P. MacKenzie, F. T. Scott and others for smaller col- lections of critical material which made possible the solution of cer- tain problems encountered ; Gerhard H. Dieke and P. H. Timberlake, who have read portions of the manuscript and offered valuable suggestions. Thanks are also due Mrs. Rhoda F. Mislove for her painstaking labors in preparing hundreds of dissections of specimens, both male and female, and for assistance in the preparation of the distributional maps. It should be noted that in preparing the drawings showing spot patterns, the elytra have been drawn as plane surfaces to avoid the effect of foreshortening; thus the tips of the elytra appear to be separated. * This is the fifth contribution to be published by the Smithsonian Institution under the Thomas Lincoln Casey Fund. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, No. 11 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 MATERIAL In all, something over 11,000 specimens of Hippodanua have been examined. The National collection, with the collections of Thos. L. Casey, H. F. Wickham, and the Brooklyn Museum of Arts and Sciences, contains nearly 9,000 specimens, including representatives of all known species and subspecies. Next in importance is the Dobzhansky collection of more than 2,000, rich in material from the far West and from Mexico and Central America. Other small but important lots were made available by interested colleagues. It is to be regretted that no adequate samples of the various palae- arctic species are available to the writer and therefore this paper must confine itself to the New World components of the genus. HISTORICAL Dejean, in his “Catalogue des Coléoptéres,” 3d edition, p. 456, 1836, introduced the generic name Hippodama into literature. Although he credits the name to Chevrolat, it appears that Dejean’s use ante- dates that of Chevrolat. Eleven species, with synonyms, were in- cluded, but no type species was designated. Mulsant (1850) accepted the genus in approximately the same sense as Dejean’s and also failed to designate a genotype. Crotch (1874) selected Coccinella 13- punctata L., as genotype. This, through synonymy, was one of the originally included species. Prior to 1919, all workers who dealt with species in the genus Hippodamia Dejean were concerned solely with the external ap- pearance of the individuals. This species differed from that one because of the presence or absence of a spot on the elytron. The result was chaos, for, without exaggeration, it may be stated that except for a few it is absolutely impossible to identify a “described” species unless one has access to the type material. In his 1919 paper (Journ. New York Ent. Soc., vol. 27, pp. 162- 174) P. H. Timberlake laid the foundation for an adequate classifica- tion of the genus as it exists in the New World. Four groups of species were recognized at that time. Of these the present writer accepts three with the same limits placed upon them by Timberlake, divides his fourth group into two coordinate groups, and proposes. another and new group for a recently described species. THE GENUS HIPPODAMIA The genus Hippodamia is a not large genus of Coccinellidae whose species show an inordinate amount of variation in the pronotal and NO. II HIPPODAMIA—CHAPIN 3 elytral markings. Its species are found throughout the holarctic region, but because of a lack of material from the palaearctic area the present study is confined to the species of the New World. It may be defined as containing those species of Coccinellidae in which the femora extend beyond the sides of the body when directed outwardly, in which the pronotum is without basal marginal bead and with nearly straight anterior margin, with claws toothed at middle, with first tarsal segment of male rarely dilated, with two spurs on each middle and hind tibia and without metacoxal arcs. The sclerotized portions of the male genitalia conform to a pattern which is found in most parts of the family. The following description of the organ as it is in H. parenthesis (Say) serves well as an example. The terms employed are those used by Dobzhansky (following Ver- hoeff) and have been adopted because they appear to have been more widely used than any of the other terminologies. Synonymous terms are given in parenthesis. The sipho (aedeagus, median lobe) is in the form of a tube, U- shaped when relaxed and at rest, straight when extended during copulation. At its basal end it is hinged to the free extremity of the trabes. To accommodate this attachment, the basal extremity is flattened and widened, somewhat rounded and with a minute median notch. The sperm duct enters at a pore on the outer side just in front of the flattened portion and somewhat in front of this pore but on the inner side there is a pronounced bump or protuberance. The basal third of the sipho is heavily sclerotized and pigmented. At the beginning of the middle third, the inner side is not sclerotized but membranous, which tends to increase the flexibility of the organ. Near the apex, the inner side again becomes completely sclerotized while the outer side is membranous and furnished with a lateral, triangular or rounded flap on either side. The extreme apex is very slender and terminates in delicate fingerlike processes. The basal plates (tegmen, theca) are fused into a ring through which the sipho passes. This ring is broad above and very narrow beneath, at which point the trabes is attached. The paramera are articulated at either side of the pemis. The penis (posterior lobe of theca) is a more or less triangular sclerite, intimately fused with the basal plates and lying between the paramera. Its apex is acute and barbed. With the ventral alae, it forms the guide for directing the stpho during copulation. The paramera (lateral lobes) are paired structures arising from the basal plates, somewhat finger-shaped and furnished in their apical thirds with moderately long hairs. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 The ventral alae are paired structures lying along the margins of the penis and attached to that sclerite by membrane. Their other margins are free and apically each is extended in a slender process. The ventral alae and penis together form a three-sided channel through which the sipho slides. The trabes (strut) is short and somewhat triangular, is attached at its base by an articulation to the basal plates, and at its apex by mus- cles to the proximal end of the sipho at its base. It functions in the ex- tension and retraction of the sipho. The sclerotized portions of the female genitalia are of little use in separating closely related species but may be used in defining the groups of species. The description given below is generalized and does not apply in its entirety to any one species. The bursa copulatrix (uterus) is a more or less elongate sac aris- ing just behind the dorsal and ventral plates (sternite and tergite VIII). It is usually unlined but in the tredecimpunctata group it bears on its inner wall patches of short conical spines. Leading away from the bursa copulatrix at or near its upper end is the sperm duct, a delicate tube whose union with the bursa is strengthened by the heavily sclerotized accessory piece. The form of this accessory piece varies somewhat with the groups of species. At the other end of the sperm duct is the very heavily sclerotized sausage-shaped receptaculum senunis. Its shape is quite constant in the species of Hippodamia, but in many genera of Coccinellidae it offers excellent specific characters. GROUP I—TREDECIMPUNCTATA This group is characterized by the somewhat oblong rather than oval body form (due chiefly to the relatively greater width of the pronotum) ; in the male sex by the strong development of the ventral alae of the penis, and by the small, subtriangular, closely approximated dorsal flaps of the sipho ; and in the female sex by the numerous fine dark sclerotized denticles which line portions of the bursa copulatrix. The tredecimpunctata group contains four species, all of which are represented in the Western Hemisphere. The typical species, 1. tredecimpunctata (1.), breaks up into three subspecies, two of which, H. tr. tredecimpunctata (L.) and H. tr. timberlakei Capra, are from the Old World. The third, H. tr. tibialis (Say), is widely distributed over the northern half of North America. Closely related to the above is the usually misidentified H. americana Crotch, whose range, ac- cording to our present knowledge, is restricted to the area from NO. II HIPPODAMIA—CHAPIN 5 Hudson Bay south to Lake Superior and west to Saskatchewan. Two other species, closely allied inter se but distinctly separated from tre- decimpunctata and americana, belong in this group. These are H. falcigera Crotch from Canada and the northern Rocky Mountains area and H. washington Timberlake from the States of Washington and Oregon. HIPPODAMIA TREDECIM-PUNCTATA (L.) Fics. 1-9, 29, 48-55, 227 Coccinella 13-punctata LINNE, 1758, Syst. Nat., ed. 10, p. 360. Hippodamia tredecim-punctata MuLsant, 1850, Species Coléopt. Trim. Sécuri- palpes, p. 10; 1866, Monogr. Coccinellides, pt. 1, p. 8. Hippodamia 13-punctata Crorcu, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 368; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 77; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, p. 50. Coccinella tibialis Say, 1824, Journ. Acad. Nat. Sci. Philadelphia, vol. 4, p. 94. Hippodamia tibialis TIMBERLAKE, 1919, Journ. New York Ent. Soc., vol. 27, p. 165; Korschefsky, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 331. Hippodamia tibialis subsp. timberlakei Capra, 1931, Boll. Soc. Ent. Italiana, vol. 63, p. 17. This species with its subspecies inhabits the holarctic region. The typical subspecies is distributed generally over Europe and western Asia and the subspecies H. tr. timberlakei Capra in Japan, China, and eastern Siberia. These subspecies need no further discussion in this paper. The remaining subspecies, H. tr. tibialis (Say), is North American and is widely distributed in the area north of the 37th parallel of latitude. Rarely is it found as far south as South Carolina. HIPPODAMIA TR. TIBIALIS Say Type locality —‘‘Missouri.” Characterized by-the disk of the pronotum being largely black, without oblique white marks and with the elytral pattern of the full component of spots. .. H. tr. tibialis is in general a very constant subspecies, showing only slight variation in its color pattern, not at all comparable to that displayed by the typical subspecies in Europe. Individuals having two or more of the spots coalesced are rare, and in the series of more than 550 individuals that is before me, there is no specimen which lacks completely any one of the normally occurring spots. Through the kindness of Dr. J. C. Chamberlin, more than 300 specimens of this species from Matanuska, Alaska, have been available for study. The lateral prolongations of the ventral alae are moderately long 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 and rounded at their tips, the marginal prolongations are somewhat longer, bent outward at their middle and terminate in acute, some- what feebly hooked apices. The penis is broadly triangular at its apex, which is not reflexed. The denticles which occur on the inner wall of the bursa copulatrix are distinctly grouped in four longitudinal bands. HIPPODAMIA AMERICANA Crotch Fics. 10, 30, 56-57, 228 Hippodamia americana CrotcH, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 368; Wickham, 1894, Can. Ent., vol. 26, p. 306; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, p. 52, fig. 39; Korschefsky, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 338. Type locality—Hudson Bay. Crotch described this species, without designating a holotype, as from “Kansas, Hudson Bay.” In his 1910 paper, Johnson set-Hud- son Bay as the type locality and that fixation is accepted here. The Kansas record certainly refers to another species (spuria Lec.?). Little can be said concerning the variation in the spot pattern on the elytra. The linking of spot 3 with its counterpart across the suture at right angles to the elongation of spot 4 appears to be an important characteristic of this species. The absence of spot 2 and the tendency of spots 1, 5, 4, and 6 to join to form a longitudinal stripe suggests a close relationship between americana and falcigera. The lateral prolongations of the ventral alae are short, and abruptly rounded, the marginal prolongations are longer, more slender, somewhat bent outward and terminate in simple subacute apices. The penis is acutely triangular at its apex, which is reflected at its extreme tip. Denticles of the bursa copulatrix arranged much as in tr. tibialis. There is a single male specimen of this species in the national col- lection which was taken by Hubbard and Schwarz on Whitefish Point, Lake Superior, Mich. A female from Waskesu, Saskatchewan, June 10, 1938, C. Shaw, collector, was made available for study by the kindness of F. T. Scott. No other specimens have been seen by me. HIPPODAMIA FALCIGERA Crotch Fics. II, 31, 58-62, 229 Hippodamia falcigera Crorcu, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 368; Casey, 1899, Journ. New York Ent. Soc. vol. 7, p. 81; 1908, Can. Ent., vol. 40, p. 399; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, p. 55. NO. II HIPPODAMIA—CHAPIN 7 Hippodamia sinuata subsp. albertana Casey, 1924, Mem. Coleopt., vol. 11, p. 157. Ceratomegilla cottlei’ NUNENMACHER, 1934, Pan-Pacific Ent., vol. 10, p. 20. Type locality—Slave Lake, Hudson Bay. Nine specimens available to me for study come from Edmonton, Alberta (including the type material of H. s. albertana), Yellowstone Park, Wyo. (type locality of C. cottlei), Moscow, Idaho, Hudson Bay and Stewart River, Yukon Territory. This species does not ap- pear to vary greatly in its markings or size; one specimen shows an unusual lack of pigment in the pronotum and another has the discal vittae of the elytra broken in two parts. The extension of spot 4 as a sutural stripe and the absence of spots 2 and 3 appear to be characteristic. The lateral prolongations of the ventral alae are long, well sclero- tized and terminate in strong, outwardly directed hooks. The marginal prolongations are straight and acute. The penis is broadly rounded with a narrow truncate apical termination. The denticles of the bursa copulatrix form a single patch instead of being arranged in four bands. HIPPODAMIA WASHINGTONI Timberlake Fics. 12, 33, 63-67, 230 Hippodamia washingtoni TIMBERLAKE, 1939, Proc. Hawaiian Ent. Soc., vol. 10, p. 265. Type locality—Longmire Spring, Mount Rainier, Wash. Twenty-three specimens, including two, paratypes, are available for study. Of these, 15 were collected at Hoquiam, Wash., four in the Blue Mountains of Oregon, and two on Mount Hood, Oreg. The pigmentation of the pronotum varies only slightly in the series of specimens before me, but there is a considerable range in the elytral pattern in the Hoquiam series. No evidence of subspeciation can be detected, however, as the series is evenly graded from heavy to light pigmentation. This is the only species of the group which occurs minus all its elytral spots. In spotted individuals, it appears to be characteristic for spots 4 and 3 to be linked with their counterparts to form an inverted T-shaped spot. Spot 2 is usually present. The lateral prolongations of the ventral alae are long, as in falcigera, but are only slightly turned outward at their tips. The penis is more elongate and its apical termination is broader than in falcigera. The denticles of the bursa copulatrix are in a single patch. GROUP II—PARENTHESIS The parenthesis group is characterized by the rather small oval form, by the single median marginal pale spot on the base of the pro- 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 notum, by the frequent linkage of spots 4 and 3, the usual suppression of spot 2 and the usual linkage of spots 5, 4, and 6; in the male sex by the low, elongate dorsal flaps of the sipho and in the female sex by the thin collar at the distal end of the accessory piece beyond the limit of heavy sclerotization. The group contains four species, all of which are of North Ameri- can origin and no one of which is known to occur naturally in the Old World. These, in chronological order of their description, are parenthesis (Say), widely distributed from the Atlantic to the Pacific, lunatomaculata Mots., apicalis Casey, and expurgata Casey, the last three in western North America. Two of these appear to be separable into two subspecies each. HIPPODAMIA PARENTHESIS (Say) Fics. 13, 34, 68-73, 231 Coccinella parenthesis Say, 1824, Journ. Acad. Nat. Sci. Philadelphia, vol. 4, Dp. 93. Adonia parenthesis MULSANT, 1850, Species Coléopt. Trim. Sécuripalpes, p. 40. Hippodamia parenthesis CrotcH, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 368; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 81; Leng. 1903, loc. cit., vol. 11, p. 44; Casey, 1908, Can. Ent., vol. 40, p. 399; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, p. 52; Timberlake, 1919, Journ. New York Ent. Soc., vol. 27, p. 165; Korschefsky, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 343. Coccinella tridens Kirpy, 1837, in Richardson, Fauna Boreali-Americana, pt. 4, p. 229; Crotch, 1874, Revision Coccinellidae, p. 97. Type locality.—“United States.” Notwithstanding the fact that this is a common species over a wide range, the variation in the color pattern is less than in some of the other geographically more restricted species. It may be noted that there is only a slight tendency for spot 1 to join with the sub- apical lunule (spots 5, 4, and 6) to form a longitudinal discal stripe, and there is no tendency at all for the subapical lunule to be extended in the direction of the apical angle. Also, the reduction of pigmenta- tion of the elytra is not coordinated with a reduction of pigmentation of the pronotum. On the contrary, the specimen before me that shows the least pigment in the elytra shows at the same time an extension of pigment in the pronotum which completely obliterates the median basal pale spot. This particular specimen is the only one of the more than 300 specimens which completely lacks the spot. The species may be immediately recognized by the barbed apex of the penis. NO. II HIPPODAMIA—CHAPIN 9 The range of this species extends from the north Atlantic seaboard west to Alaska, British Columbia, and California and it is the only one of the group known to live east of the Mississippi River. In the east it occurs as far south as South Carolina and to the north into Ontario, Canada. A definite type locality was not given in the original publication, and the type specimen is lost. HIPPODAMIA APICALIS Casey Fics. 14, 35, 74-86, 232 Hippodamia apicalis Casry, 1899, Journ. New York Ent. Soc., vol. 7, p. 81; 1908, Can. Ent., vol. 40, p. 399; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, p. 54. Hippodamia lunatomaculata subsp. or var. apicalis TimMBERLAKE, I919, Journ. New York Ent. Soc., vol. 27, p. 166. Hippodamia lunatomarginata ab. apicalis KoRSCHEFSKY, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 342. Hippodamia lengi JOHNSON, 1910, Carnegie Inst. Washington, Publ. 122, p. 55; Timberlake, 1919, Journ. New York Ent. Soc., vol. 27, p. 167. Adalia nigromaculata NUNENMACHER, 1934, Pan-Pacific Ent., vol. 10, p. 20. (= engi Johnson.) The apex of the penis is simple and the apical portion is not set off from the main portion by a sudden constriction producing sharp lateral angles as in H. expurgata Casey. Roughly, the distribution of this species is in a horseshoe-shaped area extending from Colorado north to British Columbia and thence south through California. As yet no specimen has been seen from Arizona or New Mexico, where the related H. cxpurgata Casey is abundant. HIPPODAMIA A. APICALIS Casey Type locality—Reno, Nev. This subspecies is characterized, in its more heavily pigmented forms, by a strong tendency for the humeral spot and subapical lunule to unite and form a longitudinal discal stripe. Further, the reduction of the pigmented areas of the elytra is in general coordinated with a similar reduction of pigment of the pronotum. Lastly, except in rare cases, the elytral suture at apex carries a trace of dark pigment even in specimens where the pigment reduction has been the greatest. The range of this subspecies is that of the species except for its apparent absence from southern California. HIPPODAMIA A. LENGI Johnson Type locality —California (San Diego County ?). In this subspecies there is no tendency visible toward the forma- 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 tion of longitudinal discal stripes. Instead, the suture is dark through- out its length and in specimens with intensified pigment, the humeral spots are joined with the scutellar spot. Nowhere else in this group of species does one find a black sutural stripe. As in the typical subspecies, the pigmentation of the pronotum is coordinated with that of the elytra. At present this subspecies is known only from San Diego County, Calif. A short series from Jacumba is available for study. HIPPODAMIA LUNATOMACULATA Motschulsky Fics. 16, 36, 87, 97-106, 234 Hippodamia lunatomaculata MotscHvutsky, 1845, Bull. Acad. Nat. Sci., Moscow, vol. 18, p. 382, pl. 7, figs. 8, 8’; Timberlake, 1919, Journ. New York Ent. Soc., vol. 27, p. 167 (part). Hippodamia parenthesis CasEy, 1899, Journ. New York Ent. Soc., vol. 7, p. 81 (part) ; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, p..53 (“‘sub- species of Oregon”). Hippodamia lunatomarginata KorscHEFSKY, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 342. Hippodamia lunatomaculata dobzhanskyi, new subspecies. The range of this species is from the San Francisco Bay region of California north into Alaska. In external appearance it is not to be distinguished from H. parenthesis (Say) in all specimens. The two are, however, amply separated by the form of the male genital organ. The apical portion of the penis in this species is set off from the main portion by a sudden constriction, and its extreme apex is slightly enlarged. As in H. apicalis, two subspecies may be distinguished. HIPPODAMIA L. LUNATOMACULATA Motschulsky Type locality California, vicinity of San Francisco Bay. The original description of this species is as follow: 78. Hippodamia lunatomaculata m. Tab. VII, fig. 8, 8’. H. septemlunata ? Eschsch. Dej. Cat. Oblonga, nigra, nitida; ore, antennis, capitis thoraceque limbo, elytrisque testaceis, his maculis nigris septem 1, 4, I, I. Long. 2} lign.—larg. 14 lign. Par sa taille et ses couleurs cette espéce ressemble a la H. mutabilis, mais elle est plus grande. Les élytres sont testacées avec une tache noire commune sous l’écusson, une autre de la meme couleur a l’angle huméral, une troisiéme sur le milieu et une tache allongée en forme de lunule vers l’extrémité. Sur les bords latéraux des segments de l’abdomen on voit des taches blanches. Elle se trouve en Californie et je crois que c’est a cette espéce qu'il faut rapporter l’H. septemlunata du Catalogue du Comte Dejean, au moins c’est la NO. II HIPPODAMIA——-CHAPIN II seule espéce de ce genre que j’ai pu trouver parmi les Californiens de feu Eschscholtz. Eschscholtz visited California twice, both times as naturalist at- tached to the expeditions of Otto von Kotzebue. On the first voyage, 1815-1818, von Kotzebue took the Rurik into San Francisco Bay, where she stayed from October 1 to November 1, 1816. She departed on the latter date for the Hawaiian Islands. On the second voyage, 1823-1826, the Predpriatie dropped anchor at the same place on Sep- tember 27, 1824, and remained there until the 25th of November of that same year. During this period, field trips were made as far south as Santa Clara and as far north as Ross Colony (Fort Ross), 80 miles north of San Francisco Bay. An expedition was also made up the Sacramento River for some miles. Therefore the actual type locality of Hippodamia lunatomaculata Mots. is within 80 miles of San Francisco Bay. Among the specimens submitted to me for study by F. T. Scott, there is a single specimen from Half Moon Bay, Calif. It is fortu- nately a male, and an examination showed it to belong, not to paren- thesis (Say) which it superficially resembles, but to the species which lives mainly along the coasts of Oregon and Washington. Its color pattern (fig. 102) compares very favorably with that illustrated by Motschulsky for the type of lunatomaculata (fig. 87). With this specimen I associate a series of 16 specimens taken by Th. Dobzhansky at Willows, Calif. These individuals are characterized by the heavy pigmentation of the pronotum and the unusually light pigmentation of the elytra, and I consider them also to be true H. lunatomaculata Mots. With more material available it may become evident that the Willows population should be set apart under a new name. HIPPODAMIA L. DOBZHANSKYI, new subspecies Type locality—Port Angeles, Wash. Type (male) and 20 paratypes (df and 92) collected at the type locality August 12, 1934, by Th. Dobzhansky, U.S.N.M. No. 57891. Paratypes in collections of Th. Dobzhansky, G. H. Dieke, F. T. Scott, J. E. Blum, and G. P. Mackenzie. orm and structure of typical H. lunatomaculata Mots. but dif- fering from that form in the notable reduction of pigment in the pro- notum coupled with a notable extension of the pigment in the elytra. This is the dominant form in the coastal areas of Oregon and Wash- ington. To the north, the species is known from Vancouver Island. I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 HIPPODAMIA EXPURGATA Casey Fics. 15, 32, 88-96, 233 Hippodamia parenthesis subsp. expurgata CaseEy, 1908, Can. Ent., vol. 40, p. 400. Hippodamia lunatomaculata subsp. or var. expurgata TIMBERLAKE, 1919, Journ. New York Ent. Soc., vol. 27, p. 166. Hippodamia lunatomarginata ab. expurgata KorsCHEFSKY, 1932, Junk-Schenk- ling, Coleopt. Catalogus, pars 120, Coccinellidae IT, p. 342. Type locality.—Boulder, Colo. The color pattern of H. expurgata parallels that of H. apicalis ex- actly as the color pattern of H. lunatomaculata parallels that of H. parenthesis. There is the same tendency to form longitudinal discal stripes in the heavily pigmented forms. The forms with reduced pigmentation show one small point of difference: the dark area at the extreme tip of the suture is usually absent. There also appears to be no correlation between the degree of pigmentation of the elytra and that of the pronotum, which remains moderately heavily pig- mented throughout the range of variation. In the more than 260 speci- mens of this species examined, there is no evidence of subspeciation apparent. In normal individuals of this species, sides of the penis are parallel nearly to the apex. At about apical fifth, they converge rapidly but without forming acute angles as in H. lunatomaculata. The extreme apex is sharply triangular and is turned up at right angles to the main axis of the penis. Certain individual males from series of otherwise normal individuals have the apical portion of the penis somewhat attenuate as in /7. apicalis. The degree of elongation does not seem to be constant and it is possible that these are hybrids between expurgata and apicalis. This species is essentially a southwestern one with its center of distribution in Utah and Arizona. It is also known from Nevada, Idaho, Wyoming, South Dakota, Nebraska, Colorado, and New Mexico. GROUP III—GLACIALIS Two species are the members of a group which is characterized by certain conformations of the male genitalia. The penis carries, near its apex, a transverse wrinkle which has become elevated to form a sharp, high, backwardly directed crest. The margin of the crest is, in H. quinquesignata, entire and slightly rounded. In H. glacialis the crest is deeply and broadly emarginate. The sipho is also character- istic in that it is strongly swollen or inflated behind the apical flaps. NOs) II HIPPODAMIA—-CHAPIN 13 The apical flaps are reduced in size in H. quinquesignata but large in H. glacialis. In either case, they are directed forward rather than outward. Hippodanua quinquesignata (Kby.) embraces, in addition to the typical subspecies, two subspecies inhabiting the West Coast, both of which are characterized by immaculate elytra. H. glacialis (F.) is represented in North America by four subspecies, two of limited distribution and two of wider range. HIPPODAMIA QUINQUESIGNATA (Kirby) Fics. 19, 37, 127-146, 237 Coccinella quinquesignata Kirsy, 1837, in Richardson, Fauna Boreali-Americana, VOlrAN DE 230; pls 7 fies I. Hippodamia quinquesignata MutsAnt, 1850, Species Coléopt. Trim. Sécuri- palpes, p. 15; 1866, Monogr. Coccinellides, pt. 1, p. 10; Crotch, 1873, Trans. Amer. Ent. Soc., vol 4, p. 366; 1874, Revision Coccinellidae, p. 95; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 78; Leng, 1903, loc. cit., vol. II, p. 40, pl. 4; Casey, 1908, Can. Ent., vol. 40, p. 395; Timberlake, 1910, Journ. New York Ent. Soc., vol. 27, p. 171; Korschefsky, 1932, Junk- Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 344. “Hemisphaerica n. gen., typical species quinquesignata Kirby” Hope, 1840, Coleopt. Man., pt. 3, p. 157. Hippodamia mulsanti Leconte, 1852, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 131; Crotch, 1873, Trans. Amer. Ent. Soc. vol. 4, p. 366; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 78; Timberlake, ro1g9, loc. cit., vol. 27, p. 169. (= quinquesignata (Kby.).) Hippodamia ambigua Leconte, 1852, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 131; Crotch, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 366; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 79; Timberlake, 1919, loc. cit., vol. 27,) p. 172. Hippodamia punctulata Leconte, 1852, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 131; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 79; Timber- lake, 1919, loc. cit., vol. 27, p. 172. Hippodamia leporina MuLsAnt, 1856, Opusc. Ent., vol. 7, p. 135; Korschefsky, 1932, Junk-Schenkling, Coieopt. Catalogus, pars 120, Coccinellidae II, p. 344. (= quinquesignata (Kby.).) Hippodamia obliqua Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 790; Timberlake, 1919, loc. cit., vol. 27, p. 172. (=ambigua Lec.) Hippodamia subsimilis Casry, 1899, Journ. New York Ent. Soc., vol. 7, p. 79; Timberlake, 1919, loc. cit., vol. 27, p. 172. (= uteana Csy.) Hippodamia vernix Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 79; Timberlake, 1919, loc. cit., vol. 27, p. 171. (= uteana Csy.) Hippodamia uteana Casey, 1908, Can. Ent., vol. 40, p. 397; Timberlake, 1910, Journ. New York Ent. Soc., vol. 27, p. 172. Hippodamia uteana quadraria Casey, 1924, Mem. Coleopt., vol. 11, p. 156. (= uteana Csy.) Hippodamia convergens pugetana CAsry, 1924, loc. cit., p. 156. 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 H. quinquesignata (Kirby) with its subspecies has essentially a western and northern distribution. Typical specimens predominate in the material seen from Michigan northwestward to Alaska. South from Alaska and more or less following the mountains the maculate forms occur, reaching as far south as Arizona and New Mexico and as far west as central Washington, eastern Oregon, Nevada, and eastern California. The Olympic peninsula of Washington is also to be included in this area. It is possible in long series to differentiate the more lightly marked variety pugetana Casey of the Washington district from the more heavily marked variety uteana Casey of Utah, Nevada, and eastern California. In general, this species shows a gradual reduction in pigment of the elytra to the west and a gradual increase in pronotal pigment to the south. The available specimens of this species number more than 1,200. Several very interesting population samples assist greatly in the under- standing of the taxonomic problems involved. The more important of these samples are: Mount Logan, Utah, 9,713 it., May 13, 1939, Knowlton and Nye, 68 specimens. All specimens heavily maculate, a mixture of typical quinquesignata (Kby.) and uteana Csy. Argus Mountains, Calif., 5,500 ft., May 30, 1933, Dobzhansky, 41 specimens. Most of the specimens are uteana Csy., a few typical quinquesignata (Kby.). Summit of Peavine Peak, 8,270 ft., near Reno, Nev., January 16, 1923, H. S. Barber, 188 specimens. Most specimens heavily maculate, none immaculate; a few typical quinquesignata (Kby.) but most uteana Csy. Port Angeles, Wash., August 12, 1934, Dobzhansky, 72 specimens.. Thirty-nine percent (28 specimens) of the lot is a mixture of q. punctulata and q. ambigua. This is the most northerly record for q. punctulata Lec. The remaining individuals are maculate forms of various types, only 3 or less than 4 percent of the whole being typical quinquesignata. Modesto, Calif., fall of 1928, A. O. Larson, 60 specimens. All gq. punctulata Lec. except for four faintly dotted specimens. Willows, Calif., June 23, 1932, Dobzhansky, 40 specimens. All but one are q. punctulata Lec. Willamette Valley, Oreg., February 5, 1931, A. O. Larson, 47 specimens. Eight specimens show traces (or more) of maculation ( =obliqua Csy.) ; the rest are H. g. ambigua Lec. In the Casey collection, a series of 59 individuals from Fairfield, NOs it HIPPODAMIA—CHAPIN I5 Wash.’ (apparently part of the large lot studied by Johnson in 1910) has been divided between species as follows: 19 specimens form the type series of H. convergens pugetana, 16 were identified as H. vernix Csy., 11 as H. caseyi Johnson, 10 as H. spuria Lec., 2 as H. americana Cr., and a single specimen as H. uteana Csy. Disregarding for the present the 23 specimens which belong to spuria and caseyi (ameri- cana of Casey=spuria Lec.), we find nearly every possible spot pat- tern except those of typical quinquesignata and the immaculate sub- species ambigua and punctulata. It appears significant that in the series of 15,415 specimens analyzed by Johnson, there were no im- maculate specimens. As the material available to me shows that there is no geographic correlation in the distribution of members of the several types of maculation and as there appears no other means of separating these types, I have reduced H. pugetana and H. uteana to the rank of variety and have maintained the immaculate West Coast forms, H. ambigua and H. punctulata, as subspecies. HIPPODAMIA Q. QUINQUESIGNATA (Kirby) Type locality.—‘Lat. 65.” From a study of the itinerary of Richardson’s journey, I conclude that this refers to a place near Great Bear Lake. Specimens agreeing with the original description show a union of spots 4, 1, and 3 to form a transverse bar near the bases of the elytra, the absence of spot 2, the union of spots 4 and 5 to form a transverse subapical bar on each elytron, and the presence of spot 6. The pro- notum is furnished with oblique discal marks. This is the dominant form in the northern part of the range of the species and occurs frequently throughout the range except that it is rare on the Pacific coast. HIPPODAMIA Q. QUINQUESIGNATA var. PUGETANA Casey Type locality —Fairfield, Wash. ? The type specimen has the following pattern: Spots 4 and 3 united, *If my assumption is correct that Colonel Casey received this material from Mr. Johnson, there is some question whether these specimens came from Fair- field or Kamiack Butte. On page 25 of his 1910 paper, Johnson mentions lots of specimens of 1,406 from Fairfield and 15,415 from Kamiack Butte. Two pages later he states “One large lot taken from the top of Marsh Hill, Fair- field, Washington, contained 15,415 individuals... .” It seems unlikely that Johnson counted two separate lots of 15,415 specimens each, 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 spot I is absent, spot 2 feebly developed, spots 4 and 5 well developed, spot 6 similar to spot 2. Pronotal oblique marks well developed. This variety as well as the next, occurs sporadically throughout the range of the spotted forms of the species. HIPPODAMIA Q. QUINQUESIGNATA var. UTEANA Casey Type locality—Sevier Lake, Utah. Similar to the last except that spot I is present and well developed, spot 2 absent, spots 4 and 5 usually united, and with the pronotal marks reduced or absent. A commonly met-with variety in Utah and eastern California but by no means confined to that area. HIPPODAMIA Q. AMBIGUA Leconte Type locality —“‘California and Oregon.” Characterized by the absence of all elytral spots except spot 4 and by the presence of heavy oblique discal marks on the pronotum. Western Oregon is the center of distribution of this subspecies. Specimens are found on the eastern shore of Puget Sound and oc- casionally in northwestern California and southern Washington. HIPPODAMIA Q. PUNCTULATA Leconte Type locality—San Francisco, Calif. Similar to the last in elytral pattern but with the pronotal markings completely or almost completely lacking. This is the southern counterpart of H. q. ambigua. Abundant in western California and the Channel Islands, it is met with occasionally as far north as Port Angeles, Wash. : HIPPODAMIA GLACIALIS (Fabricius) Fics. 17-18, 40, 45, 107-126, 235-236 Coccinella glacialis Fasricius, 1775, Syst. Ent., p. 80. Hippodamia glacialis Mutsant, 1850, Species Coléopt. Trim. Sécuripalpes, p. 18; 1866, Monogr. Coccinellides, pt. 1, p. 12; Crotch, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 367; 1874, Revision Coccinellidae, p. 95; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 79; Leng, 1903, loc. cit., vol. 11, p. 41; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, p. 19; Timber- lake, 1919, Journ. New York Ent. Soc., vol. 27, p. 171. Hippodamia extensa MuLsant, 1850, Species Coléopt. Trim. Sécuripalpes p. 17; 1866, Monogr. Coccinellides, pt. 1, p. 11; Crotch, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 366; 1874, Revision Coccinellidae, p. 95; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 79; Leng, 1903, loc. cit., vol. 11, p. 41; Timberlake, 1910, loc. cit., vol. 27, p. 171. NO. II HIPPODAMIA—CHAPIN 7 Hippodamia lecontei Mutsant, 1850, Species Coléopt. Trim. Sécuripalpes, ap- pendix, p. 1010; 1866, Monogr. Coccinellides, pt. 1, p. 9; Crotch, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 366; Casey, 1919, Journ. New York Ent. Soc., vol. 7, p. 78; 1908, Can. Ent., vol. 40, p. 396. Hippodamia convergens var. extensa JOHNSON, 1910, Carnegie Inst. Washington, Publ. 122, p. 23. Hippodamia convergens var. lecontei JOHNSON, 1910, loc. cit., p. 23. ?Hippodamia convergens var. pseudoglacialis JOHNSON, 1910, loc. cit., p. 23. ?Hippodamia hoppingt NUNENMACHER, 10934, Pan-Pacific Ent., vol. 10, p. 21. Hippodamia glacialis mackenziei, new subspecies. Unfortunately it is not possible for the writer to place exactly the two forms Hippodamia convergens var. pseudoglacialis Johnson and Hippodamia hoppingi Nunenmacher. The first of these is apparently mentioned only once in Johnson’s paper and the entire description is as follows: “Spots 4, I, 4+5, 6, pseudoglacialis (new variety) ; New Mexico and northward.” No type material is mentioned as preserved in any collection. Hippodamia hoppingi was described from a short series of specimens from Mt. Stillman, California. There is nothing in the description to indicate whether the species belongs in the glacialis or in the convergens group. The assignment of the two names to the synonomy of H. glacialis is not based on accurate knowledge. I am unable to separate the three “species” glacialis F., lecontei Muls., and extensa Muls. in a satisfactory manner. All the male speci- mens of extensa available to me have been dissected and the emargina- tion of the transverse crest of the penis does not differ from that in specimens of lecontei. As there appears to be no other character of importance by which the two forms may be separated, I have reduced the status of extensa to the rank of a subspecies. When specimens of glacialis from eastern United States are com- pared with specimens of lJecontei from New Mexico, the two may usually be separated according as the depth of the emargination of the crest is less (glacialis) or greater (/econtet) than half of the total length of the crest and apical portion of the penis. However, in a series of specimens taken at Watch Hill, R. I., there is noticeable variation from much less than half to nearly half, and in another series from Rocky Ford, Colo., both types with intergrades occur. There is also a difference to be noticed in the length of the slender tip of the sipho as compared with the length of the dorsal flap. These measurements must be taken from wet or balsam preparations if they are to be of value. In H. glacialis, the length of the dorsal flap is generally noticeably less than half as long as the apical portion of the sipho (measured from the origin of the flap to the apex). On the 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 contrary, the dorsal flap in H. lecontei is generally a little more than half the length of the corresponding portion of the sipho. If the length of the flap is expressed as a percentage of the length of the apical portion of the sipho, the values for H. glacialis are from 35.7 percent to 40 percent in New England and along the Atlantic coast, to 46.1 percent at Clemson College, S. C., and Lake Okoboji, Iowa. For H. lecontei, values of from 45.8 percent to 52.6 percent were measured in a series from Rocky Ford, Colo., other specimens from Idaho and Arizona falling between these limits. The value of the proportion rises again in the far West, reaching a maximum of 57.1 percent in a specimen of H. extensa from Alameda County, Calif., and in a specimen of the melanic H. mackenziei from Glacier Lodge, Inyo County, Calif. In view of the variations above noted, I am considering H. glacialis (F.) as a widely distributed North American species which breaks up into four reasonably well-defined subspecies. HIPPODAMIA G. GLACIALIS (Fabricius) Type locality —“America boreali.” Characterized by the frequent absence of spots 4, 1, and 2, the habitual absence of spot 3, the habitual linkage of spots 4 and 5, and the presence of spot 6, which is often linked with spot 4. The large majority of specimens will have only spots 4 plus 5 and 6 and with spot 2 feeble or absent. Pronotum usually with oblique discal marks. Widely distributed over the eastern half of North America, from southeastern Canada south to South Carolina, west to North Dakota, and south through Colorado to Oklahoma and Alabama. HIPPODAMIA G. LECONTEI Mulsant Type locality—Santa Fe, N. Mex. Given erroneously as “Santa Fe de Bogota” in the original pub- lication. Compared with the typical subspecies, characterized by a general reduction of pigment of the elytra and by the frequent union of spots 4, 1, and 3 to form a basal transverse bar across the elytra. Pronotal markings absent or feebly defined. This subspecies occurs in the area immediately west of that oc- cupied by H. g. glacialis, that is, from Colorado to Idaho, Utah, and Arizona and south into New Mexico. NO. II HIPPODAMIA—CHAPIN 19g HIPPODAMIA G. EXTENSA Mulsant Type locality.—*‘Californie septentrionale.”’ Characterized by the entire absence of spots 2, 4, 5, and 6, and by the frequent linking of spots 4, 1, and 3 to form a basal transverse bar. Pronotum without oblique discal marks. This subspecies appears to be confined to the coastal area of California in the vicinity of San Francisco. Most specimens seen are from Alameda County. HIPPODAMIA G. MACKENZIEI, new subspecies Type locality—Inyo County, Calif. Type (male) and paratypes (both sexes) from Glacier Lodge, August 2-8, 1942, G. P. Mackenzie, paratypes (both sexes) from Independence, June 17, 1937, J. E. Blum, U.S.N.M. No. 57892. Paratypes in collections of G. P. Mackenzie, J. E. Blum, Th. Dobz- hansky, F. T. Scott, and G. H. Dieke. Form and structure as in H. g. glacialis (Fabr.) but with a general increase in pigmentation. In the series available for study, more than half of the specimens are unusually heavily pigmented. In the darkest individual, spots 4, I, 3, 4, 5, and 6 are united to form a butterfly- shaped spot covering about two-thirds of the elytra, with spot 2 separate and distinct. The first reduction in this pigmented area, as seen in the series before me, is the freeing of spot 6. Further reduc- tion eliminates spot 2 and gives a pattern similar to H. g. lecontei. Apparently this form is confined to the high mountain area in eastern California. GROUP IV—CONVERGENS The species which I have placed together to form group IV, while quite diverse as to elytral pattern, agree in the structure of the male genitalia. The penis is broadly or narrowly triangular and always somewhat attenuate at apex. The tips of the ventral alae project slightly at the sides of the penis near the tip and the paramera are moderate in size and never longer than the penis. The sipho is slender, without a pronounced swelling before the dorsal flaps which are directed forward and upward and somewhat blunt. No species of the group has a transverse crest as in the preceding group but in certain individuals of H. quindecim-maculata Muls. there is a fine wrinkle which may be homologous with that crest. The female geni- talia are simple, without denticles on the inner wall of the bursa or other conspicuous modifications. In the present paper five species are assigned to group IV. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 HIPPODAMIA QUINDECIM-MACULATA Mulsant Fics. 20, 42, 162-166, 238 Hippodamia quindecim-maculata MULSANT, 1850, Species Coléopt. Trim. Sécuri- palpes, p. 20; 1866, Monogr. Coccinellides, pt. I, p. 12. Hippodamia 15-maculata CrotcH, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 366; Casey, 1908, Can. Ent., vol. 40, p. 398; Timberlake, 1919, Journ. New York Emits SOG VOle 27.5 pa hogs Hippodamia convergens subsp. 15-maculata LENG, 1903, Journ. New York Ent. Soc., vol. 11, p. 42; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, pe 27 Flippodamia quinquedecimmaculata KorsCHEFSKY, 10932, Junk-Schenkling, Coléopt. Catalogus, pars 120, Coccinellidae II, p. 343. 4; 8 Type locality —‘Les bords du Missouri, dans l’Amérique septen- trionale.” This species appears to be neither widespread in its distribution nor abundant in any part of its range. Twenty specimens are available for my study, eleven in the Casey collection and nine in the main National Museum collection. The spot pattern of this species is unusual in the genus in that there is a well-marked tendency toward the loss of spot 4, which is always reduced and frequently absent. Spots 2 and 3, and 4 and 5 are often united, 1 and 6 are always (?) present. The most characteristic feature of the male genitalia is the un- usually long and attenuated apex of the penis. Most of the males examined have no trace of transverse wrinkle where the transverse crest of the species of the glacialis group occurs, but in one from Batchawaung Bay, Lake Superior, this wrinkle is well defined. How- ever, this specimen was prepared with the aid of potassium hydroxide and there appears to have been a certain collapse of parts. The majority of the specimens examined were taken near St. Louis, Mo. Single specimens from various localities extend the range from Lake Superior south to about 40 miles south of St. Louis, and from Wisconsin and Illinois west to Denver, Colo. HIPPODAMIA MOESTA Leconte Fics. 22, 38, 177-186, 241 Hippodamia moesta Leconte, 1854, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 19; Crotch, 1874, Revision Coccinellidae, p. 97; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 78; Leng, 1903, loc. cit., vol. 11, p. 40; Timber- lake, 1910, loc. cit., vol. 27, p. 168; Korschefsky, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 342. /lippodamia lecontei var. moesta Crorcu, 1873, Trans. Amer. Ent. Soc., vol. 4, Pp. 367. NO. II HIPPODAMIA—CHAPIN 21 Hippodamia convergens subsp. “moesta JOHNSON, 1910, Carnegie Inst. Wash- ington, Publ. 122, p. 27. Hippodamia bowditchi JOHNSON, 1910, loc. cit., p. 45. Hippodamia politissima CAsEy, 1899, Journ. New York Ent. Soc., vol. 7, p. 80; 1908, Can. Ent., vol. 40, p. 395. Hippodamia ambigua race politissima LENG, 1903, Journ. New York Ent. Soc., vol. II, p: 41. Hippodamia convergens ab. politissima KorsCHEFSKY, 10932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 330. This species, which ranges from Colorado, north to Montana, west to British Columbia, and south along the Pacific coast to California, breaks up into three well-marked subspecies. A melanic subspecies (moesta) inhabits the West Coast region from Vancouver Island south to northern California. A maculate subspecies (bowditchi) cen- ters its distribution in Montana and ranges south to Colorado and west to British Columbia. A third and immaculate subspecies (politis- sima), infrequent in collections, is known from Santa Cruz and San Luis Obispo, Calif. FH. moesta Lec. is evidently rather closely related to H. quin- decim-maculata Muls., and sufficient material may show that the two should be merged as one. HIPPODAMIA M. MOESTA Leconte Type locality —Prairie Paso, Oreg. The most extreme case of melanism found in the*genus Hippodamia and comparable only to Coccinella prolongata bridwelli Nun. The average specimen is entirely black above except for the front angles of the pronotum and a pale spot on the elytral margin at about apical fifth. This last-mentioned spot is all that remains of the pale area separating spot 6 from the combined spots 4 and 5. Very few integrating specimens have been seen but one from Blair’s Ranch, Redwood Creek, Humboldt County, Calif., is of more than passing interest. Here the area which would be normally pale in the maculate form is dark red but not sufficiently dark to conceal the normal black markings of the species. This is one of several specimens taken at Blair’s Ranch, June 1903, by H. S. Barber. These were found among the individuals of a colony of Chrysomela scripta which infested the willows along Redwood Creek. The Hippodamia was apparently feeding on the young of the Chrysomelid. ® HIPPODAMIA M. BOWDITCHI Johnson Type locality—St. Maries, Idaho. This might be considered as the normal form of the species, that 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 is, the form comparable to the other species of the genus. Essentially its spot pattern is that of H. quinquesignata (Kby.). Spots 4, 1, and 3 are joined to form a heavy transverse basal bar, spot 2 almost always is absent, spots 4 and 5 always (?) joined, and spot 6 is present and discrete. Through the kindness of Maj. W. L. Jellison, of the Rocky Moun- tain Laboratory at Hamilton, Mont., the writer has available for study a good series of specimens of this subspecies. These were taken from willows along the Flathead River in the La Salle district, Mont. This is one of the most constant forms to be found in the entire genus, there being no cases of complete suppression of any spot except number 2. Specimens from Fort Collins and Denver, Colo., Heywood’s Cor- ner, Fernie, and Trinity Valley, British Columbia, and numerous localities in Montana have been seen. No specimen has yet been seen from Vancouver Island. HIPPODAMIA M. POLITISSIMA Casey Type locality—Santa Cruz, Calif. In the absence of evidence to the contrary, the writer is considering this form as a subspecies of moesta characterized by the complete sup- pression of elytral markings. Two specimens have been seen, both males, and dissections show them to belong to moesta. More material should be searched for along the coast of California, south of San Francisco. The true relationship of this form was first pointed out by Timberlake, during one of his visits to the National Museum. HIPPODAMIA CONVERGENS Guérin FIGs. 21, 47, 147-161, 239-240 Hippodamia convergens GUERIN, 1842, Icon. Regne Animal, vol. 7, (1829-1844), p. 321; Mulsant, 1850, Species Coléopt. Trim. Sécuripalpes, p. 22; 1866, Monogr. Coccinellides, pt. 1, p. 14; Leconte, 1852, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 130; Crotch, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 367; Gorham, 1891, Biol. Centr.-Amer., Coleopt., vol. 7, p. 153, pl. 8, figs. 22-24; Weise, 1895, Ann. Soc. Ent. Belgique, vol. 39, p. 125; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 80; Leng, 1903, loc. cit., vol. 11, p. 40, fig. 4; Casey, 1908, Can. Ent., vol. 40, p. 398; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, p. 21 et seq.; Timberlake, 1919, Journ. New York Ent. Soc., vol. 27, p. 168; Korschefsky, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 338; Timberlake, 1943, Hawaiian Planters’ Record, vol. 47, p. 11; Bull. Exp. Sta. Hawaiian Sugar Planters’ Assoc., Ent. Ser., Bull. 22, p. 11 (reprint of above, without change of pagination). Hippodamia juncta Casry, 1899, Journ. New York Ent. Soc., vol. 7, p. 80. NO. II HIPPODAMIA—-CHAPIN 23 Hippodamia modesta MELSHIMER, 1847, Proc. Acad. Nat. Sci. Philadelphia, vol. 3, p. 178; Leconte, 1852, loc. cit., vol. 6, p. 130. Hippodamia obsoleta Crorcu, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 367; Casey, 1908, Can. Ent., vol. 40, p. 398. Hippodamia praticola Mutsant, 1850, Species Coléopt. Trim. Sécuripalpes, p. 23; Weise, 1895, Ann. Soc. Ent. Belgique, vol. 30, p. 125. Type locality—Mexico and California. H. convergens Guér. is by far the commonest and most widespread species of the genus in the New World and is probably the most abundant species of the whole family in that same region. It is generally distributed over southern Canada, the whole United States, Mexico, and Central America as far south as Honduras. Also it has been introduced into Cuba. Notwithstanding its wide distribution, there is a surprising lack of variation in the spot pattern. Linkage of spots is rare and never extensive ; rather, the prevailing type of variation is in the opposite direction, toward the reduction of pigment. In the course of examin- ing about 6,000 specimens of this species, nearly every conceivable variation dealing with reduction or loss of one or more spots was en- countered. It would be footless to illustrate all these variations, just as it would be to name them, as is done by certain European writers. Such names merely clutter up the literature to no good end. Two hibernating lots of this species have been available for study. Both lots were composed only of individuals of H. convergens Guér., although at first glance the larger lot seemed to contain numerous specimens of H. quinquesignata Kby. Dissection proved these smaller individuals to be merely depauperate and otherwise abnormal convergens. A large lot was collected in December 1945 from a hibernating mass found near the Smithsonian Institution Solar Observing Station at Tyrone, N. Mex., and sent in through the courtesy of Dr. C. G. Abbot. After the fragmentary individuals were discarded, 2,856 specimens were preserved for study. All types of elytral maculation, from immaculate forms to those showing the full component of spots, were observed. The large number of depauperate individuals present was remarkably high; a condition perhaps related to the apparently high percentage of mermithid parasitism. A second and smaller lot of some 200 specimens was collected September 3, 1945, by Dr. L. G. Henbest on Capulin Volcano, 30 miles south of Raton, N. Mex. Very few of these were undersized, and only one individual was seen to harbor a mermithid. 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 HIPPODAMIA CASEYI Johnson ‘ Fics. 24, 39, 187-194, 243 Hippodamia caseyi JOHNSON, 1910, Carnegie Inst. Washington, Publ. 122, Pp. 21, 33; Casey, 1911, Mem. Coleopt., vol. 2, p. 250; Casey, 1924, loc. cit., vol. 11, p. 155. Hippodamia lecontei ab. caseyi KoRSCHEFSKY, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae IT, p. 342. Type locality —Fairfield, Wash.? There is some doubt as to the exact type locality. None is given definitely but in Johnson’s paper the name is directly connected only with specimens from Fairfield. Material identified as this species is on the basis of a series of II specimens labeled “Fairfield WS” which I believe are part of the lot mentioned by Casey (1911, p. 250) as received by him from Johnson. The distribution of H. caseyi appears to be from Colorado, north to Montana, thence west to Washington and Oregon, and south into California. The spot pattern is rather similar to that of H. convergens, and most individuals would pass for that species or for H. quinquesignata. A large percentage of individuals show a linking of spots 4, 1, and 3, or 4 and 3, and an even greater percentage than in convergens have spots 4 and 5 joined. The short and broad penis of this species offers a ready means of identifying male specimens. Unfortunately, no differences in the female genital organs have been noted for species of this group. Two hibernating lots of H. caseyi were collected and submitted for examination by Maj. W. L. Jellison. There was a mass of about goo specimens taken on August 5, 1944, on Blue Nose Peak, Lemhi County, Idaho. The other was a smaller lot of about 250 found on Deer Mountain, Ravalli County, Mont., August 27 of the same year. In these lots the specimens showing a linkage of spots } and 3 were nearly half of the total. Those showing linkage of spots 4 and 5 were comparatively few. HIPPODAMIA OREGONENSIS Crotch Fics. 23, 46, 170-176, 242 Hippodamia oregonensis, CrotcH, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 367; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 81; Leng, 1903, loc. cit., vol. 11, p. 42; Korschefsky, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 343. Hippodamia dispar Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 79. Hippodamia puncticollis Casry, 1899, loc. cit., vol. 7, p. 78. NO. II HIPPODAMIA—CHAPIN 25 Hippodamia quinquesignata ab. puncticollis KoRSCHEFSKY, 1932, Junk-Schenk- ling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 344. Hippodamia liliputana Casry, 1908, Can. Ent., vol. 40, p. 397. Hippodamia lilliputana CAsry, 1910, Can. Ent., vol. 42, p. 109 (emendation of liliputana). Hippodamia quinquesignata ab. liliputana KorscHEFSKyY, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 344. Hippodamia cockerelli JOHNSON, 1910, Carnegie Inst. Washington, Publ. 122, p. 49; Timberlake, 1919, Journ. New York Ent. Soc., vol. 27, p. 167. This species is rare in collections and all specimens studied by me are either from Colorado or from the Pacific Northwest. I have seen in all 31 specimens, 3 in the Casey collection, 3 in the Dobzhansky collection, and 25 in the main collection of the National Museum. Twelve are from Washington State or British Columbia and the rest from various localities in Colorado. In this species the white discal markings on the pronotum are usually absent, but in occasional specimens they are represented by minute white points. Since in the Colorado series every specimen, except the type of cockerelli Johnson, has the spot 6 reduced or absent and in the Washington-British Columbia series all specimens have this spot strongly developed, I have accepted these lots as repre- senting two subspecies. For the present I am accepting Hippodamua cockerelli Johnson as representing a third subspecies of H. oregonensis. Only the type specimen is available to me for study, and there have been reported but two other specimens. It is obvious that no conclusions drawn from such a small series can be final. HIPPODAMIA 0. OREGONENSIS Crotch Type locality —Oregon. There is very little variation in the series before me. Spots 4, 1, and 3 are always united to form a transverse basal band, spot 2 is always absent, spots 4 and 5 always joined, spot 6 usually free but occasionally joined to 4. The specimens in the collection are from Mount Yakima and the Olympic Mountains of Washington and from Mount Todd, British Columbia. HIPPODAMIA O. DISPAR Casey Type locality — Colorado. As in the preceding subspecies, spots 4, 1, and 3 are always joined, and spot 2 is always absent. Usually spots 4 and 5 are joined, but occasionally the single spot is so small as to suggest the complete 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 absence of spot 4. Spot 6 is present as a small black point on 5 of the 16 specimens. The available material is from Silverton (9 specimens), Gothic (2 specimens), Delta County (2 specimens), above Ouray (1 speci- men), and Leavenworth Valley (1 specimen). The remaining speci- men is from Colorado with no further data. HIPPODAMIA 0. COCKERELLI Johnson Type locality —Sangre de Cristo Range, Saguache County, Colo. This subspecies (if future collections indicate that it deserves sub- specific rank) is distinguished from the other subspecies by the com- plete linkage of all aS: number 2 being absent as usual in this species. The type is the only specimen of this form that is available to me for study. Johnson reports two other specimens; they are from Yel- lowstone Park, Wyo., and are now in the Bowditch collection. GROUP V—SINUATA Hippodamia sinuata Muls. is withdrawn from its association with the species of the convergens group to form a fifth group in the genus. Here the penis is very broad and flat, widest at about apical third where the sides are sharply angulate and the apex is submucronate. Each paramere is relatively large, surpassing the apex of the penis by from one-third to one-fourth of its length. The elytral spots are frequently linked to form longitudinal vittae. Nearly a thousand specimens have been examined in the course of this study. For the present I am considering this group to consist of a single species which breaks up into five subspecies and which ranges from Alaska into Mexico and east to Iowa. The present species is one of the most interesting of the genus from the standpoint of its variability and the problems of speciation pre- sented. More material from known ecological habitats will be neces- sary in order to complete the picture, but for the present certain facts may be noted. The shape of the penis seems to vary somewhat as in the species H. glacialis (F.). Generally, from north to south in its range, the slender apical portion of the penis becomes more elongate and this character is associated with a strengthening of the lateral angles of the penis. The northern type is found among specimens taken along the coast down into California as far as Martinez, while the southern form is projected into the north inland as far as Colorado. The differences are not clean-cut in the middle area, and many intergrading specimens are found. NO. II HIPPODAMIA—CHAPIN 27 In a series of 15 specimens of H. sinuata disjuncta Timb. collected at Loveland, Colo., June 26, 1935, by Th. Dobzhansky, there was a single male specimen with deformed genitalia (fig. 28) of a peculiar and interesting type. The acetabula for the attachment of the para- meres, normally lateral, are situated dorsally, and the parameres, normally capable of independent movement, are joined rigidly by a yoke across their bases so that they are required to move together and only in the vertical plane. Further, the penis carries a transverse crest reminiscent of that in H. quinquesignata (Kby.). HIPPODAMIA SINUATA Mulsant Fics. 25-26, 28, 41, 43, 195-226, 244-245 Hippodamia sinuata Mutsant, 1850, Species Coléopt. Trim. Sécuripalpes, p. 1011; Crotch, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 365; 1874, Revision Coccinellidae, p. 96; Casey, 1809, Journ. New York Ent. Soc., vol. 7, p. 81; Leng, 1903, loc. cit., vol. 11, p. 40, pl. 4; Casey, 1908, Can. Ent., vol. 4o, p. 398; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, p. 50, fig. 35; Timberlake, 1919, Journ. New York Ent. Soc., vol. 27, p. 167; Korschefsky, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 344. Hippodamia spuria Leconte, 1861, Proc. Acad. Nat. Sci. Philadelphia, vol. 13, p. 358; Mulsant, 1866, Monogr. Coccinellides, pt. I, p. 15; Crotch, 1873, Trans. Amer. Ent. Soc., vol. 4, p. 365; 1874, Revision Coccinellidae, p. 96; Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 80; Leng, 1903, loc. cit. vol. 11, p. 42, pl. 4; Casey, 1908, Can. Ent., vol. 40, p. 309; Johnson, 1910, Carnegie Inst. Washington, Publ. 122, p. 46, figs. 29-31; Timberlake, 1919, Journ. New York Ent. Soc., vol. 27, p. 168; Korschefsky, 1932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coccinellidae II, p. 345. Hippodamia trivittata Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 80; Korschefsky, 10932, Junk-Schenkling, Coleopt. Catalogus, pars 120, Coc- cinellidae II, p. 345. (= sinuata Muls.) Hippodamia crotchi Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 80; 1908, Can. Ent., vol. 40, p. 399. Hippodamia complex Casey, 1899, Journ. New York Ent. Soc., vol. 7, p. 80. (= spuria Lec.) Hippodamia sinuata disjuncta TIMBERLAKE, 1919, Journ. New York Ent. Soc., vol. 27, p. 169. Hippodamia americana fontinalis CASEY, 1924, Mem. Coleopt., vol. 11, p. 156. (= crotchi Csy.) HIPPODAMIA SIN. SINUATA Mulsant Type locality —California. The spot pattern is rather difficult of interpretation. Spot 4 is continued along the suture from the scutellum nearly to the apex and in some specimens appears to involv¢ spot 3. Other specimens show no expansion at basal fourth of the suture that can be considered as 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106° spot 3; therefore I consider spot 3 as unstable and often absent. Spot 2 seems always to be absent. Each elytron bears a broad longitudinal vitta composed of spots I, 5, 4, and 6. The range of this subspecies is from the San Francisco Bay region south into Mexico (Durango). HIPPODAMIA SIN. CROTCHI Casey Type locality—Lake County, Calif. At first glance hardly more than a variety of H. sin. sinuata, but as the form is restricted in its distribution it seems best to treat it as a subspecies. Typically, the spot pattern is as follows: Spots 3 and 3 joined to form a single sutural spot covering about basal fourth of suture ; spot 2 absent ; spots I, 5, and 4 joined to form an inverted Y with long stem and short arms; spot 6 isolated and always (?) present. This is the dominant form in New Mexico and Arizona. It appears in Lake County, Calif., a locality on the periphery of the distribution of H. sin. sinuata. Inland and to the north ‘it is frequently met with in Colorado, Utah, and Wyoming, there mixed with H. sin. disjuncta. HIPPODAMIA SIN. DISJUNCTA Timberlake Type locality—Utah (Salt Lake and Murray). In this and the following subspecies, there is a complete separation of spots 1 and 5. In the case of H. sin. disjuncta, spot 2 (as in all subspecies of H. sinuata) is missing, spot 3 often missing but oc- casionally spot 4 is extended along, the suture and widened at basal fourth of suture to suggest presence of this spot, spots 4, 5, and 6 usually isolated, 4 and 5 frequently joined, 4, 5, and 6 rarely joined. Pigmentation light as compared with H. sin. spuria. The dominant form in Utah, Wyoming, Idaho, and eastern Oregon and eastern Washington. Frequently seen in collections from Colorado and northern California. HIPPODAMIA SIN. SPURIA Leconte Type locality —Oregon. Through the kindness of Dr. P. J. Darlington, the type of H. spuria Leconte has been compared with Oregon specimens and its identity with the heavily pigmented form is established. This subspecies is the heavily pigmented counterpart of H. sin. disjuncta. The arrangement and. linkage of spots is the same in the two forms except that spot 3 is almost always evident as a posterior NO. II HIPPODAMIA—CHAPIN 29 expansion of spot 4 and very rarely there is a trace of linkage between spots 4 and 3 and spot 5. Linkages involving spots 4, 5, and 6 are more frequent here. This is the West Coast form and is apparently confined to the area west of the coast range from British Columbia south to Port Angeles, Wash. HIPPODAMIA SIN. STRAMINEA, new subspecies Type locality—Klamath River, Calif. Type (male) and paratypes (both sexes), U.S.N.M. No. 57893. Paratypes in collections of Th. Dobzhansky, F. T. Scott, G. H. Dieke, J. E. Blum, and G. P. Mackenzie. The Dobzhansky collection contains a series of 25 specimens from the Klamath River area (Klamath River, Klamath Glen, and For- tuna), of which 20 are without trace of spots on the elytra except for the very feeble development of spot 4. The remaining five are very lightly to lightly pigmented. The lot is suggestive of the col- lection of H. lunatomaculata Mots. made by the same author at Willows, Calif. In the absence of adequate evidence to the contrary I am treating this lot as an interesting but highly restricted subspecies. GROUP VI—KOEBELEI This group contains the single aberrant species H. koebelei Tim- berlake, recently described from Mexico. It is similar in body form and general appearance to H. convergens Guérin of Group IV. It is however distinct from all other known species of the genus in that the first segments of the front and middle tarsi of the male are broadly dilated. The third antennal segment is simple and not triangularly expanded as in other genera of the Hippodamiini which have dilated tarsal segments. Further, the male genitalia are quite distinct, having the following peculiarities: Each paramere has the dorsal internal edge margined by a membranous lobe; a lesser but similar lobe is along the ventral internal edge. The ventral alae are connected apically with the penis by a loose membranous area. The tip of the sipho bears two lateral processes which are cylindrical and terminate in bladderlike structures and are situated on either side of the median line above and between the flaps which are lateral. HIPPODAMIA KOEBELEI Timberlake Fics. 27, 44, 167-169, 246 Hippodamia koebelei TIMBERLAKE, 1942, Proc. Ent. Soc. Washington, vol. 44, Pp. 39; 1943, Hawaiian Planter’s Record, vol. 47, p. 11; Bull. Exp. Sta. 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Hawaiian Sugar Planters Association, Ent. Ser., Bull. 22, p. 11 (reprint of above, without change of pagination). Type locality —Mexico City, Mexico. Except for the differences in tarsal structure and genitalia of male noted above, individuals of this species can hardly be distingushed from immaculate specimens of H. convergens Guérin. Usually the pronotal markings in HH. koebelei are more finely drawn and even throughout. The emargination of the fifth visible sternite of the male is wider and deeper in H. koebelei than in H. convergens, a condition coordinated with the relatively greater size of the genitalia in the former. In addition to a portion of the type material, more than 50 speci- mens of this species have been examined by the writer, most of which were intercepted at quarantine along the Mexican-United States border. Exact localities for intercepted material cannot be established, but a statement in a letter of January 31, 1940, from J. B. R. Leary, Supervisor, District 2, Mexican Border, to O. D. Deputy, Super- visor, Mexican Border Service, Bureau of Entomology and Plant Quarantine, casts some light on the subject. I quote: “Inquiry in the Nuevo Laredo market, however, reveals that lettuce comes from San Luis Potosi, S. L. P. and Ramos Arispe, Coah., during the winter. In the summer it is shipped from Celaya, Gto.” Specimens from definite localities, other than those in the type series, have been seen from Amecameca, March 9g, 1938, and Toluca, March 1, 1938, both State of Mexico, collected by Th. Dobzhansky. A specimen recorded in the original description as from Mexico City appears to have been taken at Puebla, State of Puebla, May 27, 1922, by E. G. Smyth. EXPLANATION OF PLATES PLATE I Anatomy or Hippodamia tredecin-punctata (L.) (SPANDAU, GERMANY). Tyre Species oF Hippodamia, AS DESIGNATED BY CROTCH, 1873 Fic. 1. Dorsal view, showing conventional numeration of the elytral spots. 2. Ventral view of abdomen of male. 3. Ventral view of abdomen of female. 4. Aedeagus, dorsal view. BP, basal piece; P, penis; PA, paramere; VA, ventral ala. 5. Aedeagus, lateral view (front paramere dissected away). TR, trabes. 6. Aedeagus, ventral view. 7. Sipho. SC, siphonal capsule; SF, dorsal flaps. 8. Sclerotized portion of female genital system. BC, bursa copulatrix; IN, infundibulum accessory piece; SD, sperm duct; RS, receptaculum seminis; AG, accessory gland. NO. Fic. Fic. Fic. Fic. Fic. II 14. 15. 25. 26 HIPPODAMIA—CHAPIN 31 PENIS AND PARAMERES PLATE 2 . H. tr. tibialis (Say). Jerome, Idaho, August 26, 1930, D. F. Fox. . H. americana Crotch. Whitefish Point, Lake Superior, Mich., Hub- bard and Schwarz collection. . H. falcigera Crotch. Hudson Bay, Hubbard and Schwarz collection. 12. H. washingtoni Timberlake. Hoquiam, Wash., May 1, 1903, H. E. Burke. PLATE 3 . H. parenthesis (Say). Adams County, Iowa, November 1936. H. apicalis Casey. Independence, Calif., July 11, H. F. Wickham. H. expurgata Casey. Salt Lake, Utah, June 13, Hubbard and Schwarz collection. . H. lunatomaculata Motschulsky. Port Angeles, Wash., August 12, 1934, Th. Dobzhansky. (Type of H. /. dobzhanskyi n. subsp.). PLATE 4 . H. glacialis (Fabricius). Watch Hill, R. I., July 22, 1900, W. Robinson. . H. glacialis (Fabricius). Variation in penis. A, B, C, Watch Hill, R. I. D, E, F, Rocky Ford, Colo. . H. quinquesignata (Kirby). Duluth, Minn. . H. quindecim-maculata Mulsant. St. Louis, Mo., September 13, 1891, H. Soltau. PLATE 5 . H. convergens Guérin. Barnesville, Ga., May 29, 1932, T. H. Bissell. . H. moesta Leconte. Nanaimo, Vancouver Island, May 13, 1897, Taylor. . H. oregonensis Crotch. Olympic Mts., Wash. August 1932, Th. Dobzhansky. . H. caseyi Johnson. Moscow, Idaho, July 10, R. C. Shannon. PLATE 6 H. sinuata Mulsant. Martinez, Calif., December 12, 1882, Turner. . H. sinuata Mulsant. Variation in penis. A, Martinez, Calif., December 12, 1882, Turner. B, Maxwell, N. Mex., May 4, 1916, D. J. Caffrey. C, Puyallup, Wash., June 17, 1916, T. H. Scheffer. D, Greeley, Colo., July 28, 1930. E, Flagstaff, Ariz., July 16, B. P. Clark. F, Durango, Dgo., Mexico, November 1927, F. C. Bishopp. 27. H. koebelei Timberlake. Ex Mexico at Laredo, Tex., December 16, 28 1939, U. S. D. A. Interception Laredo No. 18648. _ H. sinuata Mulsant. Abnormal specimen from Loveland, Colo., June 26, 1935, Th. Dobzhansky. Fic. Fic. Fic. 20. Le 52. 55- SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 SIPHO PLATE 7 H. tr. tibialis (Say). Half Moon Bay, Calif., June 24, 1932, Th. Dobzhansky. H. americana Crotch. Whitefish Point, Lake Superior, Mich., Hub- bard and Schwarz collection. H. falcigera Crotch. Yellowstone, Wyo., July 1935, J. E. Blum. H. expurgata Casey. Salt Lake, Utah, June 13, Hubbard and Schwarz collection. : H. washingtoni Timberlake. Hoquiam, Wash., May 27, 1917, H. G. Dyar. . H. parenthesis (Say). West Point, N. Y., May 17, 1909, W. Robinson. . H. apicalis Casey. Paonia, Colo., June 14, 1904, E. S. G. Titus. . H, lunatomaculata Motschulsky. Corvallis, Oreg., July 22, 1931. . H, quinquesignata (Kirby). Milner, Idaho, July 20, 1929, L. B. Jones. PLATE 8 . H. moesta Leconte. Nanaimo, Vancouver Island, May 13, 1897, Taylor. . H. caseyi Johnson. Little Baldy Mts., Utah, 8,780 feet, May 13, 1939, Knowlton and Nye. . H. g. lecontei Mulsant. Victor, Idaho, August 7, 1939, Th. Dobzhansky. . H. sinuata Mulsant. Martinez, Calif., December 12, 1882, Turner. . H. quindecim-maculata Mulsant. Denver, Colo., August 16, 1808, BayaOslar. 43. H. sinuata Mulsant. Durango, Dgo., Mexico, November 1927, F. C. Bishopp. . H. koebelei Timberlake. Ex Mexico at Laredo, Tex., January 27, 1940, U. S. D. A. Interception Laredo No. 19379. . H. g. glacialis (Fabricius). Agawam, Mass., June 27, 1916, D. A. Ricker. 40..H. oregonensis Crotch. Mount Yakima, Wash., 6,000 feet, September 16, 1922, E. J. Newcomer. . H, convergens Guérin. Greeley, Colo., August 4, 1930. Spot PATTERNS PLATE 9 . H. tr. tibialis (Say). Nulato, Alaska, June 14, 1916, Harrington. . H. tr. tibialis (Say). Whitefish Point, Lake Superior, Mich., Hub- bard and Schwarz collection. H. tr. tibialis (Say). Oregon, Koebele. H. tr. tibialis (Say). Truro, Nova Scotia, October 1, 1919. H. tr. tibialis (Say). St. John, New Brunswick, June 23, 1901, W. McIntosh. H, tr. tibialis (Say). Aztec, N. Mex., May 5, C. F. Baker. H. tr. tibialis (Say). Muscatine, Iowa, May 109, 1917, C. E. Smith. H. tr. tibialis (Say). Burley, Idaho, July 24, 1931. H. americana Crotgh. Waskesu, Saskatchewan, June 10, 1938, C. Shaw. NOs If Fic. HIPPODA MIA—CHAPIN 33 57. H. americana Crotch. Whitefish Point, Lake Superior, Mich., Hub- bard and Schwarz collection. 58. H. falcigera Crotch. 59. H. falcigera Crotch. Hudson Bay, Hubbard and Schwarz collection. Edmonton, Alberta, June 15, 1920, F. S. Carr. (Type of H. albertana Casey.) 60. H. falcigera Crotch. 61. H. falcigera Crotch. Edmonton, Alberta, aihie 15, 1920. Yellowstone Park, Wyo., July 1935, J. E. Blum. (Topotype of Ceratomegilla cottlei Nunenmacher.) 62. H. falcigera Crotch. Moscow, Idaho, October 2, 1927, H. A. Walker. 63. H. washingtoni Timberlake. Blue Mts., Oreg., June 7, 1938, H. P. Lanchester. 64. H. washingtoni Timberlake. Longmire Springs, Mount Ranier, Wash., 2,700 feet, July 19, 1935, H. P. Lanchester. ( Paratype.) 65. H. washingtoni Timberlake. Hoquiam, Wash., May 1, 1903, H. E. Burke (Hopk. U. S. 1844a). 66. H. washingtont Timberlake. Hoquiam, Wash., May 1, 1903, H. E. Burke (Hopk. U. S. 1844a). 67. H. washingtoni Timberlake. Hoquiam, Wash., May 1, 1903, H. E. Burke (Hopk. U. S. 1844a). PLATE I0 68. H. parenthesis (Say). Rockaway Beach, L. I., N. Y., June 20, 1904, Brooklyn Museum collection. 69. H. parenthesis (Say). West Point, N. Y., June 14, 1909, W. Robinson. 70. H. parenthesis (Say). Glen Echo, Md., July 12, 1930, J. C. Bridwell. 71. H. parenthesis (Say). Antelope Valley, Calif.. May 1925, R. E. Campbell. 72. H. parenthesis (Say). Proberta, Calif., June 23, 1932, Th. Dobzhansky. 73. H. parenthesis (Say). Spearfish, S. Dak., August 1, 1938, Th. Dobz- hansky. 74. H. a. apicalis Casey. Schwarz collection. 75. H. a. apicalis Casey. 76. H. a. apicalis Casey. 77. H. a. apicalis Casey. Dobzhansky. 78. H. a. apicalis Casey. 79. H..a. apicalis Casey. 80. H. a. apicalis Casey. 81. H. a. apicalis Casey. hansky. 82. H. a. apicalis Casey. Dobzhansky. 83. H. a. lengit Johnson. 84. H. a. lengi Johnson. Morrison. 85. H. a. lengi Johnson. Morrison. Lake Tahoe, Calif., July 12, Hubbard and Reno, Ney., Casey collection. (Type.) Truckee, Calif., Casey collection. (Paratype.) Klamath Agency, Oreg., June 20, 1932, Th. Madras, Oreg., June 19, 1932, Th. Dobzhansky. Maupin, Oreg., June 19, 1932, Th. Dobzhansky. Madras, Oreg., June 19, 1932, Th. Dobzhansky. Terrebonne, Oreg., June 19, 1932, Th. Dobz- Klamath Agency, Oreg., June 20, 1932, Th. San Diego County, Calif., Coquillett. (Paratype.) Near Jacumba, San Diego County, Calif., H. Near Jacumba, San Diego County, Calif., H. 34 Fic. 86. 87. 88. I0I. 102, 103. 104 105 106 G. 107. 108 109 TIO ene . SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 H. a, lengi Johnson. Near Jacumba, San Diego County, Calif., H. Morrison. H. lunatomaculata Motschulsky. Original figure from Bull. Acad. Nat. Sci., Moscow, vol. 18, pl. 7, fig. 8, 1845. (Enlarged.) PLATE II ? H. expurgata Casey. Bright Angel, Grand Canyon, Ariz., July to, Barber and Schwarz collection. . H. expurgata Casey. Prescott, Ariz., July 20, Barber and Schwarz collection. . H. expurgata Casey. Williams, Ariz., May 30, Barber and Schwarz collection. . H, expurgata Casey. Wasatch, Utah, June 27, Hubbard and Schwarz collection. . H. expurgata Casey. Bright Angel, Grand Canyon, Ariz., November 7, Barber and Schwarz collection, . LH. expurgata Casey. Fort Robinson, Nebr., July 30, 1935, Th. Dobz- hansky. 4. H. expurgata Casey. Boulder, Colo., Casey collection. (Paratype.) . H. expurgata Casey. Boulder, Colo., Casey collection. (Type.) . H. expurgata Casey. Ucross, Wyo., August 2, 1935, Th. Dobzhansky. . H. |. dobzhanskyi, new subspecies. Willamette Valley, Oreg., Feb- ruary 5, 1931, A. O. Larson. . H. 1. dobzhanskyi, new subspecies. Oregon, Koebele. . H. |. dobzhanskyi, new subspecies. Oregon, Koebele. T00. H, |. dobzhanskyi, new subspecies. Port Angeles, Wash., August 12, 1934, Th. Dobzhansky. (Type.) H. 1. dobzhanskyi, new subspecies. Port Angeles, Wash., August 12, 1934, Th. Dobzhansky. (Paratype. ) H. 1. lunatomaculata Motschulsky. Half Moon Bay, Calif., Scott. H. 1. lunatomaculata Motschulsky. Willows, Calif., June 23, 1932. Th. Dobzhansky. H. |. lunatomaculata. Motschulsky. Willows, Calif., June 23, 10932, Th. Dobzhansky. H. |. lunatomaculata Motschulsky. Willows, Calif., June 23, 1932, Th. Dobzhansky. H. |. Ilunatomaculata Motschulsky. Willows, Calif., June 23, 1932, Th. Dobzhansky. PLATE 12 H. g. glacialis (Fabricius). Springfield, Mass., Geo. Dimmock. H. g. glacialis (Fabricius). Orange, N. J., H. Soltau. H. g. glacialis (Fabricius). Riverhead, L. I., N. Y., July 15, 1912, Russell. H. g. glacialis (Fabricius). “Texas,” C. V. Riley collection. H. g. glacialis (Fabricius). Vienna, Va., August 25, 1o11, C. W. Hooker. NO. Fic. II I12 113 114. 115. 116. 117. 118. II9. 120. 121. 122. 123. 124. 125. 126. 135 136 137 138 HIPPODAMIA—CHAPIN 35 . H. g. glacialis (Fabricius). Ipswich, Mass., August 1922, D. H. Blake. . H. g. glacialis (Fabricius). Boulder, Colo., Casey collection. (Type of abducens Casey.) H. g. glacialis (Fabricius). “Montana,” C. V. Riley collection. H. g. extensa Mulsant. Alameda County, Calif., J. E. Blum. H. g. extensa Mulsant. Alameda County, Calif., J. E. Blum. H. g. lecontei Mulsant. Albuquerque, N. Mex., W. Robinson. H. g. lecontei Mulsant. Seligman, Ariz., H. F. Wickham. H. g. lecontei Mulsant. Seligman, Ariz., H. F. Wickham. H. g. lecontei Mulsant. Albuquerque, N. Mex., W. Robinson. H. g. lecontei Mulsant. Seligman, Ariz., H. F. Wickham. H. g. mackensziei, new subspecies. Glacier Lodge, Inyo County, Calif., August 2, 1942, Mackenzie. (Paratype. ) H. g. mackenziei, new subspecies. Glacier Lodge, Inyo County, Calif., August 3, 1942, Mackenzie. (Type.) H. g. mackensziei, new subspecies. Glacier Lodge, Inyo County, Calif., August 4, 1942, Mackenzie. (Paratype.) H. g. mackensziei, new subspecies. Glacier Lodge, Inyo County, Calif., August 3, 1942, Mackenzie. (Paratype. ) H. g. mackenziei, new subspecies. Glacier Lodge, Inyo County, Calif., August 3, 1942, Mackenzie. (Paratype.) PLATE 13 . H. q. quinquesignata (Kirby). Bear Lake, British Columbia, July 21, 1903, R. P. Currie. . H. q. quinquesignata (Kirby). Whitefish Point, Lake Superior, Mich., Hubbard and Schwarz collection. . H. q. quinquesignata (Kirby). Locality unknown, Levette collection. (Type of H. subsimilis Casey.) . LH. q. quinquesignata (Kirby). Green River, Wyo., Casey collection. (Type of H. vernix Casey.) . H. q. quinquesignata (Kirby). Boulder, Colo., Casey collection. (Type of H. coccinea Casey.) . H. q. quinquesignata (Kirby). Sevier Lake, Utah, July, Wickham. (Type of H. uteana Casey.) . H. q. quinquesignata (Kirby). Edmonton, Alberta, June 7, 1910, F. S. Carr, Casey collection. (Type of H. quadraria Casey.) . H. q. quinquesignata (Kirby). Fairfield, Wash., Casey collection. (Type of H. c. pugetana Casey.) . H. q. quinquesignata (Kirby). Kaslo, British Columbia, July 2, 1903, Rage. (Currie: . H. q. quinquesignata (Kirby). Santa Rosa, Calif.,. Casey. collection. (Type of H. obliqua Casey.) . H. q. ambigua Leconte. Willamette, Oreg., February 5, 1931, A. O. Larson. . H. q. ambigua Leconte: Willamette, Oreg., February 5, 1931, A. O. Larson, 36 Fic. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 139. H. q. ambigua Leconte. Willamette, Oreg., February 5, 1931, A. O. Larson. 140. H. q. ambigua Leconte. Willamette, Oreg., February 5, 1931, A. O. Larson. H. q. ambigua Leconte. Willamette, Oreg., February 5, 1931, A. O. Larson. . H. q. punctulata Leconte. Modesto, Calif., fall of 1928, A. O. Larson. . H. q. punctulata Leconte. Modesto, Calif., fall of 1928, A. O. Larson. . H. q. punctulata Leconte. Modesto, Calif., fall of 1928, A. O. Larson. . H. q. punctulata Leconte. Modesto, Calif., fall of 1928, A. O. Larson. . H. q. punctulata Leconte. Modesto, Calif., fall of 1928, A. O. Larson. PLATE 14 . H. convergens Guérin. Totonicapan, Guatemala, February 11, 1938, Th. Dobzhansky. . H. convergens Guérin. Half Moon Bay, Calif., June 24, 1932, Th. Dobzhansky. . H. convergens Guérin. Half Moon Bay, Calif., June 24, 1932, Th. Dobzhansky. . H. convergens Guérin. Yosemite Valley, Calif., 4,000 feet, June 11, 1932, Th. Dobzhansky. . H. convergens Guérin. California, Casey collection. (Type of H. juncta Casey.) . H. convergens Guérin. Yosemite Valley, Calif., 4,000 feet, June 11, 1932, Th. Dobzhansky. . H. convergens Guérin. Yosemite Valley, Calif., 4,000 feet, June 11, 1932, Th. Dobzhansky. . H. convergens Guérin. Yosemite Valley, Calif., 4,000 feet, June 11, 1932, Th. Dobzhansky. . H. convergens Guérin. San Luis Obispo, Calif., May 15, 1932, Th. Dobzhansky. . H. convergens Guérin. Puebla, Mexico, May 209, 1932, E. G. Smythe. 57. H. convergens Guérin. Yosemite Valley, Calif., 4,000 feet, June 11, 1932, Th. Dobzhansky. . H. convergens Guérin. Yosemite Valley, Calif., 4,000 feet, June 11, 1932, Th. Dobzhansky. 59. H. convergens Guérin. Yosemite Valley, Calif., 4,000 feet, June 11, 1932, Th. Dobzhansky. . H. convergens Guérin. San Juan Mts., Colo., July 20, 1935, Th. Dobzhansky. . H. convergens Guérin. Yosemite Valley, Calif., 4,000 feet, June 11, 1932, Th. Dobzhansky. H. quindecim-maculata Mulsant. St. Louis, Mo., September 13, 1891, H. Soltau collection. H.. quindecim-maculata Mulsant. Sioux City, Iowa, September 20, 1920, C. N. Ainslie. H. quindecim-maculata Mulsant. Sioux City, Iowa, October 12, 1922, C. N. Ainslie. NO. Fic. Fic. II 165 166 167 168 169 HIPPODAMIA—CHAPIN 37 . H. quindectm-maculata Mulsant. Chelsea, Iowa, July 19, M. P. Somes. . H. quindecim-maculata Mulsant. Batchawaung Bay, Ontario, August, Hubbard and Schwarz collection. PLATE 15 . H. koecbelei Timberlake. Oaxaca, Mexico, September 18, L. O. Howard. (Paratype. ) . H. koebelei Timberlake. Puebla, Mexico, May 27, 1922, E. G. Smythe. (Paratype. ) . H. koebelei Timberlake. Mexico City, Mexico, May 22, 1922, E. G. Smythe. (Paratype.) . H. o. cockerelli Johnson. Cottonwood Gulch, Saguache County, Colo., August 4, 1887, T. D. A. Cockerell. (Type.) . H. 0. oregonensis Crotch. Mount Todd, British Columbia, Hubbard and Schwarz collection. . H. 0. oregonensis Crotch. Canadian Rocky Mts., Casey collection. (Type of H. puncticollis Casey.) . H. o. dispar Casey. Colorado, Casey collection. (Type of lilliputana Casey. ) . H. o. dispar Casey. Silverton, Colo., July 12, 1903. . H. o. dispar Casey. Colorado, Casey collection. (Type of H. dispar Casey.) . H. o. dispar Casey. Silverton, Colo., July 5, 1903. m. moesta Leconte. Seattle, Wash., S. Bethel. m. moesta Leconte. Seattle, Wash., C. V. Riley collection. . moesta Leconte. Nanaimo, Vancouver Island, May 13, Taylor. . m. moesta Leconte. Blair’s Ranch, Humboldt County, Calif., June 12, 1903, H. S. Barber. . H. m. moesta Leconte. Blair’s Ranch, Humboldt County, Calif., June 9, 1903, H. S. Barber. . H. m. bowditchi Johnson. Fort Collins, Colo., June 2, 1904, E. S. G. Titus. . H. m. bowditcht Johnson. Kalispell, Mont. June 16-20, H. F. Wickham. . H. m. bowditchi Johnson. Missoula, Mont., August 2, 1904. . H. m. bowditchi Johnson. Denver, Colo., July 7, Hubbard and Schwarz collection. . H. m., politissima Casey. Santa Cruz, Calif., Casey collection. (Type.) mo = PLATE 16 . H. caseyi Johnson. Little Baldy Mts., Utah, 8,780 feet, May 13, 19309, Knowlton and Nye. . H. caseyt Johnson. Little Baldy Mts., Utah, 8,780 feet, May 13, 1939, Knowlton and Nye. . H. caseyi Johnson. Little Baldy Mts., Utah, 8,780 feet, May 13, 19309, Knowlton and Nye. . H. caseyi Johnson. Little Baldy Mts., Utah, 8,780 feet, May 13, 1939, Knowlton and Nye. 38 Ere IOI. 192. 193. 194. 195. 196. 197. 108. 199. 200. 201. 202. 203. 204. 205. 206. 207. 208. 209. 210. 211. li ak 213. 214. 215. 216. 2178 218. 210. 220. 221. 222, 223. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 H. caseyi Johnson. Mica Peak, Idaho, J. C. Merrill. . caseyt Johnson. Little Baldy Mts., Utah, May 13, 19309. . caseyt Johnson. Fairfield, Wash., Casey collection. A H. caseyt Johnson. Fairfield, Wash., Casey collection. H H . sin. sinuata Mulsant. H. sin. sinuata Mulsant. F. C. Bishopp. H. sin. sinuata Mulsant. Blaisdell. H. sin. sinuata Mulsant. trivittata Casey.) H. sin. sinwata Mulsant. E. C. Van Dyke. H. sin. sinuata Mulsant. H. sin. sinuata Mulsant. seum collection. H. sin. sinuata Mulsant. Durango, Dgo., Mexico, H. F. Wickham. Durango, Dgo., Mexico, November 1927, Vine Hill, Calif., September 25, 1909, F. E. Near Napa Junction, Calif. (Type of H. Alameda County, Calif., October 19, 1902, “California.” Martinez, Calif., December, Brooklyn Mu- Gilroy, Calif., August 8, 1908, L. J. Condit. H. sin. disjuncta Timberlake. Murray, Utah, May 20, 1914. (Type.) H, sin, disjuncta Timberlake. Richfield, Utah, August 16, 1927, E. W. Davis. H. sin. disjuncta Timberlake. Murray, Utah, August 22, 1913. (Para- type. ) H, sin. disjuncta Timberlake. Salt Lake, Utah, July 30, 1915. (Para- © type.) H. sin. spuria Leconte. of H. complex Casey.) PLATE 17 British Columbia, Casey collection. (Type H. sin. spuria Leconte. Port Angeles, Wash., August 12, 1934, Th. Dobzhansky. H, sin. spuria Leconte. Nanaimo, Vancouver Island, Taylor. H. sin. spuria Leconte. Scio, Oreg., June 2, 1931. H. sin. spuria Leconte. Corvallis, Oreg., August 14, 1931. H. sin. spuria Leconte. Buckhorn. H. sin. spuria Leconte. Buckhorn. Amity, Oreg., February 24, 1930, W. J. Amity, Oreg., February 24, 1930, W. J. HH. sin. spuria Leconte. Corvallis, Oreg., July 20, 1931. H. sin. spuria Leconte. Dever, Oreg., July 18, 1931. H. sin. crotchi Casey. Lake County, Calif., Casey collection. (Type.) H. sin. crotchi Casey. Jemez Springs, N. Mex., John Woodgate, Casey collection. (Type of H. fontinalis Casey.) sin. crotchi Casey. Cheyenne, Wyo., April 22, H. Soltau. sin. crotchi. Casey. Denver, Colo. sin. crotchi Casey. Sante Fe, N. Mex., June, T. D. A. Cockerell. sin. crotchi Casey. New Mexico, Hubbard and Schwarz collection. sin. crotchi Casey. Cheyenne, Wyo., July 29, 1939, Th. Dobzhansky. sin. straminea, new subspecies. Klamath River, Calif., August 5, 1939, Th. Dobzhansky. (Paratype.) NO. Fic. Fic. Fic. Fic. Tic. 227. 228. 220. 230. 231. 232. 233. 234. 2350 236. 237. 238. 239. 240. 241. 242. 243. 244. 245. 246. i. Biba, aye A eomsmsemes Teas by by Sy by By Se my my by HIPPODAMIA—CHAPIN sin, straminea, new subspecies. Klamath River, Calif., August 5, 1939, Th. Dobzhansky. ( Paratype.) sin, straminea, new subspecies. Klamath River, Calif., Th. Dobz- hansky. (Type.) sin. straminea, new subspecies. Klamath River, Calif., Th. Dobz- hansky. (Paratype.) GEOGRAPHICAL DISTRIBUTION PLaTE 18 . tr. tibialis (Say). . americana Crotch. . falcigera Crotch. washingtoni Timberlake. PLATE 19 . parenthesis (Say). . apicalis Casey. . expurgata Casey. . lunatomaculata Motschulsky. PLATE 20 . glacialis (Fabricius). . glacialis (Fabricius). . quinquesignata (Kirby). quindecim-maculata Mulsant. PLATE 21 . convergens Guérin. (United States.) . convergens Guérin. (Mexico and Central America.) . moesta Leconte. . oregonensis Crotch. PLATE 22 . caseyit Johnson. sinuata Mulsant. . sinuata Mulsant. . koebelet Timberlake. ot) ‘ 1) ier ke) tut odo ii i esitaat | e | ie are steel Wid Watinaden rr ar ; i : “ ah iy ue beg ee 2 a AE B46, ; Lica " ‘Salt, ri i Me a ay iia yee AYE) ‘ajinid. 4 ey i wie | cy pearcengetn | Mo Haken 3, Asean’ \ Tye “chnvey’: ESREEN * by | Ah f rit yee lot he Motor aneyiateh, “cua aaa ; ee alte At i eee nae aetna t wotynidaayy AL og i ; : P I (ye i a (% rar rey OBE ray , qh ON i Coy PNET? ir arad us \ ae SORE i ef ’ ae i yn k +] i eis ‘ on ly, Re ys Cia) reser tiihallt li, bee pla tt sar Gee) sioymgas 7D Cvehorhoer tit: CA certs LN ae) y eure lL) abaghy f Mite miidel) Woon, ay: 7 be bit) DLO Be ty ; A ees Taemider IY, rohnlyeraterinteyh ne my ant eR sl Se st gy ( ratte boris PY yy tS) ah ae Cit wrote. ei bie Ooty Mi ALON) TA eS (OG aaa merit sa De in Rei nook SST i # i ig eo wees | a Hoa] eat 1aHOgS ; 4) At, 7 4 if, Levant ka) t ¥ ' i ee eh 3% Ligh sigmuictol, 9 ri wea et Des, ; ¥ baht MPA hakyh Raat A eng scien arty kin scl ii seh saath | Sptne y 38 Vit bi bd wnt , ha j j ‘ dak au ia ¥ Ve. a M - f Ss ( A hs fed ES lta we he cu Te a a 4 ae (ceevaioe, Wis Vey a Sl ia ah tS re Ly Mi t Deven Cele sas ean ged. Cisiapal Suara ei, & MMien,, haa a ath a} Pad ee ae ae ie eee Morn i Chitral mint “heat ‘ # ft AM, pneebye ths Saayy A Chey: af if Vs hie -dualy a, FOLIA, = . BE sangre Ne we wa? i peut, Ape amie Eg Thien (Parsing) ora) a SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 1 HIPPODAMIA (For explanation, see p. 30.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 2 TIBIALIS 9 AMERICANA /0 Li FALCIGERA // WASHING TON/ 12 HIPPODAMIA (For explanation, see p. 31.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 3 it PARENTHES/S / 3 APICALIS / 4 il I N\ \ | ‘| y I" ] | 1 \ vin Hi nl \ ih EXPURGATA / to] LUNATOMACULATA / 6 HIPPODAMIA (For explanation, see p. 31.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 4 OM A 44 GLACIALIS 17 GLACIALIS QUINQUE SIGNATA /9 QUINDECIMMACULATA 20 HIPPODAMIA j (For explanation, see p. 31.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 5 ai y \ CONVERGENS Zi MOESTA 22 OREGONENSIS 23 CASEY 24 HIPPODAMIA (For explanation, sce p. 31.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 6 SINUATA 25 SINUATA 26 KOEBELE! eT ABNORMALITY 2a HIPPODAMIA (For explanation, see p. 31.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 7 )\ J/ 32 33 34 NN 3S 36 HIPPODAMIA (For explanation, see p. 32.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 8 . 39 38 40 42 43 4/ ‘ 46 45 47 HIPPODAMIA (For explanation, see p. 32.) eSB wes VOL. 106, NO. 11, PL. 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS ; HIPPODAMIA PP. 33-34.) SMITHSONIAN MISCELLANEOUS COLLECTIONS SMITHSONIAN MISCELLANEOUS COLLBETIONS VOL. 106, NO. 11, PL. 12 HiPPODAMIA (For explanation, see pp. 34-35.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 13 142 HiPPODAMIA (For explanation, see pp. 35-36.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 15 x; 3 i /80 Sk BE @& 182 = 183 ) 184 185 S786 HiIPPODAMIA (For explanation, see p. 37.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 16 204 206 HIPPODAMIA 203 (For explanation, see pp. 37-38.) > SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 11, PL. 17 HiIPPODAMIA (For explanation, see pp. 38-30.) (‘6 -d aas ‘uoneueldxa 10,7) VINVOOddIH 40 NOILNEINLSIG IwoIHdvas0aD INOLONIHS VM VeIDNITVS 5 4, ---— = el — em } — ar arc se ' ba ee SITVIGIL °AL } ae aL 81 “Id ‘TI ‘ON ‘90T “10A SNOIL931109 SNOANVTISOSIN NVINOSHLINWS 6T “Id ‘TT ‘ON ‘90T “10A \ (‘6€ -d vas ‘uoryeurjdxe 10,7) VINVOOdd!IH 40 NOILNEIYLSIG IwOIHdvuYDOAD + - JANSNVHZE0OO 7 += VLY INIVWOLPNIT 7 +-/9NI7T V SITVIISY V ba — —— wench 2) VLYIUNAXF i 2 --=--3e7L ! 1 \ ' b een | a ae ania ¥ SNOIL937109 SNOSNV1I139SIW NVINOSHLIWS (6€ ‘d vas ‘uorjzeue{dxa 10,7) VINVGOdd!IH 40 NOILNSIYLSIG IvOIHdveDSo3adD + - VLVYINLINAd 0 ® - ynIIGWY O * = VLYNSISINONINO 0 | ; a We Py =e + - FHZNINIVN 9 « - SITVIDV79 9 + - VSNFLXF 9 * - /FLNOI37 9 p ae oe See 02 “Id ‘IL ‘ON ‘90T “10A SNOI1931109 SNOSNV1I13Z0SIN NVINOSHLIWS (‘6€ -d vas ‘uoryeuetdxea 10,7) VIN VOOddIH JO NOILNGIYLSIG WolHdvyeySosap @ - VPWISSILITOS W = evdsid O a ee « - /HILIOMOG W @- /7734FHIOD O +- SISNINOITIO O SNIDAFIANOD be mee pepehiws oo ae MOE te e SNOIL9371109 SNOANVTIZ9SIW NVINOSHLIWS IZ “Id ‘TL ‘ON ‘90T “10A (6 “d vas ‘uorjeuejdxe 10,7) VINVGOddIH 40 NOILNGINLSIG IvVOIHdVeDOaD 13 73830H : © -VININVYLS § + -/H2L0WD S$ - vlsnds $ ' S- So ed i meee ¢ -KLINITSIG S$ + - VLYNNIS S ret { Re rere a os <= ZZ “Id ‘IT ‘ON ‘90T “10A SNOIL9371109 SNOANVTISOSIN NVINOSHLIWS y apart aa hy 2 , i, te _ SMITHSONIAN MISCELLANEOUS COLLECTIONS | aes unas VOLUME 106, NUMBER 12 FROM SIAM. Ne BY cy H. G. DEIGNAN Associate Curator, Division of Birds é Ze | _ULS. National Museum ~ del eeRON 38356) city OF WASHINGTON : PUBLISHED BY THE SMITHSONIAN INSTITUTION — etreihs JUNE 24, 1946 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 12 DESCRIPTIONS OF TWO NEW LEAFBIRDS FROM SIAM BY H. G. DEIGNAN Associate Curator, Division of Birds U. S. National Museum @ 00200800, iy aM PsOSSS ITV Se NGTO (PUBLICATION 3856) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JUNE 24, 1946 The Lord Baltimore Press BALTIMORE, MD., U. & A. aed DESCRIPTIONS OF TWO NEW LEAFBIRDS FROM SIAM By H. G. DEIGNAN Associate Curator, Division of Birds U. S. National Museum I Study of fresh-plumaged (September—February) adult golden- hooded leafbirds from Siam has shown that the form of the Peninsula is a well-marked intermediate between cochinchinensis of Siam proper and moluccensis of Pattani and the Malay States. I suggest that it be called CHLOROPSIS COCHINCHINENSIS SERI-THAI, new subspecies Type.—vU.S.N.M. No. 330477, adult male, collected at Ban Tha Lo, southwest of Surat Thani or Ban Don (ca. lat. 9°5’ N., long. 99°15’ E.), Peninsular Siam, on September 23, 1931, by Hugh M. Smith (original number 5000). Diagnosis —The adult male may be known by its having the greens of the plumage somewhat deeper than those of cochinchinensis, but paler than those of moluccensis; by having the golden suffusion over the occiput and nape stronger than in cochinchinensis.and distinctly invading the upper back as in moluccensis; by having the golden gorget of the upper breast obsolescent, scarcely differentiated from the clear yellow of the lower throat and the green of the lower breast (in moluccensis the gorget is verditer green, wholly free of golden suffusion ) ; and by having the bill slightly larger than that of cochin- chinensis, slightly smaller than that of moluccensis. The adult female has the greens of the plumage and the size of the bill intermediate between those of cochinchinensis and moluccensis and the golden suffusion over occiput and nape as strong and almost as extensive as in the male (this suffusion is hardly apparent in cochinchinensis). Range.—Peninsular Siam, with the exception of Pattani Province, north at least as far as the Isthmus of Kra. Fresh-plumaged adults examined.—C. c. cochinchinensis, 23 oo (including 5 topotypes), 14 9? (including 2 topotypes) ; C. c. seri- SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 12 Zz, SMITHSONIAN MISCELLANEOUS COLLECTIONS ® vot. 106 thai, 8 6S (including 3 topotypes), 5 99 (including 2 topotypes) ; C. c. moluccensis, 6 ¢% (including 4 topotypes), 1 9; C. c. ictero- cephala, 2 33, I &. Remarks—The new race is named in honor of the Thai Resistance (Seri-Thai). C. c. moluccensis has been synonymized with icterocephala of Sumatra, but is separable by having the verditer-green gorget of the upper breast obsolescent, scarcely differentiated from the clear yellow of the lower throat and the green of the lower breast (not strongly indicated as in icterocephala), and by having the yellow of front, forecrown, and sides of head and neck more greenish, less golden, less extensive posteriorly on the crown, and less clearly demarcated from the coppery gold of the occiput and nape. The original reference for moluccensis will be: Chloropsis Moluccensis J. E. Gray, Zoological Miscellany, No. 1, pp. 3-4, 1831 (“Molucca”; type locality here corrected to Malacca). II The orange-fronted leafbird of northern Siam and the adjacent parts of Burma is not aurifrons, as all authors have hitherto recorded it, but a well-defined intermediate between aurifrons (Cachar, by restriction) and inornata (eastern Siam). I propose to name it CHLOROPSIS AURIFRONS PRIDII, new subspecies Type.—U.S.N.M. No. 311538, adult male, collected on the lower slopes of Doi Ang Ka or Doi Inthanon (ca. lat. 18°35’ N., long. 98°30’ E.), northwestern Siam, on December 9, 1928, by Hugh M. Smith (original number 2752). Diagnosis ——The adult of either sex may be known by its having the gorget bordering the black throat patch only about half as broad as the gorget of aurifrons, by having it of a more yellow, less orange, color, and by having almost no golden suffusion adjacent to the black on the sides of the head and neck. C. a. inornata in turn differs from pridii by its complete lack both of the gorget and of the golden suffusion on the sides of head and neck. Range.—Northern Siam, in the west, south to Muang Tak (Rahaeng); Karenni; southern Burma, south in Tenasserim to Amherst. Adults examined.—C. a. hodgsoni, 3; C. a. aurifrons, 9 (including 5 topotypes) ; C. a. pridii, 15 (including 1 topotype) ; C. a. iornata, 23 (including the type and one other topotype). NO. 12 TWO NEW LEAFBIRDS FROM SIAM—DEIGNAN 3 Remarks.—This form is named in honor of Pridi Bhanomyong (Luang Pradit Manudharm), leader of the Thai Resistance. Hume and Davison (Stray Feathers, vol. 6, pp. 326-327, 1878) have completely described this bird from Tenasserim, but failed to name it because immature birds were “scarcely separable” from those of the Himalayas. The birds from Hill Tipperah that appeared to them to be “intermediate between Tenasserim and Himalayan speci- mens” were probably true aurifrons. Himalayan examples seem to belong to a recognizable race, dis- tinguished from aurifrons by having both bill and wing rather longer and perhaps by having a slightly more extensive orange frontal patch. They must be called hodgsoni, for which the original reference will be: Phyllornis hodgsoni Goutp, The Birds of Asia, vol. 3, pt: 13, pl. 15 and text, May 1861 (Northern India, Himalayas, etc.; type locality here restricted to Nepal). For the loan of comparative material of this species, I am indebted to Dr. Ernst Mayr and the authorities of the American Museum of Natural History. one / Syitsinieys! pe apUneis inarpanvari thw tngatiee i 3 lor vrata th yee), oganeth = ot fistych sud, caipianat: aio tt pith bedtime a sols rent “ald eyeqae y toot” otadw abilld ontretnnth 6 of Aeresquantech, deroca®’ AUREL sos), abnish oD pet 00 ; eta SOREN bas. seixteona ly mapugiod, pi sad Arar oi wa rh a tel ae . “ithe: aoe ai aergeo, 3,0) yaledaps casia eglqeuamea’ § pagel, aeultins: aunty, bun igh Aloe meric), Wd avers Sie et Fez Janyou} omieme ay incon, stant ath dp nd a oth Mt sous GAT ipaigizo, apf dotlyy 40) noun toi bray pela at iy, Wow sinktionin eh a ‘ase inca . hatoitiest, oyu wii ivd 5 ate epgaleepld, seta] aia, he y a) bis fabigy: grt {ly #51 jet 2hitt atl taitsinet ‘Aves ee oe va ; jo mcitenhl aspera’ adt to esttoodtug ont. as tysbl by atta ig $ a 5 hyo lal VEN} ee Teh un ste tiesy | F CArh say: that ae or Lge ae haa . 4 : bias { Aa 2 oi Teo Oe Oe Vries iS é & = - ~ | u "aT rot alt PUES De ee «2 ee na LA IN OD TTS WT TT lina vy eS ; ails li . } ‘Dh 4, Lak Mie (ty Hale, cor chia on r ft mA ps a, | ip’ pT mY CMa (s ee Hs oviges Shih oh Teen AW j ; ite wet rip henneyem wy ie ' bidet ano ute La oR, a ty te Ts st i] SOLANA Ce ALE Let Bir a ie | ee ssenad Pd Lae, peel ig a We aa PORTS ee Jseenh "f a Ae OCR PIM ad OO) m F Lbeset | Gi, VR Bees Le 4 AS ae ain tear bale Way ake oe ey Lace of the Ronan ; ‘< ote ‘iene yah a ¢ Mi: ayn Patel ao ie) SOR re oh ue Wet, sith ce “i iy sae?) Nevo) petite: Satan. iia ames Thad al eS iyi yh wee sh ot yvtyhornea: 8; 3 p Oe ia Ltai it ; ea a are a ane inking i oe 1) Be ’ aii 08 VA eae Us a Wg efhili wepy a a en pe ay 4 -. oe ai o¥ Sih a abs a | RRL Be oe a Pi Cogs a Lt ae dy Re eee oy en, eg =: te Sree aes wt es ee Sate SPST - —_ _ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 13 oA NEW CARNIVOROUS DINOSAUR FROM ee THE LANCE FORMATION OF i | MONTANA Evite Four Puares) BY CH ARLES’ W. GILMOR E v3 | Curator, Division of Vertebrate Paleontology ; U.S: National Museum (PUBLICATION 3857) he CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION _ SEPTEMBER 12, 1946 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 13 A NEW CARNIVOROUS DINOSAUR FROM THE LANCE FORMATION OF MONTANA (WitH Four PLATEs) BY CHARLES W. GILMORE Curator, Division of Vertebrate Paleontology U. S. National Museum (PUBLICATION 3857) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION SEPTEMBER 12, 1946 any Pens ‘ The Bord Baltimore (Dress BALTIMORE, MD., U. 8. A. \ i i 4 i i ne Ph ee Ms 7 * 4 ‘a _ aw We, hy a . me Sd he Pe mess ay PoANN, yy we s ’ ‘ : wate, 2 : a % “ ev i - ‘ sn a” . f : 1 ; } ‘ . \ t . +/ * ‘ ¢ \ 4 fe » on, ’ ‘ ‘ « e . ‘ 2 - - - ‘ sf i m4 ~ . - . ‘ A NEW CARNIVOROUS DINOSAUR FROM THE LANCE FORMATION OF MONTANA By CHARLES W. GILMORE1!1 Curator, Division of Vertebrate Paleontology, U. S. National Museum (WirTH 4 PLATES) In 1942 an expedition from the Cleveland Museum of Natural History, under the leadership of Dr. D. H. Dunkle, explored the Lance formation in Carter County, Mont. Among the important discoveries resulting were the well-preserved skull and jaws of a carnivorous dinosaur. The specimen clearly pertains to the Deinodontidae, a family pre- viously represented in the Lance fauna only by the gigantic Tyranno- saurus. That other deinodonts occurred in this fauna has long been known from scattered teeth and phalangeal bones found from time to time during the 50 years or more that the Lance has been searched for fossil remains. The present discovery of an identifiable specimen is therefore of considerable interest. Through the courtesy of Dr. Dunkle this specimen has been placed in my hands for study and description. Family DEINODONTIDAE Cope The Deinodontidae includes all of the large Upper Cretaceous carnivorous dinosaurs such as Deinodon, Tyrannosaurus, Gorgo- saurus, and Albertosaurus. The affinities of the present specimen appear to be in the genus Gorgosaurus, as shown by many close similarities to G. libratus and G. sternbergi, thus greatly extending the geologic range of Gorgosaurus from the Belly River into the Lance. On the basis of its geologic occurrence, smaller size, and slight differences in skull structure, it becomes necessary to describe it as a new species, for which the name G. lancensis is proposed. The specific name is in reference to the Lance formation in which the type specimen was found. 1 Published posthumously. Mr. Gilmore died September 27, 1945. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 13 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Although following Matthew and Brown (1922) in their concep- tion of the availability and scope of the family Deinodontidae, it is well to point out that the genotype, Deinodon horridus, was founded by Leidy (1856) exclusively upon detached teeth, a fact well known to both of the above authorities, but who make the dogmatic statement that the family Deinodontidae was “based upon an unquestionably valid genus.” With this conclusion I disagree for, with the accumu- lating evidence of later years, it becomes more and more apparent that genera of Upper Cretaceous carnivorous Dinosauria cannot be dif- ferentiated on tooth characters alone. It therefore appears highly probable that the use of the genus Deinodon may have to be aban- doned and with it, following nomenclatural procedure, the family term Deinodontidae. Genus GORGOSAURUS Lambe, 1917 Genotype.—Gorgosaurus libratus. GORGOSAURUS LANCENSIS, new species Type.—C.M.N.H. No. 7541. Skull and lower jaws. Collected by D. H. Dunkle, August 16, 1942. Locality SE, sec: 11, R: 61 E., 'T.'4 S., ‘Sand "Creek, Carter County, Mont. Horizon.—Upper Cretaceous, Lance formation, Hell Creek member. The type and only specimen is a nearly complete skull with tightly articulated lower jaws. It has suffered the loss of the squamosal, postfrontal, quadratojugal, and the upper half of the quadrate of the right side; there is some disarrangement of the bones of the palate, and many of the teeth are either missing or incomplete. Otherwise the skull is in well-preserved condition. The coalescence of many of the sutures strongly suggests the adult age of the individual. So close is the union between upper and lower jaws that not only is the lower dentition wholly concealed, but the very grave danger of doing permanent injury to the specimen prevents any attempt to separate them. In size the skull is smaller than the cranium of the type of Gorgo- saurus sternbergi with which, in several respects, it appears to have the nearest affinities. Unfortunately for the present purposes the type of G. sternbergi has never been adequately described, and its original specific characterization was very meager. Furthermore, a single figure of the mounted skeleton published by Matthew and Brown NOT 13) NEW DINOSAUR FROM MONTANA—GILMORE 3 (1923, fig. 4) constitutes the only published illustration of this type. Since the skeleton is on exhibition and under glass that cannot be readily removed, direct comparison or study of the actual skull is practically out of the question at this time. In plate 4, figure 2, is shown a side view of the skull enlarged from the one and only nega- tive of this specimen and generously placed at my disposal by Dr. E. H. Colbert. Matthew and Brown (1923, p. 7) distinguished Gorgosaurus stern- bergi as follows: “It is of smaller size and more slender proportions than the G. libratus. The jaws are much less massive and the muzzle is more slender, the maxilla more elongate and shallow, the orbital fenestra more circular, the tibia is considerably longer than the femur.” In most of the skull characters mentioned, the cranium is in agreement with the specimen now before me. In slenderness of the muzzle it is intermediate between G. libratus and G. sternbergi. The dental formula appears to be the same in all three species. It should be pointed out, however, that this apparent slenderness of muzzle in the type of G. sternbergi (see pl. 4, fig. 2) is greatly ex- aggerated by the downward crushing of the nasal region. Restored to its normal profile this slenderness becomes much less noticeable. In its smaller size, elongate shallow maxillary, and rounder orbital fenestra, G. lancensis appears to have its closest affinities with G. sternbergi. There are some minor differences in skull structure, but none is regarded of specific significance. Such differences as are observed in the lateral temporal fenestral region are clearly due to postmortem distortion. The quadrate having been crushed forward and upward narrows the fenestra and otherwise alters the natural angulation of the elements composing this part of the skull. In the absence of detailed knowledge of the cranial characteristics of the two described species, it is quite impossible at this time, on structural features of the skull alone, properly to characterize the three species now assigned to Gorgosaurus, but in view of the great interval of time that has elapsed between Belly River and Lance, one appears justified in believing that it is too great a span in time for a species to pass unchanged from one to the other, and that with the discovery of additional materials characters will eventually be found that will adequately distinguish them. The genus Gorgosaurus was established on an adequate specimen, consisting of the greater portion of an articulated skeleton, but the skull was badly crushed and broken with much of the top portion missing. Lambe (1917) described’ in detail such parts of the type skull as were available, and although today there are several well- 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 preserved skulls of this genus known, a description of a complete cranium has not yet appeared. Because of this discrepancy in the literature the present specimen is here described in as much detail as its condition will permit. DESCRIPTION Parietal—The combined parietals are relatively short and unite with the frontals by a nearly straight transverse suture. Their upper surfaces at the center rise into a thin, sharp-edged, antero-posterior sagittal crest and a transverse supraoccipital crest. The latter rises high above the level of the skull top. Its slightly thickened upper border, viewed from the rear, presents a sinuous outline divided at the center by the truncated end of the longitudinal crest. Whereas the sides of the supraoccipital crest converge slightly from top toward the bottom in G. libratus, as shown by the skull of A.M.N.H. No. 5434, there is a strong overhang of the upper half of the lateral border when viewed from the rear. At midheight the supraoccipital crest has a greatest transverse diameter of 116 mm.; at the bottom, which rests upon the exoccipital and paraoccipital bones, a long, tapering process extends outward to increase suddenly the transverse diameter to 144 mm. or more. This process is received in a slotlike depression of the squamosal at its junction with the paraoccipital. Above the top of the supraoccipital the parietal shows a triangular-shaped thickening that articulates directly with the heavy median part of the supraoccipital, probably functioning in the same capacity as the heavy, bluntly pointed posterior projection that stands out so prominently on the Antrodemus skull. In front of the supratemporal crest the parietals are strongly con- tracted transversely between the supratemporal fossa, again widening at their anterior junction with the frontals, as shown in plate 2. Be- tween the fossa they have a least diameter of 55 mm. The greatest length of the parietals at the center is 68 mm. Frontal——The frontal area in ,Gorgosaurus lancensis, owing to the coalescence of many of the sutural contacts, can be interpreted only somewhat uncertainly. The union with the parietal and postfrontal is quite clear, but the outline and anterior extent of the frontals where they meet the lacrimal and nasal bones cannot be surely determined. The median suture separating the paired frontals is clearly discernible on the anterior third of their length, but posteriorly the fusion is so complete as to leave no trace of the line of contact. On the right side, between the hornlike projections of the lacrimals, an offset in the bone surface extends backward and outward from the median NOI 3 NEW DINOSAUR FROM MONTANA—GILMORE 5 suture, and is thought to represent the line of lateral union between the frontal, nasal, and prefrontal of that side. If this interpretation is correct, the combined frontals end anteriorly in a point deep within a V-shaped notch in the nasals as shown in plate 2. Contributory evidence as to the correctness of this interpretation is found in a skull of Gorgosaurus libratus, U.S.N.M. No. 12814, which shows a closely similar arrangement of these elements. That the frontal is coalesced with the prefrontal latterly appears to be indicated on the left side by a short sutural edge that extends inward and backward from the border of the orbital notch. If this is the posterior end of the pre- frontal, it occupies a position similar to that of the prefrontal in Antrodemus, and quite unlike the more central position in the Tyran- nosaurus skull. A wedge-shaped process of the frontal extends out- ward to fill the interspace between the postfrontal and prefrontal bones, but it appears to be excluded from participation in the orbital border by the articulation of these two elements at the bottom of the notch. On the right side of the skull there is no trace of the prefrontal suture. The sagittal crest extends forward from the midline of the parietal on to the frontal, reaching its highest elevation on the posterior fourth of that bone. Anteriorly this crest drops rapidly downward to the level of the skull surface. It is estimated that the nasal bones at the center had a greatest length of about 112 mm. Nasal——The nasal bones, as in other members of the Deino- dontidae, are especially long and narrow, with transversely rounded dorsal surfaces anteriorly, but flattening out on the posterior third. The central areas are coalesced into a single rugose element that is suturally separate anteriorly. The contact with the frontals is broad, not narrow as in Tyrannosaurus; the pointed ends of the combined frontals being received in a deep V-shaped notch on the midline, as shown in plate 2. Anteriorly the nasals have bifurcated ends, a heavy upper process that extends forward above the nares to meet an ascending process of the premaxillary which it underlaps, and a much weaker process that extends forward and downward along the upper surface of the maxillary border to meet an ascending process of the premaxillary and thus exclude the maxillary bone from par- ticipation in the boundary of the external nares. The lateral contacts of the nasal with the maxillary, lacrimal, and prefrontal bones have the sutures coalesced, and their course can be only approximately determined. A row of small foramina on the side of the left nasal, running posteriorly from the border of the nares, suggests that the 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 roughened areas may have had a chitinous hornlike covering.. The greatest length of the nasals is 388 mm. Postorbital—tThe bone here designated postorbital in all probability represents the coalesced postfrontal and postorbital. Since no trace of their sutural union can be detected in this specimen they are de- scribed as a single element. The postorbital is a triradiate bone with short, heavy inner-directed process that articulates with the frontal by a strong suture; a second slender, tapering process that extends back- ward and downward is received in a depressed groove on the outer side of the squamosal, thus forming the upper temporal bar; the third, the longer one of the three, also a tapering process, extends downward and laps along the anterior side of the jugal to form the wide bar separating the orbit and lateral temporal fossa. From the superior border to the end of the inferior branch it measures 114 mm. Its greatest antero-posterior diameter is 145 mm. There is no “post- frontal rugosity” as in Tyrannosaurus and the notch separating this bone from the lacrimal rugosity is shallow. At the bottom of this notch it appears to be in contact with the prefrontal, thus excluding the frontal from the orbital border. Prefrontal—The narrow area on the top of the skull, lateral to the frontal and wedged in between the lacrimal frontal and nasal bones, is here tentatively regarded as pertaining to the prefrontal (see pl. 2). If correctly determined, it has a greater dorsal exposure than in Tyrannosaurus, and it differs further in being in contact with the postfrontal. Lacrimal.—The lacrimal of Gorgosaurus lancensis resembles those of Antrodemus and Ceratosaurus in having a stout, blunt-edged ele- vation rising from its outer superior surface. It may have supported, as first suggested by Osborn (1903), “something in the nature of a low dermal horn.” A similar horn-projection is present on the skulls of G. libratus and G. sternbergi, but is absent in Tyrannosaurus rex. On the external side the base of this “hornlike” projection is exca- vated, and the bone is still further lightened by internal cavities, as shown by the incomplete right element which lacks its top (see pl. 2). The sutural contacts of the lacrimal with the adjacent, prefrontal (?), frontal, and maxillary bones cannot be accurately traced in this specimen. Posteriorly it is separated from the postorbital by a narrow notch. Ventrally it unites with the jugal, the posterior angle being received in a notch in the top of that bone, but anteriorly these two bones appear to lap one another. Premaxillary—The premaxillary, as in other deinodonts, is re- duced in size. It carries either four or five teeth, a point that cannot NO; £3 NEW DINOSAUR FROM MONTANA—GILMORE il be certainly determined by this specimen owing to the obliteration of the premaxillo-maxillary suture. In all probability it will be found there are four premaxillary teeth as in Gorgosaurus libratus, so clearly demonstrated by Lambe (1917). The slender superior process rising from the upper anterior border, with its counterpart of the opposite side, meets the nasals above the midlength of the external nares. Maxillary—Roughly triangular in outline, the maxillary is deeply excavated posteriorly by the large antiorbital fenestra. The posterior process that extends backward to meet the jugal is especially long and slender, its tapering termination ending below the center of the orbital opening as it does in G. sternbergi. Above, it unites with the nasal and lacrimal, but in this specimen most of the sutural contacts are obscured by the coalescence of the bones, as is the union with the premaxillary. On the left side of the mid- lateral surface of the maxilla adjacent to the position of the pre- maxillo-maxillary suture there is evidence of a small opening, which in Tyrannosaurus Osborn designates the third antiorbital fenestra. It appears to be absent on the right side of this specimen. The maxil- lary is excluded from participation in the boundary of the nares by a slender downwardly directed process of the nasal. The base of the ascending maxillary process is perforated by the small second antiorbital fenestra. The surface of the bone in front of this fenestra is smooth and depressed, forming a marginal tract at a lower level than the general surface of the skull at this point. Viewed from the side the dental border is sinuous, with a row of dental foramina. The greatest length of the maxillary is about 405 mm., of which 280 mm. is tooth bearing. There are clearly 13 teeth and possibly 1 more if the premaxillo-maxillary suture cuts between the first and second U-shaped teeth of the front series as Lambe (1917) found it to do in the type of G. libratus. This point cannot be deter- mined in this specimen owing to the coalescence of this suture. Jugal—The jugal has the usual triradiate shape, but is more slender and elongate antero-posteriorly than in Tyrannosaurus. It extends forward as a tapering process to join the maxillary by squamous union. It ‘is underlapped for nearly half its length by the slender posterior process of the maxilla that terminates below the center of the orbit. This anterior process of the jugal is perforated by a large oval-shaped jugal fenestra, ventral to its union with the lacri- mal. There is a notch in the upper border of the jugal for the recep- tion of the lower posterior end of the lacrimal, but anterior to this notch it overlaps the side of the descending lacrimal. The tapering superior process unites by squamous union with the descending 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 process of the postorbital, the two forming the postorbital bar which separates the orbit and the lateral temporal fenestra. Posteriorly the jugal is overlapped by a fingerlike process from the quadratojugal, as shown in plate 1. In this specimen crushing has broken this process in two so that its full posterior extent is not to be observed (see pl. 1). The lower border, anterior to the forward termination of the quadratojugal, has a flattened projection that extends prominently downward as a blunt process (see pl. 1). That this is a normal de- velopment is demonstrated by its presence on the opposite side. This process appears to be absent in G. sternbergi but is present in G. li- bratus. The jugal forms all of the lower boundary of the orbit and little, if any, of the postero-ventral boundary of the large antiorbital fenestra. The greatest height of the jugal is 170 mm. Squamosal.—The right squamosal is missing but the left is almost complete, and the open sutures give a clear picture of its extent and relationship to the surrounding elements (see pls. 2 and 3). Externally the forwardly directed bar is overlapped on the outside by the pos- terior process of the postorbital, the two forming the upper supratem- poral bar; an inwardly directed process passes in front of the high vertical supraoccipital crest to lap along the parietal on the postero- medial side of the supratemporal fossa ; posteriorly the squamosal rests against the front of the paraoccipital process; on the external ventral side a slender process curves forward and downward to cap the top of the quadratojugal. Ventrally it articulates with the quadrate, but this side is hidden by enveloping matrix. Quadratojugal—tThe lateral view of the skull (see pl. 1) shows clearly how the quadratojugal and quadrate bones have been crushed forward, thus closing the lower part of the lateral temporal fenestra to a narrow slotlike opening. As shown by the overlapping broken ends of the lower anterior process, this fossa should be at the least 26 mm. wider at the bottom than as shown in plate 1. The anterior border of the quadratojugal within the fossa is incomplete, so that its full extent and proper outline cannot be determined. Immediately below midlength, on the outer side, the surface is indented by a shallow rounded depression. Internally it unites by suture with the quadrate except at the point of the large quadrate foramen. The proximal end is capped by a descending process of the squamosal. The greatest length of the inferior border is estimated to be about 95 mm.; the greatest height 110 mm. The sutures on the inner side with the quadrate are coalesced and cannot be traced. Quadrate-—Both quadrate bones are so much hidden by matrix, or closely enveloped by the squamosal above and the mandible below, NO. 13 NEW DINOSAUR FROM MONTANA—GILMORE 9 that little is to be seen of these elements except from the rear. The large vertical ovate quadrate foramen lies between the quadratojugal externally and the quadrate internally. Below this foramen the distal portion of the quadrate rapidly widens toward the outside where it is coalesced with the quadratojugal. The left quadrate at the distal end has a greatest transverse diameter of 67 mm. The bilobed distal extremity fits closely into the cotyloid depression of the mandible. On the inner side slightly above the distal end a process extending inward and forward articulates with the pterygoid, but its full forward development is hidden in the adhering matrix. Occipital region.—The coalescence of most of the bones of the occip- ital region of the present skull makes a detailed study of its several elements very unsatisfactory. Viewed from the rear the median upper part of the occiput is dominated by the vertically and horizontally expanded supraoccipital crest formed wholly by the parietal. Medially it envelopes the stout supraoccipital; its lower outer border is con- tinued outward as a slender, pointed process that rests on the upper border of the paraoccipital process in a transverse depression crossing the back of the squamosal. The upper end of the supraoccipital is squarely truncate, with its surface rugosely roughened. This surface, in conjunction with the lateral pits, probably gave attachment for the ligamentum nuchae. Laterally and ventrally its sutural limits cannot be traced. For that reason it is not known whether the supraoccipital participated in the boundary of the foramen magnum or not. The exoccipitals contribute extensively to the formation of the occipital condyle as in Antrodemus valens. Their dorsal connection with the supraoccipital can no longer be traced. The surfaces lateral to the foramen magnum are perforated by two foramina, the larger and most posterior one being for the passage of the twelfth or hypo- glossal nerve; the smaller and more anterior one for the eleventh or accessory nerve. The paraoccipitals or opisthotics coalesce toward the median line with the exoccipitals and below the latter turn downward to join the basioccipital by an oblique suture. Both paraoccipital processes lack their outer ends, though the left element is sufficiently complete to show it was expanded and in contact with the posterior surface of the squamosal. The basioccipital contribution to the occipital condyle looks strongly downward. This articular surface, when viewed from below, is sub- triangular in outline as contrasted with the oval shape in Tyranno- 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 saurus. The condyle, as in nearly all Dinosauria, is inclined ventrally in relation to the longer horizontal axis of the skull. Anteriorly, where the basioccipital meets the broad basisphenoid, strong, diverging basioccipital processes extend forward and down- ward to a point 44 mm. below the level of the occipital condyle. Laterally, they articulate with the exoccipitals and anteriorly with the basisphenoid. The median area between the descending processes is shallowly concave transversely. In this specimen this whole region has been considerably flattened by vertical crushing. The basisphenoid viewed from below is subrectangular in outline with the greatest diameter transverse. Posteriorly it unites with the basioccipital, anteriorly with the pterygoids. The smooth surfaces of the enlarged basisphenoid processes that articulate with the ptery- goids indicate this joint to have been movable. These processes are joined transyersely by a wide septum of bone as in Tyrannosaurus. The posterior surface of the basisphenoid is hollowed out with a single opening near the center that leads upward and backward toward the brain cavity. This is identified as the internal carotid foramen. A second opening posterior to the carotid foramen has every appearance of being abnormal; it is asymmetrically placed and the adjacent sur- faces suggest an unhealthy condition of the bone. The sides of the basisphenoid are hidden in the enveloping matrix. Palate.—Although the type skull has much of the palate present, postmortem crushing, coalescence of the sutures, and adhering matrix have made it impossible at this time to give a complete account of the structural details of this important part of the cranium. The palate shown in plate 3 is an attempt to give the ventral view of the pterygoids in combination with the palatines, which in this skull are visible only in the antiorbital fenestra and from the dorsal side. Since there is ample opportunity here for a difference of opinion as to their proper interpretation, this view should be used with caution. The pterygoids, as in Tyrannosaurus, unite with the basipterygoid processes by movable joints. They unite laterally with the ectoptery- goids and near the posterior end a process extends upward, outward, and backward to meet the quadrate, but the full extent of this sutural contact is partly hidden in the matrix. The broad outward extension of the pterygoid that meets the ectopterygoids in relative extent and outline closely resembles those of Tyrannosaurus as illustrated by Osborn (1912, fig. 5). How much of this winglike expansion is ectopterygoid cannot be determined because of the coalescence of the sutures. The ectopterygoid portion appears to be perforated by an ovate opening as in Tyrannosauras. The internal barlike parts of the NOS Ls NEW DINOSAUR FROM MONTANA—GILMORE inal pterygoids extend forward with their inner borders parallel, and with their widest surfaces strongly inclined upward from the horizontal, indicating a high but narrow palatal roof. They do not meet on the median line. This bar portion gradually widens toward the front, its anterior end is bifurcated. The outer or heaviest branch meets the palatine by a straight transverse suture; the inner branch, a slender, flattened extension of the median side, turns gradually inward to abut the inward extension of the palatine, thus inclosing a subtriangular vacuity between the two bones at this point. In Tyrannosaurus the vacuity is absent, the slender inner process extending forward tight along the median site of the palatine to meet the vomer. The vomer in G. lancensis has been tentatively identified as extending forward from the palatine to disappear in the matrix between the closely appressed jaws (see v, pl. 1). Mention should be made of the doubtful nature of the exact point of sutural union between the pterygoid and palatine. This suture as tentatively determined is based on the presence on the ventral or palatal surface of a coalesced suture. This determination, however, needs additional confirmation. As preserved the main portions of the palatine bones have been crushed upward to a nearly perpendicular position, as best shown in the left antiorbital fenestra (see /, pl. I). The right palatine occupies a similar position in the opposite fenestra but it is less well preserved. A skull of G. libratus, U.S.N.M. No. 12814, shows a similar position of the left palatine. The outer end of the palatine that articulates with the maxillary is extended forward as a slender tapering process, and posteriorly a much shorter process contacts the jugal and possibly the lacrimal. The heavy central portion of the palatine, shown best in the left anti- orbital fenestra (pl. 1), stands nearly vertical. Although it is quite evident that in their normal relationships the palatines would meet on the medial line of the palate, it is also clear they must have been steeply inclined upward toward the center of the palate; otherwise, their great length would cause them to overlap strongly if brought to a horizontal position. On the dorsal surface close to the palatine- maxillary suture the palatine is perforated by two small ovate open- ings, the smaller one being anterior. In Tyrannosaurus Osborn (1912) shows only one opening through the palatine. The inner end of the palatine is extended forward as a relatively wide tapering process that disappears from view in the matrix now filling the skull anterior to the large antiorbital fenestra. The anterior portion of this process may represent the posterior end of the vomer, as shown in plate 1. This suggestion is made on the basis of the presence of a 12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 diagonal break crossing this process in practically the same manner on both sides of the palate. If the conclusion reached here is correct, the internal narial opening is bounded principally by the palatine, only slightly by the maxillary and vomer, and the vomers would be excluded from contact with the pterygoids as they are in Platecarpus and Cyclura. Until confirmatory evidence is available, however, the suggestions made should be used with caution for it would seem highly improbable that the palate of Gorgosaurus would differ so radically from that of Tyrannosaurus and Ceratosaurus, both of which have the pterygoids and vomers articu- lating on the median line. EXTERNAL OPENINGS IN THE SKULL Besides the orbital and narial openings there are four other fenestra on the side of the skull. The orbit is subround, having a greatest ver- tical diameter of 122 mm., a greatest antero-posterior diameter of 95 mm. Dorsally the upper boundary is formed by the lacrimal pre- frontal and postfrontal. The frontal does not appear to participate in its boundary. Anteriorly the lacrimal forms the whole border; ven- trally it is bounded by the jugal, and posteriorly by the postorbital. The first antiorbital fenestra is the largest opening in the skull, measuring 153 mm. antero-posteriorly, with a greatest diameter dorso- ventrally of 111 mm. Its upper boundary is formed by the lacrimal and maxillary, anteriorly by the maxillary, and ventrally by the maxillary, lacrimal, and jugal. The small second antiorbital fenestra lies wholly in the maxillary bone. The external nares are elongate oval in outline, and have a greatest antero-posterior diameter of 83 mm., a vertical diameter of 32 mm. The boundaries are about equally divided between the premaxillary and nasal bones; the maxillary does not participate. The latero-temporal fossa is much narrowed by the pushing forward of the quadrate and quadratojugal bones, so that a true conception of its proper outline is not to be obtained from this specimen. Above it is bounded by the posttemporal bar formed by the united processes of the squamosal and postorbital bones; anteriorly by the jugal and descending process of the postorbital ; ventrally by the quadratojugal, and posteriorly by the quadratojugal and squamosal. The greatest vertical extent of this opening is about 152 mm. The jugal foramen is an ovate opening that perforates the jugal immediately below its junction with the lacrimal. Its antero-posterior diameter measures 29 mm. NOE) LES NEW DINOSAUR FROM MONTANA—GILMORE 13 The supratemporal fossa is bounded anteriorly by the parietal and postfrontal + postorbital ; externally by the postorbital and squamosal ; posteriorly by the squamosal and internally by the parietal and squamosal. The quadrate foramen lies between the quadrate and quadratojugal and looks directly backward. Its longest diameter dorso-ventrally measures 44 mm. in length and 18 mm. wide at the center. Mandible——Although the lower jaws are completely preserved, they are so closely articulated with the skull that much of their :de- tailed structure is hidden from view. In so far as comparison can be made, except for their smaller size, the jaws are in close agreement with those of Gorgosaurus libratus. Viewed from below the anterior ends of the rami are in close apposition, having a combined width of 55 mm. In a posterior direc- tion they rapidly diverge (see pl. 3), until across their posterior ends they have a spread of 330 mm. from side to side. The dentary, as in other deinodonts, has a receding anterior end as contrasted with the more squarely truncate ends of the megalosaurians Ceratosaurus and Antrodemus. Posteriorly the dentary articulates above with the large surangular and below with the angular, under- lapping the one and overlapping the other. In front the ventral border of the dentary is heavy and rounded, gradually narrowing transversely toward its posterior extremity which laps along the outer side of the angular as a thin fingerlike extension. The inside of the posterior part of the dentary is hidden from view by the overlying splenial. The surangular in Gorgosaurus has been so completely described by Lambe (1917, pp. 13 to 16) that it seems unnecessary to repeat the description here. The angular is clearly defined on both rami, and is almost wholly confined to the outer aspect of the ramus, being slightly visible on the inner side near its anterior end where it underlaps the prearticular and the posterior extremity of the splenial. In outline it is roughly triangular in shape, overlapping the surangular above and extending posteriorly to a sharp point that is separated internally and posteriorly from the prearticular by the interposition of a long tapering process that extends forward from the articular. Anterior to this process it comes in direct contact with the prearticular. On the anterior end it sends forward a small fingerlike process that is received between the splenial and dentary on the ventral surface. On the lateral surface at the point of junction with the dentary and surangular, there is a small elongate foramen that leads forward to the inside of the dentary. This foramen in position corresponds to the large oval opening in the 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Ceratosaurus ramus. The left angular has a greatest length of 174 mm., a greatest width of 52 mm. The coronoid, as in Tyrannosaurus, is a small, thin bone that is confined to the forward part of the pterygoid fossa wedged in between the prearticular and surangular bones, and only visible on the inside of the ramus. TABLE I1.—Comparative measurements of skulls 29 3S = 2% poole Oe Sia 37, 855 SZ IZ SOH Bah 3Ou Ss eS mm, mm mM. mm, mm Length of skull, pmx. to mand. condyle.. 570 678 980 1355 Length of skull, pmx. to occipital condyle 572 Ae fare 925 1210 Height of skull, supraoc. crest to mand. CONAYVIE Se ayeriiecigth ecm Sopra aaa tare 190 205 seh 303 635 Width of skull across quadratojugals... ... ae a 380 835 Kengthiotupper dentitionessaqaemeenee 300 340 ae 495 ae FenothwotmMoweln aware emer cae 570 690 950 999E =1205 Height of muzzle above last tooth...... 17I 160 Fane 260 The prearticular is a long curved element that extends anteriorly from the articular and, with the angular, contributes to the formation of the lower posterior mandibular border. At midlength it curves strongly upward, terminating in a pointed end that is wedged in be- tween the upper posterior end of the splenial and the surangular. This half of the bone is relatively thin transversely; narrowest at the center, it gradually widens toward both ends. It is almost identical in shape with the prearticular of Tyrannosaurus as figured by Osborn (1912, fig. 18), but in Gorgosaurus lancensis the contact with the surangular is much stronger. The prearticular-articular contact at the posterior end is partly coalesced, but on the external side of this end where the tapering process separates it from the surangular the suture on both jaws is to be clearly observed. The articular forms the posterior portion of the ramus, but owing to the coalescence of most of the sutures and the covering of the superior surfaces by the closely articulated quadrate, the articular bones in this specimen cannot be delimited. Internally, it develops a wide flattened process that extends inward at nearly right angles to the length of the ramus. Ventrally a long tapering process extends forward between the posterior ends of the prearticular and angular, a condition previously noted in Albertosaurus by Lambe (1904, pl. 1). s NO. 13 NEW DINOSAUR FROM MONTANA—GILMORE 15 Measured from the inner end of the articular process to the outside of the ramus, this posterior end of the jaw has a greatest diameter of II5\ mm. Hyoid—Embedded in the matrix covering the midpalatal region a slender, subround, sinuous bone, with slightly enlarged ends, is identified as one of the hyoid elements. From end to end in a straight line it has a length of about 234 mm., greatest diameter of the pos- terior end, 13.5 mm. Its position as found im situ is shown in plate 3, h. Dentition—The upper dentition of Gorgosaurus lancensis consists of 37 teeth, of which the crowns of more than half are present in the type specimen, 13 on the left and 14 on the right. Owing to the coales- cence of the premaxilla-maxillary suture, the number of teeth carried by each of these bones cannot be positively determined. In the front of the series there are Io teeth of reduced size, rodlike and with flattened sides. As far as these teeth can be examined, they appear to be in full accord with those described by Leidy (1868) under the genus name Aublysodon. On similarity of structure alone all these teeth might be regarded as premaxillary, but Lambe (1917, p. 16) has pointed out that ‘“‘one of the principal distinctive characters of the dentition of Gorgosaurus is the similarity of the first maxillary tooth in size and shape to those of the premaxilla.”’ If the same con- dition prevails in the specimen under consideration, there are 4 premaxillary and 14 maxillary in this species, the same as in G. libratus. The maxillary teeth are of the usual compressed trenchant type, irregular in size, with serrated edges and backward curvature near the pointed end. The irregularities in size of adjoining teeth in the maxillary series are due to the different degrees of extrusion, and they are not always in agreement on the two sides of the skull. On the left side the third, fifth, seventh, and eleventh are larger than those fore and aft, whereas on the right side the third, sixth, eighth, and tenth are the largest teeth. On this side there is a gradual decrease in size from the tenth backward. The fourteenth tooth is especially small. The teeth of the lower jaw are completely hidden by the skull, so complete is their articulation. The dental formula of 18 upper teeth on a side is in full accord with the type of Gorgosaurus libratus as distinguishing it from Tyranno- saurus with 16 upper, Albertosaurus with 15, and Dromaeosaurus with 12. At this time it does not appear possible to determine genera of the Deinodontidae by detached and scattered teeth. 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 TABLE 2.—Dentitions of Deinodontidae compared Maxillary Premaxillary Lower jaw Gorgosaurus lancensis .......2....eee0es 14 4 as GRUDTGIUS aeeR ers Go Ee ee 13 4 14 GoSerNDETOT sone conse mie oe steelers austen 15? ? ae DT AONNOSQUTUSENEA erie i tae) iets sieialcieie ore ste 12 4 13 to 14 Albertosaurus sarcophagus ...........+-- 12 a 14 to 15 DISCUSSION OF UPPER CRETACEOUS DEINODONTS Arranged in chronological order the following genera and species of deinodont dinosaurs have been named as occurring in the Lance formation of North America: Aublysodon amplus Marsh, 1892; A. cristatus Marsh, 1892; Manospondylus gigas Cope, 1892; Tyran- nosaurus rex Osborn, 1905; and Dynamosaurus imperiosus Osborn, 1902. The two species of Aublysodon named by Marsh were each based on a single anterior tooth, and on the assumption that the types were inadequate Matthew and Brown (1922) referred both to “? Tyrannosaurus sp.” At this same time these authorities pointed out that the genus Aublysodon had to be abandoned on the ground of priority as being a synonym of Deinodon. Manospondylus gigas Cope, based on two worn vertebral centra, is regarded indeterminate by Osborn (1916), a decision concurred in by Matthew and Brown (1922). Tyrannosaurus rex is a thoroughly good genus and species, based on an adequate specimen. Dynamo- saurus imperiosus was abandoned by Osborn as a synonym of Tyran- nosaurus rex. This brief review of the named deinodonts from the Lance shows that Tyrannosaurus rex is the only one of the five named species that has a clear title. On geological position alone it would be to the Edmonton Alberto- saurus that one would first turn in looking for the progenitor of the Lance Gorgosaurus lancensis, but here again adequate comparison cannot be made owing to the fragmentary nature of the only known skulls of A. sarcophagus. The genus Albertosaurus has not been satisfactorily characterized, and Matthew and Brown (1922, p. 374) have already suggested that it may be identical with Deinodon. Men- tion should also be made of A. arctunguis Parks (1928), a species based on a partial skeleton without skull. Parks distinguished it from A. sarcophagus on differences found in the pes and pelvic bones, but since there is the possibility of wrong association of these parts of the original Osborn types, this species at the present time is also of uncertain value. In the Belly River formation the genus Gorgosaurus is the single known representative of the Deinodontidae. Although G. libratus and NOZ ES NEW DINOSAUR FROM MONTANA—GILMORE W7, G. sternbergi are both established on articulated skeletons, Matthew and Brown (1922, p. 383) have pointed out there is nothing in the dentition to separate Gorgosaurus from Deinodon and that, as far as the teeth are concerned, they would seem to be the same genus. For the present, however, I fully concur with their final recommenda- tion that “pending discovery of adequate topotypes in the Judith River beds the identity has not been finally and conclusively proven. There may be differences in the skull.” Although not prepared at this time to recommend the abandonment of Deinodon as an indeterminate genus, accumulating evidence offers little hope of its eventual stabilization. Thus, in the present paper, both Deinodon and Gorgo- Saurus are regarded as valid genera. In the eastern United States the deinodonts are represented by Dryptosaurus aquilunguis from the Upper Greensand of New Jersey, and two doubtfully referred species, D. medius and D. potens, from the Arundel of Maryland and the District of Columbia, respectively. Originally described as Laelaps by Cope (1866), Marsh (1877) found it to be preoccupied and proposed the name Dryptosaurus to replace it. In 1890 this same authority proposed and characterized the family Dryptosauridae. Should the genus Deinodon eventually be aban- doned, thus rendering the family name Deinodontidae untenable, the New Jersey family name is available to replace it. This brief review of the large Upper Cretaceous carnivorous Dinosauria focuses attention on the very unsatisfactory state of our knowledge concerning the nomenclatural status of many of the in- cluded forms, and the great need for the recovery of topotypic ma- terials if the family is ever to be placed on a solid foundation. The geologic distribution of the Deinodontidae as known at the present time is graphically shown in the accompanying table. TABLE 3.—Geologic distribution of the Deinodontidae Formation Western Upper Cretaceous Lance Tyrannosaurus rex Osborn Gorgosaurus lancensis Gilmore Edmonton Albertosaurus sarcophagus (Cope) A. arctunguis Parks Belly River and Judith River Gorgosaurus libratus Lambe G. sternbergi Matthew and Brown Deinodon horridus Leidy Eastern Upper Cretaceous Hornerstown Dryptosaurus aquilunguis (Cope) D. ? medius (Marsh) Arundel D. ? potens (Lull) 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 REFERENCES Brown, BARNUM. 1907. The Hell Creek beds of the Upper Cretaceous of Montana. Bull. Amer. Mus. Nat. Hist., vol. 23, pp. 823-845. Gore he) 1866. Remarks on dinosaur remains from New Jersey. Proc. Acad. Nat. Sci. Philadelphia, June, pp. 275-270. GILMORE, CHARLES W. 1920. Osteology of the carnivorous Dinosauria in the United States National Museum, with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus. U. S. Nat. Mus. Bull. 110, pp. 1-155, 30) pls.) 143 text figs. Jalas, @5 12 1908. On certain genera and species of carnivorous dinosaurs, with special reference to Ceratosaurus nasicornis Marsh. Proc. U. S. Nat. Mus., vol. 35, pp. 351-3606, text figs. I-4. LAMBE, LAWRENCE M. 1904. On Dryptosaurus incrassatus (Cope) from the Edmonton series of the Northwest Territories. Contr. Can. Paleont. Geol. Surv. Canadas vole.3) pt 3) pp) 5-25) plac: 1917. The Cretaceous theropodous dinosaur Gorgosaurus. Geol. Surv. Canada, Mem. 100, No. 83, Geol. Ser., pp. 1-80, text figs. 1-84. Lerpy, J. 1856. Notice of remains of extinct reptiles and fishes, discovered by Dr. F. V. Hayden in the Bad Lands of the Judith River, Nebraska Territory. Proc. Acad. Nat. Sci. Philadelphia, vol. 8, pp. 72-73. MarsH, O. C. 1877. Notice of a new and gigantic dinosaur. Amer. Journ. Sci. (3), vol. 14, pp. 87-88. 1892. Notes on Mesozoic vertebrate fossils. Amer. Journ. Sci., vol. 44, pp. 174, 175, pl. 3, figs. 4-6. MatTrHew, W. D. 1920. Canadian dinosaurs. Nat. Hist., vol. 20, No. 5, pp. 541, 544. MatrHew, W. D., and Brown, BAaRNUM. 1922. The family Deinodontidae, with notice of a new genus from the Cre- taceous of Alberta. Bull. Amer. Mus. Nat. Hist., vol. 46, art. 6, pp. 367-385, fig. 1. 1923. Preliminary notices of skeletons and skulls of Deinodontidae from the Cretaceous of Alberta. Amer. Mus. Nov., No. 89, pp. I-10, figs. I-5. Osporn, H. F. 1903. The skull of Creosaurus. Bull. Amer. Mus. Nat. Hist., vol. 19, art. 31, pp. 697-701. 1905. Tyrannosaurus and other Cretaceous carnivorous dinosaurs. Bull. Amer. Mus. Nat. Hist., vol. 21, art. 14, pp. 259-265. 1912. Crania of Tyrannosaurus and Allosaurus. Mem. Amer. Mus. Nat. Hist., vol. 1, pt. I, pp. 3-30, pls. 1-4. 1916. Skeletal adaptations of Ornitholestes, Struthiomimus, Tyrannosaurus. Bull. Amer. Mus. Nat. Hist., vol. 35, pp. 733-771, 3 pls., 21 figs. mNOS 13 NEW DINOSAUR FROM MONTANA—GILMORE 19 Parks, W. A. 1928. Albertosaurus arctunguis, a new species of theropodous dinosaur from the Edmonton Formation of Alberta. Univ. Toronto Stud., Geol. Ser., No. 25, pp. 1-42, pl. 1. EXPLANATION OF PLATES PLATE I Skull and jaws of Gorgosaurus lancensis viewed from the left side. Type. C.M.N.H. No. 7541. p, palatine; v, vomer. About 4 natural size. PLATE 2 Skull of Gorgosaurus lancensis viewed from above. Type. C.M.N.H. No. 7541. About } natural size. PLATE 3 Skull and jaws of Gorgosaurus lancensis. Palatal view. Type. C.M.N.H. No. 7541. h, hyoid. About 4 natural size. PLATE 4 _ Comparison of Gorgosaurus skulls Fic. 1. Skull and jaws of Gorgosaurus lancensis. Type. C.M.N.H. No. 7541. Viewed from left side (reversed). Fic. 2. Skull and jaws of Gorgosaurus sternbergi. Type. A.M.N.H. No. 5664. Viewed from the left side. Fic. 3. Skull and jaws of Gorgosaurus libratus. A.M.N.H. No. 5434. Viewed from left side. All figures about 1/Io natural size. ‘poortiennpedn iy asc , did date ota ks, cit pois i soa eet ee 4 Be ho: Hep ty) ie prams Wain: 0 i witht Ps oxy rR en Ae LE 4 | | BA TAIT HO HOITAUAIEEA ; Me ; ¢ tel WV aw pen We) yi wees ; arrive Lene: um HPD -(Sbte she See ert ‘heweahid Lienasanl sHenegh ie), pe ie “st iit a PRR SOTO? set toon ae aie Ne i. Hy rex: Wail Penwal® rare tre, tah A ie EA SLM k's sn a mt Me rh od he . brid Be Bu. cen, ee i Md ager ame mtn mont Maa una ainh ur winging 7 . " A a Ah ¥ ae ui iat hos " f ‘ Mbbetianssee « u nate \ 7.) ie | ya Oh aa tole oe . x GA wel AID) sx P wore tas JOO) DrutyouNoD ik { tN ie) ETT aay Fe ig sin Ab . ‘ rit ‘ Ci Pte one . : ae ; i! Peta r ni ai . ? 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(For ex SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 06 NOS 3 PEs GORGOSAURUS LANCENSIS (For explanation, see p. 10.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES L0G) NOA tS ye Piaae Eres 2 Gens COMPARISON OF GORGOSAURUS SKULLS (For explanation, see p. 10.) ids i | SMITHSONIAN MISCELLANEOUS COLLECTIONS Bo a VOLUME 106, NUMBER 14 A . NEW DUSSUMIERIID FISH OF THE _ GENUS JENKINSIA FROM ey: BERMUDA | | ‘ (Wire ONE Pratt) s: NSABY EN | “. LUIS RENE RIVAS Havana, Cuba \ r2 E:T out 40) fea , wet ee ee > c ee Ss eR a ae A A oe, oe re ee — IPO ee PO ee. Pe Pe = PME ye eed Neg. FS ep esas FS pees Re sy Pes otteger te eee we on Se zh Ce te Be a) Mad ri Pag Sad SE SG Nas et ae et ee . NS ea eRe Oe TS : Ses ae TA sl ree Site are 7 7 SETS ———— SES ST rte MeO! Es - . Pe ee ee ee eer bser see Sl ™ Pia ah ye ed “Le Te 7 . = - “=, = Star a Ta ie aig 2 eee Book ra ji i Deeaat . : c : (PUBLICATION 3859) { | CITY OF WASHINGTON PUBLISHED BY ‘THE SMITHSONIAN INSTITUTION NOVEMBER 22, 1946 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 14 A NEW DUSSUMIERIID FISH OF THE GENUS JENKINSIA FROM BERMUDA (WitH ONE PLATE) BY LUIS RENE RIVAS Havana, Cuba (PUBLICATION 3859) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION NOVEMBER 22, 1946 The Lord Baltimore Dress BALTIMORE, MD., U. 8. A. “wy ge js roi e Re oe i ¥ t vy . ; Me % rt fit A ‘ . \ i ; 4 ‘/* é ve n ee ‘ a , 7 - v A NEW DUSSUMIERIID FISH OF THE GENUS JENKINSIA FROM BERMUDA By LUIS RENE RIVAS Havana, Cuba (WitTH ONE PLATE) Recently, while studying dussumieriid fishes of the genus Jenkinsia from the West Indies and the Bahamas, in the United States National Museum (U.S.N.M.), it became necessary to examine also specimens from the Florida Keys, the Bermudas, and the Caribbean area, for comparative purposes. The specimens from Florida, the West Indies, the Bahamas, and the Caribbean area are all J. lamprotaenia (Gosse, 1851, p. 291, pl. 1, fig. 2). Those from Bermuda proved to be a dif- ferent species which is described as new in the present paper. The nominal species J. stolifera, described from Key West, Fla., by Jordan and Gilbert (1885, p. 25) as Dussumieria stolifera, has been shown to be a synonym of Jenkinsia lamprotaenia by Beebe and Tee-Van (1928, p. 44), Parr (1930, p. 3), and Hildebrand (im Longley and Hildebrand, 1941, p. 12). I am grateful to Dr. Leonard P. Schultz for allowing me to report on this interesting new species, and to Dr. Samuel F. ‘Hildebrand for reading the manuscript critically. The photograph reproduced in this paper was made by Gurney I. Hightower, photographer, United States National Museum. Measurements are expressed as the number of times a given part is contained either in the standard length, measured from the tip of the snout (anterior tip of upper lip) to the caudal base, or in the length of the head, from the tip. of the snout to the extreme margin of the opercular membrane. Owing to the small size of the specimens, it was found desirable to make-all measurements under magnifi- cation. The last ray, in the dorsal and anal fins, although double, was counted as a single ray. The transverse scale rows were counted from the upper end of the opercular margin to the caudal base; the longi- tudinal. rows were counted between the origins of the dorsal and pelvic fins. Throughout the description, the proportions and counts of the holotype are given first, followed in parentheses by those of the paratypes. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 14 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Genus JENKINSIA Jordan and Evermann Jenkinsia JORDAN and EverMANN, 1896, p. 418 (original diagnosis ; genotype: Dussumieria stolifera Jordan and Gilbert). JENKINSIA BERMUDANA, new species Figure 1 Green fry, dwarf herring Jenkinsia stolifera (not Dussumieria stolifera Jordan and Gilbert) Fow er, 1930, p. 145 (material; Bermuda). Jenkinsia lamprotaenia (not Clupea lamprotaenia Gosse) BEEBE and TEE-VAN, 1933, Pp. 37 (common names; field characters; diagnosis; distribution in part: Bermuda only), fig. (drawing). Hollister 1936, p. 276 (caudal skel- eton), figs. 40-44 (drawings of tail). The holotype is an adult specimen 45 mm. in standard length (U.S.N.M. No. 100546), collected by the United States Coast and Fic. 1.—Jenkinsia bermudana, new species. From the holotype, 45 mm. in standard length, U.S.N.M. No. 100546, collected off Bermuda. (Drawn by Mrs. Aime M. Awl.) Geodetic Survey Steamer Bache (Grampus), off Bermuda (Sta. 10178; see Bigelow, 1917), on February 17, 1914. Fourteen paratype specimens 33 to 48 mm. in standard length (U.S.N.M. No. 124329) were collected with the holotype. Predorsal contour convex to occiput, the rest almost straight. Body slender, its greatest depth, 5.3 (5.3 to 6.4) in standard length, 1.4 (1.4 to 1.8, usually 1.5 or 1.6) in head, 1.3 (1.2 to 1.7, usually 1.3 to 1.6) in distance between origins. of pelvic and anal fins. Head, 3.7 (3.6 to 3.9, usually 3.7 or 3.8) in standard length; its greatest width, 2.2 (2.2 to 2.4, usually 2.3) in its length. Least depth of caudal peduncle, less than eye, 3.5 (3.3 to 3.7, usually 3.4 to 3.6) in head. Eye, 2.9 (2.9 to 3.2, usually, 3.0 or 3.1) im head,.2.7 (24 taj2i9@ NO. I4 A NEW FISH FROM BERMUDA—RIVAS 3 usually 2.5 to 2.8) in distance between origins of pelvic and anal fins. Snout less than eye, 3.4 (3.3 to 3.6) in head. Interorbital, 4.6 (4.3 to 4.7) in head. Maxillary, 2.5 (2.4 to 2.6, usually 2.5) in head, not quite reaching to anterior margin of pupil. Lower jaw somewhat projecting beyond upper. Dorsal rays, 12 (11 to 13, usually 12) ; anal, 14 (14 or 15, usually 14); pectoral, 13 (13 or 14); pelvic, 8. Origin of dorsal fin nearer tip of snout than caudal base, about midway between origin of anal and posterior margin of orbit. Origin of anal fin about midway between origin of pelvic and caudal base. Origin of pelvic fin below fifth or sixth ray of dorsal; nearer anterior margin of orbit than caudal base. Distance between origin of pelvic and tip of mandible, 1.8 (1.8 or 1.9, usually 1.8) in standard length. Distance between origin of pelvic fin and caudal base, 1.2 (1.1 or 1.2) in distance between origin of pelvic and tip of mandible. Distance between origins of pelvic and anal fins, 2.4 (2.2 to 2.5, usually 2.3 or 2.4) in distance between origin of pelvic and tip of mandible. Gill rakers, 29 (27 to 30, usually 28 or 29) on lower limb of first arch. Scales nearly all lost, in about 40 transverse rows, and 6 longi- tudinal rows. General coloration (in 70 percent alcohol) yellowish brown. A lateral bright silvery streak narrower than eye from shoulder girdle to caudal base. A similar, much narrower streak along the midventral line from the base of the pelvic fins to the anus. A middorsal double series of chromatophores from the occiput to the caudal origin. Belly silvery white. Opercular plates silvery. A dark brown blotch on top of head behind eyes. Iris silvery. Fins colorless. This species differs from Jenkinsia lamprotaenia, the only other known member of the genus, in the number of gill rakers and the position of the pelvic fins, as shown in the following key. Both species are very closely related and are exactly alike in external appearance and coloration. A fairly good description of J. lamprotaenia, including references and a figure, has been given by Fowler (1944, p. 123). Gill rakers in moderate number, 19 to 24, usually 20 to 23 on lower limb of first arch. Origin of pelvic fin midway between anterior margin of orbit and caudal base, or nearer caudal base. Florida Keys, Bahamas, West Indies, and Caribbean; Seatm since ee ie eee es Poe ee Jenkinsia lamprotaenia Gill rakers more numerous, 27 to 30, usually 28 or 29 on lower limb of first arch. Origin of pelvic fin Nearer anterior margin of orbit than caudal base. IB eet Lats rho Ba A cinerea a sith contain Es Game Ce eee eee Tenkinsia bermudana 4 SMITHSONIAN ‘MISCELLANEOUS COLLECTIONS VOL. 106 LITERATURE CITED Breese, W., and TEE-VAN, J. 1928. The fishes of Port-au-Prince Bay, Haiti. Zoologica, vol. 10, No. 1, pp. 1-279, figs. 1933. Field book of the shore fishes of Bermuda, pp. i-xiv, 1-337, 343 ills. G. P. Putnam’s Sons, New York. Bicetow, H. B. 1917. Explorations of the United States Coast and Geodetic Survey Steamer “Bache” in the Western Atlantic, January-March, 1914, under the direction of the United States Bureau of Fisheries-Oceanography. Rep. U.S. Comm. Fish., 1915, app. 5, pp. I-62, map. Fow ter, H. W. 1930. Notes on Tropical American fishes. Proc. Biol. Soc. Washington, vol. 43, pp. 145-148. 1944. The fishes. Jn Results of the fifth George Vanderbilt Expedition (1941) (Bahamas, Caribbean Sea, Panama, Galapagos Archipelago and Mexican Pacific Islands). Acad. Nat. Sci. Philadelphia Monogr. No. 6, pp. 57-520, figs. 1-268, pls. 1-20. Gosse, P. H. 1851. A naturalist’s sojourn in Jamaica, pp. i-xxiv, 1-508, pls. 1-7. London. Ho ttistTeEr, G. 1936. Caudal skeleton of Bermuda shallow water fishes. I. Order Tsospon- dyli: Elopidae, Megalopidae, Albulidae, Clupeidae, Dussumieriidae, Engraulidae. Zoologica, vol. 21, pt. 4, pp. 257-290, figs. I-53. Jorpan, D. S., and EvermMann, B. W. 1896. The fishes of North:and Middle America. U.S. Nat. Mus. Bull. 50, pt. I, pp. i-lx, I-1240. Jorpan, D. S., and Gipert, C. H. 1885. Descriptions of ten new species of fishes from Key West, Florida. Proc. U. S. Nat Mus. vol. 7, pp. 24-32. LoncLey, W. H., and Hi_pEepranp, S. F. 1941. Systematic catalogue of the fishes of Tortugas, Florida, with ob- servations on color, habits, and local distribution. Pap. Tortugas Lab., vol. 34, Carnegie Inst. Washington Publ. No. 535, pp. i-xili, 1-331, pls. 1-34. Parr, A. E. 1930. Teleostean shore and shallow-water fishes from the Bahamas and Turks Island. Bull. Bingham Ocean. Coll., vol. 3, art. 4, pp. 1-148, figs. 1-38. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 14, PL. JENKINSIA BERMUDANA, NEW SPECIES A, holotype, 45 mm. in standard length, U.S.N.M. No. 100546; B-D, paratypes, 44, 37, and 33.5 mm. in standard length” U.S.N.M. No. 124320. i wate Pan "SMITHSONIAN MISCELLANEOUS COLLECTIONS Ale gong VOLUME 106, NUMBER 15 * ih "LADYBEETLES OF THE GENUS : _ EPILACHNA (SENS. LAT.) MIN ASIA ~ EUROPE, "AND AUSTRALIA “ow ITH 27 | Puares) tBY _G. H. DIEKE - (PupLication 3860) - Pe} cIry OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JANUARY 20, 1947 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 15 Thomas Lincoln Casey Fund Peo BEETEES;OF THE GENUS EPILACHNA (SENS. LAT.) IN ASIA, EUROPE, AND AUSTRALIA (WiTH 27 PLATES) BY G. H. DIEKE (PUBLICATION 3860) y CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JANUARY 20, 1947 The Lord Baltimore Press BALTIMORE, MD., U. S ds CONTENTS PAGE LIRMLR OCG REVCSIONRL «AS Staab AD OREROC OCR DIGRS: Gishcit oka & hick AE ond M1, Sarr ae a ot I The genus Epilachna and the subfamily Epilachninae....................- 5 Romparisonvor Le pulachhaand ASS: ooo, sks Rw c ess etc e sass coeueews 9 SOvriRaS “GE Ves Bako nig Pyne eae oss ped SUN ok MRE Sta oR rae te 21 RetdALK SM Olle the eho eS iyee. cc petscroialaciaeheie hale secicis oasis atone eee 21 ease oMacnne Chevrolat. o..co% 2% Wot’ We dededicjemie dnt aus sleins sok 22 EMOTO CLS LEG Cm OLOUTN Meet penne ee PE OO Te ioe re Sere 100 ae PESEASEE OTOUBT Ser eee ets vs tit eset ae) echt eee Cee was 104 EANACMLD MGW RE CLIUUISITaR: hom. events ais te See OG Mrs oS oc pee ae ene oeate 109 PIR PAUENG ETIMG OM geht Sate ea wis amie kes 24 Slaves ble 6 = OA Aleta tapes ok 113 NA Peniy eb ILtS SOUP) iicscictssapsiess ides AOE See ee eye Tol ee Cowan Cee eee s IIS MOE ioe PLO aa Ct Os hel onic tei mika 6 555s cl ta cgattersegingleie’? 124 MEO 2 IGO LES OT OUP a crohs a cscicvene oie = Ores SIRPEA ENG ol Seba Cro Pee es era Ere 133 IRR CO NUPIIGUIA VST OUD erates acct cn ciae cee cele one sieisiciove) over nye eicie Gattis quae Conte 157 NSEC HUAN CNET OUP Ne: ccs Aocraes Haier eens Mae teisen hocks sewers ois cots 163 NOD US EGU). 4 roan ts tes titre aE et hs Lads oth se ees OS Ro Ske 166 ERO UTICHLCE MAVENS CNIUIS 6 cleh « or ajshp ciey s svlagenersuaysie ous Gus ean cho si shavers stores aiels cea. Sis 169 Subcoccinella vigintiquattworpunctata (Linnaeus)..................--...-- 170 GUIEUE LESLIE PUNG LET 9 MUITINIAGCUIS)) co perso erecn tia 5G © aeseta «eres arr a sfoyai iss ae 171 List of all known Epilachninae : CINVTEN 3B cate BIS ORR Ceo ee SHES. S.A RE Se eRe aon ene ce ee Cem rh ar 172 Resend Mra e dae: lis ea tAp ra ecu hS ais. score bs b,ro) ssa a Bharode aus Gh © © a'e vermis sheete 8 imply Maes 174 Uh nee RR eae ei eae ONE coe Acne See. 5 ene | See 175 TAG CNM Se sea ccc ays Ena) ay SUA on oe CoE al Steals soy ERMA ete tees, aie 181 iii chy ae ; ales BT ey ae 4: yng A BP plates CDA Mee OA gary wea ae ay Aes) chen an ot ee LY, a at io aTr Alvin Un Naas Ant fe Rat: ; 2 Hits poe yn erick path stl ch banat, ayreelnataga o ; 2k eg a ie ah Cope GS a Wit ighaart ETE ae Ae tee telod yt Rabbis: es) rh Ban hi oy Orr Ah tatty ; hh Beary et alain dab annnaty ‘1’ ay: et A" , ‘ = ay 7 che ‘ KAA : Fiirecmelaui toh iv ae he “uae Bea Er A . Al Ae aalied lee ae? oot, eae ee | , ae (iota 7h eke Ser Hh Oh GG eye LD greens aia p41 Miers RAE Dad ee ME ies | aly ; : ’ seal cl as he iL ayy | Try arene at i i a Opt P My hs rary eh hand 1448) hae Sd CyB ¥ - ae , romnityalhgt avpel At ne , ie ita ye , Fj , mn rage ee ce | As ays Lae is a aye | Rs na abides ‘ i Thomas Lincoln Casey Fund LADY BEETLES OF THE. GENUS .EPILACHNA. (SENS, LAT).IN ASIA, EUROPE, AND AUSTRALIA? By G. H. DIEKE (WITH 27 PLATEs) INTRODUCTION About one-sixth of all the described species of the Coccinellidae belong to the subfamily Epilachninae, and almost all these to the single genus Epilachna. The members of this subfamily are readily recognized by certain morphological characteristics and are distin- guished from the rest of the Coccinellidae not only by these structural differences but also by marked differences in their food preferences. The true coccinellids are carnivorous, feeding on other insects; the epilachnines and psylloborines, on the contrary, are phytophagous, the former feeding on leaf tissue, the latter on fungus spores. The species of Epilachna feed almost exclusively on leaves of plant species belonging to Solanaceae and Cucurbitaceae, with a few species attack- ing plants of other families, particularly the Fabaceae and the Com- positae. As many of our important cultivated plants, such as potato, tomato, squash, and bean, are members of these plant families and are at times subjected to devastating attacks by species of Epilachna, the genus may be considered as economically one of the most im- portant among the beetles. Reports from China have told of the destruction of whole potato crops by Epilachna beetles (Reh, 1913). In Australia they are among the worst enemies of tomato, eggplant, and squash, and similar con- ditions prevail wherever these beetles are found. The example of Epilachna varivestis Mulsant, the Mexican bean beetle, in the United States has shown that the invasion of some species into new territory may be expected with devastating effects. A thorough knowledge of the genus is therefore of great practical importance. At the time (1837) when Chevrolat split off the genus Epilachna from the by that time unwieldy number of species of the original 1 This is the sixth contribution to be published by the Smithsonian Institu- tion under the Thomas Lincoln Casey Fund. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 15 zZ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 genus Coccinella, only isolated descriptions of species of this genus had appeared. Redtenbacher (1843) apparently was the first to call attention to the fact that the species of this genus are phytophagous in contrast to the majority of the other coccinellids that feed on plant lice. He also was the first to give a description in words of the morphological characteristics of the genus. Mulsant, in the big mono- graph “Species des Coléopteres Triméres Sécuripalpes” (1850), de- scribed a large number of new species and brought the known number from 37 to 152. He was also the first to try a systematic grouping of the species of the whole world. The only attempt to do this after Mulsant was made by Crotch in his revision of the Coccinellidae (1874), when the number of species had grown to 200. Whereas Mulsant’s monograph suffers from lengthy descriptions and the repe- tition of not very important details, Crotch’s work, on the other hand, is rather sketchy. After Crotch’s revision no further account of the genus as a whole has appeared—only descriptions of new species scattered widely over the literature. Korschefsky’s Catalog (1931) lists 457 species of Epilachna, and a few have been described since. Mulsant and Crotch’s descriptions and identifications of the species, and even more the earlier ones, are almost completely based on super- ficial characters like color and spot patterns. It has turned out since that in many genera of the Coccinellidae coloration is not a safe cri- terion for the fixation of a species, as it may vary widely in one species or be almost the same in two different species that are not even closely related. Nevertheless, most of the subsequent descriptions were chiefly based on coloration, probably partly because morphological characters that can be observed easily tend to be very uniform throughout the genus. The situation that confronts the student of the genus is therefore very unsatisfactory. In order to identify a species he must look through descriptions scattered through a large number of periodicals, many of them very difficult of access to the average entomologist. ven when he succeeds in collecting all the descriptions, and after he has taken the trouble of reading each one, he finds that in many cases it is totally impossible to identify the species from the descrip- tion because the latter may fit a number of allied species equally well, or because it may not fit the particular specimens at all, as they may deviate in unessential details of coloration from the type. I found this particularly true when, on Dr. Chapin’s suggestion, I tried to identify the extensive material from the Philippines in the NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE (os) United States National Museum. A thorough revision of the genus seems to be the only satisfactory remedy. The present paper is the first contribution to such a revision.? It deals with the species from Europe, Asia, the Indomalayan Archipelago, Australia, and the Pa- cific Islands. The material from the Philippines, which contains the Baker collection, is probably more complete than anything from this locality ever gathered together before. The material from China is also rather extensive, with long series of some species. There is considerable material from Java, the South Pacific Islands, and British India, and a more scattered representation from the rest of the region. Although a complete revision of the genus must wait until a de- tailed study of the African and American species will have been accomplished, the present paper hopes to make it possible that in future the species with which it deals may be identified without much uncertainty. That is very desirable because of the great eco- nomic importance of several species and because the inability to rec- ognize them has caused some confusion in the accounts of their biology. Unfortunately, even for the restricted region with which we are dealing, some of the species had to be left out, as I was unable to obtain them, and I have included in this paper only material on the morphology and taxonomy I have been able to check by personal ebservation. The analysis had to proceed almost exclusively along morpho- logical lines, as information on the geographic distribution and the biology of the genus is in general yet too scanty to enable one to treat the species as biological entities rather than to define them by morpho- logical characters. Such a morphological analysis, however, is the necessary prerequisite for all further studies. As many of the older descriptions, perhaps most of them, are not sufficient to enable one * Although after Crotch’s Revision no further general study of the genus Epilachna has appeared, there are a number of monographs of the faunas of certain more restricted regions that have included the genus Epilachna. How- ever, those of Europe or North America are of little help because of the very restricted number of species occurring in those regions. The home of the genus is definitely in the Tropics, and only feeble branches of it extend into the more temperate regions. Sicard (1907) gave a synopsis of the species of Madagascar, Fauvel (1903) one of New Caledonia. Mader’s “Die Evidenz der palaeark- tischen Coccinelliden” (1926) contains 29 species. However, in most cases he gives only a copy of the original description and in all cases restricts himself chiefly to coloration and maculation, which, as mentioned before, are not suffi- cient for a definition and the recognition of the various species. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 to recognize the species beyond any reasonable doubt, I have en- deavored to give new descriptions embodying such characters by which the species may be recognized without question in the future. As the type of none of these species is available at the present time, the names I have attributed to some of them may be in error. If that should prove to be the case, no great harm is done, as the error can easily be rectified if the types can be compared with the new description. As in so many other genera of the Coccinellidae, the male geni- talia have proved a great help in the study of the genus, and without them no disentanglement of some of the more closely allied species would have been possible. Very little systematic use has previously been made of the genitalia for taxonomic purposes in Epilachna. Weise (1900) described those of a few species, and even though he omitted figures, it is easy to recognize definitely the species he had before him. In later descriptions he unfortunately abandoned the use of the genitalia and only occasionally does he or other authors refer to their structure. The female genitalia also proved very valuable. I should like to express my appreciation to Dr. E. A. Chapin, curator of insects of the United States National Museum, not only for suggesting this study and making available to me the complete material on Epilachna in the National Museum, but also for con- tinued advice and helpful suggestions. My thanks are also due to H. S. Barber, of the National Museum, for helpful discussions ; and to E. T. Cresson, Jr., of the Academy of Natural Sciences of Phila- delphia, Dr. M. A. Cazier, of the American Museum of Natural His- tory, and Dr. E. C. Zimmermann, of the Bernice P. Bishop Museum, for valuable help through the loan of material from the collections in their charge. To Prof. Th. Dobzhansky I am greatly indebted for a number of specimens and some interesting discussions. After this paper had been completed I had the opportunity to study a number of additional specimens. These came partly from the Museum of Comparative Zoology and were kindly put at my dis- posal by Prof. Nathan Banks, partly from entomologists attached to the Armed Forces in India and the Pacific and made available to me through the cooperation of Dr. Chapin. The material contained a number of new or hitherto unavailable species which could .be added at the appropriate places without upsetting too much the original scope of the paper. The male genitalia of the added species, where available, are illustrated on plates 26 and 27. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 5 THE GENUS EPILACHNA AND THE SUBFAMILY EPILACHNINAE The genus Epilachna was first proposed in 1837 by Chevrolat in the third edition of the Dejean catalog of Coleoptera.* He did not give a description but only a list of species which he included in the new genus. From this its scope was perfectly defined. Hope in 1840 indicated Epilachna borealis (Fabricius) as the typical species of the genus. A few years later Redtenbacher (1843) first gave a des- cription of the characters of Epilachna in words which agreed in its main aspects with those derived from the species in Chevrolat’s list.* Nothing changed in this status except the addition of a great number of additional species until Weise (1898a) did two things. 3 Some editions bear the year 1836 on the title page. According to Barber and Bridwell (1940), the last part of the second edition which began to appear in 1833 was printed from the same type as the third edition but at a later date. This Mr. Barber concluded from that fact that though both editions were printed from the same type, in many cases the letters in the second edition were damaged while they were undamaged in the third. The second edition was destroyed by fire, and the last part, which contains the Coccinellidae, was added to the second edition after the third was printed. Chevrolat’s list contained the following valid species from the Eurasian region: chrysomelina Fabricius I1-maculata Fabricius (argus Geoffroy) 28-punctata Fabricius 26-punctata Boisduval (Dejean) signatipennis Boisduval (d’Urville) obsoleta Olivier flavicollis Olivier haemorrhoa Boisduval (d’Urville) besides a number of American (among them borealis Fabricius) and African species. 4 Redtenbacher’s description of the genus Epilachna translated from the Latin is: Mardibles multidentate; the teeth laterally serrate; the ligula conical with obtuse apex. Claws of the tarsi bifid, armed with a broad sharp tooth. Body with wings. As the only species of the genus (he restricted himself to the fauna of Austria), he listed Epilachna globosa, now called Subcoccinella 24-punctata Linnaeus, which does not fit the description of the genus at all. If we would accept Redtenbacher as the author of the genus Epilachna, according to the international code (article 30c), we would have to accept Subcoccinella 24- punctata as type of the genus, which would make it impossible to leave any of the other species in Epilachna. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 First he concluded that the genus Epilachna should be attributed to Redtenbacher, as he was the first to give its description. He sug- gested therefore that everything done preceding Redtenbacher, which means Chevrolat’s and Hope’s contributions, should be disregarded. Following this he proceeded to subdivide the genus into two parts. Those species that have their tarsal claws provided with a basal tooth he continued to call Epilachna, whereas: for those species without such a basal tooth he proposed the new genus Solanophila. In this he was followed by many workers, whereas others rejected Solano- phila as a separate genus because it was not always clearly separated from Epilachna sensu stricto. It seems now that quite apart from the question whether the genus Epilachna should have been split up into two or more separate genera, Weise’s procedure was unjustified in the light of the now generally accepted international procedure of nomenclature. Weise’s idea probably was that for the establishment of the genus, as well as of a species, the description is of primary importance. An illustration of this attitude is contained in the following passages, translated from Reitter (1908, p. 27) referring to description and types of a species. Since priority of a species can be obtained only by the accomplished de- scription of it, for the study of a species only the description should be perti- nent, for it is lasting but the type perishable and not recognizable as such under all circumstances ... Types which do not agree with the description must not be considered as such even if they originate from the author . . . The statements of the description must be taken into account in the first place, and the requisition of the type should only have the purpose of ascertaining the correctness of the statements in the description—that always has been the main purpose—and to examine on the type those properties about which the author made no statements, because at the time when he made the description the importance of such special data had not yet been recognized. With this attitude, which very likely was also that of Weise, sooner or later serious difficulties are encountered, of which the case of Epilachna is a typical example. For Redtenbacher’s description of the genus agrees neither with Chevrolat’s conception of it, as it obviously was meant to do, as species like borealis Fabricius and flavicollis Thunberg would be left out, nor with the only species that Redtenbacher himself cited. To avoid confusion resulting from such a state of affairs, the International Rules of Zoological Nomenclature were set up, which make the type (more particularly the holotype) the primary thing to which the name of a species is fixed, and simi- larly the genotype fixes permanently the name of a genus. For the NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 54 selection of the name of a genus and its type, the following articles of the International Rules apply: ARTICLE 21.—The author of a scientific name is that person who first publishes the name in connection with an indication, a definition, or a description, unless it is clear from the contents of the publication that some other person is respon- sible for said name and its indication, definition or description. This unquestionably makes Chevrolat the author of the genus, as he was the first to publish, and through the list containing valid species belonging to the genus, gave an indication which served in fact better to characterize the genus than Redtenbacher’s later description, and all authors until Weise (1898) and many since have accepted the genus in Chevrolat’s sense and not in that of Redtenbacher. For the selection of the genotype, in the case that the original author does not indicate a type species, the rules are: ARTICLE 29g.—If an author, in publishing a genus with more than one valid species fails to designate (see a) or to indicate (see b, d) its type, any sub- sequent author may select the type, and such designation is not subject to change. (Type by subsequent designation.) Hope (1840) explicitly designated Epilachna borealis (Fabricius ) as the typical species of the genus. As Hope fulfilled all other re- quirements of the code, his designation is perfectly valid and binding. When a valid genus is divided into two or more genera, the Inter- national Rules provide: ARTICLE 29.—If a genus is divided into two or more restricted genera, its valid name must be retained for one of the restricted genera. If a type was originally established for said genus, the generic name is retained for the re- stricted genus containing said type. When Weise in 18098 divided the old genus Epilachna into two genera, he retained the name Epilachna for those species that do have a basal tooth on their tarsal claws and gave the new name Solanophila to the species without such a tooth. The division of the genus was made solely on the basis of the presence or absence of this basal tooth. However, as borealis Fabricius, the type of Epi- lachna, has toothless claws, article 29 makes it mandatory to leave the name Epilachna with the restricted genus containing the type that Weise had, however, called Solanophila. Therefore, Solanophila Weise and Epilachna Chevrolat are identical; in other words, Solanophila must be regarded as a synonym of Epilachna and cannot be used for any other genus. So far*this touches only the question of whether Solanophila is a 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 valid name, not whether the separation of the old genus Epilachna into two or more genera is justifiable. The subdivision of such a bulky genus with almost 500 species is certainly desirable. The ques- tion of whether it can be carried out without ambiguity can, of course, be decided only after an examination of the species of the whole world. An examination of the species of Europe, Asia, and Australia indicates a natural separation into two well-characterized genera with approximately the same boundaries in this restricted region as Weise proposed for his two genera Epilachna and Solanophila. Besides the presence and absence of the tooth on the tarsal claws on which Weise based his separation exclusively, the species with toothed claws (with rare exceptions relegated to the genus Afidenta) have the sixth visible abdominal segment of the female divided longitudinally, while the species with toothless claws show usually no sign of such a separa- tion. If the biology of Epilachna admirabilis Crotch is typical for the rest of the toothless part, there is also an important difference in life habits, as admirabilis hibernates as larva, whereas as far as is known the toothed species hibernate as adults. If, also, the American species are taken into consideration, the matter is less simple, and an inclusion of the African species may complicate matters even more. The American species E. borealis (Fabricius), the genotype of Epilachna, has toothless claws but has the sixth abdominal segment of the female divided. The question of which of the Eurasian subdivisions should retain the name E#7- lachna rests therefore on whether the structure of the claws or the structure of the female sixth segment is considered more important to make it the key character of the two genera. In order to change as little as possible in the present nomenclature, I shall adopt the division of the sixth segment of the female as the distinguishing character, which makes the toothed species congeneric with the type borealis, so that they will retain the name Epilachna. For the Eura- sian species with undivided sixth female segment and toothless claws (equivalent to Weise’s Solanophila), I propose the new name Afissa.® 5It is quite likely that a closer examination of the American and African species will make it necessary to subdivide into more than two genera or that no subdivision at all should be carried out. In that case Afissa may at least be retained as a subgenus. Epilachna varivestis Mulsant would belong to Afissa. There are also South American Epilachna with toothed claws like the Asiatic ones. * NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 9 The genera of the subfamily Epilachninae in Europe, Asia, and Australia may now be characterized as follows: GENERA OF EPILACHNINAE (WITH GENOTYPES) ee |) 2 5 | 8| & s ro] ~ 3 Q g = ~ r= > 3 8 ao] Peesolee nell ees z os 3 ret i hema el Re aba 8 2 a 8 8 @ = A} Character Ss ar herarte Sarl hee eres a) 8] 82 SUSE] 8) a] sf] se 3 s 8 3 2 eet S 5 ss| 23 2 1838/22/58] .5/ 33 =. S38 88 | se] ese] S81ss] cs ze ES FAS SN] 8s =S xs $s < <3) x 7 |O |g |Q |S Sixth abdominal segment of female (s-split, BrETICACE) clot econ cic ta siete Sake eyecare Sins e s e e e e ry Tarsal claws (s-single, d-double, ¢-with tooth, u-without tooth)............... d-t | d-t* d-u d-u | s-t | d-u | s-u | d-u Epipleurae (h-horizontal, v-vertical, u- without, g-with grooves).............. h-u | h-u h-u, g | h-u | h-g | v-u | ? ? Punctation of elytra (s-simple, m-mixed Aneand Coarseyese CIS ee od Le m m m Ss Ss m ? m Distribution (E-Europe, A-Asiatic Conti- nent, P-Polynesia, A u-Australia, J-Indo- EAI malayan Archipelago, Ch-China)....... A P. re AL E E Gholi Ghat * Rarely without tooth (haematomelas, malkini). COMPARISON OF EPILACHNA AND AFISSA Before giving the description of the individual species, it seems useful to summarize briefly those structural characters most useful for classification and identification. They are naturally those that vary from species to species but stay constant within one species, and these are chiefly emphasized in the following synopsis. It is based entirely on an analysis of Asiatic, Australian, and European species and there- fore does not represent the properties of the complete genera for which an analysis similar to that carried out in this paper would be required also for the American and African species, which has not yet been completed. The two genera are taken together because they have many features in common and therefore unnecessary duplication can be avoided. We have thus essentially the old genus Epilachna as understood before Weise divided it in 1898. This synopsis also serves to define the use of the various terms. The properties of the other genera of the Epilachninae (characterized in the preceding table), which contain each only one or two species, can then most IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 conveniently be summarized by stating in which points they differ from Epilachna. Afissa is separated from FE pilachna by the absence of the basal tooth on the tarsal claw and by the fact that the sixth abdominal segment of the female is not split. The genus Epilachna is much more uniform than Afissa, and so much finer structural details must be employed to characterize the species in the former. With reasonable care there is not much difficulty in identifying a given species of Afissa even from a single specimen, whereas in Epilachna that is often entirely impossible from the external appearance alone and often presents difficulties even when the genitalia are available. For that reason the difficulties in identifying the species from the often inadequate original descriptions have been considerable in some groups of Epilachna, and several cases had to be left for future research when the types become available. However, only the names of such species are in doubt. The characters given in the descriptions and figures will per- mit a definite recognition of the species at any time. The shape varies from elongate-oval and rather depressed to hemi- spherical and very convex, with the extremes occurring in Afissa. Whereas the shape is a very useful character for the quick recognition of some species or groups of species, it is less suited for the separation of closely allied species, since it varies gradually. Quantitative mea- surements of the shape have been omitted because dried specimens are often distorted, but the illustrations give an indication of the out- line of the species. The length varies from 5 to 10 mm. in Epilachna and from 3 to 12 mm. in Afissa. Mulsant lays great stress on the way the elytra are rounded at their base and shoulder for his classification of the species of Epilachna. I have found this a very vague character, and the fact that Mulsant himself has placed members of the same species in two different classes shows that it is not of great practical value. Some species have a regularly oval shape with the greatest width in the middle; others are attenuated behind, with the greatest width well before the middle. The abdomen presents some very important identification charac- ters, some of which seem to have been overlooked hitherto. The first ventral segment ° shows the abdominal lines, which differ somewhat ®In taxonomic papers it has been the custom to number the abdominal seg- ments by calling the visible segments Nos. 1 to 5. In the Epilachninae, usually but not always, also the sixth ventral segment is visible in both sexes. If this nomenclature is adopted there is no correspondence with the numbering of the dorsal segments or with the state of affairs in other groups of insects. The NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE Il in the various species. In Epilachna they are usually complete, i.e., they reach in a complete arc from base to base (fig. 5A). They are called subcomplete if their outer part does not quite reach the base. They are usually subterminal, i.e., they reach to within about one-fifth of the apical margin of the first segment. The exact extent of their reaching toward the apex is variable within a species, as what appears as the apical margin of an abdominal segment in a dried specimen apparently depends on the state of contraction of the abdomen when the specimen died, and the treatment of the specimen after death. In some species, however, the plates reach only to the middle of the first segment or a little beyond it. This is mentioned specifically in the de- scription of the species. The shape of the abdominal lines may be a symmetric arc with almost uniform curvature. More often the sides, particularly the outer ones, are nearly straight. Occasionally the plates are angulate, i.e., they present a more or less well-defined angle at or near their apex. In Epilachna strong departures from the normal shape are rare and gradual, and the shape varies somewhat within a species, so that the abdominal lines are not particularly useful for the characterization of the species. In Afissa the variety is much greater and the structure of the abdominal lines may be used often with advantage. The punctation of the first segment in the middle between the abdominal lines usually consists of fine punctures more or less as on the rest of the abdomen. However, in some Afissa species (admirabilis, chapini, etc.) these punctures are rather big circles diminishing in size from base to apex. The fifth and sixth abdominal segments present the most useful characters; they usually differ in the two sexes. On the whole the hind margin of the fifth segment tends to be more concave in the male than in the female. In Epilachna it is truncate to quite pronouncedly concave (enneasticta) in the male, whereas in the female it is most often truncate, occasionally with a process in the middle (chry- first dorsal segment is always that which carries the first pair of stigmas, and it appears that in the Epilachninae the first two true ventral segments are fragmentary and appear as a membrane and the sclerotized edge of the abdomen. The first visible segment is therefore the third true ventral segment, and in morphological papers it is customary to number the segments in this way. In order to conform to both systems of nomenclature in the present paper, when a segment is called the “fifth ventral segment” the numbering is that of the visible segments, whereas when it is called “sternite VII” it is numbered to conform to the true numbering of sternites and tergites. In other words, the first ven- tral segment is the same as sternite III, etc. For the description of exterior parts, the first way of numbering is customary, whereas the second is used for the description of genital segments. I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 somelina, fig. 5 H) which makes it appear biconcave, only very rarely (enneasticta) slightly concave. In Afissa the fifth segment may have a similar structure as in Epilachna but often is convex in both male and female, usually but not always less so in the male. In the convexa group it may be considerably modified in the male, and this may affect even the fourth and third segments. The sixth segment almost always protrudes distinctly from under the fifthein both sexes (exception chapini group of Afissa). It is quite different in character from the preceding five segments. Whereas the first five segments are more or less firmly joined together along their whole width, the sixth segment is loosely joined to the fifth so that it can move freely, and it may be more or less protracted from under the fifth. In Epilachna it is always split longitudinally into two halves in the female; in Afissa it is always without such division and is usually then narrower and more convex than in the male, but may be emarginate as much as to appear bilobed (convexa). The male sixth segment. in Epilachna may have a uniform convex hind margin (fig. 5 A) or may be subtruncate or emarginate (fig. 5B) and in some species has a definite deep notch (fig. 5D). In Afissa again the variety is larger with the greatest modification in the convexa group.” 7In the average museum specimen the structure of the apex of the abdomen is usually obscured by dirt and the terminal pubescence. Often the abdomen is bent so that its tip is obscured by the elytra, and of course in the European style of mounting the abdomen is completely invisible. The most satisfactory way to remedy this is to remove the abdomen and mount it separately on a piece of cardboard. Often the base of the cardboard triangle on which the specimen is mounted will be satisfactory; otherwise a separate triangle or rectangle may be attached to the same pin under the specimen. In some speci- mens the removal of the abdomen can be accomplished without relaxing by inserting the point of a dissecting needle or better a lancet point between abdomen and metasternum. Usually, however, it will be safer to relax the specimen first, which can be done quickly by boiling it in water for a few minutes. The abdomen is then put in relaxing fluid and the dirt can be re- moved gently with a dissecting needle or a small hard brush. The abdomen is then also ready for the removal of the genitalia. For purposes of dissection or for any other purpose, specimens collected in alcohol or put in alcohol or relaxing fluid immediately after death and then dehydrated and degreased before mounting are far superior to those mounted and dried in the conventional way. Specimens prepared in the former manner will be just as good as fresh specimens no matter how many years have elapsed. Specimens that were not degreased sometimes are so much covered with a mixture of grease and dirt that all structural details and even the colors are hidden. Immersing such specimens for a few days in a degreasing fluid such as carbon tetrachloride, chloroform, or benzene, will usually improve them tremendously. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 13 In judging the apex of the sixth male segment, it is important not to confuse it with the apex of the seventh segment, which occasionally slightly protrudes. ; The normal form of the mandibles in Epilachna is as follows (fig. 1 A-C) : The basal tooth present in the other subfamilies of the Coccinellidae is always absent in the Epilachninae. There is an apical tooth and two lateral teeth, the former larger than the latter. The Fic. 1—Mandibles. A, Epilachna niponica; B, E. diffinis; C, E. argus; D, Subcoccinella 24-punc- tata; E, Afissa flavicollis; F, A. dwmerili; G, A. coccinelloides; H, Cynegetis impunctata. apical tooth seen from in front shows that it consists of three parts, with the lower and upper parts less well developed than the middle. In addition, the lateral edge of the mandibles beyond the two lateral teeth is covered with a number of very small teeth (dentules), which also cover the edge of the three main teeth. There does not seem to be a great deal of variation (not every species has been examined). The lateral teeth are sometimes smaller compared to the apical teeth in some species than in others. In chrysomelina the second lateral 2 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 tooth is considerably smaller than the first, while in the other species they are of about equal size. In judging the structure of the mandibles one must keep in ming that individuals which have done much feeding may have some of the details worn off. Therefore it does not seem advisable to use such details as the shape and size of the dentules for diagnostic purposes. In Afissa the general structure of the mandibles is similar to that in E-pilachna, but the individual variations are greater. The dentules usually are less pronounced and often,absent. The two lateral teeth may be of the same size as the apical tooth (flavicollis, fig. 1 E), or they may be smaller (admirabilis). One of the lateral teeth may be missing (dumerili, fig. 1 F) or even both (coccinelloides, fig. 1 G). In the latter case the mandibles resemble superficially those of the subfamily Coccinellinae, although a closer inspection reveals that we are dealing with a modified form of Epilachninae mandibles. (The basal tooth is always absent.) Species which hardly can be told apart in external appearance may show the extremes in mandible structure. In such cases the mandibles are a good diagnostic character. However, they hardly ever can be examined sufficiently without dissection or at least relaxing. It is reasonable to assume that a pronounced dif- ference in mandible structure is caused by a difference in food habits. The genus Afissa would furnish some interesting examples for test- ing this assumption. At present, however, nothing is known about the food habits of such species. Afidenta bisquadripunctata Schonherr has a peculiar mandible structure different from that of any of the other species of the subfamily. The tarsal claws of species of both genera are always bifid. In the genus Epilachna, at least in our restricted region, they are almost always provided with a basal tooth, which is usually triangular (fig. 2A). In chrysomelina it is quadratic (fig. 2B), and in gutta- topustulata it is quadratic and has a small toothlike process on the inner apical corner (fig. 2C). In the species of Afissa the basal tooth is always missing (fig. 2E,F). The inner claw in both genera is wider than the outer one. In the majority of the Afissa species the two inner claws of one tarsus are well separated from each other (fig. 2E). In a number of species, however, they touch each other, and in such cases the inner claw is exceptionally wide (fig. 2 F, G). Cases like this are mentioned specifically in the descriptions. The elytra of the whole subfamily Epilachninae are covered with pubescence of varying length and denseness. In general the pubes- NO. I5 * LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 15 cence is light of color on the light parts of the elytra and black on the dark parts. There are, however, many exceptions from this rule. Cases where the pubescence is light gray all over are not infrequent. Caution must, however, be observed in using this character for sep- arating allied species, as several cases are known .(sparsa, niponica, diffinis) where specimens occur with the uniformly light pubescence, together with those with the normal coloring of the pubescence. The specimens with uniformly light pubescence usually can be recognized «666 vo ¢ Ere? 2—Claws: A, Epilachna diffinis; B, E. chrysomelina; C, E. rhea cane! D, Afidenta mimetica; E, Afissa flavicollis; F, G, A. quadricollis. immediately by the fact that the maculation stands out less pro- nouncedly than in normal specimens. Under the microscope care must be taken that reflections from the hairs do not mask the true ‘color. Viewing the elytra at a glancing angle at fairly high mag- nifications seems to give the best results. The elytra are always covered with double punctation. There are coarse punctures with a larger number of very fine punctures inter- spersed. The denseness of the punctation and the depth of the indi- vidual punctures vary, but this fact is usually not mentioned in the 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 descriptions unless extremes are encountered or variations useful for the separation of closely allied species. Occasionally the coarse or the fine punctures are almost obsolete. The apical angle of the elytra may be rounded or show a definite angle (fig. 3). This is often useful for the separation of species other- wise very similar. In many cases, however, the angle is so indistinct that a decision on whether it is there or not is doubtful. The epipleurae in general show no particular diagnostic characters except in a few species of Afissa (chapini, magna) where they show definite and deep cavities for the reception of the tips of the hind femora. Such cavities occur also in Cynegetis and were used as one of the chief criteria that separated Cynegetis and Epilachna. The color and maculation of the various species are subject to great variations within a species, and at the same time, in some groups, there are often no systematic differences between different Fic. 3.—Tips of elytra. A, angle distinct (sparsa); B, angle rounded (28-punctata). species. As a rule in Epilachna the infraspecific variations are larger than the interspecific ones, whereas in Afissa in general the inter- specific variability seems to be greater. No reliance at all can be put on the color of the under side. It may vary from completely light to completely dark within a species. In the intermediate stages the sides of the metasternum are dark with the rest of the under side light, then the whole metasternum and parts of the abdomen get dark. The under side of the prothorax and the sides of the abdomen usually keep their light color last. The legs and mouth parts behave in the same way as the rest of the under side. The original descriptions of most of the Epilachna and Afissa species were almost exclusively based on the color pattern of the upper side. The fact that this is so variable within a species and so similar in different species has caused a great deal of confusion and presents the chief difficulties to a rational analysis of the two genera. As mentioned before, the situation is much worse in Epilachna than in Afissa. Neither the number of the elytral spots nor their size, NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 17 shape, or position can be regarded as safe characters for the identi- fication of species, and a similar situation prevails with respect to the pronotal pattern. Though it is true that in many cases conclu- sions derived from the spot pattern will obscure rather than clarify the relations between the species, the spot pattern is still one of the most valuable criteria even for the analysis of the genus Epilachna. It may not be possible in many cases to recognize a single specimen by its spot pattern, but the trend and range of the variations are quite characteristic for the species, particularly if one restricts himself to a definite locality. Details of the maculation and its variation will be discussed under the various groups. Male genitalia——As in so many other groups of the Coccinellidae and other beetle families, the male genitalia have proved an excellent criterion for the separation of closely allied species, and without their help the identification of many species and the separation of some of the closely allied groups would be hopeless. In the genus Epi- lachna the majority of the species have very similar male genitalia, and so finer details must often be made use of for their separation. In Afissa there is so much greater variety of forms that usually even a superficial inspection will be sufficient for positive identification. First a brief description of the normal form of the male genitalia in Epilachna will be given, with those details of their structure empha- sized which are particularly useful for diagnostic purposes, and attention will be called to the deviations that may occur from this normal form. Details of this will be found in the figures of the various species and under the various groups of Afissa. The three main parts of the male genitalia are the penis, the paramera, and the sipho (fig. 123). The penis is a slender tube, straight or almost straight for most of its length, with a lengthwise seam along the middle of its under side.*® Near its end the penis is almost always curved up, and the shape of this curvature is often characteristic for the particular 8 The terminology is that of Verhoeff and Dobzhansky. During copulation the genitalia are extended from the tip of the abdomen so that the middle lobe or penis is lowest and the paramera just above it. In the drawings they are oriented as if the paramera would stick out horizontally and the terms “above” and “below” are used in the descriptions with this position in mind. Actually they point more or less downward. When the genitalia are retracted in the abdomen they lie on their side in the left part of the abdomen. The sipho is led through the penis and in the rest position protrudes just a little from the orifice near the end of the penis. In the active position it can be extended. so that it protrudes considerably. 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 species. At the very tip the penis may curve suddenly again so that a small hook is formed, or it may have a straight point. The sipho is led through the penis and emerges through an orifice, the basal edge of which usually is formed by the edges of the seam gradually diverging apart. The apical end of the orifice is usually rounded. The length and shape of the part of the penis beyond the orifice constitute important characters. On the upper side of the penis near its base between the roots of the paramera there is a very thin knifelike ridge called the basal knife edge. It is very prominent in some species, rudimentary or completely absent in others. Its size seems to be somewhat variable within one species. In most species of Epilachna there are erect hairs on the upper side of the penis. The extent and length of this pubescence often presents a good character. Toothlike transverse ridges on the upper side of the penis present in a few species (niponica, diffinis, dentulata) are very useful for the identification of those species. The paramera are two flattened elongate appendages attached at the upper side of the base of the penis. There is not much variety in their shape in the majority of Epilachna species. They have usually about the same length as the penis. When their length is significantly different (e.g., argus), this difference can be used for the identification of the species. In most species the paramera have a short thorn at their apex (called apical thorn). The presence of this thorn may sometimes be obscured by the dense hairs that line the end parts of the paramera. Occasionally these hairs are of exceptional length and help then to characterize the species. The sipho is a long slender tube of considerably smaller diameter than the penis but of greater length. Normally in Epilachna it is curved near its base through an angle of less then 90° and is ap- proximately straight from the basal bend to the apex. In the gutta- topustulata group (fig. 144) it has a basal bend of about 180° and is greatly modified in the enneasticta group (fig. 146). Whenever departures from the normal shape occur they are mentioned specifi- cally in the individual descriptions. The characters most useful for the separation of closely allied species lie at the apex of the sipho. This most often ends in a relatively sharp straight point, just before which, on the side, is‘a small oval orifice through which the semen is ejected during copulation (fig. 118). However, the apex may have a sharp notch (28-punctata, fig. 119) or be emarginate (emarginata, fig. 125) or enlarged (chrysomelina, fig. 145) or otherwise modified. In Afissa there is no such uniformity in the structure of the male NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 19 genitalia and every part may be greatly modified. Details about this will be found in the notes on the various groups of Afissa and the descriptions and figures of the species. Female genitalia..—After Verhoeff (1895) and a few others had described the female genitalia of a few isolated coccinellids, Dobzhan- sky (1924) gave a synopsis of the genital structure of the females of 55 species of Coccinellidae and showed how the variety of struc- ® Because often a positive identification in the Coccinellidae is impossible without an inspection of the genitalia, a few notes on their dissection may be helpful to those unfamiliar with this technique. The technique is essentially that used by workers in the United States National Museum, with a few modifications adapted to the particular situation in Epilachna. First the abdomen is removed as described on p. 12 and, if the whole speci- men was not relaxed before, relaxed in boiling water and transferred to a . watch glass with relaxing fluid under a binocular microscope at about 10 to 20 times magnification or under a hand lens mounted so as to leave both hands free. The only instruments needed besides a pair of light forceps with a sharp point are two dissecting needles, one with a straight point, the other with a lancet-shaped point with as sharp an edge as possible. An iridectomy knife will be better but is rather expensive and can be dispensed with. Male: Make an incision at the left edge of the abdomen and lift the dorsal membranes. The male genitalia are then exposed and can be extracted with the forceps. They should then be cleaned of all nonsclerotized matter with the help of the dissecting needles. The sipho is better pulled out of the penis. The relaxing fluid will dissolve a greasy layer which otherwise might be present. Both parts, as well as the abdomen, are then left to dry and mounted on card- board pieces so as to permit an inspection from all sides. These cardboard pieces are kept on the same pin as the specimen. Female: For the extraction of the female genitalia, relax the abdomen and place it in relaxing fluid as for the male genitalia. Then sever the connection between sternites and tergites VIII and the membranes that hold the genitalia to these segments. The genitalia surrounded by nonsclerotized matter can then be taken out. The soft matter prevents a clear view of the genitalia and must be removed. For this purpose, boil them in a microtest tube in a 9-percent solution of KOH. If the soft parts of the genitalia are to be preserved, about 15 seconds is sufficient, and a few trials will give the best time. If only the sclerotized parts are wanted, a much longer boiling time is permissible, which will destroy all the soft parts. Aiter boiling, place the genitalia for about 10 minutes in a watch glass with water to wash out the KOH. Then transfer successively to 95-percent alcohol (10 minutes), absolute alcohol (5 minutes), and xylol (2 minutes). After this place them on microscope slide in Canada balsam and cover with cover glass. If the inner parts are important, a bath in 7o-percent alcohol between water and 95-percent alcohol may be advisable to avoid distortion. Many prefer to clear the parts in oil of cloves after the 95-percent alcohol and transfer directly to the balsam, thus omitting the absolute alcohol and xylol. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 tures can be used for taxonomic purposes. In a later paper (1926) he described the female genitalia of three*species of Epilachna and figured one. Since then only isolated examples have been recorded and so the structure of the female genitalia of the Epilachninae is still virtually unknown. Dissection of the females of a few species showed soon that the female genitalia in the Epilachninae are ex- tremely useful for the classification of the species and that they supplement admirably the use of the male genitalia. A brief descrip- tion of the structure of the parts most important for us follows. The female genitalia consist of the inner part, usually more or less soft, and the outer parts which are well sclerotized. The former consist of the ovaries, oviducts, vagina, bursa copulatrix, and receptac- ulum seminis. These parts are often in very bad condition in imperfectly preserved specimens, and their real structure cannot easily be ascertained in such specimens. They are therefore of less practical value for the identification of specimens and are left out of consideration in what follows. Figure 189 shows vagina, bursa copu- latrix, and receptaculum seminis of E. deyrollu. The receptaculum is the most sclerotized of the inner parts and can always be recog- nized. It is colorless and seems to show little variation in structure. Many species have the same ball-shaped form of the bursa as shown in figure 189. In others it is more elongate. The outer parts, always heavily sclerotized and therefore. easily obtainable in good condition, are modifications of the abdominal segments IX and X. Tergite IX is divided in two parts, each folded double, and envelops the sides of the sternite IX. It has hardly any distinguishing marks. Tergite X, however, varies its shape considerably from species to species. It is an undivided segment, convex or pointed apically in most species of Epilachna, but with a truncate apical margin in the guttatopustulata group. In Afissa the apical margin may be convex, truncate to deeply emarginate, or folded over. It also differs from species to species by the degree of pig- mentization. It may be a clear membrane with only the sides dark, or its apical margin may be more or less broadly pigmented. Sternite IX is divided in half longitudinally and consists therefore of two plates, called genital plates. The shape of these is very varied and forms the most useful character for the separation of species. The figures give a better indication of the shapes than a description. Particular attention should be drawn to the inner margin of the plates in Epilachna. Near the base there is usually a notch or indentation the shape of which is characteristic for the species and often furnishes a criterion for the separation of closely allied species. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 21 The usefulness of the genitalia for the classification, in contrast to their use as mere aids for identification, is due to the fact that their structure is supposed to have been less subject to external in- fluences than other parts of the body. It is therefore believed a good indication of the natural relationship of the species. A classi- fication based to a large extent on the structure of the genitalia may be considered therefore as being a close approach to the natural classi- fication. In Epilachna and Afissa changes in the structure of the female genitalia can be very effectively used for the definition of natural groups. The changes in the genitalia are usually accompanied by changes in other parts of the body. Figures 181-218 are photographs of typical cases. In judging the pictures it must be kept in mind that the relative position of sternite IX and tergite X is not significant, as these two parts are movable with respect to each other. Secondary sexual characters—The sexes can be told apart most easily by the structure of the abdomen, particularly the fifth and sixth segments. This has been discussed above. In such species of Afissa as convexa or complicata the differences are striking; in others much closer observation is needed to recognize them. As in most insect groups the female in the average is larger than the male. Often dried specimens are found in which the abdomen is bent strongly up so that the last segments disappear completely from view with the abdomen and elytra in place. Such specimens almost invariably are females, as the male genitalia give the abdomen a greater rigidity that prevents the curling up. SOURCE OF MATERIAL The following abbreviations indicate the collections from which the particular specimens were obtained: AMNH American Museum of Natural History, New York City. BBM Bernice P. Bishop Museum, Honolulu, T. H. D Author’s collection. MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Mass. NM United States National Museum, Washington, D. C. PA Academy of Natural Sciences of Philadelphia, Philadelphia, Pa. REMARKS ON THE FIGURES A representation of the spot patterns of beetles as convex as the majority of the Coccinellidae cannot be made without considerable distortion. The figures accompanying this paper were made either 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 from camera-lucida drawings or tracings from photographs with the specimen seen from directly above. The distortion near the margin is then quite pronounced, and round spots may appear as very narrow ellipses. However, care was taken to bring out whether a spot actually reaches the margin or not, even if that could not be seen in a view from directly above. The outline of the pronotum is determined chiefly by its relative inclination to the elytra in the particular specimen. Differences in the outline of the pronotum in the figures do not reflect differences in its actual shape. The scale to which the figures are drawn is indicated as often as necessary. The drawings of all male genitalia and of a considerable portion of the spot patterns were made by J. S. Spurbeck, to whom ac- knowledgment is due for the care which he bestowed on them. Figures 7-116 represent spot patterns; figures 117-180, 219-226, male genitalia; and figures 181-218, female genitalia. , Genus EPILACHNA Chevrolat Genotype: Epilachna borealis (Fabricius). The morphological properties of the genus in general have been brought out sufficiently in the preceding section. Its distribution on all continents centers in the tropical parts, and although some species penetrate into the temperate regions, the representation there is poor. Accordingly, in the Eurasian-Australian region the greatest number of species are found on the southern parts of the Asiatic Continent and the Indomalayan Archipelago. This region is so well isolated from the east, west, and north by oceans, deserts, and mountains that almost all the species occurring there are confined within its limits. Only two species are found in the warmer parts of Europe. One, chrysomelina, a species of African origin, has extended its range as far as eastern Siberia but is ap- parently found only rarely south of the Asiatic mountain barrier. The other species found in Europe (argus) is a Mediterranean species of limited distribution. A few of the species have a very wide distribution. E. sparsa (Herbst) with its subspecies occurs prac- tically in the whole region where climatic conditions are favorable for its development. Other species have a much more limited dis- tribution, though more material is needed before a clear picture of the geographical distribution of the various species can be obtained. This is necessary also before we know in the majority of cases how NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 23 a species is broken up into races or subspecies. The terrain with the many islands and mountain barriers is very favorable for the development of such subspecies, and there are indications that they do occur, but only in the case of sparsa, and perhaps on a more limited scale a few others, is the material sufficient for such an analysis to be attempted successfully. To the northeast the distribution is limited by climatic factors. Only two species reach Japan (sparsa and niponica), and northern Japan and northern China harbor only one species (niponica). The number of species also decreases to the south, probably because most species were unable to cross the water barriers between the chains of islands. The fauna of New Guinea and Australia contains a few ubiquitous species but otherwise is quite distinct from that of the islands farther north. No Epilachna has been observed in New Zealand. Two species, sparsa 26-punctata Boisduval and 28-punctata Fabricius, reach as far east over the Pacific Islands as Samoa. How- ever, it is very likely that these species were introducede by man to these islands from the west. This is supported in the case of 28- punctata by the fact that although it is common in Samoa it seems to be completely absent from the Fiji group and from New Caledonia, and a natural migration from Australia would not have bypassed these islands. No Epilachna species has been recorded from Guam or the neighboring groups or on any island east of Samoa.?° Epilachna boisduvali, which also occurs on Fiji and Samoa, may have reached there in much earlier times, as the races occurring on Fiji (fijiensis Crotch) and Samoa (samoana, new) seem to be different from those occurring farther west. A rational classification of the Epilachna species is not easy because of the great uniformity of practically all characters among the ma- jority of species. A subdivision into a few more or less well-defined groups is feasible, but it does not help a great deal for practical identification, because by far the largest number of species belong to one of the groups. The various groups which are subject to revision when more species become known may be identified as follows: A. 28-punctata group. Elytra normally each with 6 to 14 spots, which may partly coalesce. Pronotum spotless or with a modifica- tion of the normal 7-spotted pattern. Male genitalia with penis an 10] am indebted for information on the Pacific distribution of Epilachna to Dr. E. C. Zimmermann, of the Bishop Museum, who, with his associates, has made extensive collections in the Pacific islands. 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 elongate tube more or less curved up near end. Sipho with a bend of less than go° near base. Female genitalia with tergite X apically convex or pointed. Species Nos. 1-32. B. Miscellaneous. Species that do not fit into any of the other groups are lumped together here. Each probably should be regarded as a separate group. (The position of some of these species is doubtful, as only one sex is known, and the separation from the preceding group is not quite clear.) Species Nos. 33-41. C. Ennaesticta group. Elytra each with six spots, pronotum as in A. Male genitalia with penis broad and short, sipho greatly deformed. Species Nos. 42-44. D. Guttatopustulata group. Species with a dark ground color. Tergite X of female truncate or broadly convex. Species Nos. 45-48. The groups B-D present no difficulties for the separation of their species. The 28-punctata group, however, contains a large number of closely allied species, many of them so similar to one another that even with the help of the genitalia a proper limitation of the species is not always easy. A definite recognition of the species from the original description is often impossible. The name labels attached to them in some of the collections I have seen looked as if they had been distributed at random. Very likely the synonymy of many species will have to be revised ultimately with the help of the types. References to species in this group in the literature must always be regarded with caution, as a great deal of confusion has existed. The spot pattern of pronotum and elytra of this group is very variable within one species, often much more so than the systematic differences between different species. The variations follow always a definite pattern that may be characteristic for the species. For an understanding of the interrelations of the various species a discussion of the spot pattern is very essential. . The species of group C and some of group B follow the same elytral pattern, but the pronotum is usually spotless. The spot pattern both of pronotum and of the elytra of groups A and C can be regarded as derived from the same basic pattern. This may be true also for the other groups, but the connection there is not so obvious. Pronotum.—The basic pronotal pattern is that of fig. 4E with seven separate spots which will be numbered throughout the paper as indicated in the figure. This pattern may be modified either by a decrease in the black pigment or by an increase. In the former case (fig. 4 A-D) some of the spots are missing or are smaller or NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 25 less black, and in the extreme case, which is rather common, the pronotum is entirely spotless. When there is an excess of pigment (fig. 4 G-L) the spots are enlarged and flow together, and in the extreme case the pronotum is entirely black with only a narrow light margin at the front and sides. Various other combinations and intermediate forms not shown in the figures also occur. All observed spot patterns of the species of group A belong clearly in this category. The individual range of variation within one species may be considerable, as for instance in 28-punctata (observed ETE Fic. 4.—Maculation of pronotum of Epilachna species. extremes, fig. 4 A-I) or niponica (fig. 4 H to K). However, the speci- mens from a given locality seem to show much less variation. Care is therefore needed in evaluating the significance of the pronotal spot pattern in the descriptions. It is quite possible that specimens of the same species obtained from localities different from those used for the descriptions may show variations either in the direction of more or less pigment. In a few species (libera, signatipennis) the spots have a tendency to become very hazy. FE. boisduvali seems to have a spot pattern that does not fit into the scheme of figure 4. Elytra—The fundamental elytral spot pattern consists of six dark spots on each elytron arranged as indicated in plate 1, figure 8. The exact position, shape, and size of the spots varies considerably, 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Fic. 5.—Structure of abdomen in Epilachna and Afissa. A, abdominal lines complete, subterminal, sixth segment ¢ convex (E. dentulata); B, ¢ sixth segment emarginate (E. diffinis); C, concave (E. indistincta) ; D, notched (E. guttatopustulata) ; E, abdominal lines rounded, complete (E. chrysomelina); F, terminal, incomplete (Afissa chapini) ; G-I, sixth segment of 2 of Epilachna species (G, 28-punctata; H, chrysomelina; I, pytho). 7 Sa Fic. 6.—Sixth abdominal segment of female. A, Afissa alternans; B, A. grayi. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 27 but none of these spots is ever absent except in rare exceptional cases. They are called therefore persistent spots and numberéd * I to 6 as indicated in figure 7. This fundamental spot pattern may be modified by the addition of one to eight extra spots on each elytron, or by the enlargement and confluence of several spots. In either case the modification proceeds according to definite rules. e The additional spots are called nonpersistent spots because their presence, even in the same species, is very variable. If the full num- ber of spots, 14 on each elytron, is reached, we have again a very stable form of very characteristic appearance (figs. 7, 12, 19, 21-23, etc.). The nonpersistent spots are identified by the letters a to h as indicated in figure 7. Since their position varies only slightly, there is in general no trouble in identifying these spots even if only a few are present. Their size and pigmentation, however, are very variable. In some cases they are much smaller than the persistent spots (fig. 9) or they are definitely less dark. In other cases they have substantially the same size and development as the persistent spots (figs. 21, 22). There are species that so far as known are always 12-spotted, i.e., without nonpersistent spots (boisduvali, argus, diffinis, etc.), and others like 28-punctata which always or nearly always carry the full number 28, but in general the number and development of the nonpersistent spots may vary within one species between the two extremes. The development of the non- persistent spots can therefore never be regarded as a specific charac- ter. Often the variations may be extreme if specimens of the whole range of the species are considered, but individuals from one locality are more or less constant (e.g., sparsa 26-punctata on the Pacific islands). Then the species is broken up into a number of subspecies which differ by the development of the spot pattern. These subspecies may merge gradually one into another, as seems the case for sparsa, or they may be quite distinct without transitional forms. A few cases are known where only the 12-spotted and the 28-spotted forms exist, without any intermediates. Whether in the evolution of the genus the development proceeded from the 12-spotted to the 28-spotted form or vice-versa can at Sresent only be speculated upon. The material suggests strongly another possibility—that there were originally two geographically 11 This manner of numbering is that adopted throughout by Mulsant and followed by Weise, at least in his later papers. Other authors have used dif- ferent ways of numbering the spots. 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 separated species, one with 12, the other with 28 spots, which gradually intermingled and produced the variety of forms by cross- breeding, followed possibly by further evolution. The second way in which the spot pattern is modified in Epilachna is by the enlargement and confluence of spots. The extent to which this occurs is again very variable within one species, but the ten- dency for a certain form of variation may be characteristic for the species. The most frequent variation is probably a lateral enlarge- ment of spots 3 and 4 until they coalesce. This is occasionally ob- served in sparsa on the Indonesian islands (fig. 17). If No. 3 is also joined to No. 5 we have the typical spot pattern of signatipennis Boisduval (figs. 18, 50). Spots 1 and 2 may also coalesce as in libera (figs. 59 and 60). E. libera shows a few of the nonpersistent spots, which usually seem to be absent in those species showing great ten- dencies to coalescence. When this process of coalescence is completed we have a pattern like that of solomonensis (fig. 61), consisting of two transverse fasciae, the basal one due to the coalescence of spots I and 2, the discal one of 4, 3, and 5, and in addition spot 6. In solo- monensis, for instance, the origin of the completed pattern as a modi- fication of the fundamental 12-spotted pattern is little apparent, but in other species all the intermediates occur. This type of pattern with two fasciae is recognizable by the fact that the discal fascia is bent back near the suture so that a V is formed there. The pattern of parafasciata (fig. 62) has also two fasciae with an apical spot, but the discal fascia does not show this V-like bend. The exact relation of this with the 12-spotted pattern is not quite clear. Possibly spot No. 5 moved forward on the suture. Another type of coalescence is illustrated by figures 39 and 40, in which the spots are still separate but show the tendency of variation, and figure 41, where the coalescence is complete. Spot 2 is still quite unaffected. If this is also enlarged along the margin and toward No. 3 we get a pattern like that of delesserti (fig, 58), which appears as five light spots on each elytron on a dark background. Whether a pattern like that of haemorrhoa (figs. 63, 64) is directly related to the 12-spotted pattern is problematical, although in light specimens it is resolved into several dark spots on a light background. The similarity of the genitalia of haemorrhoa and species of grotip A makes this probable. Even more uncertain is the relation of the pattern of the species of the other groups to the 12-spotted ground form even though the form ftricincta of guttatopustulata (fig. 72) is very similar in appearance to solomonensis or parafasciata. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 29 There is a third type of modification found frequently among the coccinellids but rarely among the Oriental species of Epilachna. This is the darkening of the general background of the elytra, often with a lighter ring around each spot. Such ocellate forms are common with chrysomelina, particularly among the African races. It occurs in ocellata apparently as the ground form (fig. 38). On the Fiji Islands such a darkening of part of the elytra occurs without the light rings, so that in the extreme form part of the elytra is completely black (fig. 16). This form apparently occurs, however, only as an aberration. Because of the great similarity between many of the species, a key in the usual form would be of little help. A study of the figures will make a quick orientation possible, and a comparison with the individual descriptions will be necessary for the exact identification. All the descriptions should be used together with the figures, because for those characters (like spot pattern, shape, etc.) expressed in the figures, the descriptions are meant chiefly to call attention to the most important facts and are not to be regarded as an alternate to the figures. 1. EPILACHNA SPARSA (Herbst) Ficures 8-18, 117, 181 Coccinella sparsa Herpst, 1786, p. 160. Abdomen.—Abdominal plates subcomplete, reaching to within one-fifth of the apical margin of the first segment. Fifth segment, male, apical margin truncate or slightly concave; female, truncate. Sixth segment, male, distinctly ** emarginate; female, split. Male genitalia (fig. 117).—Penis seen in profile, under side with a bulge beyond the middle followed by an emargination, after which the penis curves up into an apical hook; upper side with a wide, bladelike vertical ridge beginning at the foot of the paramera (basal knife edge). Second half with two rows of hairs, 0.1 to 0.2 mm. long. Seen from below, closed tube with seam along the middle, orifice elongately diamond-shaped; penis immediately after the orifice less than half the width it had before the orifice, and then almost with parallel sides until it curves up. Paramera about 1.3 mm. long with apical thorn and covered with hairs slightly shorter than those on 12 There are some specimens from Java that have the apical margin of the male sixth abdominal segment ertire, without an emargination. These speci- mens cannot be distinguished in any other way from those showing the emargination. 3 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 penis on the last 0.2 mm. of its length. Sipho gently curved near the base, from then on nearly straight, ending in a point. The orifice oval, on the outside just before the point. Female genitalia—Plates (fig. 181) on the inner margin about one-fourth distance from base, an excavation on the under side with a sharp dark edge toward apex. By the shape of this excavation the female specimens of sparsa can be distinguished from neighboring species. Length.—6 to 7.5 mm. Apical angle of elytra usually distinct; mandibles with trilobed apical and two lateral teeth and many dentules. Color and maculation—Varying between wide limits, both among the specimens found at one locality and between the various geo- graphical races, to be described in detail below. The under side may vary from completely light yellowish red, including legs and appendages of head, to dark brown to black on metasternum and large part of abdomen. The ground color of the upper side is red. The pronotum varies in maculation from the spotless form to the form with all spots well developed and beginning to coalesce (fig. 4 A-E, H). Forms with the pronotum largely black, due to the confluence of the spots such as occur in F. niponica, 28-punctata, and similar species, have not been observed. The elytra varies between the form with 6 spots on each elytron and the other extreme with all 14 spots present. Rarely is one of the persistent spots missing, but practically any number of spots between 6 and 14 may be expected with greatly varying sizes of the individual spots. In some instances the confluence of several spots takes place (figs. 17, 18). The most common forms and the range of variability are very different in the various localities where the species is found, and details will be given below. Type locality.—East Indies. Distribution—More than 500 specimens from a wide range of localities permit a fairly clear idea of the distribution. On the Asiatic Continent it ranges from India to North China, with the exact western limits unknown. Its limitation to the north and east is probably determined by the fact that it needs an average yearly temperature of 15°C. or more for its development. It occurs also in southern and central Japan. In the Indian Archipelago it is known from Sumatra, Java, Lombok, and Amboina, and probably is found on many other islands. In the subspecies 26-punctata, it occurs on | New Guinea and Australia and extends to the Fiji and Samoa Islands, where it reaches its eastern limit. It is not known from the NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 31 Philippines and probably is replaced there by E. philippinensis, which may be a subspecies of sparsa. Because of its confusion with other species, all reports in the literature on this and related species must be regarded as unreliable. Epilachna sparsa (Herbst) has been regarded by all authors as a synonym or variety of E. 28-punctata (Fabricius). What Fabricius had before him when he described Coccinella 28-punctata cannot be ascertained now without the type, which apparently has not been pre- served. Weise and others identified E. 28-punctata with a species that is obviously different from Epilachna sparsa (Herbst). Accord- ingly, the latter becomes a valid name. The figure that accompanies Herbst’s meager description (fig. 11) gives much more information of the species than the description itself. It shows that the spots are of very unequal size and that spots a and f are missing. Specimens like this are very common in India, whereas what is usually known as E. 28-punctata never shows this kind of spot pattern. What is called E. 28-punctata in the literature may refer to the true 28-punctata but probably more often to E. sparsa and sometimes to several other related species. (See also the comments under FE. 28-punctata.) E. sparsa also goes under the name E. 28-maculata Motschulsky, which, however, is very probably a name belonging to a different species, E. niponica. (See also comments under that species.) Epilachna sparsa is very variable in appearance, and this variation has resulted in the description of some of its forms under separate names. EF. gradaria Mulsant, with its varieties addita, vieta, socors, congressa, and stolida, and E. territa Mulsant are probably only forms of E. sparsa. It is quite likely that several other Mulsant species will fall in the same category, although the descriptions are not definite enough to settle this. In collections I have seen F. sparsa under the names taeniata Mulsant, dodecastigma Mulsant,'* pusillanima Mul- sant, indica Mulsant, and others. The status of these different forms is of great interest. Of course, we may be dealing here with a number of closely related but distinct species. However, this does not seem very likely. There is no def- inite structural character by which such species might be separated. The genitalia are alike, though not completely identical in all forms. The differences in the genitalia that are found, such as the width of 13 F. dodecastigma (usually misspelled dodecostigma) is credited in the lit- erature, including the Korschefsky Catalog, to Mulsant. However, the species was described by Wiedemann (1823). Mulsant (1850, p. 789) omitted the reference to Wiedemann’s description, which omission he later corrected (1853, p. 248). 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 the bladelike ridge on the upper side of the base of the penis or the exact shape of the profile of the penis, seem to vary gradually and are not correlated with features in the external appearance. To take coloration and spot pattern as a basis for separation into species as has been done in the past leads immediately to difficulties, since almost all intermediates occur. The other alternative—that we are dealing with one variable species that is broken up into a number of geographical races or subspecies— seems much more likely to represent the real state of affairs and is in agreement with the known facts, although much more material is needed, preferably supplemented by biological observations and breed- ing experiments, before we have anything like a complete picture. Some of these subspecies are sharply defined; others partly over- lap and can be defined only statistically. The separation of Epilachna sparsa from species with similar ap- pearance can best be made by the male genitalia and often also by the female genitalia. Details are given under the various species. In many cases the separation may be carried out from external charac- ters alone. Usually this can be done readily, but great care must be taken, as many of these characters are so variable that they need checking with the genitalia. After a brief characterization of the principal subspecies out- standing enough to deserve separate names, a list of the localities of the specimens seen by me is presented with a characterization of subspecies in those localities. It hardly needs emphasis that when such a characterization is based only on a few specimens it must be con- sidered as only tentative. EPILACHNA SPARSA subsp. SPARSA (Herbst) Epilachna sparsa var. gradaria Mulsant 1850, is the extreme form, with the following variations which have the nonpersistent spots indicated : additay Miulsant) te: she jac citer b, h wictan Miutcant. paises f SUCOVSP NMUISAlIL Meine. pees baGede hn congressa Mulsant ......... g Epilachna gradaria Mulsant seems to be the form of sparsa occur- ring in India, with only the 12 persistent spots present, which are arranged as in figure 8. No. 1 is well below the scutellum and No. 4 is separated from the margin. The Mulsant varieties, according to the original descriptions, have, in addition to the 12 persistent spots, NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 33 the nonpersistent spots indicated by the letters in the listing above. Two others occurring frequently are figured in figures 9 and 10. Spots I and 2 are usually much smaller than the other persistent spots, which is also true in Herbst’s original drawing. The varietal names of Mulsant should not be continued, as the differences are insignifi- cant and there would be a great number of others having similar spot combinations which would have equal rights for separate names. Those specimens with the full number 28 of spots or one or two missing should be regarded as the nominal form of sparsa. E. sparsa Sparsa and sparsa var. gradaria‘are therefore only two extreme forms of the same subspecies with all the intermediates occurring. Material examined.—164 specimens. India: Southern India, Co- imbatore, February 10, 19, 22, June, September 7, November 6, 7, 10, 12, December 1, 1945; Malabar, Walayar Forest, 700 feet, No- vember 15, 1945 (P. S. Nathan, D); Punjab, Jullundur; Punjab and United Prov., June-October (R. L. Woglum, NM); United Prov., Dehra Dun, July 10, 1944 (J. Unyal, D); Assam, Shillong, July 1945 (J. Unyal, D) ; Chabua, Doom Dooma, April 8, 28, 1945, Io miles north of Tinsukia, March 29, 1944 (D. E. Hardy, NM) ; Bengal, Barackpore, April 1, 1944 (D. E. Hardy, NM); Mangalore (J. C. Bridwell, NM). Northern India, Kooloo and Ambala (Carleton, MCZ). Remarks.—E pilachna sparsa in India shows a great deal of varia- tion. The pronotum varies from spotless to all seven spots well developed, with all intermediates. The elytral spot pattern may have only 12 persistent spots or the full number 28 with all intermediates occurring. Sometimes the persistent and nonpersistent spots are equally well developed, in other cases the nonpersistent spots are definitely smaller. Almost any number of nonpersistent spots can be found present, from one on each elytron to the full number eight with practically any combination. Spot a seems to be the one most easily disappearing and usually d is the last one to go. There is not sufficient material to decide whether there are any systematic variations with locality. The specimens from Punjab are smaller than those from either farther south or north and have lighter coloration with very few specimens having the full number of spots. The specimens from northern India (Kooloo and Ambala) approach subspecies orientalis by having spots ab3d approximately in one line. 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 EPILACHNA SPARSA subsp. ORIENTALIS, new subspecies FIGURE I2 This subspecies, which occurs in China and Japan, has the full number, 28, of elytral spots. Spot 1 is with its most forward part back of the tip of the scutellum, and No. 4 is usually free from the margin. In this respect it differs from the insular races and ap- proaches sparsa sparsa. The spots are usually fully developed, but the persistent spots are most often considerably bigger than the non- persistent ones. There is great diversity in size. Spots cb3d lie approximately in line. This latter fact immediately allows a separa- tion from F. niponica Lewis and E. dentulata, two different species with which it overlaps partly in distribution. This subspecies is usually called in the literature E. 28-maculata Motschulsky, and often 28-punctata Fabricius. It seems quite certain that Motschulsky’s 28-maculata refers to a different species (see under niponica Lewis). The new name sparsa orientalis is selected for it. Pronotum usually with seven distinct spots, of which No. 7 or Nos. 5 to 7 may be missing. Under side light, with sides of metasternum usually black. The dark color may be spread to most of the abdomen, the sides of which, however, always seem to stay lighter. Material examined——There are more than 70 specimens from China and Japan. Type: U.S.N.M. No. 57106, from China, Fukien Prov., Fuchow- Ming Chiang, June 1926 (F. P. Metcalf). Paratypes: 8 specimens. Same data as type. China: Kiang-su Prov., Shanghai, August 29, 1919 (H. F. Loomis, 1 specimen) ; Nanking, August 14, 1919 (H. F. Loomis, 5 specimens) ; Soochow (C. F. Wu, 2 specimens) ; Chingkiang, July 19, 1924 (J. F. Illing- worth, 2 specimens, BBM); Shensi Prov., Ching-ling Mountains, May-June 1904 (E. Blackwelder, I specimen); Szechwan Prov., Kuifu, October 1930, November 5, 1930 (2 specimens) ; Hunnan border, south of Suifu, June 10, 1929 (D. C. Graham, I specimen) ; between Chentu and Kuanhsien, July 2-5, 1924 (D. C. Graham, I specimen) ; Anhwei Prov., Taipingshien, October 1932; Kiuhua Shan, September 1932 (G. Liu, MCZ, 23 specimens) ; A. Koebele, I specimen (no further data); Hong Kong (1 specimen, BBM) ; Yen-Ping (I specimen). Japan: Gifu Pref., Mitsukuri (5 specimens) ; Kobe (C. F. Baker, 1 specimen) ; Kiushu (1 specimen). For additional records, see under biology. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 35 EpILACHNA SPARSA var. CINEREA, new variety This variety is distinguished from the common form of sparsa orientalis by the pubescence of all the elytra being gray, including the black spots. No other significant differences were noted. It overlaps in distribution with sparsa orientalis, and so it must be re- garded as a variety of the latter with more localized distribution. Maculation—Thorax usually as in figure 4 E, spots 3 and 4 often united. Elytra.—All 28 spots present and usually separate. Occasionally spot I touches the suture, and often spots h and g are coalescent. Material examined.—32 specimens (NM, D). Type: U.S.N.M. No. 57107, from China, Szechwan, near Kuan- hsien, alt. 2,000-4,000 feet, August 1933 (D. C. Graham). Paratypes: 24, from the following localities, all China, Szechwan Prov. (D. C. Graham): Ningyuenfu, alt. 6,000-6,200 feet, July 31- August 5, 1928, and Kuanhsien, alt. 2,000-3,000 feet (14 specimens) ; Suifu, alt. 1,200 feet, September 25, 1923, June 6, 1929. Also China, Kiang-su Prov., Nanking, August 1919 (H. F. Loomis). Additional specimens: China: Kiang-su Prov., Shanghai, August 29, 1919 (H. F. Loomis) ; Fukien Prov., Foochow, September 29, 1914 (C. R. Kellogg) ; between Yachow and Tatsienlu, alt. 2,200- 8,000 feet, July 7-14, 1930 (D. C. Graham, A. Koebele). India: Assam, Shillong, July 1945 (J. Unyal, D). Remarks.—There are some specimens of E. sparsa sparsa in which the gray pubescence of the red parts of the elytra partly transgresses on the black spots. They are intermediate between the regular forms and the variety cinerea. EPILACHNA SPARSA subsp. TERRITA Mulsant E[pilachna] territa MuLSANT, 1850, p. 787. What Mulsant described as E. territa appears to be the 12-spotted form of the subspecies of sparsa prevalent in Java. It differs from gradaria by its larger size, more elongate form, and the fact that spot t is farther advanced, so that a line tangent to the basal margins of these spots goes approximately through the apex of the scutellum. Spot 4 is widened transversely and is joined to the margin. Pro- notum usually spotless, occasionally with up to all seven spots present though only faintly developed. Figures 13 and 14 show forms with some nonpersistent spots present. Many other combinations occur, but usually the nonper- sistent spots are much smaller than the persistent ones. The fully 36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 developed form with all 28 spots has not been found. Spot a is usually absent. There were only two specimens with spot a present, both of which, however, had some other spot absent. The more fully de- veloped specimens with only spot a absent approach very closely subspecies 26-punctata Boisduval. There are in Java, however, also specimens occurring simultaneously with sparsa territa that have the shape of sparsa sparsa but the maculation of sparsa territa, and there are others that are both in shape and maculation like sparsa Sparsa. Type locality.—Java. Material exanined.—65 specimens. Buitenzorg, April-December 1896 (D. G. Fairchild, NM); March-April 1909 (Bryant and Pal- mer, NM); Tjibodas, Mount Gede, April tg09 (Bryant and Palmer, NM). EPILACHNA SPARSA subsp. 26-PUNCTATA (Boisduval) C[occinella] vigintisexpunctata BorsDUVAL, 1835, p. 590. The specimens from Australia and the Pacific Islands agree very well with the detailed description of E. 26-punctata given by Mulsant (1850, p, 838). The localities (Australia, New Guinea, Java), which Mulsant quotes for 26-punctata, agree, so that there is little doubt that these specimens do represent FE. 26-punctata (Boisduval). (The Java specimens are here referred to subspecies territa.) This sub- species is very constant in appearance and very well characterized. Spot a is invariably missing, but all the other spots present except in rare exceptions. Spots I and 4 situated as in territa Mulsant. A line touching the basal margins of spot I goes approximately through the apex of the scutellum, spot 4 widened and joined to the margin. Pronotum most often with four spots (I to 4), the middle ones frequently united or almost united. Occasionally only two spots (1 and 2) and often six or all seven. All the elytral spots are about equally well developed. The under side is light except the sides of the metasternum which are black. Often the dark color spreads to the middle of the basal abdominal segments. Material examined —More than 260 specimens from the following localities : Australia: Queensland, Babinda (J. F. Illingworth, NM, BBM) ; New South Wales (Edwards coll., AMNH), 5 specimens; Victoria (Koebele, NM ; Edwards coll., AMNH), 1 specimen. Fiji Islands, more than 250 specimens (mostly BBM): Vitilevu Island, Suva Bay, Hsivithule-Tailevu, Korovou-Tailevu, July 16, NO. 15 LADYBEETLES OF THE GENUS EPILACH NA—DIEKE 37 1923, August 3, 16, 1937 (O. H. Swezey, G. R. Wilder, J. M. Valentine, BBM) ; Suva Bay, December 1915 (J. C. Bridwell, NM) ; ‘Ways Island, Yasawa group, July 15-19, 1937 (H. St. John, BBM) ; Kadavu Island, Ndavingiele, April, 27-30, 1941 (N. L. H. Krauss) ; Moala Island, July 13, 1924 (E. H. Bryan, Jr., BBM); Totoya Island, July 15, 1924 (E. H. Bryan, Jr., BBM); Matuku Island, July 4, 1924 (E. H. Bryan, Jr., BBM); Lau Island, August 5, 31, September, 1924 (E. H. Bryan, Jr., BBM); Ovalau Island, Tha- wathi, 600-800 feet, July 16, 1938 (E. C. Zimmermann, BBM). Samoa Islands: Savaii Island, below 1,000 feet, April 30, 1924 (E. H. Bryan, Jr., BBM) ; Upolu Island, Afiamalu, Apia, June 4-14, 1940, 1,300 feet on pumpkin, 2,200 feet on Solanum nigrum (O. H. Swezey, BBM); Tutuila Island, February, March, and September 5, 1923, December 18, April 8 (E.H. Bryan, Jr., D. T. Fullaway, H. C. Kellers, BBM) ; Ofu, Manua group, February 27, 1926 (A.F. Judd, BBM). This is the easternmost locality from which any specimen of FE. sparsa or any other Oriental species has been recorded. EPILACHNA SPARSA var. NIGRESCENS, new variety FIGURE 16 Among the Fiji material are a few specimens of a very peculiar color aberration. The larger part of the elytra is solidly black. This black part extends from the line formed by spots bc3d to the apex, so that only spots 1 and 2 are free and surrounded by the yellow back- ground. In the apical black part only a few traces along the suture are left light. The color of pronotum and the under side is normal. Two females show this extreme, while two males show the black color less perfectly spread out so that the original black spots in the apical part still can be recognized. Type.—Bernice P. Bishop Museum, Honolulu, from Fiji. SYNOPSIS OF GEOGRAPHICAL DISTRIBUTION AND VARIATION OF E. SPARSA India: See under sparsa sparsa. A female from Ceylon differs so much in the genital plates that it probably belongs to a different species. Sumatra (24 specimens): Brastagi, May 1927 (F. J. Meggitt, NM); Siantar, 1937 (Mann, NGS-SI Exp., NM); R. Weber, (AMNH) ; Deli, 1912 (De Bussy, NM) ; Tandjong Poera (NM) ; East Sumatra, 1917 (Knab, NM). All specimens with spotless pronotum and 6-spotted elytra, except 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 two which have a few of the nonpersistent spots faintly indicated. In some of the Sumatra specimens the persistent spots are very un- equal in size, e.g., I-4 normal, 5 and 6 very small, practically absent. One specimen, Parpat, (coll. Mann, 1937, NM) has spots 4+3+5 united. Siam: Mekami River, February 3, 1928 (A. Mackie, NM) 12- spotted form. Indochina: Tonkin, Thai Ha, June (MNH), like orientalis (3 specimens) ; with spot a missing (2 specimens) ; with spots 1 and 4 on pronotum (I specimen); Cochin China, Bienhoa Prov., Trang Bom Arboretum, 30 miles northwest of Saigon, July 18, 1932 (M. Poilane, NM, 2 specimens) ; Annam, Hue (NM), 12-spotted, 3+4 united. Indochina (NM), 26-spotted. The Indochina specimens seem to form the transition between the subspecies sparsa and orientalis. Malay: Larat, December 1907, 3 specimens (BBM), with 26 spots (a missing), spots small, cb3d not on straight line, pronotum with spots I-4. China and Japan: See under sparsa orientalis. Java: See under sparsa territa Mulsant. Lombok: Sapit, 7 specimens (NM), very much like 26-punctata (Boisduval) ; pronotum usually spotless. Amboina: 3 specimens (Buitenbos, NM), 12-spotted with large spots. As all the specimens are females, the identification is not quite certain. Batjan Island (between Celebes and New Guinea, lat. 1°S., long. 127.5°E.): I specimen (NM), 12-spotted with rather large spots ;, pronotum spotless. Australia, New Guinea, Pacific islands: See sparsa 26-punctata (Boisduval). Borneo: Sandakan, 1 specimen (Baker, NM), 12-spotted with rather small spots. BIOLOGY The biology of Epilachna sparsa furnishes a typical example for the biology of similar species that apparently do not differ very much in this respect. It has been studied in more or less detail by several investigators in various countries, but was reported usually under the name 28-punctata (Fabricius). A report in which there can be no doubt about the identification of the species is that by Takahashi (1932), who made extensive studies on E. sparsa orientalis in Japan. Some of his results are as follows: NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 39 Climatic preferences: In Japan E. sparsa lives in the warmer parts and finds its distribution limit roughly where the mean annual tem- perature is 15°C. (69°F.), the mean maximum 19°C, (76°F.), and the mean minimum 9°C. (48°F.). Where it is colder the species cannot develop. This brings the northern limit in Japan to just above latitude 36° (north of Tokyo), higher in protected places, much lower in the mountains. E. sparsa has two generations in most of Japan, three in the south. It hibernates as adult beginning in October and emerges the end of April or beginning of May. It lays eggs in clusters on the under side of leaves, total number of eggs laid per female (in laboratory) 192 to 2,272, average 1,161. The larva emerges after 3 to 8 days and spends 14 to 26 days (four instars) until pupation. Pupal period 3 to 5 days. Total period from egg to emergence of adult 20 to 46 days. The higher the temperature, the shorter the development period. Adults developing from the spring generations are on the average larger than those of the fall generation. This is attributed to the greater moisture content of the food plants during spring. The food plants both for adults and larva are, in the order of preference: Physalis alkekengi L. Solanum nigrum L. Solanum tuberosum L. (potato). Solanum melongena esculentum Nees (eggplant). Solanum ovigerum Dan. Lycopersicum esculentum Mill. (tomato). Datura alba Nees. Cucumis sativus L. (cucumber). Actinostemma lobatum racemosum Mak. In Japan the chief damage is to potatoes and eggplants. The adult causes more injury than the larva. This contradicts the observations of other workers and may be due to too few cases. For control, arsenate of lead is recommended besides a cover crop of ground cherries (Physalis) on which the beetles gather gregariously in spring and can be destroyed. Reports from more tropical countries differ from the above report chiefly by the shorter development period, e.g., for Kwantung Prov., China (Chue, 1928), the development period from egg to adult averaged 16.4 days, with a mean temperature of 28.6°C., and there are 5 or 6 generations a year. Here the chief damage is done to eggplants, and cucurbitaceous plants are not attacked in the field when other food is available. 40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Figures and descriptions of egg, larva, and pupa will be found in Takahashi’s paper (1932). 2. EPILACHNA PHILIPPINENSIS, new species FIGURES 10, 118 General habitus like sparsa. Abdomen.—Abdominal lines subterminal and complete, outside margin straight. Fifth segment, male, hind margin slightly concave; female truncate. Sixth segment, male, hind margin slightly emar- ginate ;‘female, segment split lengthwise. Male genitalia——Penis seen in profile, distinctly bent upward at about three-fourths of its length so that both the upper and lower margin show this upward bend. Thickness slightly increases just after the bend and then decreases; ends in apical hook. Two short rows of hairs 0.1 to 0.2 mm. long on upper side after bend, occasionally a few hairs also on the basal end. Paramera, with apical thorn, length about 1.5 mm.; sipho ends in sharp point as in sparsa. Female genitalia—Plates, length 0.53 mm., greatest width 0.33 mm. Differ from those of sparsa only in having the notch on the inner edge less deep. Length—5.6 to 7.2 mm. Tips of elytra with distinct angle. Maculation.—Pronotum usually immaculate, occasionally spots I and 2 present and traces of others; elytra usually with the full num- ber 28 of spots, spot a usually small and often missing. Occasionally others of the nonpersistent spots missing or very small. Spots cb3d do not form approximately a straight line. The arrangement of spots is very much as shown in figure 19, but spot d is most often an elongate rectangle. In a few specimens spots 3 and 4 flow together. Material examined—Type: U.S.N.M. No. 57108, from Philip- pine Islands, Luzon, Nueva Vizcaya Prov., Imugan (R. C. McGregor). 55 paratypes and 22 other specimens in U. S. National Museum, all from Luzon, from the following localities: Nueva Vizcaya Prov., Balite Pass, Imugan; Laguna Prov., Bay; Bataan Prov., Limay, November 8, 1924; Ilocos Norte Prov., Bangui, Solsona, December 1923; Ilocos Sur Prov.; Nueva Ecija Prov. (all R. C. McGregor). Also from Luzon: Mount Maquiling, Los Bafios, January Io, 1910, and Mount Banajao (all Baker); Lanao (C. V. Piper). From Manila: 1909 (G. Compere, W. A. Stanton), 1911 (C. V. Piper). Remarks.—This species is very close to sparsa and might possibly be considered a subspecies of it that has developed so far away from NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 41 the main species as to form a distinct species now. It differs from sparsa by the upward bend of the male penis, by which the two species can always be distinguished. The maculation of philippinensis is very much like that of sparsa. The usual form has the full number 28 of elytral spots. However, cb3d do not lie on a straight line as in sparsa orientalis. The nonpersistent spots are often weaker than the per- sistent ones, or partly or completely absent. In the latter case we have the subspecies remota. There are a few specimens where spots 3 and 4 are confluent as in figure 17. The nonpersistent spots are, however, also partly or completely present in these cases. For the relations to doryca Boisduval, see the note to that species. The fully spotted form of philippinensis seems to be confined to Luzon. E. sparsa has not been found at all in the Philippine Islands, and so it is probably safe to decide between the two species in cases of doubt from the locality alone without dissection of the genitalia, which otherwise is the only safe criterion. EF. philippinensis is very similar in external appearance to E. dentulata, which also occurs on Luzon. Itither the male or female genitalia permit an unambiguous separation of these two species. With a little care they can also be distinguished by their external appearance. FE. dentulata is more rounded and convex and has the tips of the elytra definitely rounded. Spot 4 reaches the side margin in dentulata except in some very lightly colored specimens from Indochina, whereas in philippinensis it never seems to touch the margin. EPILACHNA PHILIPPINENSIS subsp. REMOTA, new subspecies FIGURE 20 This subspecies is identical with philippinensis s. str. in the male and female genitalia and in all morphological characters. It differs by the absence of all the nonpersistent spots on the elytra and so there are only six spots on each elytron. It seems to occur on the outlying islands rather than on Luzon. Some of the few specimens known from Luzon have a few of the nonpersistent spots present, and form thus transitions between this and the typical subspecies. Material examined.—26 specimens (all NM). Type: U.S.N.M. No. 57109, from Philippine Islands, Mindanao, Davao Prov., Mati, June 1927 (R. C. McGregor). 13 paratypes and 12 other specimens: From same locality as type, March and June 1927 (2 specimens). Negros Island (8 specimens) : Victorias, 1927 (W. D. Pierce) ; Cuernos Mountains (Baker) ; May Igtr (C. V. Piper). Others from: Mindanao, May 1g11 (C. V. : 42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Piper) ; Manila, 1911 (C. V. Piper) ; Cadiz, on Datura alba; Pala- wan Island, Puerto Princesa (Baker); Sulu Islands, Jolo, July 24 (A. Duyag, McGregor); Bohol Island, Bilar, September 1923 (McGregor) ; Luzén, Los Bafios (Baker) ; Luzén, Nueva Vizcaya Prov., Balite Pass (McGregor). One specimen from New Guinea (G. Compere) has spots 3 and 4 confluent. EPILACHNA PHILIPPINENSIS subsp. AUSTRALICA, new subspecies Specimens from Australia are known which, though they re- semble in maculation dark specimens of E. 28-punctata much more than the typical E. philippinensis, have genitalia that cannot be distin- guished from those of E. philippinensis. They are therefore tentatively identified as a subspecies of philippinensis, but more material is needed before any definite conclusion can be reached as to relation between philippinensis australica, sparsa 26-punctata, and 28-punc- tata, which all occur in Australia and are so similar in appearance that they have been confused completely in the previous literature. The maculation is very similar to that of figure 22, but spots d and g are actually on the suture. The pronotum has a black disk not resolved into spots with a narrow margin at apex and base and a wider one at the sides light in all but one of the specimens. In this one, which is lighter all around, pronotal spots I to 4 flow together but are recognizable as separate spots, and No. 7 is also present, small and separated from the others; spots 5 and 6 are absent. All elytral spots are present including a, which is always missing in sparsa 26-punctata. Material examined.—Type: U.S.N.M. No. 57110, Australia: New South Wales, Richmond River. 4 paratypes all from Australia: New South Wales, Richmond River, Botany Bay (NM, AMNH). 3. EPILACHNA 28-PUNCTATA (Fabricius) FIGURES 21, 22, 119 Coccinella 28-punctata FaBRIcIus, 1775, p. 84. Abdomen.—Abdominal lines subcomplete, reaching to within one- fourth of hind margin of first segment, outer margin nearly straight. Fifth segment, hind margin, male, slightly concave to truncate; female, truncate. Sixth segment, male, entire, convex; female, split lengthwise. + NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 43 Male genitalia-—Penis seen in profile, curved upward near apex in gentle uniform curvature; no apical hook. Very narrow basal knife ridge; two rows of sparse hairs (about 0.2 mm. long) from middle to close to apex. Paramera length 1.25 mm. with apical spine. Sipho with tip rounded and with deep notch (fig. 119). Female genitalia (as in fig. 182).—Genital plates length 0.6 mm., greatest width 0.37 mm. On the inner edge near the base an almost circular piece is cut out (diameter about 0.1 mm). Length—6 to 7 mm. Apex of elytra rounded. Color and maculation.—In general all 28 spots are present. They may be small points (fig. 21) to large spots covering most of the area (fig. 22). The prenotum may vary from spotless to a pattern like that shown in figure 41. Persistent and nonpersistent elytral spots are in general equally well developed. There is not the tendency of spot a to disappear or other nonpersistent spots to be very small as generally observed in sparsa. However, occasionally some spots are absent. The under side may vary from completely light to almost completely black. Occasionally in dark specimens spots 3 and 4 or 3 and 5 lightly joined (Samoa). The following characters are most useful to distinguish FE. 28- punctata from species with similar appearance: Rounded apical angle of elytra; spots cb3d never on a straight line; genitalia of male, par- ticularly tip of sipho; female, inner indenture of plates. Material examined.—78 specimens, all with 28 spots except when specifically mentioned otherwise, from the following localities: Ceylon (Weise, 1900) ; Ceram Island, Piroe (Mann, 1937, NGS- SI Exp., NM, 1 specimen) with spots e, f, g, absent; New Guinea, northeastern Papua, Mount Lamington, alt. 1,300-1,500 feet (C. T. McNamara) ; Solomon Islands, Guadalcanal Island, 1927 (MNH, 18 specimens), January-February 1920 (J. A. Kusche, BBM) ; Fulakora Island (Ch. Bignell, NM, 3 specimens) ; Bougainville Island, Sep- tember 1944 (L. F. Gunther and R. Cross, NM). The Solomon Islands specimens, especially those of Guadalcanal, have a spotless pronotum, very small elytral spots (as in fig. 21), the apical h often missing, under side completely light, and size rather small. The Bougainville specimens are somewhat darker with pronotal spots 3 and 4 present, the nonpersistent spots of the elytra smaller than the persistent ones and one or two usually missing. Australia: Northern Territory, Darwin, Stapleton (G. F. Hill, NM, 3 specimens) ; Roper River (N. B. Tisdale) ; Behn River (E. Kimberley, BBM, 3 specimens) ; Queensland, Gordonsvale, N. W. (J. F. Illingworth, BBM, NM); Redlynch, N. W., August 14, 1938; 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Cairns (BBM); Brisbane (NM); Townsville, February 5, 1945 (B. Malkin); New South Wales, Illawarra (H. Peterson, NM); North Ryde, January 10, 1932 (R. W. Burrell, NM; A. Koebele, NM). In general the specimens from Northern Territory and North Queensland are very light with small spots similar to the Guadalcanal specimens. Occasionally spot a is missing. The Roper River speci- mens are somewhat darker, one with spot e missing. The specimens from South Queensland and New South Wales are dark with large spots like the Samoan specimens. A number of Australian specimens without any specific locality data are similar. From Samoa (28 specimens): Upolu Island, Apia, January 1925 (G. H. Hopkins via Higgins, NM) ; Apia-Aleisa Road, 1,300 feet, on pumpkin, June 4, 1940 (O. H. Swezey, E. C. Zimmermann, BBM) ; Tapatapae, July 16, 1940, on pumpkin (O. H. Swezey, BBM) ; Tutuila Island, February (D. T. Fallaway, BBM); Savaii Island, Salailua, May 19, 1924 (E. H. Bryan, Jr.,. BBM). Remarks.—As all the Samoan specimens have large spots, the space between them is small compared to the size of the spots (fig. 22); pronotum as in figures 4 E and 41. Under side, except part of epipleurae, completely black. Legs and last segments of antennae and maxillary palpi also black. It is peculiar that while 28-punctata is common in Samoa, it seems to be absent from the Fiji Islands, which lie between Samoa and Australia and the Solo- mons where the species also occurs. It will be very difficult to obtain complete certainty about the identity of Coccinella 28-punctata of Fabricius in the absence of the original type. The original description is merely: ‘“‘C. coleoptris rubris: punctis nigris viginti octo. Habitat Tranquebariae. Dr. Koenig.”” This would fit any of the species with 28 spots. The type locality, Tranquebar, is located on the east coast of India south of Madras in latitude 11°N. In the literature since Fabricius, the name 28-punctata has been used for at least five different species, which could not be separated by their external appearance alone. In the present paper a choice has been made that disturbs the existing nomenclature as little as possible and is not in contradiction with any -of the known facts, but there is no definite proof of its correct- ness. The only previous reference in the literature that allows a positive recognition of the species with which the author was dealing is that of Weise (1900), who described the male genitalia of Ceylon specimens which he called 28-punctata (Fabricius). This descrip- tion agrees very well with the genitalia of the specimens from Aus- tralia and the Pacific area that I have identified with 23-punctata NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 45 in the present paper and does not agree at all with the genitalia of E. sparsa (Herbst), which always has been regarded as a synonym of 28-punctata (Fabricius) (see under sparsa). Since Ceylon is not far from the type locality of 28-punctata (Fabricius), Weise’s identi- fication seems quite reasonable, and without evidence to the con- trary it is best to assume that he was right. Although with this as- sumption it seems certain that sparsa and 28-punctata are distinct species, the identity of the Australian and Pacific specimens with those from Ceylon and India, though probable from Weise’s descrip- tion of the genitalia,* should be checked by a more careful com- parison, which I have been unable to make, as no specimens of 28- punctata from much west of New Guinea have been available to me. Should the Australian specimens prove different, they would require a new name. The synonymy of this and related species would have to be changed even more if Weise’s 1900 identification of 28-punc- tata should prove wrong. Complete certainty in these matters can never be obtained without the type, which is apparently lost, but careful examination of fairly complete material from southern India would help a great deal. As the various species can now be told apart easily by their genitalia, much of the former uncertainty should disappear. BIOLOGY Biological notes for Epilachna 28-punctata have been published by a number of workers, but in almost all cases it is impossible to be certain whether the data refer to 25-punctata or to one of the allied species. Contradictions in the reports on food preference probably find their explanation in the fact that the observers referred to different species. As far as the actual habits are concerned, there seems to be little difference between the species of this group, and most of what was found for E. sparsa will probably apply to 28- punctata. It is obviously a tropical species and feeds on solanaceous and cucurbitaceous plants. It has been reported from Australia (New South Wales, Olliff, 1890) as one of the worst enemies of potatoes, pumpkins, and tomatoes. Judged from the data given by the authors and the present knowl- edge of the geographical distribution of the various species, it is 14In a later paper Weise (1923a) professes ignorance about the identity of the Australian and Indian specimens of 28-punctata. Unfortunately, he made then no use of the genitalia, and from his remarks it seems very probable that he must have had then specimens of sparsa that he compared with specimens of 28-punctata from Queensland. 4 46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 likely that the following biological accounts supposedly of E. 28- punctata refer to the species indicated : Olliff, 1890: Australia, New South Wales—28-punctata, possibly philip- pinensis australica. Fletcher, 1914: South India—sparsa. y Chue, 1928: South China—sparsa or dentulata. Temperley, 1928: Australia, Queensland—sparsa 26-punctata. 4. EPILACHNA DENTULATA, new species FIGURES 23-25, 120, 182 Abdomen.—Abdominal lines complete, or nearly so, subterminal, angulate or subangulate. Fifth segment, male, hind margin slightly concave with a very slight median process; female, truncate. Sixth segment, subtruncate to convex in male, split in female. Male genitalia—FPenis in profile, length 1.4 mm., with apical end gently curved on both upper and lower edge. Total bend about 45°. No apical hook. Seen from below, orifice elongately oval. Penis at apex of orifice about 0.1 mm. wide, length from orifice to apex 0.2 mm., uniformly narrowed to point. On upper side with 3 to 6 low transverse ridges, which appear as small teeth seen in profile. These occasionally may practically disappear. Paramera 1.4 mm. long, of nearly equal width (0.15 mm.), with apical thorn. Rim covered with hairs on apical half (about 0.2 mm. long). Sipho of normal shape, ending in point. Orifice on side, subapical, oval. The shape of the male genitalia is the same as those of 28-punc- tata. They differ by the presence of the dentules and the structure of the tip of the sipho, which is more like that of sparsa and philip- pinensis (see fig. 118). Female genitalia—Very much like those of 28-punctata. The borders of the notch near base of inner edge dark. Length.—6.5 to 7.2 mm. Apical angle of elytra rounded. Color and maculation—Upper side brownish red. Pronotum varying from spotless to one with all seven spots present. Pubescence light gray, dark on the spots. Elytra with all 28 spots arranged as in- 28-punctata. Indochina specimens with small spots, Philippine specimens with larger ones. Material examined.—30 specimens. Type: U.S.N.M. No. 57111, from Cochin China, Trang Bom, 30 miles northwest of Saigon (M. Poilane, 1932). 17 paratypes, 5 from the same locality, others from: Cochin China, Bienhoa Prov., Trang Bom Arboretum, August 12, 1932, September NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE | 47 1, 1932 (M. Poilane), and 58 km. from Colonial R.-20, alt. 100 meters, October 26, 1932 (M. Poilane, NM). Philippine Islands: Bataan Prov., Limay, November 1924 (R. C. McGregor, NM); Lanao, March-June 1911 (C. V. Piper, NM); Los Bafios (Baker). Additional specimens from: China, Soochow (?), Foochow. Cochin China, Bienhoa Prov., Trang Bom Arboretum, September 4-9, 1932 (M. Poilane, NM). Flores Island, Larantuka (female might be 28-punctata) (F. Knab, 1917, NM.). Philippine Islands: Luzon, Ilocos Sur (McGregor, NM); Luzon, January 16, 1920 (MNH); Manila, December 1924 (McGregor, NM). Remarks—tThis species is very close to 28-punctata (Fabricius). It can positively be distinguished from it by the differences in the tip of the sipho and the ridges on the penis, although the latter may occasionally tend to become obsolete. The two species do not seem to overlap geographically, and further data may disclose that dentu- lata should be regarded as a subspecies of 28-punctata. If it is true that the two species do not overlap geographically, the locality of a specimen will give the best clue as to which species a specimen belongs. This will help especially in the case of females, which are other- wise difficult to separate. The other species with 28 spots occurring in the same region are E. sparsa, E. philippinensis, and Afidenta mimetica. From all three it can be definitely separated by the genitalia and from the first two in addition by the fact that the tips of the elytra are rounded and do not show a definite angle. From Afidenta mimetica it differs by the maculation of the pronotum (if any) and the structure of the claws and abdomen. EPILACHNA DENTULATA subsp. PARVINOTATA, new subspecies FIGURES 24, 25 This is the 12-spotted form of dentulata. The structural charac- ters including the genitalia are the same, but there are only six spots on each elytron. Since the geographical distribution of the two forms does not seem to overlap substantially, we may regard them as true subspecies. The maculation of the elytra of parvinotata shows considerable variation with locality, and so it might be subdivided further into several subspecies. The pronotum is spotless in all but the Indian specimens, which have spots 1 to 4 and 7. The elytral spots are very variable in size (see figs. 24 and 25, which show the extremes). The front tangent 48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 to spot 1 usually passes the suture well behind the tip of the scutel- lum, but when the spots are strongly developed it may cut the scutel- lem near the apical end. Spots I and 5 in that case touch the suture. No. 4 touches the side margin or almost so. Material examined.—More than 60 specimens (NM). Type: U.S.N.M. No. 57112, Philippine Islands, Mindanao, Da- pitan (Baker). Paratypes: From the Philippines: Mindoro Island, Port Galera (McGregor) ; Samar Island, June 8, 1924 (McGregor) ; Sulu Islands, Bofigoa, July 1924 (A. Duyag); Negros Occidental, Cadiz, De- cember 4, 1929; Manapla, June 25, 1929, on cane; Victorias, August 25, 1928, on Physalis. From Celebes Island, Macassar. Further specimens from: Lower Siam: Trong (W. L. Abbott), spot I practically absent. Borneo: Sandakan (Baker), Banguey Is- land (to north of Borneo). Moluccas: Batjan Island, August-Sep- tember; Sula Island (Doherty); these specimens have the spots very large. Java-Sumatra: Padang. India: Sikkim Prov., Darjeel- ing, June, 2 specimens both with rather large elytral spots and pronotum with spots 1-4 and 7. This subspecies of dentulata seems to occur over a wide territory. I have found this form in collections under the names taeniata, do- decostigma, sparsa, pusillanima, and others. It is more than likely that the forms from the Indian or the Western Archipelago have been described previously under one of these or possibly another name. However, at present only guesswork could say which one. Therefore it is possible that after examination of the types the synonymy of FE. dentulata will have to be revised. 5. EPILACHNA REDUCTA, new species Ficure 26 Abdomen.—Abdominal lines complete, subangulate, reaching to within one-fourth to one-fifth of hind margin, variable. Fifth segment male, hind margin truncate to slightly concave; female, truncate. Sixth segment, male, truncate or slightly convex; female, split, apical corners distinct. Abdominal segments with shallow impressions on sides. Male genitalia—Penis seen in profile, 1.15 mm. long, widened near base, basal knife edge present. Middle part with parallel edges 1.15 mm. wide, bent upward near apex through angle of nearly 9o°. No hairs on penis. Seen from below, uniformly narrowing from base to apex, width at base 0.2 mm., at apex of orifice 0.07 mm. ending NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 49 in sharp point. Paramera 1.11 mm. long, gently bent outward near base, slightly widened toward apex, maximum width 0.13 mm. Apical thorn present but not very prominent. Rim of apical third covered with light blond hairs (about 0.2 mm. long). Sipho normal shape, tip emarginate. Female genitalia—Apex of tergite X broadly truncate, shape of genital plates like 28-punctata, notch near base of inner edge narrow and deep. Length—6 to 6.5 mm. Tips of elytra rounded. Color and maculation—Upper side light brownish red. Pronotum black with light edges (as in fig. 4K, a little toward 4J). Elytra with 13 spots each, a missing or possibly united with No. 2. Spots large and showing a tendency to be elongated. Under side except head black, epipleurae, tarsi, antennae, and mouth parts light, femora and tibiae black with light tips. Material examined.—7 specimens, as follows: Type: U.S.N.M. No. 57113, Philippine Islands, Luzon, Benguet Prov., Baguio, (Baker). 6 paratypes, same locality (Baker, G. G. Haslam, McGregor, NM). All specimens are very much alike in color and maculation except one which has lighter tibiae. Remarks.—E. reducta can easily be recognized by its elytral mark- ings. It is more slender than sparsa, 28-punctata, and allied species, from which it is also easily separated by its genitalia. In the struc- ture of the male genitalia it is closest to E. emarginata (fig. 125). The genitalia of these two species are practically identical in struc- ture, but the two species differ considerably in external appearance. 6. EPILACHNA EMARGINATA, new species FIGURES 34, 125, 195 Abdomen.—Abdominal lines complete, subterminal. Fifth seg- ment with hind margin truncate in both sexes. Sixth segment, male, hind margin slightly emarginate; female, split; apical angles sharp. Male genitalia—Penis seen in profile, 1.4 mm. long, with rudi- mentary basal knife edge; curvature uniform, very sparse pubescence with only a few hairs present. Paramera 1.3 mm. long with apical thorn, slender (greatest width little over 0.1 mm.), sparsely pubescent with light hairs on apical third. Sipho with tip emarginate. Female genitalia.—Plates 0.43 mm. long, greatest width 0.27 mm. ; lateral edge almost semicircular; notch on inner edge near base, pointing somewhat backward. 50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Length—6 mm. Tips of elytra with distinct angles. Color and maculation—Upper side red, pronotum spotless, elytra each with six black spots. No. 1 usually very small, in one specimen absent. (Fig. 34 was made from a specimen that had this spot un- usually large.) The other spots rounded, No. 4 touching the margin in specimens where the spots are large. Under side light, metasternum darker. Material examined.—6 specimens, as follows: Type: U.S.N.M. No. 57114, Philippine Islands, Samar Island, May 30, 1924 (McGregor). 5 paratypes: Samar Island, May 26, 1924 (McGregor, NM), and Bho (NM). Remarks.—The male genitalia of this species are very similar to those of E. reducta, which is the only other species with the tip of the sipho emarginate. EPILACHNA EMARGINATA subsp. ALTERA, new subspecies This is the multipunctate form of E. emarginata that, as in several other species, seems to occur as subspecies on Luzon and also on Mindanao. All specimens have all the spots except a. No. 1 is of normal size. Material examined—Type: U.S.N.M. No. 57115, Philippine Islands, Luzon, Mount Banajao (Baker). Paratype: Luzon, Benguet, Irisan, June 1903 (R. C. McGregor) ; Los Bafios (Baker, MCZ). Additional specimens: Mindanao, Zamboanga (MCZ). 7. EPILACHNA NIPONICA Lewis FIGURES 27, 28, 122 Epilachna niponica LEwts, 1806, p. 23. ? Epilachna 28-maculata MotTscHULSKy, 1857, p. 40. Abdomen.—Abdominal plates complete, subterminal, rounded. Fifth segment, male, hind margin mildly concave; female, truncate with a depression in the middle of apical half. Sixth segment, male, emarginate ; female, split. Male genitalia.—Penis seen in profile, small basal knife ridge. Penis tube straight for the major part with parallel edges, 1.7 mm. long. Curved near apex through approximately go°, curvature strongest just before tip, without, however, forming a hook. Upper side with hairs from about one-third to five-sixths from the base. Three to six (usually about 4) teeth leaning backward toward base NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 51 on upper side, variable in size. Penis seen from below, tube gradually narrowed from base to apex. Part beyond orifice short (0.2 mm.) and wide (0.17 mm. at apex of orifice narrowing uniformly toward apical point). Paramera 1.6 mm. long, 0.1 mm. wide, sub- parallel for apical two-thirds, gently curved down near apex, with apical spine. A single sparse row of hairs on apical two-thirds with traces of additional hairs. More densely clothed with hairs immediately near apex. Sipho profile of tip, see figure 122 (different from that of any other species). Female genitaliaa—Very similar to those of E. sparsa, possibly a little wider. Length of plates 0.53 mm., greatest width 0.37 mm. Notch near base of inner margin shallow, elongately diamond-shaped. Length—6.5 to8 mm. Tips of elytra rounded. Color and maculation—Upper side light reddish brown; head often with two confluent dark spots at middle of base. Pronotum with spots 3, 4, and 7 united in a median spot (fig. 41). Spots 1 and 5 often united (fig. 4 J). However, also lighter specimens occur with practically spotless pronotum and dark ones where the pronotum is completely black except for a narrow front and side margin (fig. 4 K). Elytra with the full complement of 28 spots. The nonpersistent ones nearly as fully developed as the persistent ones. I have seen no speci- men with any spot missing. Spots cb3d never approximately on a straight line. Under side and appendages light in the lighter speci- mens with the sides of metasternum and the middle of the basal abdominal segments dark. In the darker specimens the dark color spreads over the whole under side including spots on the femora and the elytral epipleurae. However, even in very dark specimens, the depression in the middle of the fifth abdominal segment of the female is light. Distribution.—Takahashi (1932) has made a detailed study of the distribution of E. niponica Lewis in Japan. The species occurs in the northern parts and the mountainous parts of the southern sections. The southern limit is roughly where the mean annual temperature becomes higher than 15°C. This means in Japan a latitude of about 32.5°N. It seems that niponica will not hatch when the temperature is above 32°C., as the eggs perish. This may be the reason for the limit of distribution. E. sparsa needs a mean annual temperature of 15°C. or more. There is only a relatively narrow region between the isotherms 14°C. and 15°C. where both species occur. Material examined——More than 160 specimens (NM, MNH, BM), from Japan, China, and Siberia: Japan: Sapporo, May 10-11, 1928, July 24, 1927 (T. R. Gardner) ; 52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Karuizawa Mountains (NM); Takada, July 13, 1930 (H. Suguira, NM) ; Koebele (NM) ; Yokohama, Kagoshima (NM). China: Peking; Tsing Ling Shan, 65 miles northeast of Peking; Peking, summer 1913 (F. N. Meyer, NM) ; Chih-li Prov., Hsiao Wu Tai Shan (F. N. Meyer, NM) ; Pei-tai Ho near Shanhai-kuan, Chih-li Coast, August 1914 (A. DeC. Sowerby, NM); Kiang-su Prov., Soochow, N. Gist Gee (NM); Jehol Prov., Yuasa, 1936; Ming Chiang, Foochow, June 1926 (F. P. Metcalf, NM). Manchuria: N. Kirin, I-mien-po (Sowerby, NM) ; Darien, Sokin, August 1936 (Flexier). Southern Korea: Mount Chiisan, July 23-August 7, 1935 (D. M. Suk, NM). Siberia: Vladivostok, 1923 (V. Prinada, NM) ; Okeanskaya, 1923 (Corkerell, NM); Kongaus, August 1923 (Lavrushin, NM). Remarks—The nomenclature of this species has been quite con- fused, and even more so the identity of E. 28-maculata Motschulsky. Motschulsky described Epilachna 28-maculata in 1857 from speci- mens from Shimoda, Japan, and eastern Siberia. The description would fit any of the species with 28 spots and is therefore entirely inadequate for a diagnosis. Subsequent authors have applied the name 25-maculata with about equal frequency to two species, which are called in the present paper E. sparsa orientalis, new subspecies, and FE. niponica Lewis. We know enough about the fauna of Japan and eastern Asia to be certain that Motschulsky’s 28-maculata must be one of these two. Unfortunately the type locality Shimoda on Izu Peninsula is not among the localities where detailed observations on the distribution of the two species were made by Takahashi (1932), but it lies in the general region where both species may be expected. Takahashi’s observations seem to show that niponica does not occur in-the low- lying parts of the Tokyo district and therefore would not be expected to occur at Shimoda, which has an even milder climate. On the other hand, Lewis (1896) reported his 28-maculata, which is almost cer- tainly identical with niponica, from Yokohama, and, of course, Mot- schulsky might have obtained his type specimens of 28-maculata from the mountains near Shimoda, where niponica is almost certain to occur. Therefore, the type locality unfortunately gives no definite clue to the identity of Motschulsky’s 28-maculata. Motschulsky gives as second locality for his type series “Eastern Siberia,” which is beyond the range of sparsa orientalis and would therefore settle the question about the identity of 28-maculata, if we could be sure that the Siberian and Japanese specimens that Motschulsky had before NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 53 him actually belonged to the same species, which can be settled now only by a reexamination of the types. Until this has been done, I prefer provisionally to retain Lewis’s name E. niponica, which has never been applied to any but the present species, and this cannot possibly lead to any further confusion. Should Motschulsky’s type from Shimoda be found to be identical with niponica Lewis, then this name would have to be dropped as synonym of 28-maculata Motschulsky. This would still be true if the Japanese specimen should turn out to be identical with sparsa orientalis, but as holotype of 28-maculata a Siberian specimen of Motschulsky’s type series would be selected. Lewis (1896) in his list of Japanese Coccinellidae gave the de- scription of three 28-spotted species of Epilachna. His 28-punctata is E. sparsa orientalis of the present paper. From our present knowledge of the Japanese Epilachna it is apparent that Lewis’s description of 28-maculata Motschulsky and niponica Lewis very probably refer to a light and dark form of the same species (corresponding to figs. 27 and 28) so similar as not even to deserve varietal names. All the intermediates between these two forms can be observed. Lewis states in the original description of miponica: “The abdomen, fourth segment of the male is canaliculate in the middle, surface of the channel usually red, rarely black,” and this statement is repeated by Mader (1926). In all the specimens I have seen the fifth segment of the female has this property instead of the fourth segment of the male. It is clear, therefore, that miponica Lewis is synonymous with 28-maculata Motschulsky subject to a confirmation from Motschul- sky’s type. The continued use of niponica Lewis in this paper is only provisional. BIOLOGY The biology of Epilachna niponica Lewis in Japan has been studied by several observers, probably most in detail by Takahashi (1932). It has two generations in most of Japan, but only one in the northern parts (Hokkaido). Reports that it should have three generations in southern Korea are questionable. The eggs are laid in open clusters on the under side of leaves of the food plant, while sparsa has closed egg clusters. The number of eggs laid by one female under laboratory conditions was 222 to 806, average 445. The duration of the various stages is: egg 3 to I1 days: larvae 14 to 27 days (4 instars) ; pupa 4 to 8 days; egg to emergence of adult 25 to 45 days; oviposition period 17 to 4o days; end of egg laying to death 2 to 29 days (average 15). The lower limits are for the warmer, the upper for the cooler periods. 54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 The beetle hibernates as adult. Activity begins from the middle of March to the middle of April, and the beetles appear all at once. They disappear for hibernation from the beginning of October to November. E. niponica is said to injure more than 30 species of plants. Ac- cording to Takahashi, the principal food plants are: SOLANACEAE CUCURBITACEAE » 1. Solanum tuberosum L. Potato. 10. Cucumis sativus L. Cucumber. 2. Solanum melongena esculentum 11. Cucurbita muscata toonas Mak. Nees. Eggplant. Squash. 3. Solanum nigrum L. 4. Solanum ovigerum Dan. LEGUMINOSAE 5. Physalis alkekengi L. 6 7 _ Physalis angulata L 12. Phaseolus vulgaris L. Bean. . Lycopersicum esculentum Mill. 13. Glycine soja Benth. Soybean. Tomato. 8. Datura alba Nees. COMPOSITAE 9. Capsicum annurum L. Red yy. Arctium lappa L. pepper. CRUCIFERAE 15. Brassica campestris L. Nos. I and 2 are most commonly injured, then 3 and 7. No. 5 is seldom attacked and in this respect niponica differs from sparsa orientalis for this is the latter’s preferred food plant. No. 15 is eaten in autumn when there is little other food. EPILACHNA NIPONICA subsp. COALESCENS Mader FIGURES 29, 30 Epilachna 28-maculata a. coalescens MADER, 1930, p. 184. This seems a well-defined subspecies occurring in the mountains of Szechwan, China. It differs from the ground form by the con- fluence of a number of spots, which in the fully developed form takes the appearance of figure 30, with the following spots confluent: 2+4+1, b+c, 3+d+d, e+f, g+h. The pronotum is black with a narrow front and side margin black. The pubescence of the elytra is gray all over, while in the ground form it is black on the dark spots. This makes the spots much more indistinct in appearance. The male genitalia, though on the whole identical in appearance, seem to be somewhat shorter, and the paramera may be twice as wide (0.2 mm.) as in the typical Japanese specimens. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 55 Mader described this form as an aberration of E. 28-maculata. He does not mention the difference in the color of the elytral pu- bescence, but there seems little doubt that he must have this same form before him. Type locality—China, Szechwan Prov. Material examined.—s6 specimens, all from China, Szechwan Prov. (D. C. Graham, NM), (except 2 specimens from Tibet, MNH): Den Shiang Uen near Ningyuenfu, alt. 8,000-9,500 feet, August 4-6, 9-10, 1928; between Ningyuenfu and Den Shiang Uen, alt. 6,000-8,000 feet, August 6-8, 1928; Ningyuenfu, alt. 6,200 feet, July 31, 1928; near Wen Chuan, alt. 4,000-6,000 feet, May-August 1933; Uin Gin Shien, alt. 2,500-7,000 feet, July 14-15, 1928; near Fu Liu, alt. 5,000-8,200 feet, July 19-21, 1928; Mupin, alt. 5,000- 6,000 feet, July 19, 1929; Mupin, August 25-28, 1929; Tatsienlu, alt. 5,000-8,500 feet, August 7, 1923; Tatsienlu, alt. 300 feet, August 16, 1930; near Yachow, alt. 2,800-4,000 feet, July 8, 1930, June 16- 20, 1923; 35 miles west of Tatsienlu, alt. 5,000 feet, June 20, 1923; near Li To, alt. 5,000-9,000 feet, August 21, 1930; Shin Kai Si, Mount Omei, alt. 4,400 feet, July 6-16, 1934, August 1921; Si Gi Pin, Mount Omei, alt. 6,000 feet, August 6-15, 1925; Kuanhsien, alt. 2,200-5,200 feet, July 19-20, 1933; Yao Gi near Mupin, alt. 7,000 feet, July 17, 1929; near Mupin, 3,000-7,400 feet, July 1-3, 1929; near Hai Tang, alt. 6,000-8,000 feet, August 4, 1928; between Yachow and Suifu, alt. 1,000-2,500 feet, January 15-24, 1929; Yachow, alt. 2,000, July 1928. There are three specimens with uniformly gray pubescence on the elytra but with all the spots separated. China: Chih-li Prov., Pei-tai Ho near Shanhai-kuan, August 1914 (A. DeC. Sowerby, NM). 8. EPILACHNA WISSMANNI Mulsant FIGURES 31, 123, 190 E[pilachna] Wissmanni MutLsant, 1850, p. 832. Abdomen.—Abdominal lines subcomplete, subangulate, reaching to within one-fifth of the hind margin of the first segment. Fifth segment, male, hind margin very distinctly and broadly emarginate ; female, subtruncate with a faint middle process, sixth segment, male, with a deep notch; female, split. Male genitalia.—Penis seen in profile, length 2.8 mm., lower edge straight for about 2.1 mm., then gently curved up to the sharp apical point. The curved part with a distinct emargination (as in sparsa), 56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 upper edge with a distinct basal knife edge, the apical part in a gentle arc curved upward; a slight bulge corresponds to the emar- gination on the lower side. This curved part furnished with hairs. Seen from below, closed tube for 1.9 mm., then elongate diamond- shaped orifice about 0.5 mm. long beyond that narrow edge to apical point. Paramera 2.5 mm. long, compressed for apical two-thirds, widened near apex, with apical thorn and with two rows of hairs on apical third (0.1 to 0.3 mm. long). Sipho with a blunt right- angle bend near base ending in a long and slender point with a very elongate orifice. Female genitalia.——Genital plates, length 0.67 mm., width 0.44 mm., with a characteristic emargination about the middle of the inner margin. Length—g to 10 mm. Apical angle of elytra distinct. Beetle heart-shaped, strongly convex. Color and maculation.—Upper side light brownish red to brick red; pronotum spotless. Elytra with the six persistent spots rounded except No. 4, which is laterally widened to the margin. Nos. 1 and 5 separated from the suture. Common front tangent to the two No. 1 spots passing approximately through the tip of the scutellum. In addition, nonpersistent spot c present; size of it from very small to close to size of the persistent spots. In one specimen there is also a faint indication of spot d. Under side and appendages light; the elytral spot 4 reaches over three-fourths of the width of the epipleurae. Type locality.—Celebes. Material examined.—8 specimens (NM, MCZ) from Celebes: Bantimoerong (Mann, 1937) ; South Celebes, Bonthain (C. Ribble, 1882, 1883) ; North Celebes, Toli-Toli; Patutuang, Malino (Brues). 9. EPILACHNA BAKERI, new species FIGURES 31, 124, IQI A species very similar to wissmanni in shape and maculation but slightly smaller. It differs from wissmanni distinctly by the structure of the genitalia. Abdomen.—Abdominal lines somewhat variable. Fifth segment, male, hind margin concave; female, straight. Sixth segment, male, with shallow notch; female, split. Z Male genitalia—Penis seen in profile, lower edge practically straight until close to apex, then gentle curvature up to fine apical point; well-developed basal knife ridge on upper side, a little NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 57 thickened just before apex, but this sometimes hardly noticeable. Sparse growth of hair, about 0.1 mm. long on apical third. Paramera 2.0 mm. long with apical thorn. Sipho curved near base, then straight, tip compressed, one sharp edge, and on the other side the very elongate orifice. . Female genitalia——Genital plates 0.60 mm. long, greatest width 0.39 mm., with a feeble emargination on the inner edge about one- third from the base. Length—8 to 9 mm. Shape broadly heart-shaped, convex. Apical angle of elytra slightly indicated. Color and maculation—Upper side yellowish red, pronotum spot- less, elytra usually with seven spots (c in addition to the six per- _ sistent ones, the former of varying size and occasionally absent). No. I near suture but not quite touching it; a line tangent to the front margins usually cuts the scutellum in half. Occasionally elongated forward and almost touching the base; Nos. 3 and 4 transversely broadened, No. 4 touching the margin and partly visible on epipleurae. Under side light except the sides of metasternum. Material examined.—Type: U.S.N.M. No. 57116, Philippine Is- lands, Mindanao, Dapitan (Baker). 14 paratypes (NM, MCZ, D): Mindanao Island, Dapitan, Tutuan, Surigao (Baker), Cabadharan (C. S. Banks) ; Northwestern Panay Island; Polillo Island; Biliran Island (Baker). EPILACHNA BAKERI subsp. LUZONICA, new subspecies FIGURE 32 Identical in most respects with the preceding except for the fol- lowing differences, which, however, seem not sufficiently important to warrant its being considered as a separate species: The profile of the penis is slightly different. The upper edge is more nearly straight, but the lower one has an emargination before the apex and ends in a definite hook. The most conspicuous dif- ference is in the presence of additional nonpersistent spots, which occur as follows in the five available specimens (all from Luzon, P. I., coll. Baker and McGregor, NM) (for the notation of spots et. fig, 7): Type: U.S.N.M. No. 57117, Philippine Islands, Luzon, Mount Maquiling (Baker), with all spots except a, but the nonpersistent spots very small. 4 paratypes: (1) female, Mount Maquiling (Baker); spots like the type but also a present, though very faintly; (2) female, Mount 58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Maquiling (Baker) ; spots a, b, c, d, h: c large, b medium, the others small; (3) male, Nueva Vizcaya Prov., Imugan; spots, ¢ large, b and h small; (4) female, Ilocos Sur (McGregor) ; all spots except a of about the same size as the persistent spots; a also present, but very small. We have here another instance of the occurrence of two forms, one with approximately the full number, 28, of spots, and one with only 12 or 14 spots. As in several other cases, the multipunctate form occurs on Luzon, whereas we find the form with the reduced number of spots on the outlying islands. 10. EPILACHNA SEPTIMA, new species FIGURE 220 Abdomen.—Abdominal plates complete, subterminal. Fifth seg- ment, male, truncate; female, with a broad triangular process in mid- dle. Sixth segment, male, hind margin convex; female, split, apical angles of parts distinct. Male genitalia—Penis seen in profile, 1.8 mm. long, with distinct basal knife edge, straight and of approximately equal thickness for about four-fifths of its length, then bent up first gently and shortly before the end more strongly through about 90°. Apical point with a trace of a hook. On upper side with two sparse rows of hairs on middle part. Seen from below, tube about 0.25 mm. wide from base to orifice, the latter elongate. Tube closed for first 0.15 mm., then seams about 0.03 mm. apart, closing toward the middle and diverging again near the orifice. Part beyond the orifice, subparallel, slightly widening toward apex, about 0.065 mm. wide immediately beyond orifice, about 0.3 mm. from orifice to apex. Paramera 1.7 mm. long, thicker than wide except near apex, greatest width near apex 0.15 mm., no distinct apical thorn, pubescence on apical third rather short and sparse. Sipho regular shape as in figure 123, including the com- pressed part near the tip; detail of tip, figure 220. Female genitalia——Shape of plates very much as in FE. wissmanni Mulsant (fig. 190). Length 0.63 mm., greatest width 0.37 mm. Length.—6.5 to 7.0mm. Tips of elytra rounded. Color and maculation—Very much like that of the other 28- spotted species. Pronotum with all spots except 7 present but hazy. All 14 elytral spots present and more or less rounded, the persistent spots on the whole bigger than the nonpersistent ones. None of the spots touching the suture or the margin. Joint front tangent of No. I passing the suture behind the scutellum. Under side mostly light except parts of metasternum. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 59 Material examined.—Type: U.S.N.M. No. 57963, from Indochina, Annam, Phuc-son, November-December. I paratype (D), with same data. Remarks.—E. septima is so much like other 28-spotted species that it can be clearly recognized only by the genitalia. Those of the male are different from those of any other known species. There are other species with the wissmanni type of female genitalia, however, none with the same external appearance. E. septima might with superficial inspection be confused with the following species of the same general appearance known to occur in the same region: FE. sparsa, E. dentulata, Afidenta mimetica. E. sparsa has sharp elytral tips, and A. mimetica is sufficiently different by the generic characters. From FE. dentulata, septima differs defi- nitely by the structure of both the male and the female genitalia. Also, dentulata has spot 4 joined to the margin, while it is free in septima. This may be a safe criterion for separating the two species without dissection. 11. EPILACHNA DIFFINIS (Eydoux and Souleyet) FIGURES 33, I2I Coccinella diffinis Eypoux and SOULEYET, 1839, p. 267. E[pilachna] diffinis MuLSANT, 1850, p. 783. Abdomen.—Abdominal plates subterminal, not quite complete to complete; inner margin curved, outer margin nearly straight. Fifth segment, male, hind margin concave; female, truncate. Sixth seg- ment, male, with considerable emargination ; female, split. Male genitalia—Penis seen in profile, bent upward near apical fourth, on the under side with slight emargination after the bend, with apical hook. Basal knife ridge present, sparse pubescence; on middle part of upper side four transverse ridges which appear like teeth in profile. Paramera slender, 1.8 mm. long, with apical thorn and relatively short and sparse pubescence. Sipho as in sparsa, ending in sharp point with orifice oval on outside. Female genitalia.—Plates, length 0.57 mm., greatest width 0.38 mm., gentle emargination on inside edge near base. Length—7.2 to 8. mm. Shape 6val, apical angles of elytra distinct. Color and maculation—Upper side reddish, pronotum spotless or with faint indications of spots 1-4. Elytra with six spots each, none touching the margin or the suture, pubescence light gray, dark on the spots but. occasionally specimens with gray pubescence through- 60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 out (subsp. signatula Mulsant). Color of spots often brown instead of black, size variable from about 0.4 to 1.3 mm. in diameter. Under side light, except metasternum and central basal part of abdomen, which are usually darker. Remarks.—This species agrees very well with the detailed descrip- tion of diffinis given by Mulsant. The only difference is in the macu- lation of the pronotum, which according to Mulsant should have spots I and 2 present, while in the specimens I have seen it is usually spotless or, when there are spots, I to 4 are present. As the pronotal spots are extremely variable in nearly all species, not much impor- tance need be attached to this difference. Since Mulsant, 1850, re- peated by Crotch, 1874, diffinis has not been mentioned again in the lit- -erature. Weise (1913) and Schultze (1925) mention F. pusillanima Mulsant in their list of Philippine beetles. Judged from the number of cited localities, it must be one of the most abundant species of Epi- lachna in the Philippines, and I believe that their pusillanima is identical with diffinis Eydoux and Souleyet. The type locality of pusillanima is Java, and its description is very little different from that of diffinis. I have not been able to identify it with certainty but suspect that it is one of the forms of sparsa. EPILACHNA DIFFINIS subsp. SIGNATULA Mulsant E[pilachna] signatula MuLsant, 1850, p. 784. Mulsant gives.a variety, signatula Mulsant, that differs from the ground form in that the line joining the fifth and third spots, instead of passing over the callus, passes it on the inside. The specimens from Luzon are of this type. They have uniform gray pubescence all over the elytra but do not differ in the structure of the genitalia. It is reasonable to consider this form as a subspecies. Type locality—Philippine Islands, Luzon, Manila. Distribution of diffinis, including subspecies signatula—57 speci- mens (all NM): Luzon Island, Mount Maquiling and Los Banos; Bataan Prov., Limay ; Sulu Islands (17 specimens), from Jolo, Mount Budaho, and Mount Daho (McGregor) ; Banaran Island (McGregor) ; Bofigao, July 24 (A. Duyag); Basilan Island (Baker) ; Palawan Island, Puerto Princesa (Baker) ; Mindanao Island, Mount Apo, June, July (E. A. Mearns); Davao (Baker); Davao, June 1927 (McGregor) ; Dapitan (Baker) ; Iligan (Baker) ; and one from Min- danao, May 1911 (C. V. Piper). Remarks.—Epilachna diffinis can easily be distinguished from all other species by its male genitalia. The few species (dentulata and NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 61 niponica) that also have similar ridges on the upper side of the penis differ so much in other respects that no confusion is possible. 12, EPILACHNA ELONGATA, new species FIGURE 37 Abdomen.—Abdominal plates subterminal, not quite complete. Fifth segment, female, with hind margin truncate with an indistinct middle process; sixth segment split. Female genitalia—Plates, length 0.57 mm., greatest width 0.33 mm. ; inside edge with a very shallow notch near base. Outer part of apical edge straight. (This part is concave in diffinis, and this dif- ference seems to be the easiest way to tell the genitalia of the two species apart. The plates in diffinis are slightly wider.) Length—7.5 mm. Tips of elytra with quite distinct angle. Shape elongate. Color and maculation—Upper side red, pronotum spotless. Ely- tra each with six fairly large spots; No. 1 elongate and not quite touching the suture, No. 4 joined to the margin. The spots quite black in color and contrasting sharply with the red ground color. (In diffinis the spots are not so black or the ground color so bright.) Under side light, with metasternum and parts of abdomen darker. Clypeus with front edge concave. (This is slightly convex or straight in diffinis.) Type—vU.S.N.M. No. 57118, Philippine Islands, Mindanao, Iligan (Baker). Remarks.—Although this specimen, because of its narrower and more convex shape, its larger spots and brighter colors, gives an im- pression quite different from all the diffinis specimens, there are few definite characters that separate the two species. Those mentioned in the description are slight but apparently valid. Unfortunately, males which probably would permit much more definite conclusions are not available. 13. EPILACHNA MINDANAONIS, new species FIGURES 35, 126, 193 Abdomen.—Abdominal lines complete, subterminal, rounded, reaching to within 0.1 to 0.2 mm. of the hind margin of the first seg- ment. Fifth segment, male, hind margin concave; female, truncate. Sixth segment, male, slightly emarginate; female, split, tips rounded off. 62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Male genitalia——Penis seen in profile, 1.5 mm. long, curved gently up near apex, the curved part very thin. From base to the beginning of the curvature, wedge-shaped with the broad end 0.3 mm. wide near the base. Only a rudimentary basal knife edge. Just before the beginning of the curved part a tuft of long hairs (0.5 mm. long or longer), the hairy part extending only 0.2 to 0.3 mm. Paramera 1.4 mm. long, gently curved outward near base, slightly widened toward apex, no apical spine, densely covered with long (0.4 mm.) hairs on rim of apical part. Sipho ending in sharp point (as in sparsa or philip- pinensis), orifice on side at apex, just before it a short tonguelike process. Female genitalia —Length of genital plates 0.48 mm., greatest width 0.31 mm. The notch on inner edge enlarged so that it covers almost the entire edge concave inward with a sharp edge only at the basal part. This gives the plates almost crescent shape. No similar struc- ture known for any other species. Length—7 to 7.5 mm. Tips of elytra with slight indication of angle. Shape broadly oval, convex. Color and maculation.—Upper side light brownish red. Elytra each with six rounded spots; Nos. 1 and 5 smaller than the others, close to the suture but not quite touching it. Front tangent to No. I through tip of scutellum. No. 4 not touching the margin. Under side and appendages light except tip of mandibles and metasternum and sometimes part of the abdomen, which are darker. Line through middle of spots 3 and 5 touches outside of No. 2. Material examined.—Type: U.S.N.M. No. 57119, Philippine Is- lands, Mindanao, Butuan (Baker). 6 paratypes: Mindanao, Butuan, Surigao (Baker, NM); Samar Island, July 21, 1924 (McGregor). Remarks.—This species is hardly different in external appearance from many others but is very well characterized by its male and female genitalia, and it can easily be separated by them from species of similar appearance. 14. EPILACHNA DUBIOSA, new species FIGURES 36, 127 Abdomen.—Abdominal lines incomplete, subterminal. Fifth seg- ment, male, hind margin a very shallow V; female, truncate. Sixth segment, male, with decided emargination or notch; female, split, apical angles distinct. Male genitalia—Penis seen in profile, 2.1 mm. long, with pro- NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 63 nounced basal knife ridge. Upper edge gently curved, coming to a point with the indications of a tiny hook. Apical half sparsely covered with hairs about 0.3 mm. long. Seen from below, orifice elongately diamond-shaped, the part from orifice to tip a sharp ridge about 0.3 mm. long. Paramera slender, 2.0 mm. long. Seen from above, greatest width near apex, about 0.13 mm. A sharp apical ridge in- stead of the usual apical thorn, sparse pubescence on apical third. Sipho ending in blunt point compressed sidewise near apex, orifice on side oval, elongate with liplike process. The female genitalia of subspecies assamensis are described below. A female specimen from Borneo has the same structure of the geni- talia. The difference in geographical location makes it, however, less certain that this is actually the female of dubiosa. Length—7.5 mm. Shape broadly oval, convex. Tips of elytra rounded. Color and maculation—Upper side red, pronotum with only spots I and 2, faint. Elytra each with six spots: No. 1 beginning well behind the scutellum, No. 4 not touching the margin. Under side light except parts of metasternum, which are darker. Faint dark spots at the sides of the abdominal segments. Material examined.—Type: U.S.N.M. No. 57120, India, Goa, Murmugao (Bridwell). 1 female: Borneo, Sandakan (Baker, D). Remarks.—This is a species very similar in external appearance to others of the 12-spotted species but distinguished by its genitalia. ‘Its appearance does not seem to agree exactly with that of any of the described species, but there are so few distinguishing marks that it is impossible to be sure. The female specimen has much the same ex- ternal appearance as the type and the same structure of the female genitalia as subsp. assamensis. Therefore it is likely that it is actually the female of dubiosa, notwithstanding the diversity of locality. EPILACHNA DUBIOSA subsp. ASSAMENSIS, new subspecies These specimens from Assam have the same structure as FE. dubiosa including the male genitalia. However, the maculation is greatly modified in a manner very similar to that illustrated in figure 41 for E. gangetica. Female genitalia (which are presumably identical with those of E. dubiosa s. str.).—Plates, length 0.66 mm. Shape approximately as shown in figure 190 (wissmanni Mulsant) but the outer apical mar- gin more concave. The inner margin with a part cut out similar to 64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 figure 190 but immediately (0.03 mm.) after the basal part of the cut-out is a tooth that divides the cut-out into a short basal and a longer apical part. Maculation—The pronotum is spotless. Elytral spot I very elongate along the suture and almost reaching the base. Spots I+3+4+5+6 connected together as indicated in figure 41. Spot 2 not connected to No. 1 but joined by a narrow extension to No. 4. No. 6 does not reach the margin. Under side and appendages light, except the sides of metasternum, which are dark. Material examined.—Type: U.S.N.M. No. 57121, India, Assam, Chabua, July 1943 (W. L. Jellison). Paratype: Same locality, May 2, 1944 (D. E. Hardy, D). 15. EPILACHNA QUINTA, new species Abdomen.—Abdominal lines incomplete, subterminal. Fifth seg- ment, female, truncate with slight broad process in middle; sixth segment, split, apical angles distinct. Female genitalia——Plates, length 0.53 mm., greatest width 0.33 mm. Shape approximately that of figure 182 (dentulata), but the notch near the base of the inner edge more closed (as in figure 184). Length—7.0 mm. Shape broadly oval, convex; tips of elytra rounded. Color and maculation—Upper side red; pronotum with all seven spots present but of small size. Elytra each with six spots: No. 1 beginning well behind the scutellum, No. 4 not quite touching the margin. Under side light, except the tips of the mandibles and a spot in the apical corners of the metasternum dark. Type—vU.S.N.M. No. 57964, from Ceylon (Koebele). Remarks.—This species is in outward appearance so much like FE. dubiosa that it was first believed to be a female specimen of that species. However, the female genitalia are distinctly different. 16, EPILACHNA OCELLATA Redtenbacher Ficures 38, 222 Epilachna ocellata REDTENBACHER, 1844, p. 563. E[pilachna] oculea Mutsant, 1850, p. 701. Abdomen.—Abdominal lines complete, subterminal, subangulate. Fifth segment, male, slightly concave; female, truncate. Pigment absent near middle of apex, giving appearance of a deep emargination. Sixth segment, male, emarginate, sometimes the coloration suggests NO. 15 LADYBEETLES OF THE GENUS ,EPILACHNA—DIEKE 65 a longitudinal suture in the middle; female, split, apical angles of halves rounded. Male genitalia—Penis seen in profile, 1.35 mm. long, much wider than most other species of the genus, resembling the enneasticta group; greatest width 0.45 mm. about one-third from the base, then tapering off gradually except just before the apex, where it is bent up sharply into a sharp point. There are indications of a groove for the reception of the paramera. A sparse row of hairs on each side near the upper edge on the apical third but not on the part that is curved up. Seen from below, a tube for about one-fourth of its length, then opening up and trough-shaped becoming gradually narrower toward the apical point. It is completely closed for less than 0.05 mm. at the base and has beyond this a longitudinal slit 0.08 mm. wide, which becomes wider away from the base. Paramera 1.3 mm. long, spoon- shaped, with a strong emargination on the outside near the apex; the sides of the apical seventh sparsely, the apex itself more densely covered with light-colored hairs, about 0.15 mm. long. Sipho rather thick, about 0.2 mm. in middle, bent nearly in a semicircle near base, then bent outward through a right angle and from then straight to tip with diminished width; orifice oval on the elbow of the bend. Female genitalia.—Genital plates, length 0.65 mm., greatest width 0.40 mm., upper part of side margin slightly emarginate, a definite rounded process at the basal part of the inner margin just before the feeble impression. Length—6 to 7 mm. Fine punctation of elytra obsolete; apical angle rounded. Color and maculation—Upper side dirty reddish brown, head lighter. Pronotum as in figures 4 I, J, or close to K. Elytra with six dark spots, each surrounded by a lighter halo. Common front tangent to No. I passing approximately through tip of scutellum. No. 4 widened laterally to margin. Pubescence dense, light gray, dark on the spots. Under side dark, prothorax, appendages, and epipleurae light. The latter with a spot not reaching the inner margin, cor- responding to elytral spot No. 4. Type locality.—India, Kashmir. Material examined.—g4 specimens. India: Kooloo, Ambala (N. M. Carleton, MCZ) ; Allahabad (through W. M. Mann, NM); United Prov., Kumayun, Bhawaki, 5,000 feet, June 18, 1944 (J. Unyal, D). Remarks.—E. ocellata is easily separated from all the other species with 12 elytral spots by the structure of the abdomen, the ocellation of the elytra, and the genitalia, particularly those of the male, which 66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 are quite different in structure from those of any of the preceding species and approach those of the enneasticta group. Mulsant and Mader state that this species is variable. The series I have seen is, however, uniform. 17. EPILACHNA TERTIA, new species FIGURE 128 . Abdomen.—Abdominal lines complete, reaching to within one-fifth of the hind margin of the first segment, fifth segment, male, hind margin mildly concave, sixth segment, emarginate. Male genitalia.—Penis seen in profile, 1.75 mm. long, with rudi- mentary basal knife edge; rather wide (0.4 mm. at its widest part at about one-third of its length). Sparsely furnished with hairs (about 0.25 mm. long) on apical half. Seen from below, inner edges bent down, orifice very long (0.7 mm.). Paramera slender, slightly bent outward, 1.7 mm. long, with a slight apical thorn and with a ridge along their under side. Sipho with a constriction just before the tip and small oval orifice as indicated in fig. 128. Length—7 mm. Tips of elytra with indistinct angle. Color and maculation—Upper side brownish red, pronotum with black spots as in figure 4 in the type, spotless in the other specimen ; elytra each with six dark spots, front tangent to No. 1 passing through tip of scutellum or a little more in front. Nos. 3 and 4 transversely widened, No. 4 almost touching the margin, the others roundish. Under side light, parts of metasternum darker. Pubes- cence whitish gray, dark on the spots. Material examined—Type: U.S.N.M. No. 57965, from India, Assam, Doom Dooma, May 20, 1943 (D. E. Hardy). I paratype (PA), India, northern Bengal, Kurseong, June (Mason collection). The dark spots in the paratype are brownish instead of black. This may be due to the immaturity of the specimen. Remarks.—This is another one of the 12-spotted forms that can- not be referred to any of the previously described species. Its male genitalia are quite different from those of the majority of similar species. If the transversely widened shape of the middle spots is characteristic for the species, it should not be difficult to identify it by its external appearance. 18. EPILACHNA QUARTA, new species Abdomen.—Abdominal lines subcomplete, subterminal, rounded at apex. Fifth segment, male, hind margin truncate or slightly con- cave. Sixth segment, emarginate. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 67 Length—6.5 mm. Tips of elytra rounded. Color and maculation—Upper side red. Pronotum with a central spot, as in figure 4 F, arising from the coalescence of spots 3+4+7. Spots 1 and 2 are faintly indicated besides. Elytra with all six dark spots joined together producing a pattern similar to figure 41 but with the black more extended. No. 1 begins at the tip of the scutellum, is on the suture, extends obliquely backward, and is broadly joined to No. 3. No. 2 on the callus has a triangular shape. One corner touches approximately the middle of the base, the second reaches the juncture of spots I and 3, the third extends faintly along the margin to the forward tip of spot 4. Spot 2 leaves the margin narrowly light. Spot 4 touches broadly the margin and is joined to No. 3. No. 5 touches the suture and is joined to 3 and 6. No. 6 is elongate, leaving the reflexed margin light. The pattern might also be regarded as five light spots on each elytron on a black background. Pubescence light gray all over. Under side and appendages light except the apical and side parts of metasternum and the central parts of the basal abdominal segments. Type.—vU.S.N.M. No. 57122, India, Assam, 10 miles north of Tin- sukia, March 6, 1943 (D. E. Hardy). Remarks—This species has a modification of the 12-spotted pat- tern similar to that found in E. gangetica connecta and E. dubiosa assamensis. From both those species it is distinct by a different struc- ture of the male genitalia. There are distinct differences in the spot patterns of the three species, but these are of doubtful value as long as nothing is known about the variability of the pattern in all three species. E. quarta can be separated from dubiosa by the fact that the sipho has the usual shape of figures 117 and 118. The tip is, however, more pointed than is indicated in figure 118, and the orifice is likewise pointed at its apex. From gangetica connecta, quarta can be distinguished easily by external characteristics. The pubescence is gray all over the elytra in quarta, but dark on the spots in gan- getica. The tips of the elytra are rounded in quarta but distinct in gangetica. Very likely E. endomycina Gorham, andrewsi Gorham, and sus- piciosa Weise, all three from India, are related to these species. 68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 19. EPILACHNA GANGETICA Weise FIGURES 39, 40, 131, 188 E[pilachna] dorica Mutsant, 1850, p. 761. Epilachna doryca var. gangetica WEISE, 1908, p. 218. Abdomen.—Abdominal lines complete, subterminal, angulate. Fifth segment, male, hind margin truncate or slightly concave; female, truncate. Sixth segment, male, very slightly emarginate; female, split. Male genitalia—Very similar to those of sparsa Herbst. Penis seen in profile, somewhat more regularly curved, ending in a distinct hook. Basal knife edge long and wide. The emargination on the lower edge and the thickening on the upper edge just before the apex only very slightly indicated. Pubescence along the apical half. Seen from below, the part beyond the orifice not quite so narrow as in sparsa (0.07 as against 0.05 mm.). Paramera 1.8 mm. long with apical thorn. Sipho as in sparsa. Female genitalia—Genhital plates, length 0.5 mm., greatest width 0.36 mm. Notch near base of the inner margin diamond-shaped. They differ from the genitalia of sparsa chiefly by the more pro-. nounced notch; from those of doryca Boisduval by their greater width, their general shape, and the shape of the notch (see figs. 187 and 188). Length—6.5 to 7.0 mm. Tips of elytra with distinct angles. Color and maculation—Upper side red; pronotum with spots I to 4 or I to 6 (see fig. 4). Elytra each with six black spots. Nos. 1 and 5 close to the suture or actually touching it, No. 4 touching the margin. In the specimens with the least development of spots, Nos. 1, 2, 3, and 5 are subcircular, No. 2 smallest. No. 4 is widened laterally, No. 6 wedge-shaped with the thicker end toward the suture. In the darker specimens all spots, except 2, show a tendency to flow to- gether in the direction of var. connecta apparently with all intergrades occurring. Under side light, most of metasternum and part of abdomen dark. Type locality.—Southern India, Pondicherry. Material examined.—4 specimens (MNH): Southern India, Co- imbatoro District, Valparai, alt. 3,500 feet, July 7, September 3-15, 1937 (2. S. Nathan). Remarks—This species is very close to sparsa Herbst and would be difficult to separate from it by the genitalia, which are very similar. However, the tendency of the elytral spots to fuse in a very charac- teristic way makes it easy to recognize it. The separation from NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 69 doryca Boisduval, which particularly its darker forms (connecta) resemble, is probably safely made from the locality alone, as doryca is found only on New Guinea and vicinity. See also the remarks after doryca. EPILACHNA GANGETICA var. CONNECTA, new variety FIGURE 41 Differs from gangetica's. str. only in having all spots except No. 2 interconnected as indicated in figure 41. No. 6 often remains iso- lated. Pronotal spot 7 may be present in addition to the other ones. This variety resembles very closely the ground form of doryca Bois- duval and often has been confused with it. The differences in the spot patterns of the two species as indicated by the differences be- tween figures 41 and 42 are probably typical and would justify keep- ing the two species apart even without the differences in the genitalia. Material examined.—Type: AMNH., Southern India, Coimbatoro District, Valparai, alt. 3,500 feet (P. S. Nathan). 3 paratypes: I from type locality; India, Madura District, Palni Hills, Kodaikanal, alt. 5,000-7,000 feet, June 1915 (L. V. Newton, PA); Madura (U.S.N.M. No. 57123). 20. EPILACHNA DORYCA (Boisduval) FicurEs 42, 187 C[occinella] doryca BotspuvAL, 1835, p. 597. Epilachna doryca WEISE, 1902, p. 493; 1908, p. 218. Abdomen.—Abdominal lines subcomplete, angulate, reaching to within about one-fourth (about 0.15 mm.) or less of the hind margin of the first segment. Fifth segment, male, hind margin truncate to slightly concave; female, truncate. Sixth segment, male, convex, very slightly emarginate; female, split. Tergite VIII of female with a distinct shallow notch in the middle of hind margin. ' Male genitalia—Very similar to those of E. philippinensis (fig. 118). Penis seen in profile, distinctly bent upward at about three- fourths of its length so that both upper and lower margin show this upward bend. Thickness very slightly increased just after the bend and then decreasing, ending in apical hook. Small basal knife edge. Two short rows of hairs 0.1 to 0.2 mm. long after bend on upper side; a few isolated longer hairs on basal end. Paramera with apical thorn, 1.5 mm. long. Sipho ends in sharp point as in sparsu. Female genitalia——Genital plates rather narrow, length 0.5 mm., 7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 greatest width 0.31 mm. The notches near base of inner margin subrectangular in shape. Length.—5.8 to 7.0 mm. Apical angle of elytta distinct. Color and maculation.—Upper side brownish red. Pronotum im- maculate or with spots I to 4. Elytra in the typical specimens with points I, 3, 4, 5, and 6 confluent so that the elytra have somewhat the appearance of a black background with four red spots, two on the suture, one humeral surrounding black spot 2, and one lateral. The last may be very small in dark specimens or even completely disap- pear (Weise, 1902, p. 493). One of the New Guinea specimens has the basal margin black so that spot 2 also is joined to the others. In addition the lateral margin is also black. Under side and ap- pendages light with the metasternum more or less dark. Type.—Paris Museum. Type locality—Dorei (northwestern New Guinea). Remarks—The maculation may be less connected than in the typical form, and there are specimens with six entirely uncon- nected spots on each elytron. All the partly connected forms belong to var. erimensis. EPILACHNA DORYCA var. ERIMENSIS Weise Epilachna doryca var. -erimensis WEISE, 1902, p. 493. Type locality—Erima (on Astrolabe Bay, New Guinea). Under this name Weise designated those forms of Epilachna doryca Boisduval in which elytral spots I, 3, 4, 5, and 6 are not at all (rare) or only partly united. This variety is not sharply separated from the typical form, and intermediate forms occur. There is evidence that some of these have the character of subspecies. The specimens taken from one locality are all very similar among themselves but differ from those taken at different places. There seems to be a definite systematic change with the geographical location which would make the forms true subspecies, but there is not enough material to define the subspecies. Material examined (doryca s. str. and var. erimensis).—37 speci- mens (NM, MCZ, AMNH, D), as follows: 4 typical specimens, New Guinea, New Britain. 18 specimens var. erimensis, Solomon Islands, Bougainville, July- September 1944 (A. B. Gurney), on eggplant in garden. One speci- men with all spots free, the others with spots I+3 or 1+3+4 connected. 7 specimens var. erimensis, Dutch New Guinea, Hollandia, Jan- NO. 15 LADYBEETLES OF THE GENUS EPILACH NA—DIEKE yl uary, April, May, 1945 (B. Malkin). Mostly 1+3+4 connected. Also 1+3+4, and 5+6, one specimen 3+4 only, one, I1+3+4+5. 8 specimens var. erimensis, Solomon Islands, Waimoni, San Cristoval (W. M. Mann) ; all these have only spots 1+3 connected. Remarks—There has been some confusion concerning EF. doryca Boisduval and other related species. Boisduval (1835) described the species from Dorei, which is in the northwestern part of New Guinea. Mulsant (1850, p. 761) redescribed the species but took as typical specimens not Boisduval’s type, which he saw in the Paris Museum, but specimens from the East Indies, which had all the dark spots separated. Gorham (1903) reiterated the statement that the specimens from India and New Guinea were indistinguishable, whereas Weise (1908, p. 218), on the contrary, recognized them as separate forms and called the Indian forms var. gangetica, which con- ception was taken over into the Korschefsky Catalog (1931, p. 28). The genitalia of the New Guinea and Southwest Pacific forms, the true doryca, are quite distinctly different from the genitalia of the Indian specimens, so that the latter must be regarded as a separate species with the name gangetica Weise. E. gangetica Weise and dorica Mulsant are identical, but of course the latter name is pre- occupied by doryca Boisduval. The genitalia of E. doryca Boisduval are so similar to those of E. philippinensis that we may have to regard the latter as a subspecies of the former. However, the way in which the elytral spots coalesce allows us to keep the species apart. When in philippinensis and p. remota two spots coalesce we find invariably that the two coalescing spots are 3+4. In E. doryca when only two spots coalesce we find them to be 1+3, in darker specimens I1+3+4, then 1+3+4+5+6. The last is the typical form. However, two specimens from New Guinea were noted with only spots 3+4 united. This seems to strengthen the assumption that doryca and philippinensis are not distinct species. More material from the islands between New Guinea and the Philippines probably will clear up this problem. 21. EPILACHNA 11-VARIOLATA (Boisduval) FIGURES 43, 44, 136, 183 CLoccinella] undecemvariolata BotsDUVAL, 1835, p. 591. Abdomen.—Abdominal lines complete, subangulate, reaching to within one-fourth of the hind margin of the first segment. Fifth segment, male and female, truncate. Sixth segment, male, hind mar- gin convex, slightly emarginate; female, split. 72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Male genitalia—Penis seen in profile, length 1.2 mm., lower edge straight for four-fifths of its length, then curved up; upper edge with distinct basal knife ridge, then curved in a gentle arc with concave side up. Apical point without hook, penis without hairs or with an occasional hair; paramera I.1 mm. long, slightly wider and flattened toward apex, greatest width 0.15 mm., furnished on the rim of their apical third with a row of light blond hairs (about 0.2 mm. long). Sipho, tip rounded with an elongate orifice. Female genitalia.—Genital plates, length, 0.40 mm., greatest width 0.25 mm., shape not very different from that of 28-punctata, basal edge almost straight. Notch near base of inner edge narrow and deep, depth about 0.06 mm., very narrow at the edge, widening out into an oval shape. Length.—5.6 to 5.8 mm. Tips of elytra rounded. Color and maculation—Upper side yellowish brown, pronotum often darker in the middle. Elytra with the six persistent spots relatively large, rounded, dark brown; No. I on the suture reaching in front to tip of scutellum, No. 5 touching or almost touching the suture, No. 4 somewhat laterally extended so that it reaches the margin. In addition, several of the nonpersistent spots occur apparently to some- what variable extent. The specimen from Malino has only h present, others c,g,h1; c,d,g,ho; d,e,f,g,h, wp to all present except a and b. The nonpersistent spots are much smaller than the persistent ones. Under side and appendages light brown except sides of metasternum, which are dark, and the hind femora and base of abdomen, which are darkened in one speci- men only. Type locality—New Guinea. Material examined.—7 specimens, New Guinea, Hollandia, D.N.G., May 1945 (B. Malkin, NM, D); Celebes, Kawangosa, Malino (Brues, MCZ). Remarks.—This species resembles superficially 26-punctata or 28- punctata, which occur in the same region. It can be separated easily by the genitalia, particularly those of the female, and it seems to be closer to 28-punctata than to 26-punctata. Unless a larger series would show an unusual variability, the spot pattern is sufficient for separating it from the neighboring species. The position of spots 1, d, 5, and g on the suture is a good characteristic. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 73 22. EPILACHNA PYTHO Mulsant FIGurRES 45, 120, 185 E[pilachna] pytho Mutsant, 1850, p. 777. Abdomen.—Abdominal lines complete, subterminal, subsymmetri- cal. Fifth segment, male, truncate; female, truncate. Sixth segment, male, mildly emarginate; female, split, apical corners rounded. Male genitalia—Penis seen in profile, with lower edge practically straight until about 0.15 mm. from end, where it curves gently up. There is a very feeble emargination just before that. Length 2.0 mm. No apical hook. Upper side with basal knife blade well developed and reaching to beyond about half the length. Upper edge a gentle arc. Apical half with hairs not longer than 0.2 mm. Paramera 1.9 mm. long, narrow, greatest width near apex 0.15 mm., in apical half higher than wide; hairs on rim of apical third about 0.2 mm. long. Apical thorn present. Penis tube 0.18 mm. wide, seen from below at apex of orifice about 0.08 mm. wide, subparallel until the upward bend. Sipho normal (compressed near tip). Female genitalia.—Plates, greatest width 0.32 mm., length 0.53 mm. (Java specimen). Only very feebly emarginate on inner edge about one-third distance from base. The India specimen has shorter, the Borneo specimen longer, plates of about the same width and general shape. Length.—6.2 to 7.0 mm. Shape subhemispherical, tips of elytra rounded or with an indistinct angle. Color and maculation—Upper side light brownish red. Elytra each with six spots. No. 1 with its counterpart forming one rounded spot bisected by the suture, partly or totally enveloping the scutellum. No. 5 also on suture. No. 4 reaching to margin. Under side light. Parts of metasternum and middle parts of abdomen dark (Java and India specimens) ; abdomen all light in the Borneo and Philippine specimens. Type locality —Java, Sumatra. Material examined.—7 specimens (NM), as follows: 3 males, Philippine Islands, Mindanao, Davao (Baker) ; Luzdén, Paete. 4 females, Java, Buitenzorg, March 1909; Depok, 1909 (both Bryant and Palmer) ; India, Goa, Murmugao, June 1925 (J. C. Brid- well) ; Borneo, Sandakan (Baker). Remarks.—There can be little doubt that this is Mulsant’s pytho, although of course no absolute guarantee can be given that the males and females belong to the same species. They show no essential dif- ferences. The female genitalia of specimens from widely different 74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 localities are somewhat different in proportion and may indicate several subspecies. 23. EPILACHNA PERPLEXA, new species FIGURE 49 Abdomen.—Abdominal lines subcomplete, subterminal. Fifth seg- ment, male, truncate to concave. Fifth segment, female, subtruncate with a slight process in the middle. Sixth segment, split, with the apical corners rounded. Female genitalia—Plates, 0.47 mm. long, greatest width (0.29 mm.) slightly beyond the middle; rounded at base and apex, inner edge nearly straight, with a very slight emargination at about two- fifths distance from the base. Length—6 to 6.5 mm. Apex of elytra rounded. Coarse and fine punctation of elytra both very distinct. Color and maculation—Upper side red. Pronotum with a discal spot, which is due to the confluence of spots 3+4+7 (fig. 4F). Elytra each with six black spots, Nos. 1 and 5 on the suture, forming with their counterparts two common round spots, No. 1 partly en- veloping the scutellum. No. 4 broadly touching the side margin, laterally widened and just touching No. 3 in one specimen. No. 3 subcircular. No. 6 also widened and touching the margin. Pubescence light gray, dark on the spots. Under side and appendages light, except most of metasternum, the middle of the basal part of abdomen, the epipleurae adjacent to spot 4, and the tips of the mandibles dark. Material examined.—Type: U.S.N.M. No. 57124, Siam, at Karen settlement on headwaters of Me Ka, south of Moi Chiengda, March 16,1927). -G) Deienan): Paratype (D): India, Madura. Remarks.—This species is very close to pytho Mulsant. The dif- ferences are chiefly in the greater extent of the black pigment on pronotum and elytra. The female genitalia in the two are similar, though there are differences in the exact shape of the plates and the shape of the emargination. In the absence of the male the exact status of perplexa will have to be left indefinite. 24. EPILACHNA PYTHARGA, new species Ficures 46, 186 Abdomen.—Abdominal lines complete, subterminal, subangulate. Fifth segment, female, truncate. Sixth segment, split, apical angles of parts distinct. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 75 Female genitalia—Plates, 0.43 mm. long, greatest width 0.28 mm. Shape oval. Notch near base of inner edge deep, widening away from the edge. Length—6 to 6.2 mm. Tips of elytra rounded or with indistinct angles. Color and maculation.—Like pytho except that spot No. 5 only touches the suture and is not on it. Material examined —Type: U.S.N.M. No. 57125, Philippine Is- lands, Luzon, Mount Banajao (Baker). 2 paratypes (NM, D): One, same data as type; the other Luzon, Mount Maquiling (Baker). Remarks.—This species resembles superficially E. pytho Mulsant. It is somewhat smaller, a little more elongate. It can be recognized by the fact that spot No. 5 does not form with its counterpart one spot bisected by the suture, as in pytho, but just touches the suture. The female genitalia of the two species (figs. 185 and 186) are quite distinct and will give a definite separation in cases of doubt. There are several specimens from Mindanao, also females, which have a similar appearance and similar structure of the female geni- talia but are considerably larger in size. In the absence of the males, it seems impossible to say whether they belong to the same or to a related species. They come from Davao (Baker, NM) and Mount Apo, Galog River, 6,000 feet, November (C. F. Clagg, MCZ). 25. EPILACHNA HEMISPHERICA, new species FIGURES 47, 130 Abdomen.—Abdominal lines complete, subterminal. Fifth segment, male, hind margin slightly concave; female, approximately straight. Sixth segment, male, truncate; female, split. Male genitalia—Penis seen in profile, about 1.7 mm. long, gently curved up in the middle on both upper and lower edges, sharply bent up just before the sharp apical point. Basal knife edge practically absent. Apical three-fifths clothed with long (about 0.5 mm.) and dense blond hairs. Seen from below, gently tapering from base to apex, about 0.27 mm. wide in middle; long, oval-shaped orifice. Para- mera 1.7 mm. long, flattened and of almost uniform width (maximum 0.2 mm.). Apical thorn absent or only rudimentary. Apical three- fifths of rim sparsely and apex densely clothed with long hairs (about 0.4 mm. long). Sipho ending in straight point, orifice subterminal at side with liplike process (as in fig. 126). Female gemtalia.—Plates, 0.53 mm. long, 0.38 mm. wide; shape 76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 almost rectangular with the basal side rounded off. A slight process on inner edge near base. Length—7 to 9.5 mm. Apical angles of elytra indistinct. Color and maculation.—Upper side yellowish red to brick red. Pronotum spotless. Elytra each with six black spots, Nos. I and 5 on the suture, No. 4 touching the margin and reaching beyond the middle of the width of the epipleurae. No. 1 practically reaching the base and thus enveloping the scutellum, which remains light. Material examined.—z24 specimens (NM), as follows: Type: U.S.N.M. No. 57126, from Philippine Islands, Luzon, Mount Maquiling (Baker). 17 paratypes from the type locality and the following other local- ities in the Philippine Islands: Luzon, Laguna Prov., Mount Banajao (Ube, Baker, McGregor); Majayjay, June 25, 1928, March 31, 1929 (McGregor) ; Tayabas Prov., Malinao (Baker) ; Lucban, Ta- yabas (McGregor) ; Bho. 6 additional specimens: Mindanao Island, Davao, Iligan, Surigao (all Baker). 26. EPILACHNA SIGNATIPENNIS (Boisduval) FIGURES 50, 132, 184 C[occinella] signatipennis BoIsDUVAL, 1835, p. 593. E[pilachna] signatipennis MuLSANT, 1850, p. 764. Abdomen.—Abdominal lines complete, rounded, reaching to within one-third of hind margin of first segment. Fifth segment, male, hind margin truncate or slightly concave; female, with a very slight, broad process. Sixth segment, male, convex; female, split, with distinct apical angles. Male genitalia—Penis seen in profile, 1.1 mm. long, gently curved up, with sparse hairs (about 0.3 mm. long) even on basal part. Basal knife edge rudimentary. Seen from below, about 0.25 mm. wide, tube split open in middle with seams apart for most of the length. Triangular beyond orifice. Paramera 1.0 mm. long, slightly bent outward near base, then of almost uniform width (0.12 mm.) to apex, with small apical thorn; sparse pubescence on apical half. Sipho with tip strongly emarginate or bifurcate. Female genitalia—Plates, length 0.5 mm., greatest width 0.3 mm.; sides and base rounded. Notch near base of inner edge nearly a complete circle. Length.—5.6 to 7.5 mm. Tips of elytra subrounded. Color and maculation—Upper side yellowish red to brick red. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE IE Pronotum with the sides lighter, usually spotless, but occasionally with a median spot (3+4+7). Very dark specimens have most of the pronotum black except the sides. Elytra with six spots, which are partly confluent. In the lighter specimens only Nos. 3 and 4 united to form a transverse band, broadly touching the margin. Nos. 2 and 6 isolated, more or less rounded, No. 1 on the suture forming with its counterpart on the other elytron one rounded spot, which reaches forward to about the tip of the scutellum. No. 5 rounded, touching or almost touching the suture. Figure 50 shows an average specimen. In the darkest specimens, No. 1 extends to the base, No. 2 touches base and margin. No. 5 is also united with No. 4 and touches broadly the suture. Occasionally there is a narrow sutural margin left light. No. 6 has a tendency to touch Nos. 4 and 5. The apices of the elytra also may be dark (spot /) and this apical spot most often is con- nected with No. 6. Under side from almost completely light to almost completely black. Type locality—New Guinea and Waigiu Island. Material examined.—133 specimens, New Guinea (G. Compere, NM; H. Edwards, AMNH), Stephansort, Mount Hagen, December 10, 1944 (D. G. Hall) ; New Britain, Mafor Island (also called Mefur or Numfor Island, lat. 1°S., long. 135°E. It is one of the group of Schouten Islands at the mouth of Geelvink Bay, northwestern New Guinea, which are also sometimes called Mysore Islands). There is a good series (114 specimens) from Hollandia, Dutch New Guinea, January, May, 1945 (B. Malkin, NM, D). There is not a great deal of variation in the spot pattern in this series, which is like figure 50, except that usually the apical spot h is present and united with No. 6. The pronotum is usually spotless but occasionally shows the discal spot faintly to distinctly (as in fig. 4F). Its width is never more than one-fifth of the width of the pronotum. Remarks.—E. signatipennis can easily be distinguished from other species either by the type of maculation or the male and female geni- talia. Light specimens resemble some forms of sparsa but have spot I always on the suture. Dark specimens resemble superficially solo- monensis, new. 27. EPILACHNA DIVERSA, new species Ficures 48, 134, 196 Abdomen.—Abdominal lines complete, subterminal, broadly rounded. Fifth segment, male, hind margin slightly concave; female, 6 78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 truncate. Sixth segment, male, very slightly emarginate ; female, split. apical corners narrowly rounded. Male genitalia—Penis seen in profile, from base to tip 1.1 mm. long, gently curved up until about 0.4 mm. from end, where it is bent suddenly almost 90° ; and just before tip it is bent a little more. Upper side more uniformly curved until just before tip. Sparsely clothed with rather long hairs (about 0.35 mm.) on the basal two- thirds. Basal knife edge indistinct. Seen from below, rather broad, 0.4 mm. near base, slightly narrowing toward the orifice, which is at the bend. A longitudinal slit in the middle about 0.06 mm. wide at the base and gradually narrowing so that near two-thirds distance toward orifice the inner edges touch. Orifice rounded, about 0.16 mm. in diameter. Part beyond orifice a triangle 0.2 mm. long, base 0.15 mm. wide. Paramera 1.0 mm. long, subparallel 0.18 mm. wide, with apical thorn; apical half clothed with light blond hairs about 0.2 mm. long. Sipho long, shape of a question mark, orifice terminal. Female genitalia—Apical margin of tergite X convex; plates broad, length 0.42 mm., greatest width 0.45 mm. Emarginate at apical ends where stylus is attached. The notch near the base of inner edge hardly affects the dorsal side so that it appears as an oval depression on the ventral side. Length—6.2 to 7.0 mm. Tips of elytra rounded. Color and maculation—Upper side light reddish brown, head, pronotum, and scutellum black. Elytra each with six large black spots, making the distance between the spots much smaller than the diameter of the spots. No. 1 touching base and suture, No. 5 on the suture, No. 4 touching the margin. Under side black, mouth parts, antennae, tarsi, epipleurae, and tips of femora and tibiae light. Material examined.—Type: U.S.N.M. No. 57127, Philippine Is- lands, Luzon (R. C. McGregor). Paratype: Luzon, Benguet Prov., Baguio (Baker). Remarks.—This species is easily recognized by its external ap- pearance. Its genitalia are also distinctive and considerably different from the usual type of this group. EPILACHNA DIVERSA subsp. TAYABASA, new subspecies This subspecies is based on a specimen with the same structure of the genitalia as diversas. str. It differs by the coloration and macu- lation, upper side light brownish red, head and pronotum completely light. Elytral spots much smaller than in diversa s. str., all more or less rounded, No. 4 touching the margin broadly, No. 5 just touching the suture. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 79 Type—vU.S.N.M. No. 57966, from Philippine Islands, Tayabas, Malinao (Baker). 28. EPILACHNA BOISDUVALI Mulsant FIGURES 5I, 52, 135, 198 E[pilachna] Boisduvali MuLSANT, 1850, p. 765. Abdomen.—Abdominal lines subterminal, complete, and forming a regular arc. Fifth segment in both sexes subtruncate to slightly concave. Sixth segment, male, subtruncate with a slight emargina- tion; female, split. Male genitalia.—Penis seen in profile, a straight, slightly tapering, approximately parallel tube for nearly three-fourths of its length, then bent upward, and bent a little more just before the apex which is a point, with two rows of sparse hairs on the middle half. Seen from below, tube closed for about 0.9 to 1.0 mm., then opening, the sides of the orifice with rounded edges; about 0.2 mm. wide in mid- dle. Paramera 1.1 mm. long with small apical thorn; greatest width inside near rim 0.2 mm., rim covered with hairs on apical half. Sipho with bifurcate tip. Female genitalia—Plates, oval with convex inside edge; length 0.58 mm., greatest width 0.48 mm. Tenth tergite with emargination at apex, strip along each side with brown pigment, middle clear. Length.—7 to 7.6 mm. Tips of elytra rounded. Color and maculation—Upper side yellowish red. Pronotum in the Australian specimens with a central spot, which in lighter speci- mens can be seen to originate from the confluence and expansion of the spots 3, 4, and 7. The Philippine specimens have the pronotum spotless or the spots only very faintly indicated. Elytra each with six spots. Nos. I, 3, and 5 have usually approximately the same dis- tance from the suture. However, in some Philippine specimens, No. 3 is definitely farther from the suture than 1 and 5 (fig. 51). Tan- gent to No. 1 goes about through tip of scutellum or a little in front of it. No. 4 transversely widened and linked with margin. Under side light, the sides of the metasternum and often the whole metasternum and part of the abdomen darker. Type locality—Australia (?). Material examined.—t12 specimens, as follows: Australia: Queensland, Cairns, Babinda, October 1920 (J. F. Iil- ingworth, BBM). Philippine Islands: Luzon, Bataan Prov., Mount Banajao (Baker) ; Mount Maquiling (Baker) ; Limay, San Andales; Mindanao Island. 80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Bukidnon Prov., Tangcolan (Baker, NM); Davao (C. S. Banks, MCZ); Tayabas Island, Lucban, May 1926 (McGregor, NM); Leyte Island, Jaro, October 1915 (MCZ). Sumatra: Pagacamba (C. T. and B. B. Brues, MCZ), large elytral spots; disk of pronotum dark. Remarks.—The Philippine specimens agree very well with the description and figure of E. nativitatis Arrow (1900) from Christmas Island. As there are only very few species with the third spot not much farther from the suture than the first and fifth, the probability is very great that nativitatis Arrow is identical with boisduvali Mul- sant. The only difference between the Australian and Philippine specimens seems to be the absence of the pronotal spot in the latter, which is too insignificant even to consider them a separate subspecies. Should this difference, however, be consistent and the identity with Arrow’s species be confirmed, the Philippine and Christmas Island specimens might be separated from the Australian ones as subsp. nativitatis Arrow. EPILACHNA BOISDUVALI subsp. CHABUANA, new subspecies A pair of specimens of Epilachna boisduvali from Assam, India, are sufficiently remote from the typical form to deserve to be classi- fied at least provisionally as a new subspecies. Both male and female genitalia are indistinguishable in structure from those of the typical boisduvali. In appearance the Assam speci- mens present the following differences: Size smaller (length 5.5 to 6.0 mm. as against 7 to 7.6 mm. of the typical form), elytral spots relatively smaller. Pronotum of the female with four large not very distinct spots arranged in a transverse row. The individual spots cover longitudinally about the central two-thirds. Color darker, which may, however, be due to the method of preservation. Type.—U.S.N.M. No. 57128, India, Assam, Chabua, April 20, 1943 (D. E. Hardy). One paratype, same data. Remarks.—These two specimens are of great interest, as they show that E. boisduvali is a very widespread species, ranging from northern India to the Fiji and Samoa Islands. The female specimen is so far the only example in which the arrangement of spots on the pronotum does not fit into the standard pattern of figure 4. It resembles much more the pattern of Afidenta mimetica. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 81 EPILACHNA BOISDUVALI subsp. FIJIENSIS Crotch FIGURE 53 Epilachna Montrouszieri var. fijiensis CrotcH, 1874, p. 89. This form differs from the typical specimens only in a somewhat larger sizejand an enlargement of the elytral spots. No. 3 is sub- wedged-shaped, with the narrow end toward the suture, and No. 5 is triangular, with the base parallel to the suture. Also the pronotal spot is more developed. Material examined.—5 specimens, as follows: Fiji Islands: Ovalau Island, Levula (Abbott, MCZ) ; Thawati, July 12, 1938; Viti Levu Island, near Nandarivatu, 2,500 feet, September 15, 1938 (E. C. Zimmermann, BBM). New Hebrides: Espiritu Santo Island, September 1944 (K. L. Knight, NM). This specimen is different from the Fiji specimens in that spots 3 and 4 are joined to form an oblique fascia which does not reach the suture. It may be the representative of a new subspecies. EPILACHNA BOISDUVALI subsp. SAMOANA, new subspecies FIGURE 54 Only one female of this form is available. Except for the elytral pattern, it resembles E. boisduvali in practically all respects, including the female genitalia. It is therefore likely that this form is a sub- species or perhaps even only a variety of E. boisduvali. The elytral spots tend to be longitudinally enlarged in contrast with the trend to transversal enlargement present in boisduvali s. str. and its sub- species fijiensis, and this would ordinarily suggest a separate species. In the absence of more material, particularly males, no definite deci- sion can be reached. Color and maculation—Upper side reddish yellow; pronotum with a basal dark spot, which represents the coalescence of the usual spots 3, 4, and 7. Elytra with the dark spots more or less longitudi- nally elongated. This is most evident in Nos. 2 and 5, least in 1 and 4. In the latter a tendency to lateral dilatation toward the margin is apparent. Spot 3 is considerably farther away from the suture than Nos. I and 5. Nos. 3 and 5 touch. Type-——Female, Samoa Islands, Tutuila, Vaitogi, February 1930 (D. T. Fullaway, BBM). 82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 29. EPILACHNA MOULTONI subsp. MANUSENSIS Korschefsky Epilachna moultom subsp. manusensis KORSCHEFSKY, 1933a, p. 236. Abdomen.—Abdominal lines complete, subterminal, inner edge broadly rounded, outer one straight. Fifth segment, hind margin truncate or slightly concave. Sixth segment, male, hind margin con- cave; female, split, apices with rounded angles. Male gemtalia—Very similar to those of E. boisduvali (fig. 135) with no significant differences. Female genitalia—Of the same general shape as those of E. bois- duvali Mulsant (fig. 198). Length 0.56 mm., greatest width 0.50 mm. There is a sharp narrow transverse depression near the base of the inner edge which seems more indistinct in boisduvah. Tergite X with pigment divided on a strip on each side. Length—6 to 7 mm. Tips of elytra rounded. Color and maculation.—Upper side yellowish red. Pronotum spot- less. Elytra each with the following black spots: Nos. 1 and 2 united into one broad spot reaching from near the suture to the callus and touching the base in the darker specimens, but not the side margin; scutellum left light; No. 3 small; No. 4 broadened and reaching to the middle of the epipleurae; No. 5 on the suture; No. 6 rounded and relatively smaller. Spot 3 separate in the lighter specimens, joined to No. 4 and No. 5 in the darker ones. Pubescence yellowish, dark on the spots. Under side and appendages light except for tips of mandibles, parts of mesosternum and metasternum, and the middle of the abdominal segments, which are often dark. Type——Museum Bremen and Korschefsky collection. Type locality—Admiralty Islands, Station Manus. Material examined.—g specimens (NM, D): Admiralty Islands, December 23, 1944 (P. T. Riberd). Remarks.—The differences in structure between FE. moultoni manu- sensis and boisduvali are so slight that eventually moultoni may turn out to be a subspecies of boisduvali. The confluence of spots I and 2 permits an easy identification of moultont. EPILACHNA MOULTONI subsp. MOULTONI Crotch Epilachna Moultoni Crorcu, 1874, p. 89. Three specimens of moultoni probably belong to the typical form, although two of them would have to be classed under subspecies manusensis Korschefsky if the author’s definition of this subspecies is accepted. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 83 Korschefsky classified all specimens with spot 3 completely miss- ing as moultont moultoni Crotch, whereas all the forms with spot 3 present were called by him subsp. manusensis. In subsp. manusensis, spots 3, 4, and 5 may be partly or completely united. The three specimens from the Solomon Islands differ from those from the Admiralty Islands listed under subspecies manusensis Kor- schefsky by their larger size (length 8 mm.), by the fact that the apex of the abdominal lines is more rounded, and by the smaller size or absence of spot 3. Even though spot 3 is present in the specimens from Guadalcanal, they resemble so much in general appearance the specimen from San Cristoval, which conforms entirely to Crotch’s description, that they must be considered as belonging to moultoni s. str. Unfortunately, Crotch does not mention the size of E. moultoni in the original description. Type locality—New Caledonia. Material examined.—3 specimens, all females, from Solomon Is- lands: San Cristoval, Pamua (W. M. Mann, MCZ), spot 3 missing completely, 4 large, 5 and 6 small; Guadalcanal, August 15, 1943 (P. W. Oman, NM), spot 3 present, small, spot 4 touching the suture. 30. EPILACHNA BAGUIANA, new species FIGURES 55, 133, 192 Abdomen.—Abdominal lines entire, subterminal, subangulate. Fifth segment in both sexes with hind margin subtruncate. Sixth segment, male, hind margin convex, slightly flattened; female, split. Male genitalia—Smaller than in most other species. Penis seen in profile, 0.95 mm. long, only a rudimentary basal knife edge; upper edge curved almost from the base in a wide arc concave side up, point without hook, hairs on basal half. Seen from below, tube nar- rowing regularly from base to apex, closed part short, about 0.35 mim. long; orifice ovaloid, about 0.35 mm. long. Paramera 0.9 mm. long, slightly bent outward near base, from then on flattened with parallel sides 0.15 mm. wide. Rim of apical third with light blond hairs (about 0.2 mm. long). Sipho with tip rounded, orifice oval. Female genitalia——General shape and size as in E. sparsa and allied species. A triangular notch on the basal half of the inner edge distinguishes baguiana from all other known species. Length—7 to 8 mm. Sutural angles of elytra subdistinct. Color and maculation—Upper side brownish or yellowish red. Pronotum with all seven black spots (fig. 4G), No. 7 small or often 84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 absent, I and 5 (resp. 2 and 6) confluent. Elytra each with six large spots, No. 3 not appreciably farther from the suture than I and 5. The dimensions of the spots much larger than the distance between them, No. I smallest, 4 not touching the margin. Under side mostly dark, except prothorax and the two last sternites. Epi- pleurae and appendages, except tip of mandibles, light. Femora with elongate dark spots in middle. Material examined.—Type: U.S.N.M. No. 57129, Philippine Is- lands, Luzon Island, Baguio, Benguet (Baker). 4 paratypes (NM): Same locality (G. G. Haslam, Baker). One additional specimen: Mindanao Island, Zamboanga (MCZ). Remarks.—This species is well characterized by its spot pattern and its male and female genitalia, and so confusion with any other species is not likely. 31. EPILACHNA SEXTA, new species FIGURE 219 Abdomen.—Abdominal lines complete, subterminal, subangular, outside edge straight. Fifth segment, male, hind margin straight or slightly concave ; female, slightly convex. Sixth segment, male, emar- ginate; female, split, tips rounded. Male genitalia—Penis seen in profile, 1.5 mm. long, only trace of basal knife edge; widest at base, nearly straight for basal two-thirds, upper edge then curved to the apical point, which points straight up ; lower edge bent at two-thirds of its length and straight again until before the apex where it is bent up through practically go°, covered with hairs (0.4 to 0.5 mm. long) on its middle portion. Seen from below, tube widest at base and becoming narrower toward orifice, which is elongate; width at center of orifice about 0.15 mm.; part beyond orifice narrow, about 0.03 mm. wide, subparallel but slightly widening toward apex. Length from orifice to apex about 0.3 mm. Paramera 1.5 mm. long, greatest width (near apex) 0.15 mm., slightly curved outward, with blunt apical thorn; rim of apex and apical half of outside covered with blond hairs. Sipho, regular shape (fig. 117), orifice oval, subterminal; apical 0.4 mm. compressed. Female genitalia—Plates, 0.65 mm. long, greatest width 0.40 mm. ; shape roughly that of ‘figure 186. On the inside edge about one-third from base a strong depression, tergite X pointed. Length—o.65 to 0.85 mm. Tips of elytra rounded. Color and maculation—Upper side light brownish red, head and pronotum spotless. Elytra with six black spots. Nos. 1 and 5 on NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 85 the suture in the darker specimens, not quite reaching the suture in the lighter ones. No. 2 rounded, No. 4 broadly on the margin in the darker specimens, just reaching the margin in the lighter ones. Under side light except tips of mandibles, part of metasternum and the middle part of the first four abdominal segments, which are.dark. In the lighter specimens abdomen completely light. Material examined.—Type: Museum of Comparative Zoology, from Celebes, Molino (Brues). 6 paratypes (MCZ, D, all Brues) from the type locality and from Celebes, Langsa (one marked July 1937). Remarks.—This is another one of the 12-spotted species that are practically unseparable from related species except by their genitalia. 32. EPILACHNA LIBERA, new species FIGURES 59, 60, 138 Abdomen.—Abdominal lines complete, rounded, reaching to within one-fifth of the hind margin of the first segment. Fifth segment, male, truncate ; female, slightly convex. Sixth segment, male, convex ; female, split, apices rectangular. Male genitalia.—(Inasmuch as the only male specimen is apparently a somewhat distorted immature specimen, the actual shape of the undistorted genitalia might be slightly different.) Penis seen in profile, 1.7 mm. long. Basal knife edge present. Lower edge straight until shortly before apex, where it is strongly curved up through about go°. A slight emargination just before this bend and a slight bulge on the corresponding part of the upper edge. From two-fifths to four-fifths of its length the upper edge densely covered with two rows of long (0.6 mm.) light blond hairs. Penis seen from below, tube with elongate orifice without definite front margin. The part beyond the orifice long (0.5.mm. or longer), constricted immediately beyond orifice, then widened again, to about 0.1 mm. and then gradually narrowing to a blunt point. Paramera 1.6 mm. long, widened toward apex (maximum width 0.2 mm.), densely covered with rather long hair. Sipho with orifice small, terminal. Female genitalia.—Genital plates, length 0.60 mm., greatest width 0.36 mm., shape like 28-punctata. Notch near base of inner edge transversely oval, not cut all the way through; edges of the notch darkened. Length. to 8.5 mm. Tips of elytra with angles distinct. Color and maculation—Upper side brownish red. Pronotum with 86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 hazy dark spots, the lateral ones, Nos. 1 and 5, better developed than the central ones. Elytra with the persistent spots large and often ynited. No. 1, on the suture, triangular, enveloping the scutellum. Nos. 3 and 4 united in all three specimens, No. 5 rounded, practically touching the suture; No. 6 laterally widened, slightly bent, with con- cave side to rear. Spots 2, 4, and 6 practically touch the margin and leave only the narrow reflexed part light. Spots I and 2 may be united, forming a broad fascia with a light spot on each side of the scutellum enclosed, -spots 4+3+5 are completely united in two of the specimens. In addition to these persistent spots, several of the nonpersistent spots may also be present: d+e and g+h. In one specimen e is united with g+h. Pubescence light gray all over. Under side and appendages light, metasternum black, basal part of abdomen and sometimes other parts of under side dark. Material examined—Type: U.S.N.M. No. 57130, China; Szech- wan Prov., Kuanhsien to Uen Chuan, July 4-7, 1924 (D. C. Graham). 2 paratypes: Shin Kai Si, Mount Omei near Kiating, alt. 4,400 feet, 1921; and near Kuanhsien, alt. 2,000-4,000 feet, August 1933 (D. C. Graham, NM). Remarks.—This species is easily recognized by its maculation, which does not coalesce in this particular way in any other species known to me, by the uniformly light gray pubescence also in the black spots (this is shared by some other forms, such as sparsa and niponica, occurring in the same territory), and particularly by the structure of the male genitalia. 33. EPILACHNA DELESSERTII (Guérin) Ficures 58, 137 Coccinella (epilachna) Delessertii GUERIN, 1840, p. 42. E[pilachna] Delessertii MULSANT, 1850, p. 747. Abdomen.—Abdominal lines complete in the two females; in- complete, reaching only to the hind margin of the first segment in the male; subterminal. Fifth segment, male, slightly concave; female, truncate with depression in the middle. Sixth segment, male, emar- ginate; female, split, apical corners rounded. Male gemtalia.—Penis seen in profile, with a distinct bend at about two-thirds of its length, this bend more distinct on the under side than on the upper side; bent again about 0.2 mm. before the end, without, however, forming a definite hook. No basal knife edge, sparsely pubescent between the two bends. Seen from below, orifice elongate with pointed apical margin at about the position of apical NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 87. bend. Paramera 1.5 mm. long, gradually widening toward apex and slightly bent down on apical third; greatest width about 0.14 mm., no apical thorn; pubescence on apical third rather short. Sipho, bent mildly at about one-third of its length, bent outward about 0.35 mm. from end. Orifice small, oval, about in the middle of the side of this last part. Female genitalia—Plates rounded at basal end, more or less trun- cate at apical end. No notch on inside edge. Length—7 to 8.5 mm. Elytra prominently margined, a subapical depression near the margin which makes the reflexed part very wide there. Tips of elytra rounded. Color and maculation.—Upper side brownish red. The elytra with all the spots interconnected so that a grating results that can best be described by considering the ground color of the elytra black, each with five large red spots 2, 2, 1, the first four touching neither suture nor margin, the fifth apical. Under side light, metasternum, part of mesosternum and prosternum, and abdomen, except its sides, dark. Epipleurae partly dark. Type locality —East Indies, Plateau of Nilgherry. Material exanuned.—3 specimens: India: Madura district, Kodai- kanal. Palni Hills, alt. 5,000-7,000 feet, June 1915 (L. V. Newton, C. Leigh, PA) ; Trichinopoli. Remarks—tThe difference in the abdominal lines between male and female is peculiar and rather pronounced. Since the specimens are alike in all other respects, it is reasonably sure that they belong to the same species. Whether this is really a sexual difference or an accidental variation only more material can decide. 34. EPILACHNA SOLOMONENSIS, new species FIGURES 61, 139 Abdomen.—Abdominal lines complete, subterminal, somewhat flat- tened at apex. Fifth segment, male, hind margin mildly concave; female, truncate. Sixth segment, male, subtruncate; female, split. Male genitalia——Both male and female genitalia very much like those of E. 28-punctata (Fabricius). There are differences, but they would come out only in an actual comparison of the specimens. Penis seen in profile, 1.4 mm. long, gently curved toward apex and ending in sharp point. Small but distinct basal knife edge. Penis with sparse hairs on most of its upper side. Seen from below, the usual tube, orifice with oval apex ; at the apex of the orifice the penis is 0.13 mm. wide, distinctly wider than for 28-punctata (which has there only a 88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 e width of 0.09 mm.). Paramera 1.4 mm. long, about constant width (0.I mm.), somewhat narrower just before apex ; apical thorn present, rim of apical half with hairs (about 0.2 mm. long). Sipho, tip with the same kind of notch as in 28-punctata. Female genitalia—vVery similar to figure 182. Length.—7.5 mm. Tips of elytra rounded. Color and maculation.—Upper side red, pronotum spotless, elytra with a basal fascia consisting of spots 1 and 2, extended farther back at the suture; scutellum light, sometimes with a dark border. Spots 4+3+5 form a discal fascia bent back in form of a V near the suture. Spot 6 widened and touching the margin. Pubescence short, blond; black on the dark spots. Under side and appendages light, except tip of mandibles, metasternum, and outer parts of epipleurae where the spots reach them. Material examined.—Type: U.S.N.M. No. 57131, Fulakora, Solo- mon Islands (Ch. Bignell). 5 paratypes: Solomon Islands, Guadalcanal Island, July 1927 (AMNH), 1920 (J. A. Kusche, BBM); Bougainville, July-Sep- tember 1944 (A. B. Gurney, D). Remarks.—This species is in all its morphological characters very close to 28-punctata Fabricius, particularly the notched apex of the sipho, which has not been observed in any other species. How- ever, the type of maculation immediately distinguishes the species. There is superficial resemblance to E. guttatopustulata tricincta Montrouzier, and the specimens were found labeled as such in one of the collections. The two species can, of course, immediately be separated by the genitalia. Actually, however, the superficial ap- pearance is sufficient to keep them apart. E. solomonensis is less elongate and more convex than ¢ricincta. Furthermore, in tricincta the discal fascia does not have the V-shaped bend near the suture and is besides narrowly interrupted*by the suture. The apical spot is in the apical angle of the elytra, whereas in solomonensts it is definitely removed from it. Finally, the part near the margin between the two fasciae is usually considerably lighter than the rest of the elytra in tricincta, as it is the remnant of the yellow spot in guttatopustulata. Epilachna parafasciata, the only other species of Epilachna with un- questionable fasciae, does not have the V-shaped bend of the discal fascia and besides has a different geographical distribution. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 89 35. EPILACHNA PARAFASCIATA, new species FicuRES 62, 221 Abdomen.—Abdominal plates complete, reaching to within one- third of apical margin of first segment. Fifth segment, male, trun- cate or slightly concave; female, truncate with a rounded process in middle. Sixth segment, male, emarginate; female, split, corners distinct. Male genitalia (fig. 221).—Very similar to those of E. sparsa and E. gangetica. Penis seen in profile, 1.60 mm. long, with distinct basal knife edge and apical hook; apical half slenderer than in sparsa or gangetica and without a distinct bulge as in sparsa and not wedge- shaped as in gangetica; pubescence about 0.2 to 0.3 mm. long on apical third. Seen from below, wider than in sparsa and gangetica, about 0.2 mm. at the orifice, wedge-shaped from there to apex. This last is the most easily detected difference from sparsa and gangetica. Paramera 1.6 mm. long, with apical thorn. Sipho as in sparsa. Female genitalia—Plates, length 0.60. mm., greatest width 0.37 mm., shape normal. Notch near base of inner edge with rounded edges cut in much farther on the under side than on the upper side; axis pointing toward base. Length.—7 to 7.6 mm. Tips of elytra with distinct angle. Color and maculation—Upper side light brownish red; head and pronotum spotless; elytra with two transverse fasciae and an apical spot (No. 6), which is widened and touches the margin. The first fascia touching the base, but not quite reaching the side margin and leaving the scutellum light ; wider at the suture, but only slightly so, not suddenly widened there. Discal fascia almost straight from margin to margin. Somewhat constricted about one-third distance from the margin (indicating the separation of spots 3 and 4), the front margin practically going straight across the suture. Under side and appendages light, mesosternum and metasternum, the middle of the first four abdominal segments, and the epipleurae where the mid- dle fascia reaches them, dark. Material examined —Type: U.S.N.M. No. 57132, from Java, Tyi- bodas, Mount Gede, alt. 9,500 feet, April 1909 (Bryant and Palmer). 7 paratypes (NM, MCZ, D): two from type locality (alt. not given); one from Buitenzorg, Java, March 1909 (Bryant and Palmer) ; four from Tjiboda (T. Barbour). Remarks.—This species, although very similar to others in struc- tural characteristics, can be easily recognized by its external appear- ance. The eight available specimens show no appreciable variation, go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 and there is no known Epilachna species with which it can be con- fused even with a superficial inspection. It is, however, very similar in appearance to Afissa orthofasciata, from which it can easily be separated by the generic characters and the genitalia. 36. EPILACHNA HAEMORRHOA Boisduval Ficures 63, 64, 140, 194 C[occinella] haemorrhoa BotspuvaL, 1835, p. 599. Abdomen.—Abdominal lines complete, reaching to within one- fourth of the hind margin of the first segment. Fifth segment, male, hind margin truncate or slightly concave ; female, with a longitudinal ridge in the middle of apical half, hind margin subtruncate. Sixth segment, male, hind margin, convex ; female, split, each half rounded. Male genitalia—Penis seen in profile, 1.43 mm. long, no basal knife edge, wide at base, then becoming much narrower and curved up in a gentle arc. Sparsely covered with hairs (about 0.2 mm. long) where it becomes narrower. Seen from below, with elongately dia- mond-shaped orifice with the sides rounded. Width of penis at widest part of orifice 0.16 mm., at apex of orifice 0.10 mm., part be- yond orifice about 0.33 mm. long gradually tapering to point and curved up. Paramera 1.3 mm. long, seen from above gently curved outward, slender, widest (0.13 mm.) near apex, without apical thorn, with rather short hairs on apical part. Sipho, normal shape, compressed laterally at apex, orifice oval, subterminal on side. Female genitalia—Plates together heart-shaped with heart-shaped notch near base of inner edge. In general, shape of genitalia similar to those of signatipennis Boisduval. Length.—5 to 7.5 mm. Apical angle of elytra slightly protruding inward. Basal tooth of claws triangular; epipleurae reaching prac- tically to apex, hollowed in front half. Color and maculation.—Upper side, head, scutellum, and pronotum reddish brown, sides of latter yellowish; elytra dark brown or black with a common apical red spot. This spot covers about one-fourth to one-third of the length, has its front margin arched, and usually reaches less far forward at the suture than at the margin. Hairs on elytra uniformly light gray. Under side, in the lighter specimens mostly light brown, with metasternum, hind tibiae and most of the epipleurae dark. In the darker specimens the dark color spreads so that only the mouth parts, the prothorax, the sides of the basal four abdominal segments, and the entire fifth and sixth remain light. The legs in such dark specimens are black, except the front femora ex- NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE gI clusive of their tips and small spots at the base of the middle and hind femora, which remain light.1° (The specimens from Mount Misum, New Guinea, are of this kind. The red apical spot of the elytra may be reduced then to a very small size.) Type.—Paris Museum. Type locality —Dorei, northwestern New Guinea. Material examined.—11 specimens: New Guinea, Finschhafen (NM, D) ; Mount Misum, Morobe Distr. (MCZ). Remarks.——The male genitalia of this species are very similar to those of E. mjoebergi and E. mindanaonis. It differs from these species by the type of maculation and from the latter also by the female genitalia. EPILACHNA HAEMORRHOA HOLLANDIAE, new subspecies There is a good series from Hollandia, Dutch New Guinea, which although showing considerable variation is lighter than any of typical specimens from farther west (type locality) or farther east (Finsch- hafen). The male and female genitalia show no significant differ- ences. The average size is smaller. It seems justified to consider these forms as a separate subspecies. The darkest hollandiae specimens have besides the apical red spot, which occupies about the apical third of the elytra, a humeral spot extending backward to about one-third of the length of the elytra and inward about one-half of their width. It is directed slightly inward so that the margin behind the shoulder remains dark. The under side is entirely light, with only small traces of dark left. Most specimens of the series have elytra with the light coloring more extended. Progressively the entire front margin becomes light, then also a narrow longitudinal strip at the suture from a little be- hind the scutellum to somewhat in front of the apical spot. If the red is even more extended the ground color of the elytra appears red with a few dark spots. In that case the humeral spot is united with the apical one so that the pattern consists of a marginal spot at about the middle of the margin and a longitudinal elongated spot reaching from the scutellum to the suture behind the middle but leaving the middle of the suture red. This longitudinal spot is then broken up into two or three, one, the largest, on the disk on the suture 15 The coloration of these specimens is what Weise (1903, p. 229) describes as typical for E. fulvimana Weise, which may therefore be identical with haemorrhoa Boisduval, as the other distinguishing characters are all rather vague. The type locality of E. fulvimana is New Guinea, Huon Gulf, Saddle Mountain, Simbang. Q2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 behind the middle, and a smaller one immediately behind the scutel- lum. If the scutellar dark spot is absent we have what Weise de- scribed as var. dissoluta (Weise, 1902, p. 491). The lightest specimen from Hollandia I have seen has no trace of the dark coloration left. Material examined.—Type: U.S.N.M. No. 57967, from Hollandia, Dutch New Guinea. 84 paratypes from the same locality, January, April, May, 1945 (B. Malkin, NM, D). Remarks.—It will be interesting to see whether the transition to the typical form is gradual.*® 37. EPILACHNA MJOEBERGI Weise Ficures 66, 142 Epilachna Mjoebergi WEISE, 1923a, p. 131. Abdomen.—Abdominal lines complete, reaching to within one- fourth of the apical margin of the first segment. Fifth segment, male, truncate to slightly concave. Sixth segment, slightly emarginate. Male genitalia—Penis seen in profile, 1.6 mm. long, with distinct 16 From New Guinea and the neighboring islands a number of species have been described having the elytra dark with one or more reddish spots. Many of the descriptions are such that the species cannot be recognized from them. Some of these species also have lighter forms, for which the ground color of the elytra is red. On the whole these species do not form a very homogeneous group, and characters can be found by which they can be separated. A few notes about these species may be helpful. Type locality is New Guinea unless differently indicated. E. haemorrhoa Boisduval: See p. 90. E. fulvimana Weise: Very probably a dark form of haemorrhoa. See p. QI. E. kampeni Weise: Possibly a variety of haemorrhoa. E. antiqua Weise: Ranges from a light form with the six normal dark spots to a form with practically completely black elytra. E. biroi Weise: See p. 106. E. cirunigra, new: See p. 107. Close to biroi, but pubescence black on black parts of elytra, while it is gray in biroi. E. haematomelas Boisduval: Claws without tooth, elytra with a shoulder and apical spot. E. malkini, new: Claws like haematomelas, but smaller. Only subapical red spot present. See p. 108. E. papuensis Crotch: Might be identical with malkini. E. aruensis Crotch: Aru. “Elytra with two rounded yellow spots, one basal and one subapical, neither reaching the margin.” E. sobrina Harold: Salwatty. “Almost exactly like E. arvensis in colora- tion but hemispherical, more pubescent, and with the apical spot much smaller.” NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 93 basal knife edge, which reaches practically to middle. Apical bend uniform through about go°. No apical hook. Sparsely pubescent on basal two-thirds. Seen from below, orifice very elongate, width of penis at apex of orifice 0.1 mm., length beyond it 0.33 mm., grad- ually tapering to point. Paramera 1.65 mm. long, very gently bent outward near base, gradually increasing in width, greatest width 0.17 mm. near apex. On the under side of apical third a ridge near the outside edge instead of apical thorn. Pubescence on rim of apical fourth about 0.15 mm. long. Sipho, shape normal, compressed just before apex, orifice elongately oval, subapical on outside. (Genitalia very similar to those of mindanaonis and haemorrhoa, but penis narrower. ) Length—7 mm. Tips of elytra rounded. Color and maculation—Upper side. light brownish red, sides of pronotum yellowish. Elytra with the following dark spots: A basal fascia consisting of spots 1 and 2 of the normal pattern not quite reaching the side margin, widest at the suture. Scutellum light. Spot 3 near the suture connected with spot I, spot 4 wide, touching the margin farther back than No. 3. In addition, a common spot on the suture somewhat farther back than No. 5 usually is prolonged back- ward to the apical angle. This spot connected lightly with No. 4 at the latter’s back inside corner and along the margin of the elytra so that a subapical light area is completely enclosed by a dark region. Pubescence short, light gray all over. Under side and appendages light except tips of mandibles, metasternum and the outside of the apical part of the epipleurae, which are darker. Type.—National Museum, Stockholm. Type locality.—Australia, Queensland, Malanda. Material examined.—2 males: Australia, New South Wales, Innis- fail (J. F. Illingsworth, BBM). Remarks.—Weise’s description of the single type specimen gives spots I and 2 separated. Otherwise the two specimens agree well with the original description except in unessential details of the coloration of the under side. The affinity of this species is not quite clear. The spot pattern cannot be clearly recognized as a modification of the normal 12- spotted pattern. Spot No. 6 is either on the suture or absent, which is not observed in any of the 12-spotted species. The male genitalia are very closely related to those of E. mindanaonts and E. haemorrhoa. 94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 38. EPILACHNA DEYROLLII Crotch Ficures 65, 141, 189 Epilachna Deyrollii Crotcu, 1874, p. 78. Abdomen.—Abdominal plates complete, reaching to within 0.7 mm. of hind margin of first segment. Fifth segment, male, hind margin slightly concave ; female, wider, truncate with a slight middle process. Sixth segment, male, slightly emarginate; female, split, inner edges diverging so as to form a notch in the middle of the segment. Male genitalia.—Penis seen in profile, 1,34 mm. long, straight tube for three-fourths of its length, about 0.21 mm. in middle, gently curved up near end and again sharply turned up immediately before tip, which is a sharp point. Sparsely pubescent on apical half. A lateral depression on each side occupying the last fourth. Seen from below, flattened tube about 0.3 mm. wide, closed at base but split open immediately behind it and the edges gradually becoming wider apart. Apex open, slightly wider, edges rounded. Paramera 1.3 mm. long, becoming gradually narrower from base (width 0.2 mm.) to apex (width 0.1 mm.) and slightlyp bent down. Flattened, with apical thorn present and sparse pubescence on apical third. Sipho slightly bent, compressed near apex; orifice oval, small at base of compressed part on the narrow edge about 0.15 mm. from tip. Female genitalia (see fig. 189, which shows, besides the highly sclerotized parts, also the ball-shaped bursa copulatrix and the re- ceptaculum seminis).—Plates 0.6 mm. long. Greatest width 0.38 mm. near its apical fourth, where there is a distinct corner. Inner edge with a gentle emargination on the basal third. Length—8.5 to 10 mm. Tips of elytra rounded. Epipleurae of elytra wide, nearly vertical near apex. The reflexed margin of the elytra wide near apex with a more or less distinct longitudinal plica. Mandibles with the two lateral teeth distinctly smaller than the apical one (as in fig. 1 C). Color and maculation—Upper side red, pronotum with an indis- tinct dark spot on the sides ; scutellum black ; elytra with three fasciae, the first touching the base and dilated along the suture, the second just before the middle and dilated at the margin and suture, and a subapical one also dilated near margin and suture. The suture black except for the apical part beyond the third fascia, the margin black between the second and third fascia. Pubescence light gray all over. Under side mostly black, apical part of abdomen, basal third of ely- tral epipleurae, and appendages light. The basal parts of the femora and tips of mandibles usually darker. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 95 Type locality—India, Darjeeling. Material examined.—2 specimens (PA): India, northern Bengal, Kurseong, June. Remarks.—This species has its genitalia more different from the majority of species of this group than any so far considered, and it shows other points of divergence, as for example the form of the epipleurae. Its elytral pattern can be regarded as a modification of the normal 12-spotted pattern, with spots 1+2 forming the first fascia, spots 3+4 the second, and 5+6 the third. It has not been mentioned in the literature since its original description by Crotch. 39. EPILACHNA LAESICOLLIS Mulsant FIGURES 75, 143 E[pilachna] laesicollis MULSANT, 1850, p. 735. Abdomen.—Abdominal lines subterminal, complete. Fifth seg- ment, male, hind margin slightly concave. Sixth segment with a deep notch. Male genitalia——Penis seen in profile, 2.5 mm. long, upper edge departing little from straight line until near the apical bend. Basal knife edge slight though distinct; middle part covered with hairs about 0.3 mm. long. Lower edge characterized by a prominent sharp triangular ridge, which has not been observed in any other species. Tip with a sharp barb on the upper side. Seen from below, orifice just behind the triangular ridge more or less diamond-shaped, 0.4 mm. long. The part of penis beyond the orifice very narrow. Para- mera 2.2 mm. long, slender, slightly bent up, compressed laterally for basal two-thirds, with blunt apical thorn. Terminal pubescence dense, about 0.2 mm. long, a single row of longer hairs on the lower inner face for about the last 0.6 mm. Sipho, shape regular, ending in a sharp point, orifice very elongate, subterminal. Length—8 to 8.5 mm. Tips of elytra with an angle of about go’. Color and maculation—Upper side black, head and middle of the pronotum from base to apex reddish brown, the side margins nar- rowly yellowish; elytra each with seven sharply defined yellow spots, as shown in figure 75, and with the reflexed margin around the shoulder and from the second marginal spot to apex yellow. Pu- bescence short, light gray. Under side, mesosternum and most of abdomen dark, the rest, including the appendages, light. Type locality —Himalaya. Material examined.—4 specimens, all males: India: northern Ben- gal, Kurseong, June (PA; F. R. Mason, NM); Sikkim; northern India, Darjeeling, alt. 3,000-5,000 feet (L. V. Newton, NM). 96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 40. EPILACHNA ARGUS Fourcroy FIGURES 57, 149 La coccinelle argus GEOFFROY, 1762, p. 325. (No Latin name used.) C[occinella] argus Fourcroy, 1785, p. 145. Abdomen.—Abdominal plates complete, reaching to within about one-fifth (1.3 mm.) of the apical end of the first segment. Fifth segment truncate in male, convex and considerably wider in female. Sixth segment, male, gently emarginate ; female, convex, split. Male genitalia——Penis seen in profile, curving first down after the middle, then up, so that the apical half forms a gentle arc, concave side up. Seen from below, tube split open longitudinally, seams to- gether for basal two-thirds, then opening and apical third wide open ; penis without hairs; paramera very narrow (0.1 mm. wide), 1.0 mm. long, considerably shorter than penis. Covered: with hairs near apex, no apical thorn. Sipho strongly curved near base, gently so near apex in the opposite direction. Width diminishing gradually from base to apex, ending in a simple blunt point. Orifice oval on side near apex. I have examined no other Epilachna species with a similar structure of the genitalia. Female genitalia——Plates asymmetrically pear-shaped with narrow part at base. Tergite X broad, apex emarginate. No other Eurasian Epilachna has a similar structure of the female genitalia. Length—6.5 to 7.0 mm. Mandibles with apical tooth large, lateral teeth much smaller, but both approximately of same size. Lateral dentules present. Claws with triangular basal tooth. Color and maculation.—Upper side red. Pronotum spotless, elytra with standard 12-spotted pattern with rather small spots, No. I on the suture. Under side also reddish, with metasternum and middle of first four abdominal segments partly darker. All appendages reddish. Type locality.—F rance. Feeds on Cucurbitaceae. Material examined.—Specimens seen from Italy, Turino; Croatia ; Caucasus ; Algiers. Remarks.—Very close to argus, if not identical with it, is E. angus- ticollis Reiche, which occurs in Spain and southern France. It is supposed to have a larger head, narrower pronotum, and more elon- gate and less rounded elytra. According to Klemm (1930), the male genitalia are identical with those of argus, which makes it rather probable that angusticollis cannot be more than a local race of argus, NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 97 I have seen specimens labeled angusticollis from Africa, which were indistinguishable from argus. Epilachna argus seems to occur sparingly over the southern part of Europe. It has been recorded from Spain, France, all Italy, southern and western Germany as far north as Kassel, and Prussia (Reitter), Austria, and North Africa. 41. EPILACHNA CHRYSOMELINA (Fabricius) FIGuRES 56, 145, 197 Coccinella chrysomelina Fasricius, 1775, p. 82. Abdomen.—Abdominal lines complete, broadly rounded, reaching to within about two-fifths of apical margin. Fifth segment, male, truncate ; female, biconcave with process in middle (fig. 5 H). Sixth segment, male, strongly emarginate ; female, convex, split. Male genitalia.—Penis seen in profile, 1.6 mm. long, no basal knife edge ; gently bent up, with a bulge on the upper side and an emargina- tion on the lower side before the apex, furnished with hairs on side of apical half. Seen from below, narrow tube split open toward apex. Apical quarter widened and forming a deep, oval cavity. Paramera very slender, 1.5 mm. long, furnished with hairs on apical third. No apical thorn. Sipho bent nearly 180° near base, widened at apex so that the swelling prevents the sipho from being drawn through the penis. Orifice terminal, fringed with short bristles. Female genitalia—These are considerably different from those of all the other examined species, which emphasizes the isolated position of chrysomelina in the genus. The shape of the plates (ninth sternite ) is apparent from figure 197. The tenth tergite is split in the middle, in contrast to all the other species, and between the ninth sternite and the tenth tergite is another shorter segment, which according to Verhoeff must be regarded as middle part of the ninth tergite. Length.—6.5 to 7.5 mm. The mandibles in profile have three teeth of very unequal size, the apical largest, the middle one considerably smaller, and the second lateral one not very much larger than the dentules. Apical angle of elytra close to 90° but somewhat rounded. Color and maculation—Upper side red, pronotum spotless. The typical elytral spot pattern of the species in Asia and most of Europe resembles very much that of the 12-spotted pattern of the 28-punc- tata group. However, spot 3 is not significantly farther from the suture than Nos. 1 and 5. There is a great deal of variation in the spot pattern, and different local races behave quite differently in this respect. 98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Type locality —A frica. Remarks.—Epilachna chrysomelina, though outwardly resembling many of the other Epilachna species of Europe and Asia, differs very considerably in structure from all those species. Neither the male nor the female genitalia are at all similar to any of the Eurasian species. E. chrysomelina is essentially an African species and shows its greatest development into variegated forms in Equatorial and South Africa. It must have migrated into Europe and Asia and is now one of the most widely distributed beetles occurring, though with greatly different abundance, from South Africa through the Medi- terranean region, southern and middle Europe, and central and southern Asia all the way to eastern Siberia. I have seen an as yet unidentified species from Liberia, West Africa, with male genitalia very similar to those of chrysomelina, which confirms the assumption that chrysomelina is of African origin and that we must look for its nearest relatives in Africa. The subspecies and varieties of chrysomelina in Africa have been investigated extensively (see Zimmermann, 1936) but are outside the scope of the present paper. Zimmermann divides the forms oc- curring in the Mediterranean region and elsewhere in Europe and Asia into two subspecies. The typical form, which occurs in the western Mediterranean, shows a number of varieties that differ by the manner in which various spots coalesce. Many of these varieties have received names not worth recording. Some of the more important ones are: Ab. nigrescens Weise, 1879, p. 128. Two or three spots coalesce, for example: I1+2, 2+3, 3+5, 4+6, 3+6, 4+3+5. Ab. leroglyphica Sulzer, 1776, p. 31. Two separate pairs coalesce: 3+5 and 4+6. Ab. elaterti Rossi, 1794, p. 85. 3+5+6+4. Ab. furva Weise, 1879, p. 128. Like elaterii and besides 1+2. For details of these and other aberrations see: Mader, 1926, and supplement 1932; Leman, 1927; Della Beffa, 1912. Subspecies orientalis Zimmermann, 1936, p. 527. Translation of the original description: In comparison with the groundform subsp. orientalis is characterized by a flat, not hemispherical curvature of the elytra, by a denser and more closely recumbent pubescence of pronotum and elytra which makes the species appear less shiny; furthermore by the absence of connections between the light ocelli which surround each dark elytral spot and which are more or less clearly visible depending on the general coloration. Such a confluence of the ocelli or more correctly of the light zones on which alone a formation of the black NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 99 spots is possible, can take place in orientalis only at the front margin of the basal spots, and this is true also for the confluence of the black spots themselves. With the typical form, however, besides these connections, various combinations of confluences also of the four nonbasal spots may take place. As type of subsp. orientalis I assign a female from Hedera, Palestine, in the collection of O. Vogt. Distribution of subsp. orientalis according to Zimmermann: Pales- tine, Cyprus, Egypt, Arabia, eastern Asia Minor, Persia, Turkestan, Tien Shan Mountains. In North Africa the dividing line between subsp. orientalis and the typical form is at the Algeria-Tunisia border, farther north in western Asia Minor. Transition populations occur (e.g., in Constanti- nople). E. chrysomelina orientalis interbreeds in the laboratory freely with the typical form. Obviously it is not easy to tell from isolated specimens to which subspecies they belong, but a fair sample of a population is needed. However, the locality in many cases is sufficient to decide the matter. Examined by me, specimens from: North Africa, Cyprus, Laos (Tonkin), east Siberia; India, Punjab, Jullunder (1 specimen, D). Weise (1908) reports it from Kerachi, India. The northern limit of distribution in Europe is indicated by the localities Wurttemberg, Baden in southern Germany, and Bohemia (Reitter, 1911). All specimens except some of the North African ones belong to subsp. orientalis. Epilachna chrysomelina has been used extensively for genetic stud- ies (Tenenbaum, Timofeef-Ressovsky, Zarapkin, Zimmermann). The relations between the various Mediterranean races have been clarified considerably by a statistical analysis when quantitative measurements of the elements of the spot pattern were used and by controlled breeding experiments. Other Epilachna species because of their greater variability would be even better suited for such experiments, and the status of the various subspecies of E. sparsa, for example, would be greatly elucidated by such experiments, which would be extremely simple in a warm climate, but judged from the success of the chrysomelina experiments should not be unduly difficult in a laboratory in a temperate region. The chrysomelina breeding experi- ments were all carried out in Berlin under laboratory conditions. BIOLOGY Details of the habits of E. chrysomelina have been given repeatedly, most recently and in greatest detail by Klemm (1930). The observa- IOO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 tions were made under laboratory conditions (in Berlin) with speci- mens imported from Korfu, Greece. Oviposition begins 6 to 8 days after the first copulation. The eggs are deposited in clusters of 10 to 40, usually on the under side of leaves or on other rough surfaces. One female deposits up to 80 eggs a day, and a total of as many as 322 were observed. The dura- tion of the various stages is as follows (temperature 20° to 25°C., humidity 50 to 70 percent) : Days 1 eae A ie i era Ernie apy alg’ iin A BS Oh 5-6 DB AEViale hi coer ohh oe a eine tie a yc R AL eee 19-26 USCCINS Ga Gass avaivelo trem iernscere aero 4-5 AC MNS tATy Med el crcltalaecoutelersioe mate ont tooke 4-6 BC UMIStAT is ahs yc Seats 2 ato Mota cale eee ee 4-6 ACI AUMStAI 1 arc k Sle lapel ais tree rever ete are oe 5-6 Preplipai Weacaemecsch ee ete een 2-3 Papas iota ors Me aeons cbtetea neers 5-7 Ege to; adult, average)... denis antiacee cee 32-35 Number of generations per year 4, possibly more under more favor- able conditions. Copulation occurs from the second day after emer- gence for female, from the eighth for male. Optimal temperature for development 25° to 30°C. At about 15° to 20° no eggs are deposited ; below 15° no copulation takes place. Both adults and larvae feed on cucurbitaceous plants (cucumber, pumpkins, melons, as well as Bryonia and Ecballium species). Both the foliage and the fruits are attacked. In southern Europe where the species occurs no serious damage seems to be done, but from central Asia (Turkestan and Transcaspia) heavy damage is reported. Klemm also describes and figures the immature stages. ENNEASTICTA GROUP The three known species of this group hardly differ in external appearance from some of the 12-spotted species of the 28-punctata group. They can, however, be recognized easily by the structure of the male genitalia, which differs sharply from that of all the other species of the genus. The general shape of the penis (see figs. 146-148) is less elongate, and seen in profile it is shorter and thicker. Seen from below it appears as a tube for approximately the basal half, but completely open beyond. Whereas the penis tube is split open along its whole length in the other groups of Epilachna, it is completely closed with- out a seam at its basal end in the enneasticta group. Beyond the com- NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE Io! pletely closed part of the tube, the penis has a longitudinal slit of moderate width, and the apical half of the tube is completely open. The paramera extend somewhat beyond the penis. The shape of the sipho is greatly modified. This is best shown by figure 147. The other two figures represent specimens with the sipho somewhat damaged. The basal part of the sipho is strongly developed, prac- tically as long as the rest of the sipho, and laterally bordered by a wide clear membrane-like structure. The apical part, the sipho proper, is short and relatively wide. It has a constriction at about one-third of its length. The female genitalia of only one species, enneasticta, of the group are known (fig. 199). They are not significantly different from those of the majority of species of the genus, but have the apex of tergite X pointed. 42. EPILACHNA ENNEASTICTA Mulsant Ficures 67, 68, 146, 199 E[pilachna] enneasticta MULSANT, 1850, p. 769. Abdomen.—Abdominal lines complete, subterminal and somewhat angular, the outside edge straight. Fifth segment, male, hind edge concave ; female, strongly concave, in the middle emarginate almost to base. Sixth segment, male, emarginate; female, split. Male genitalia—Typical for the enneasticta group and quite dif- ferent from those of the majority of the genus. Penis seen in profile, 1.2 mm. long and much thicker than in 28-punctata group. A longi- tudinal ridge runs along almost the whole middle of its upper side, a row of hairs (about 0.2 mm. long) on each side of the ridge. Seen from below, 0.5 mm. wide, tube closed completely for the basal 0.3 mm.; the next 0.4 mm. with a rectangular longitudinal slit, about 0.07 mm. wide. Beyond that the tube is half open and widens and flattens just before the end, where it curves upward. Apical edge convex, without a notch (see, however, below). Paramera 1.6 mm. long, narrowing gradually from base to apex without apical thorn. Hairs on the apical half, those at apex very much shorter than those on the sides. Sipho, basal part elongated, about 1.4 mm. long, and well sclerotized. (In fig. 146 the basal part is broken off. Fig. 147 represents this part better, as the sipho of E. indistincta is not dis- tinctly different from that of E. enneasticta.) The sipho proper is rather wide, with a constriction at about one-third of its length and a sharp subapical notch. The basal tip of this notch is bifurcate; the apical one carries the small oval orifice. 102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Female genitalia—Apex of tergite X coming to a point. Plates 0.6 mm. long, greatest width 0.4 mm.; apical edge distinctly emar- ginate. Inner edge nearly straight. Length—7.2 to 8 mm. Tips of elytra rounded. Color and maculation.—Upper side red, pronotum yellowish at the sides. Elytra each with six black spots. No. 1 close to the suture or touching it, No. 2 on the callus rounded, No. 6 transverse with an emargination on the apical side so that it usually appears crescent shaped. No. 5 slightly transverse, almost touching the suture. Nos. 3 and 4 transverse, in lighter specimens No. 4 not touching the margin, in darker ones both widened so that they form a relatively narrow fascia reaching over on the epipleurae, but interrupted at the suture. Pubescence light gray, dark on the spots. Under side light, with most of the metasternum and the first four abdominal segments except the sides usually dark, occasionally also the mesosternum. Type locality—Java. Material examined.—7 specimens, all from Java (NM) : Buitenzorg, March 1909 (Bryant and Palmer); Mount Salak, May 15, 1909 (Bryant and Palmer) ; East Java, Lawang, 1897. Remarks—tThis species can immediately be recognized by its male genitalia, which are similar only to those of indistincta. The crescent- shaped spot No. 6 seems to be characteristic for this species, and the very strongly emarginate fifth abdominal segment of the female serves to distinguish it from all the other known species. (It is to be ex- pected that the female of imdistincta will show a similar abdominal structure. ) Among the males there is one specimen that has the apex of the penis distinctly notched, whereas the specimen from which the figure was made shows no trace of such a notch. The notched specimen also had a less distinct ridge on the upper side of the penis, and there are other less pronounced variations. Whether this should be regarded as a separate species or merely within the range of variation of en- neasticta can only be decided with much more material. The latter view is supported by another specimen that shows a faint emargina- tion at the apex of the penis and seems to be an intermediate between the two other specimens also in other respects. 43. EPILACHNA INDISTINCTA, new species FIGuRES 609, 147 Abdomen.—Abdominal lines complete, subterminal, angulate. Fifth segment, male, mildly concave. Sixth segment, apex wide, moderately concave. NOSES LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 103 Male genitalia.—Penis seen from below, 1.4 mm. long. The basal part a wide tube, completely closed for the basal 0.3 mm.; for the next 0.35 mm. with a narrow longitudinal slit, oval at its base and slightly narrowing toward its apex; from then on the penis is com- pletely open, widened at its apex and curved up. Seen in profile, with a blunt ridge on the upper side and a row of hairs on each side of it. Paramera slender, 1.4 mm. long, protruding beyond the penis, almost parallel or slightly tapering toward apex. Clothed with very short hairs at apex, somewhat longer ones (0.15 mm.) on the re- mainder of the apical third. Sipho of the shape indicated in figure 147 with the long basal piece to which is attached laterally a clear blade- like stiff membrane, damaged near the basal end in the type and the outline uncertain in the figure. The sipho proper with the constriction near its base and the subapical notch characteristic for the enneasticta group. The basal point of this notch is forked ; the apical one has the small oval orifice. Length—7 mm. Reflexed margin of elytra very distinct. Color and maculation—Upper side light yellowish brown. Sides of pronotum yellow. Elytra each with six rather small spots. Pubes- cence light gray, short, dark on the spots. Under side and appendages light brown, tips of mandibles and metasternum partly dark. Material examined.—Type: U.S.N.M. No. 57133, Sumatra, Sian- tar (Mann, 1937, N.G.S.-S.I. Exp.). 5 paratypes, all males: Sumatra, Pagaralam and Fort De Kock Geri and Bs 'B. Brues, MCZ, D). Remarks—This species is not conspicuously ‘different externally from the other 12-spotted species but can immediately be recognized by its male genitalia, which resemble those of E. enneasticta Mulsant. From the latter it can be distinguished by the absence of the lateral ridge near the apex of the penis and the fact that the longitudinal slit is oval at the base and not truncately cut off. 44. EPILACHNA SEMIFASCIATA, new species FicureEs 70, 148 Abdomen.—Abdominal lines complete, subterminal, outside straight. Fifth segment, male, hind margin mildly concave. Sixth segment the same. Male genitalia—Same general shape as the two other species of the enneasticta group, but with quite pronounced differences. Penis seen from below, wide tube (width about 0.35 mm.). Only the first 0.1 mm. completely closed, the next 0.35 mm. with a narrow longitudinal 104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 slit in the middle widened ovally near the base, less than 0.03 mm. wide near apex; beyond this a deep narrow trench, which ends near the blunt point that forms the apex of the penis. This apical third laterally bordered by a horizontal process. Seen in profile, small basal knife edge not reaching farther than the base of the paramera. Paramera 1.15 mm. long, slightly curved downward. Sipho: the base shows the same extended structure as in enneasticta and indis- tincta, and also the general shape is the same, but the tip has quite a different structure. Seen from the outside about 1.15 mm. before the end is a lateral constriction. There is a median sharp ridge of about 0.035 mm. length, and on each side a lateral ridge interrupted at the constriction. Length—5.5 mm. Tips of elytra rounded. Color and maculation—Upper side brick red; pronotum with all seven spots faintly present with hazy outlines, the middle three coa- lescent ; each elytron with six dark spots, Nos. 1 and 5 touching the suture, Nos. 4 and 6 touching the margin; all spots except No. 2 laterally widened, No. 3 most (0.8 mm. long, 1.6 mm. wide). Nos. 3 and 4 form a transverse fascia interrupted by a gap of 0.1 mm. width between 3 and 4, and one of twice 0.3 mm. at the suture. Pubescence light gray, dark on the spots. Under side partly dark, appendages, except tip of mandibles, light. Type—vU.S.N.M. No. 57134, Formosa, Musha, May 1, 1929 (K. Sato). Remarks.—The maculation of E. semtfasciata agrees very closely with that of E. subfasciata Weise (1923b, p. 182), also from For- mosa. However, Weise gives a short description of the genitalia of this species, which are quite different. GUTTATOPUSTULATA GROUP Besides guttatopustulata (Fabricius) a few other species with dark ground color have been placed here. Most of them differ from the rest of the genus by the structure of the female genitalia, which have the plates transversely oval. The male genitalia, however, are known only in two cases. E malkinz is so different, particularly in the absence of the tarsal tooth, that it probably should be put into a separate group. 45. EPILACHNA GUTTATOPUSTULATA (Fabricius) » FIGURES 7I, 144, 200 Coccinella guttatopustulata FABRICIUS, 1775, p. 87. Abdomen.—Abdominal lines complete, reaching to within about one-third of the apical margin of the first segment. Fifth segment, NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 105 male, truncate, slightly emarginate; female, slightly convex. Sixth segment, male, with a deep notch; female, split. Tergite VIII emarginate. Male genitalia—Penis seen in profile, straight tube for most of its length, decreasing gradually in width, then rather suddenly curved up. Apical point with trace of a hook. Seen from below, tube with the seam sometimes apart during its entire length, flattened and widened near the middle. From apex of orifice to point, 0.2 mm. Penis without hairs. Basal knife edge rudimentary. Paramera 1.7 mm. long, widest near the middle, narrowed considerably toward apex, without apical thorn. Apical hairs sparse, about 0.2 mm. long; also one row of hairs on edge starting just behind the widest part. Sipho long and slender, curved through 180° near base, ending in blunt point. Female genitalia——Tergite X with apical margin broadly truncate with a trace of an emargination. Genital plates obliquely widened with a not very distinct basal edge. Length—8 to 8.5 mm. Apical angle of elytra rounded; claws (fig. 2C) with quadratic tooth which has a small, toothlike process on the inside. Mandibles with the first lateral tooth slightly smaller than the apical one, the second lateral tooth smaller than the first; dentules present. Color and maculation.—Upper side, head reddish, pronotum yellow with a big central black spot reaching from base to apex. Elytra black with two subbasal spots, the inner one on the suture red, the outer one on the margin yellow. In addition a subapical red fascia convex forward and usually interrupted at the suture. Under side and appendages mostly black. Prothorax, parts of epipleurae ad- jacent to the light parts of the elytra, and the outer margin of the first four and usually the complete fifth abdominal segments light. Type locality— Australia. Material examined.—12 specimens. Australia: Tambourine Moun- tain, October 28, 1912 (H. Hacker, NM; Koebele; Bridwell coll.) ; New South Wales, Richmond River; Sydney, Grafton, O’Kelly, on sugarcane. Central York, Queensland, Tasmania, listed by Crotch. New Guinea (NM). Remarks.—This species is considerably removed from the vast majority of the other Epilachna species. Its female genitalia resemble much more those of Afissa maxima than those of any other Epi- lachna. The deep notch in the fifth abdominal segment of the male and the male genitalia also set it apart. 106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 EPILACHNA GUTTATOPUSTULATA subsp. TRICINCTA (Montrouzier) FIGURE 72 Coccinella tricincta MONTROUZIER, 1855, p. 76. Differs from the main form by the reduction of the black pig- ment. The spot of the pronotum is only faintly indicated. The two subbasal spots are united and the subapical fascia enlarged. This form gives the appearance of a yellowish-red upper side, with a dark basal and a discal fascia and an apical spot at the suture. The under side and appendages are entirely light. Type locality—Woodlark Island. Material examined.—24 specimens: New Guinea (NM); Dutch New Guinea, Hollandia, January, April, May, 1945 (B. Malkin, NM, D). EPILACHNA GUTTATOPUSTULATA subsp. TASMANICA Crotch Epilachna guttato-pustulata var tasmanica CrotcH, 1874, p. 78. Crotch’s original description is as follows: “Var. tasmanica. The yellow marginal spot is replaced by red, the black pigment is dimin- ished, the posterior fascia deeply angulate, the apical spot enlarged.” Type locality—Tasmania. Material examined.—1 specimen answering this description: New Hebrides, Espiritu Santo, September 1944 (K. L. Knight, NM). Female genitalia like those of guttatopustulata s. str. 46. EPILACHNA BIROI Weise FIGURES 73, 202 Epilachna Biréi WEISE, 1902, p. 491. Abdomen.—Abdominal lines complete, reaching to within one- third of the hind margin of the first segment, slightly angulate. Fifth segment, female, hind margin truncate with a slight process in the middle. Sixth segment, split. Female genitalia.——Plates transversely oval. The rest not avail- able. Length 0.23 mm., width 0.40 mm., inner edge convex. Length—7 mm. Apical angle of elytra rounded. Color and maculation—Upper side black, the head lighter; pro- notum with a yellowish-red side margin; elytra each with two red spots, one humeral, the other apical. Under side black, apex of abdo- men lighter ; epipleurae light in the parts corresponding to the elytral spots. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 107 Type.—Hungarian National Museum (?). Type locality—New Guinea, Sattelberg. Material examined.—1 specimen: Philippine Islands, Luzon, Manila (D). ; Remarks.—tThe only available specimen agrees well with Weise’s description of biroi. Whether it is closely related to guttatopustulata is not quite clear, as only a somewhat mutilated female is available. However, the genital plates resemble in form those of guttatopustulata much more than those of any other species, and the black ground color is another point of similarity. 47. EPILACHNA CIRUNIGRA, new species Abdomen.—Abdominal lines incomplete, the inner arc distinct and reaching to within one-fourth of the hind margin of the first seg- ment, the outer part indistinct. Fifth segment, male, hind margin slightly concave; female, convex. Sixth segment, male, with semi- circular notch ; female, split, apex of halves with nearly right angle. Male genitalia——Penis seen in profile, 1.2 mm. long, upper edge straight until shortly before tip, when it curves up suddenly through a go° angle. Narrow rudimentary basal knife edge present; two rows of hairs on middle part. Seen from below, tube split open for entire length and seams about 0.05 mm. apart. Paramera 0.95 mm. long, only about 0.1 mm. wide on apical half, slightly wider near base, rim covered with hairs on apical third. Sipho bent through 180° near base (as in fig. 144), tip rounded, slightly swollen, with orifice subterminal on side. The genitalia of E. cirunigra resemble most closely those of E. guttatopustulata Fabricius (fig. 144). They differ by the fact that the penis is more suddenly curved up near its tip, that the paramera do not have the sudden decrease in width, and that the tip of the sipho is swollen rather than constricted. Female genitalia—Plates transversely oval with the outer edge more pointed than the inner one, length 0.30 mm., width 0.35 mm. ; the inner half of the basal edge of both plates taken together forms an arc that is convex toward the apex. The shape of the plates resembles most that of figure 202 (biroi). Tergite X with strip of pigment on each side, middle clear. Length—7.5 to 8.5 mm. Tooth of claws square; tips of elytra rounded. Mandibles with the apical tooth widened with traces of doubling. Two lateral teeth, the outer one bigger above the apical tooth and its tip actually farther outward than the apical tooth. The 108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 inner one much smaller. Trace of one dentule between the two lateral teeth. Color and maculation—Upper side, head, scutellum, and middle of pronotum reddish brown, the sides yellow. Elytra, black with two big red spots on each elytron. One spot covers the shoulder and takes up the outer two-thirds of the base and the front third of the margin. Its inner edge is evenly curved. The other spot covers roughly the apical third and reaches both suture and margin. Its basal margin is convex forward. (The maculation is very similar to that of fig. 74 except that the spots are larger.) The pubescence is light gray on head and pronotum, black on the black parts, and reddish on the red parts of the elytra. Under side light brown except the tips of the mandibles and the middle of the epipleurae, which are dark. The apical part of the metasternum also darkish. Material examined —Type: U.S.N.M. No. 57968, from Hollandia, Dutch New Guinea, June 1945 (B. Malkin). 7 paratypes, from same locality as type, January, April, June, 1945 (NM, D). Remarks—tThis species is very close to E. biroi Weise but dif- fers sufficiently from Weise’s description to warrant listing as a new species. The most conspicuous difference is the black color of the pubescence on the dark parts of the elytra. E. biroi is supposed to have gray pubescence except on the red spots. The genitalia and the general structure leave no doubt that E. cirunigra belongs in the same group as E£. guttatopustulata Fabricius. 48. EPILACHNA MALKINI, new species Abdomen.—Abdominal lines complete, rounded, not reaching much beyond the middle of the first segment. Fifth segment, female, with straight or slightly convex hind edge. Sixth segment, female, emar- ginate, split. Female genitalia—Plates 0.28 mm. long, 0.32 mm. wide; inner edges straight, together with a heart-shaped notch at the very base, basal edge slightly, outer edge more strongly curved, apical edge emarginate. Length—6 mm. Tarsal claws split but without a distinct tooth. Color and maculation—Upper side, head, and pronotum reddish brown, sides of latter lighter; elytra black with a round reddish spot in the hind third which reaches the margin but not the suture. Under side and appendages light except the metasternum, the tips of the mandibles, and the front two-thirds of the epipleurae, which are darker. Pubescence uniformly whitish gray. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 109 Material examined.—Type: U.S.N.M. No. 57969, from Hollandia, Dutch New Guinea, April 1945 (Malkin). Paratype: Same data (D). Remarks—This species is markedly different from the other species of Epilachna by having the tarsal claws without a distinct tooth, and in this respect it resembles the species of Afissa. However, in all other respects it is much closer to the rest of Epilachna than Afissa. In particular the sixth abdominal segment of the female shows a distinct suture, which is always absent in Afissa. For this reason it seems best to consider E. malkini as belonging to Epilachna. A knowledge of the male will probably shed further light on this question. E. malkini is probably closely related to E. haematomelas Bois- duval, which according to Weise also has toothless claws. The latter has a humeral red spot in addition to the subapical one and has pronotum, under side and legs black. Its size is considerably larger (length 10 mm.). From the rest of the Epilachna species E. malkini is easily dis- tinguished by the toothless claws, the emargination in the sixth abdominal segment of the female, and the genital plates. These are distinguished by the fact that the notch at the inner edge, present in many Epilachna species, reaches to the basal edge, whereas it remains somewhat distant from it in the other species. AFIDENTA, new genus Genotype: Afidenta mimetica, new species. Claws bifid with a sharp basal tooth. Sixth abdominal segment of female not split lengthwise. The two known species of this genus, although quite different in appearance, have a very similar structure of the genitalia, different from that of any oriental species of Epilachna and Afissa. It is the most primitive, least elaborate form observed in any species of the subfamily Epilachninae and is much smaller in relation to the body size than other species. The establishment of a new genus for these two species seems therefore to be justified, at least provi- sionally. The structure of the tooth of the claw differs from that of Epilachna in that it has an irregular, weakly sclerotized outside edge (fig. 2D). Epilachna bisquadripunctata, although differing in some respects from the two other species, has been temporarily placed in this genus. 8 IIo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 1. AFIDENTA MIMETICA, new species FIGURES 74, 157, 201 Abdomen.—Abdominal lines complete, subterminal, nonangulate. Fifth segment with hind margin very slightly convex in both sexes. Sixth segment, male, convex ; female, emarginate, not split. Male genitalia—Smaller than in Epilachna. Penis seen in profile, 0.5 mm. long, wedge-shaped, with the upper margin nearly straight and the lower one slightly curved up. Apical point straight, not curved up. No hairs on penis, no basal knife edge. Seen from below, straight tube, pointed toward apex; split longitudinally with the seams apart. Paramera 0.5 mm. long, about 0.1 mm. wide, without apical thorn, slightly curved outwardly and with hair on the apical half. Sipho short, moderately bent near base, narrowed near apex, orifice subterminal on side, very elongate. Female genitalia——Tergite X with apical margin very convex, pigment in two strips along the side. Genital plates diagonally sub- oval, mildly emarginate on outer apical side. Length.—5.0 to 5.5 mm. Tips of elytra rounded. Mandibles with a split apical tooth and two lateral ones, the more basal one smaller. No dentules. Color and maculation——Upper side brownish red, pronotum with a transverse row of four black rounded spots near the middle or just in front of it, spaced equidistantly, these spots occasionally ob- solete. Elytra each with 14 black spots arranged as in Epilachna 28- punctata or allied species without any significant differences. Pubes- cence dense, light blond, dark on the spots. Under side and appen- dages light, sides of metasternum darker. In the Tibet specimens mesosternum and metasternum and abdomen also dark. Material examined—Type: U.S.N.M. No. 57135, Indochina: Annam Prov., Haut Donai, Col de Blao, alt. goo meters, September 30, 1932 (M. Poilane). 30 paratypes (NM, MNH, MCZ, D): China: Shantung Prov., Yen-Ping, July, September, 1917 (AMNH) ; Anhwei Prov., Taiping- shien, October 1932 (G. Liu, MCZ); Kwangsi Prov., Yao Shan, Yungshien, Peiliu, Wuchow, Pingliu, Yueling, Kweiping, Kweishien, March-May 1933 (G. Liu, MCZ); Fukien Prov., near Foochow (Kellogg, NM); Tibet (AMNH). The specimens from Anhwei, Shantung, and Tibet are dark with large spots, those from Kwangsi lighter with small spots. Remarks.—This species must have been confused hitherto with one of the 28-spotted Epilachna species, which it resembles very NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE itil closely and which occur in the same locality. From these it can be easily separated by its smaller size and particularly the spot pattern of the pronotum. The female abdomen and the male and female genitalia give, of course, infallible criteria for separating it from these Epilachna species. AFIDENTA MIMETICA SIMPLEX, new subspecies Structure and male genitalia the same as in A. mimetica s. str. Pronotum with the same transverse row of four black spots. Elytra, however, each with only the six persistent dark spots, which are rounded and arranged as the persistent spots of mumetica s. str. Type-——Museum of Comparative Zoology, from India, Kooloo (Carleton). 2. AFIDENTA MINIMA (Gorham) FiGcurReE 80 Epilachna minima GorHAM, 1894, p. 206. Abdomen.—Abdominal lines subcomplete, subterminal, apex sub- parallel to margin of first segment. Fifth segment, male, hind mar- gin truncate to slightly convex; female, convex. Sixth segment generally not protruding from under the fifth; male, convex with a slight apical emargination ; female, not split, convex. Male genitalia—Penis seen in profile, 0.35 mm. long, wedge- shaped, slightly bent upward immediately before the tip, no hairs. Seen from below, open tube ending in two sharp points. Paramera bent lightly up near base, slender, 0.35 mm. long, 0.035 mm. wide, very sparsely pubescent near apex. Sipho bent 180° near base, orifice oval on side, subterminal. Female genitalia—Tergite X wide, apical margin mildly convex, a strip 0.04 mm. wide along this margin pigmented. Genital plates irregularly diagonally oval. Length.—z2.6 to 2.9 mm. Color and maculation—Upper side light brown. Elytra with the basal and lateral margins and three spots dark. The dark margins often absent. Under side light, mesosternum and metasternum dark. Type locality—India, Belgaum. Material examined.—India: Mangalore, August 1926, August 1924, November 1926, on grass (J. C. Bridwell, NM, 11 specimens) ; 1924-1926 (J. C. Bridwell, 2 specimens) ; Goa, Murmugao, Septem- ber 1925 (J. C. Bridwell, 4 specimens) ; Dehra Dun Distr., August 14, 1944 (J. Unyal, D). I1I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Remarks.—Gorham’s original description gives the under side as black, which probably means only that he had darker specimens of this species. He reported them as found under bark in December. 3. AFIDENTA BISQUADRIPUNCTATA (Gyllenhal) FIGURE 223 Cocc[inella] bis 4-punctata GYLLENHAL, in Schonherr, 1808, p. 186. E[pilachna] herbigrada Mutsant, 1850, p. 805. Abdomen.—Abdominal lines rounded, subcomplete, reaching to within one-fourth to one-third of the hind margin of the first seg- ment. Fifth segment, male, hind margin truncate; female, wide, con- vex with a very small notch in the middle. Sixth segment, male, emarginate ; female, entire, convex. Male genitalia—Penis seen in profile, 0.93 mm. long, straight tube, thickened just before apex, appearing to end in a sharp, upturned point (this appears as a point only when seen in profile). Seen from below, subparallel tube, seams gradually diverging in last third and completely open in last fifth, apex bilobed. Paramera very thin, 0.9 mm. long, bent down slightly at base, width 0.03 mm., only slightly widened at apex, and apex sparsely clothed with hairs. Sipho bent through 180° near base, slightly widened at apex when seen side- wise, compressed when viewed in plane of bend from convex side. Orifice subterminal. Female genitalia——Plates rounded with the base somewhat flat- tened, diameter about 0.26 mm.; tergite X, wide (0.85 mm.), mildly emarginate. Length—4 to 5 mm. Tarsal claws split, both parts slender and with a distinct triangular tooth; mandibles very different from any of the other observed structures in this group. They are not flat as in the Epilachna and Afissa species, but about as thick as they are long and wide. There is a broad flattened apical tooth and two shal- low, broad, lateral teeth along the inner rim. There is another tooth not far from the more basal one of the two lateral teeth but apart from it and in a different plane. Color and maculation—Upper side brownish red; head and pro- notum spotless, elytra each with four black spots, the first near the base about halfway between shoulder and suture, the second slightly in front of the middle separated from the suture by less than its own diameter, the third one about as close to the margin and slightly farther in front. The fourth spot about one-third of the length of the elytra from the apex, about as close to the margin as to the suture. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE Lis The spots are usually rounded, the diameter about one-fourth the width of the elytra, or some may be transversely widened. Pubescence light gray, dark on the spots. Under side light except metasternum and base of abdomen, which may be darker. Type locality—East Indies; for herbigrada, Pondicherry. Material examined.—7 specimens (MCZ; D): India: Kooloo (Carlton) ; southern India, Malabar, Walayar Forest, 700 feet, No- vember 16, 1945 (Nathan). China: Kwangsi Prov., Kweiling, April 1933),(G, Liu). Remarks.—There has been some dispute as to whether bisquadri- punctata Schonherr and herbigrada Mulsant are the same species. The difference is supposed to be that in the former the spots are transversely widened while in the latter they are round. In some of the Indian specimens some of the spots are widened and the others round, which suggests that there is no difference between the two forms. The Chinese specimens would belong definitely to herbigrada. Their size is a little smaller than that of the Indian specimens. The presence of the tooth on the tarsal claws and the absence of splitting of the sixth abdominal segment of the female would put bisquadripunctata in Afidenta. The genitalia are, however, quite different from those of the two other species of Afidenta. This and the very peculiar structure of the mandibles suggests that bisquadri- punctata belongs to a new genus. In the absence of further informa- tion, however, it is left for the present in A fidenta. AFISSA, new genus Among the Eurasian genera, Afissa is quite clearly differentiated from Epilachna, and there never can be any doubt to which of the two genera a given species belongs. The species of Afissa do not form nearly so homogeneous a group as those of Epilachna, which have hardly any morphological differences. The shape and size may vary greatly: some species have quite remarkable modifications of the structure of some of the abdominal segments, others have distinct cavities in the epipleurae for the reception of the tips of the femora. The spot pattern is also more varied. The pronotum may be spot- less or have a discal spot or three spots of various shape, or the dark color may take up practically the whole surface of the pronotum, with all sorts of intermediates. The elytral pattern of some species resembles very much that of some of the Epilachna species with six spots on each elytron (see fig. 81 or 91). A few species have more than six spots; the additional spots, however, do not behave at all like the nonpersistent spots of Epilachna. Ii4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 The basal pattern for the majority of the Afissa species, however, consists of five spots arranged 2, 2, I as in figure 88. Many modifi- cations, such as the coalescence of various spots, occur, but on the whole the 5-spotted pattern can clearly be recognized. Much less is known about the variability of the spot pattern of the various species than for Epilachna, as the number of specimens available for most species is rather limited. Details about this will be found under the various groups. The variation of the spot pat- tern for many species for which fair series are available seems to be quite small (for instance, the 35 specimens of flavicollis from a very extended region show hardly any variation in the elytral pattern at all). On the other hand, it was found from the study of the genitalia that some forms with seemingly quite insignificant differ- ences in the spot pattern belong to distinctly different species. The male genitalia show a great variety of structures and form an excellent criterion for the identification of species. The chief differ- ences from those of Epilachna seem to be, when the structures seem otherwise very similar, that there is never a basal knife edge and very rarely any pubescence on the penis, and in the few cases that it is there it is very sparse. The paramera never have an apical thorn. The female genitalia also show a great variety in structure and are very good for a classification into various groups. The distribution of Afissa is less wide than that of Epilachna. It is completely absent from Europe and Australia and probably does not cross the deserts of central and western Asia. Its center of greatest development seems to be in southern China. It is well rep- resented also in India. A smaller number of species occur on the East Indian Archipelago, and only two species are known from the Philippines. It apparently does not reach New Guinea and the Pa- cific islands. Going north from its center of distribution, the num- ber of species diminishes rapidly, and only one species, admirabilis Crotch, has been reported from northern China and Japan. It never has been recorded from Manchuria and Siberia. Very little is known of the biology of Afissa species. Apparently none of the species occur as serious pests, partly because the number of their individuals does not become very large, partly because the chief food plants are of no economic importance. At least this is true for A. admirabilis, the only species the habits of which have been studied at all. A great variety of mandibular structures has been ob- served in Afissa, and it would be interesting to know whether dif- ferences in food habits go together with this. A striking difference between the habits of Afissa admirabilis and the Epilachna species NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE II5 is the fact that the former hibernates as larva. It is unknown, how- ever, whether this is true for Afissa in general. Probably only a rather small portion of the total number of Afissa species is known at present. This at least may be concluded from the fact that in the material from China available to me for study 15 of the 23 species are new, many only represented by one or two specimens. Almost all this material came from a rather limited region in Szechwan Province. It must be expected that, if careful col- lecting is done in other parts of Continental Asia with the proper climate, a similar yield in new species will result. The genus Afissa can be subdivided into several more or less well defined groups which are characterized by the type of spot pattern and the structure of the genitalia and of the abdomen. The following key tries to use the spot pattern as much as possible: 1. Elytra each with six or more spots, which may be more or less united to form transverse fasciae; in which case the fascia in middle of elytra is OentaDACk aN fOLm OL ia: W Meat, SULUTE... «ccs a-s «0 0:5 si0s Slo cas nse 04 ale 2 Elytra each with five or less spots; if these are united into transverse fasciae athe middle ‘one.is nearly straight...7.. 125 s2.8e. cos cens cet ee 3 ay | LASTaYSad eV SeaCGS eCeet aa: eo ON 00) v0 RP Ne admirabilis group Berapeliiess wthiag (Ok Mathie fede cca e us haven ig eee oe ante! cictele se fallax group ee a booinel, smGociin. DOL SEXES. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 131 ship of two forms so distant geographically just from superficial resemblance, without a comparison of the genitalia. Afissa bengalica also resembles superficially Afissa fallax (Weise). The two can easily be separated by the coarse punctation of the elytra, which is pronounced everywhere on bengalica but obsolete except near the margin on fallax. The male genitalia of these two species are entirely different. 13. AFISSA MIRABILOIDES, new species Ficure 86 Abdomen.—Abdominal lines complete, subterminal, rounded. Fifth segment, male, hind margin truncate. Sixth segment, mod- erately convex. Male genitalia——Destroyed except sipho, which is similar to that of bengalica. Length.—4.5 mm. Color and maculation—Upper side brownish red, head with a darker transverse bar; pronotum, disk dark with the sides and the lateral parts of the base lighter. Elytra each with six black spots arranged as in bengalica but larger. Nos. 1 and 5 on the suture. Under side, metasternum black, abdomen partly dark, femora slightly darkish, the rest light. Type—vU.S.N.M. No. 57140, from China, Szechwan Prov., Hua Yin Shan, 70 miles north of Chungking, alt. 2,500 feet, July 5, 1933 (D. C. Graham). See remarks under A. maculicollis (Sicard). 14. AFISSA MACULICOLLIS (Sicard) Ficure 85A Solanophila nilghirica var. maculicollis StcarD, I9I2c, p. 131. Abdomen.—Abdominal lines complete, subterminal, rounded. Fifth segment, female, wide, hind margin convex. Sixth segment, convex. Female genitalia—Genital plates flattened isosceles triangles with the inside edge as the base and the top of the triangle rounded. They resemble those of A. fallax (Weise) but are more stretched. Tergite X damaged in the one specimen I could examine. Length—4.9 mm. Color and maculation—Very similar to bengalica, with the follow- ing differences: The pronotum has only a small rounded dark spot 132 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 just before the middle, scutellum light; under side and appendages light with only the metasternum dark. Type locality—Formosa. Material examined—1 female, Formosa, Urai, April 10, 1929 (K. Sato, NM). Remarks.—This species was regarded as a variety of A. nilgirica (Weise) by Sicard. He characterized it by the following differences : “Form a little more elongate, pubescence a little denser, pronotum on the disk marked with a black spot. Four specimens from Formosa.” In view of the great geographical separation of the type localities of milgirica and maculicollis, and the fact that in Afissa distinct species are often marked by only slight differences in appearance, it seems safer to regard maculicollis as a distinct species until an examination of the male genitalia can settle the question. 15. AFISSA NILGIRICA (Weise) Solanophila nilgirica WEIsE, 1908, p. 219. According to Weise’s description this species is very similar to the preceding three species. It differs from them by a spotless pronotum and the fact (on which Weise puts some stress) that the sixth elytral spot is elongated and points directly to the fifth spot. In the other three species (and in fallax) the prolongation of the sixth spot would meet the suture well behind the fifth. The status of the four species bengalica, nilgirica, maculicollis, and murabiloides must be considered very uncertain until more ma- terial will make a comparison of both male and female genitalia of all four forms possible. They are very closely related, and all four have the inner tooth of the tarsal claw widened and touching that of the other claw. A. bengalica and mirabiloides are distinguished by the shape of the hind margin of the sixth abdominal segment, which is emarginate in bengalica and convex in murabiloides. If there should be no greater variability within each species of the maculation, the pro- notal pattern would be an easy means of separating the species. A. fallax (Weise), which also has the same spot pattern, can, however, easily be separated from bengalica and its associates by the coarse punctation of the elytra, which is pronounced everywhere on bengalica, etc., but obsolete except near the margin on fallax. The male genitalia of these two species are entirely different. The female genitalia of fallax and maculicollis are similar but more stretched in maculicollis. Type locality.—India, Nilgiri Hills. NO. 15 ‘LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 133 16. AFISSA ATYPICA, new species FIGURES OI, 156 s Abdomen.—Abdominal lines complete, subterminal, rounded. Fifth segment, male, hind margin slightly convex. Sixth segment, convex. Male genitalia——Penis seen from below, 0.8 mm. long, flattened tube, width about 0.3 mm., middle split open with the edges about 0.04 mm. apart, the apex of this slit about 0.2 mm. from the end point of penis, and the sides connected at the apex by a very narrow threadlike transverse bridge, which has a width of considerably less than 0.01 mm. Paramera flattened, 0.1 mm. wide, 0.7 mm. long, slightly curved downward near base. Apical third sparsely covered with hairs. Sipho curved 180° near base, curved into the epnosite direction into a wide hook near apex. Length—4.4 mm. Mandibles with apical and two lateral teeth of about the same size. Color and maculation.—Upper side reddish brown. Pronotum with a narrow elongate spot in the middle. Elytra each with six dark spots, none touching the suture. Nos. 1 and 2 near the base not very distinct. Nos. 3 and 4 forming with their counterparts on the ‘other elytron a nearly straight transverse band. Both transversely widened, No. 3 particularly so. No. 4 touches the margin. No. 6 hardly any farther back than No. 5, near the margin but not touching it. The inner edges of Nos. 3 and 5 equally distant from the suture. Pubescence light gray except on the spots. Under side brown. Type.—U.S.N.M. No. 57141, from India, northern Bengal, Kur- seong, June. Remarks.—tThis is the only species with six elytral spots that does not have the fifth spot on the suture (except maxima, with which it cannot possibly be confused). It also stands out by the fact that spots Nos. 5 and 6 are about equally far back, whereas usually No. 6 is con- siderably farther back. This distinguishes it immediately from all the species of Epilachna with 6 spots. It is very doubtful whether atypica actually belongs to the fallax group. III. FLAVICOLLIS GROUP Elytra each with five spots or less, which occasionally may coalesce. If a discal fascia is formed, it.is not V-shaped near the suture but goes straight across. The female genitalia are of the flavicollis type. (fig. 206) with the plates very elongate and tergite X with a very convex but simple hind margin. 134 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 This group is separated from the preceding one by the fact that the basal spot pattern (fig. 88) has five spots on each elytron. From the complicata and szechuana groups, which have this feature in common, it can easily be distinguished by the shape of the female genitalia. In these two last groups the abdomen is often modified or the elytra have cavities for the reception of the tips of the femora, features never found in the flavicollis group. Because of lack of long series we know very little about the varia- tion of the maculation. In flavicollis, the only species for which a fair series is available, the spot pattern does not show much variation. On the other hand, there are different species with practically the same spot pattern. In such cases the male genitalia are excellent criteria for recognizing the species. They are usually quite differ- entiated and permit in all known cases a clear-cut recognition of the species. The structure of the mandibles is occasionally also of use. Afissa lugubris, which shows quite a different spot pattern (seven light spots on a dark background on each elytron), has been included in this group because of the great similarity of the female genitalia. The flavicollis group contains about 20 species of small to moderate size (length 4 to 8.5 mm.). The figures of the spot patterns and those of the male genitalia will serve better for the identification of the species than any key. Because the male genitalia of a number of important species are still unknown, no attempt has been made to arrange the species of this group into a natural sequence. 17. AFISSA QUADRICOLLIS, new species FicureEs 87, 159, 209 Abdomen.—Abdominal lines subterminal, incomplete, the inner margin gently curved, the apex running almost parallel to the hind margin of the segment for a short interval, then suddenly curved about a right angle and the outer part nearly parallel to the side margin of the segment, big circular punctures in middle of seg- ment, diminishing in size toward apex. About the middle of the segment the abdominal line loses itself. Fifth segment, male, hind margin subtruncate, mildly emarginate; female, slightly convex. Sixth segment, male, mildly emarginate ; female, convex. Male genitalia—Penis seen from below, a flattened tube split open along the middle and ending in blunt point; seen in profile, almost straight. Paramera wider than usually in the flavicollis group, 0.7 mm. long, greatest width 0.15 mm., covered at the periphery of more than the apical half with long (about 0.4 mm.) blond hairs. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 135 Sipho curved strongly at base, then gently until a point where it is bent sharply by about 180°. Before this bend the width is about uniform, 0.07 mm.; at the bend it decreases suddenly to about 0.03 mm. and curved into more than a semicircle. Female genitalia—Most like those of the admirabilis group but tergite X with an even convex hind margin, not so narrow as in the flavicollis group. Length.—5.0 to 6.0 mm. Mandibles with one lateral tooth. Epi- pleurae with shallow cavities for the reception of the tips of the middle and hind femora. Color and maculation—Upper side brownish or yellowish red; pronotum with four spots in a transverse row, the inner ones big, the outer ones small. Often the outer and inner spots joined together (as in fig. 87) ; occasionally the two inner ones almost coalescent, but not in any of the specimens seen by me to such an extent that a continuous band is formed. Elytra with five spots, No. 2 crescent- shaped, emarginate by the callus with the horns pointing outside. Under side light or only slightly darkened. Material examined.—Type: U.S.N.M. No. 57142, China, Chekiang Prov., Hangchow, September 6, 1919 (H. F. Loomis). I2 paratypes from type locality and Kiang-su Prov., Soochow, 1925-1929 (C. F. Wu). Remarks.—This species, because of the crescent-shaped humeral spot bears resemblance to A. 10-maculata Redtenbacher. However, the fact that the pronotum has consistently four spots instead of three and that these specimens are found very far from the type locality of 10-maculata makes it likely that we are dealing here with an entirely different species. It is easily recognized by the peculiar shape of the sipho. The genitalia have no resemblance to those of a species identified as elvina, which is supposed to be closely related to, if not identical with, r0-maculata Redtenbacher. 18. AFISSA FLAVICOLLIS (Thunberg) Ficures 88, 158, 206 C[occinella] flavicollis THUNBERG, 1781, p. 18. Abdomen.—Abdominal lines subcomplete or complete reaching in a regular arc to within one-third to one-fourth of the hind margin of the first segment. Fifth segment, male, hind margin truncate; female, segment wider and hind margin convex. Sixth segment, male, sub- truncate ; female, broadly convex. Male genitalia—Penis seen in profile, almost straight, 1.05 mm. 136 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 long, slightly compressed at about basal third and very slightly bent up at tip. Seen from below, flattened tube about 0.13 mm. wide near base and widening slightly toward apex. Closed at base and the seams gradually diverging. Penis ending in two sharp adjacent points, but slightly divergent. Paramera 0.84 mm. long, slender subcylindrical, about 0.05 mm. wide in middle; apical third covered with long (about 0.2 mm.) hairs arranged in two opposite rows. Sipho moderately curved near base, total bend about 90°, about 0.25 mm. before tip bent sharply in the opposite sense, thinner in the bend. Female genitaha.—Tergite X elongate with strongly convex apical edge. Plates elongate, becoming narrower toward apex, stylus ter- minal, pigment chiefly near apex and on outer edge. Apical third sparsely covered with long, bristlelike hairs. Length.—5.3 to 6.5 mm. Mandibles (fig. 1 E) with a tridentate apical tooth and two prominent lateral teeth of about the same size as the apical tooth. Middle tooth on inside usually with two weak den- tules. Elytra with both coarse and fine punctures pronounced. Color and maculation—Upper side red. Pronotum often yellow, particularly at the sides. Elytra each with five rounded black spots. Neither No. 1 nor No. 3 reaching the suture, No. 4 touching the margin or almost touching. Pubescence yellowish, dark on the spots. Under side all light on the specimens from India, metasternum, abdomen, and mandibles partly dark on the Philippine specimens. Type locality —KEast Indies. Material examined—33 specimens (NM) from the following localities: India, Mormugao, Goa, June 1925, 1924-1926 (J. C. Brid- well, NM). Sumatra: Dolok Silau, 1937 (Mann, NGS-SI Exp., NM). Island of Penang (Baker). Philippine Islands (Baker, except those noted): Luzén Island, Mount Banajao, Mount Maquiling ; Benguet Prov., Irisan, June 1903 (McGregor) ; Nueva Ecija Prov., Minuli (McGregor); Mindanao Island, Surigao, Butuan, Iligan, Zamboanga; Samar Island; Biliran Island; Basilan Island; Sulu Islands, Jolo, July 1924 (A. Duyag) ; Bho, Paete. Borneo: Sandakan (Baker). Remarks.—One female specimen from Candy, Ceylon, has the mandibles of flavicollis but the pronotum with a dark discal spot as in r1-spilota. The color of this specimen is a much deeper red than that of the other specimens of flavicollis. This species, occurring from India to the Philippines, seems to be one of the most widely distributed species of Afissa. It is so similar in external appearance to the next three species that a sep- aration according to external characters only is very difficult. Those NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 137 characters given in the literature to distinguish the species, like the punctation of the elytra, the exact position of the elytral spots, the shape of the abdominal lines or the pubescence of the elytra, are often either too variable or too uncertain for clear recognition of the species. The account of these species in the literature is very con- fusing, and although there seems to be every reason to believe that Afissa flavicollis has been recognized here correctly, the identity of the other three species is less certain because of the small number of specimens available and because of the conflicting statements made by various authors. The structure of the mandibles is an excellent character in addition to the genitalia, which unfortunately are only partly known. A. flavicollis is distinguished from the three other species by the presence of the two large lateral teeth (fig. 1 E) on the mandibles. The male genitalia are quite different frgm those of coccinelloides (fig. 168) and dumerili. Those of the fourth species are unknown. The female genitalia seem to be very similar in structure to those of 11- spilota, but different from those of dumerili, while those of coccinel- loides are not yet known. A. dumerili and r1-spilota have only one lateral tooth (fig. 1 F) and A. coccinelloides none. It would be interesting to know whether differences in food habits correspond to the differences in mandible structure of these otherwise so similar species. 19. AFISSA DUMERILI (Mulsant) Ficures I F, 212 E[pilachna] Dumerili Mutsant, 1850, p. 801. This and the two following species are very similar in appearance to Afissa flavicollis. The male genitalia of A. undecemspilota are unknown. Those of the other three species permit easily a definite separation. The other properties are so much alike that it seems best to characterize them by their differences from flavicollis. Abdomen.—Like flavicollis except that the abdominal lines are somewhat wider and slightly asymmetric, steeper on the outside. Male genitalia—Penis seen from below, length 1.45 mm., split for its whole length. The two parts are touching each other about the middle. The space between them on the basal half is very elongately diamond-shaped, maximum separation 0.07 mm. Beyond the middle the two parts diverge widely, each ending in a blunt point with a small outward hook. Profile nearly uniformly wedge-shaped. Paramera about 0.8 mm. long, slightly bent near base, flattened and subparallel 138 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 for the apical two-thirds, rim of apical third with rather long (max. 0.4 mm.) hairs. Sipho about 5 mm. long, bent through 180° near base, thickened at apex, orifice terminal. The female genitalia (fig. 212) have the plates capable of being folded lengthwise. In the figure one side is open, the other folded. The shape of the plates is more triangular and approaches fallax (fig. 205) in this respect. The mandibles differ from those of flavicollis by having only one lateral tooth considerably smaller than the apical one, which is tri- dentate as in flavicollis. This type of mandible structure is found in many other species of Afissa. The claws have their inner parts touch- ing like figure 2 G. The maculation in both species is very similar, and it is not certain whether the observed differences are significant. Spot 1 is less close to the base, No. 3 is about as close to the suture as No. 1. No. 4 is free from the margin, and No. 5 is a little closer to the suture than it is to the margin. The pubescence is longer and denser and whitish gray instead of yellowish blond as in flavicollis. Type locality.—East Indies. Material examined.—15 specimens: India, northern Bengal, Kur- seong, June (2 specimens, NM, PA); Assam, Shillong, July 1945 (13 specimens J. Unyal, D). 20. AFISSA UNDECEMSPILOTA (Hope) Ficure 88A Coccinella r1-spilota Hope, 1831, p. 31. E[pilachna] undecim-spilota MULSANT 1850, p. 799. Epilachna undecim-spilota CrotcuH, 1874, p. 81. There has been some confusion about this species. Hope’s original description is: “Rubra, thorace margine testaceo elytrisque maculis decem notatis. Long. lin. 2-1/2 lat. 1-1/4.” Mulsant (1850) gave a more detailed description, according to his statement from the original type which had been sent to him by Hope. Crotch (1874) ques- tioned this and claimed Hope’s 11-spilota to be identical with Mulsant’s stephensi, stating that the type is in the British Museum. The de- scription he gave—“Oval, black, head and legs red, sides of thorax yellow ; elytra dull red, each with six black spots of which two are common ; etc.” —does not agree at all with that of Hope, who gave the color as red instead of black. It seems likely, therefore, that Mul- sant saw the original type, which subsequently may have been con- fused with another specimen, NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 139 The two female specimens before me agree fairly well with Mul- sant’s description. They have the mandible structure of dumerili (only one lateral tooth), but the structure of the female genitalia is that of flavicollis. The elytral pubescence is light gray as in dumerili; otherwise the maculation is as in flavicollis except for the pronotum, which has a discal dark spot which is transverse in the Bengal and round in the Siam specimen. Knowledge of the male genitalia is needed to clear up the remaining uncertainties about this and the preceding species. Type locality.—Nepal. Material examined.—2 females (NM): India, northern Bengal, Kurseong, June. Lower Siam, Trong (W. L. Abbott). 21. AFISSA COCCINELLOIDES, new species Figures 80, 168 Abdomen.—Abdominal plates complete, otherwise very similar to flavicollis. Fifth segment, male, truncate to emarginate. Sixth seg- ment, subtruncate. Male genitalia —Not at all similar to those of flavicollis. Penis seen from below, flattened tube about 0.25 mm. wide, about equal width throughout ; split in middle, the seams touching at apex, widest apart at about a quarter of the length. Tube ending into two long widely divergent points, slightly curved upward. Profile, lower edge approximately straight until just before apex; upper edge with a longitudinal ridge approximately in middle ; greatest width about 0.45 mm. Paramera 0.85 mm. long, much shorter than penis, slightly curved down, with long hairs (0.4 mm. long) on apical third. Sipho long, bent in semicircle near base, from then on straight ; tip widened. Length—6.2 mm. Mandibles (fig. 1G) without lateral teeth and without dentules; apical tooth bifid. Color and maculation.—As in flavicollis. Pubescence rubbed off on the type. Type.—U.S.N.M. No. 57143, Borneo, Brunei. 22. AFISSA INCAUTA (Mulsant) FIGURE 224 E[pilachna] incauta Mutsant, 1850, p. 803. Abdomen.—Abdominal lines subcomplete, reaching to about one- fourth of the hind margin of the first segment. Fifth segment, male, truncate. Sixth segment, emarginate. ’ I40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Male genitalia—Penis seen in profile, 0.95 mm. long, bent up gently at about a third of its length and again just before it ends in a sharp point; penis flattened, thickness in middle less than 0.1 mm. Seen from below, a flattened tube widening slightly from the base to about two-thirds of its length, where it has its maximum width (0.33 mm.). There is a seam along the middle of the tube ; orifice elongately diamond-shaped. Beyond the orifice there is a narrow median ridge, which is continued curving upward into the terminal point. Para- mera 0.70 mm. long, slightly curved down at base, of nearly equal width (about 0.08 mm.), covered with blond hairs on practically their whole length. These are densest and longest (0.3 mm.) near the apex. Sipho bent through 180° near base and less strongly in opposite direction near apex (as in fig. 161). Orifice subterminal on outside. The structure of the genitalia of this species is very similar to that of A. gedeensis (see fig. 161), so much so that a much closer rela- tionship between the two species must be assumed than is suggested by the external appearance alone. The genitalia, however, are suffi- ciently different to leave no doubt that two distinct species are in- volved. The chief differences are: The size of the genitalia of incauta is considerably smaller than for gedeensis, even though the body size is larger. Seen in profile, the penis of imcauta is curved up at about one-third of its length, whereas in gedeensis it is curved in the apical half. Length—5.5 mm. Tarsal claws as in figure 2 G. Color and maculation—Upper side yellowish red, head and pro- notum spotless, yellowish, elytra red, each with five black spots. Nos. 1 and 3 on the suture, the former starting at about two-thirds the length of the scutellum, No. 4 on the margin. Average diameter of spots about 0.8 mm. Pubescence yellow, but dark on the spots. Under side light except the tips of the mandibles and the sides of the metasternum, which are darker. Type locality.—Java. Material examined.—1 specimen, Sumatra, Pagaralam (Brues, MCZ). Remarks.—This specimen agrees well with Mulsant’s description of Epilachna incauta. It can be recognized among the species with five elytral spots by the fact that spots Nos. 1 and 3 are on the suture. Crotch (1874, p. 82) mentions specimens having the spots united to form basal and medial fasciae. He may have had other species such as gedeensis, the close relationship of which with incauta has already . NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE I4I been mentioned. The form of gedeensis with the elytral spots un- connected has, however, six spots on each elytron. 23. AFISSA EXPANSA, new species FIGURES 99, 160 Abdomen.—Abdominal lines complete, reaching to within one- fourth of the hind margin of the first segment. Fifth segment, male, hind margin truncate; female, convex and wider. Sixth segment, male, entire, subtruncate ; female, convex. Male genitalia——Penis seen from below, a straight tube split open in the middle; length 0.65 mm., width 0.20 mm. The side margin is sharply folded in and leaves a longitudinal slit of about 0.04 to 0.06 mm. width in the middle. The penis ends in two sharp points slightly turned up. The upper wall of the penis is a semitransparent membrane. Paramera 0.45 mm. long, somewhat widened toward apex, maximum width slightly less than 0.1 mm.; rim of apical half covered with long (0.2 mm.) light-gray hairs. Sipho short (1.2 mm. long), nearly straight until close to apex where it has a sharp bend. Seen from the outside of this bend, sipho suddenly narrowed at the bend, the tip wider again ; orifice round, terminal. Female genitalia—A. dumerili type. The plates folded and pig- ment lacking at the inner edge, length about 0.65 mm. Length.—4.4 to 4.6 mm. Color and maculation.—Upper side light reddish brown; head light to mostly dark. Pronotum with a big transverse spot, leaving only a narrow margin of the pronotum light in the darker specimens or a broad margin in the lighter ones. Elytra each with five large spots, which may partly coalesce. Spot 1 longitudinal, touching or almost touching base and suture, but leaving the scutellum light; No. 2 on the callus practically touching base and side margin. Spot No. 3 transverse, No. 4 practically touching the margin, in some of the specimens, including the type (fig. 99), 3+4 united to form a trans- verse fascia reaching from margin to suture. Spot 5 laterally widened in the darker specimens, practically touching the margin but not quite reaching the suture. Pubescence light gray, dark on the spots. Under side and appendages light with varying amounts of dark. In the lightest specimen only the metasternum and the abdomen darkish. In the darkest one metasternum and mesosternum and most of abdomen and femora black. Material examined.+-Type: U.S.N.M. No 57144, China, Szechwan Prov., Hua Yin Shan, 70 miles north of Chungking, alt. 2,500 feet, July 5, 1933. 10 142 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 7 paratypes with the same data; 1 female, near Muping, 7,000- 13,000 feet, July 6, 1929. (All specimens coll. D. C. Graham.) 24. AFISSA BICRESCENS, new species FiGuRES 96, 166, 210 Abdomen.—Abdominal lines subcomplete, subterminal, slightly un- symmetrical, outer part less curved than inner. Fifth segment, male, hind margin slightly convex; female, more strongly convex. Sixth segment, male, entire ; female, hind margin more convex. Male genitalia—Penis seen from below, straight, length about 0.5 mm., somewhat flattened tube, slit open lengthwise and ending in a split point. Width 0.11 mm. Profile straight, slender wedge with the point very slightly tilted up; upper side flat. Paramera slender, length 0.3 mm., apex clothed with sparse long hairs (up to 0.2 mm. long). Sipho short, about 0.8 mm. long, a slight bend at about one-third, from then on practically straight to tip. Orifice oval on outside, just before tip. Female genitalia—As in dumerili. Length of plates about 0.25 mm. Length.—4.0 mm. Color and maculation—Upper side light brownish red, head with a small black spot on each side next to the eye in the male, with a heart-shaped black spot reaching almost from eye to eye in the female. Pronotum with disk black and a narrow light margin all around, elytra with five black spots arranged 2, 2, 1, as in flavicollis, but greatly modified in shape. No. 1 elongate, touching the base and ex- tending along the suture, which it touches until it ends in a sharp point. No. 2 crescent-shaped with the horns pointing inward and practically touching No. 1, so that only a narrow oval spot between them remains light. No. 3 also crescent-shaped, but the horns point- ing back, touching the suture and joined to No. 4, which touches the margin. No. 5 large, reaching practically from suture to margin. Pubescence light blond, dark on the spots. Under side mostly dark. Prosternum and parts of abdomen light as usual in this group; mouth parts except tips of mandibles, epipleurae, and legs, light. Femora with black spot in middle. Type and one paratype -—U.S.N.M. No. 57145, China, Szechwan Prov., Hua Yin Shan, 70 miles north of Chungking, alt. 2,500 feet, July 5, 1933 (D. C. Graham). NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 143 25. AFISSA ELVINA (Mulsant) FIGURE 172 Epilachna Elvina Mutsant, 1853, p. 122. Abdomen.—Abdominal lines complete, rounded, reaching to within one-fourth of the hind margin of the first segment. Fifth segment, male, hind margin truncate, slightly emarginate; female, convex. Sixth segment, male, hind margin entire, convex, with long light blond hairs ; female, convex. Male genitalia—Penis seen from below, a short tube with a seam along the middle, length 0.45 mm. less than the length of the whole penis. This is followed by a flat blade about 0.2 mm. wide and only very slightly elevated at the margins. Along the middle of this blade runs a very sharp ridge which passes a little beyond the apex of the blade and forms the point of the penis. Paramera nearly straight, shorter than penis, 0.65 mm. long, 0.1 mm. wide. On apical half rim clothed sparsely with long (max. 0.4 mm.) light blond hairs. Sipho curved sharply in a half circle near base, then straight and ending in a straight blunt point. Orifice terminal. Female genitalia.—Plates elongate triangles, 0.95 mm. long, greatest width 0.27 mm. forming a sharp basal point. (More or less as in fig. 210.) Length—5.0 mm. Color and maculation—Upper side red. Pronotum with an elon- gated black spot in the middle and a smaller one on each side. Ely- tra with five large spots 2, 2,1. No. 1 touching the suture, No. 2 with an emargination produced by the callus. Nos. 3 and 5 transversely widened, almost but not quite reaching the suture. No. 4 practically touching the margin. Pubescence dense, light gray, black on the ely- tral spots. Under side and appendages red except the metasternum, most of the abdomen, and the tip of the mandibles, which are dark. Type locality—Northern Provinces, India. Material examined.—z2 specimens: India, Mussoorie, Landour, July 17, 1928 (R. Dudgeon, NM). Remarks—These specimens agree completely with Mulsant’s de- scription of elvina. This species and maculivestis Mulsant (type locality, Tibet; Mulsant, 1853, p. 261) have been regarded as syn- ohyms of Epilachna 10-maculata Redtenbacher (1844, p. 564; type locality, Kashmir). The material accessible to me is insufficient to test the correctness of this. The specimens would agree almost equally well with the descriptions of maculivestis and ro-maculata, and only an examination of the genitalia of all three forms would decide the matter. 144 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Korschefsky (1933b, p. 301) claimed elvina Mulsant to be a species distinct from 10-maculata Redtenbacher and identified some speci- mens from Formosa with it. As this is very far from the type locality, this should be regarded with caution until it is backed by more evi- dence than can be obtained just from external appearance. Afissa quadricollis resembles superficially this species, as it also has the emarginate humeral spot. It can, however, easily be sep- arated by the fact that it has four spots on the pronotum instead of three. The male genitalia of the two species are entirely different. 26. AFISSA DECEMMACULATA (Redtenbacher) Epilachna decemmaculata REDTENBACHER, 1844, p. 564. Abdomen.—Abdominal lines complete, semicircular, reaching to about one-fifth of the hind margin of the first segment. Fifth seg- ment, female, hind margin slightly convex. Sixth segment, strongly convex. Female genitalia——Shape as in figure 210. Length—4.8 mm. Color and maculation.—Upper side light reddish brown. Pronotum with three black spots; the one in the middle roughly longitudinally diamond-shaped, closer to the front than the hind margin, the lateral] ones rounded about one-fifth of the width from the side margins. Elytra with five black spots: No. 1 oval, well behind the scutellum, No. 2 crescent-shaped with the callus in the emargination; No. 3 transverse, not quite touching the suture, No. 4 oval, close to the margin but not touching it, No. 5 large, reaching almost from’suture to margin. Surface opaque. Pubescence yellowish, with black on the spots. Under side, metasternum, the first three segments of the abdomen, and the tips of the mandibles dark, the rest including the appendages light reddish brown. Type locahity—Kashmir. Material examined.—2 specimens: India, Chabua, August 1943 (D. E. Hardy, NM) ; United Prov., Kumayun, Bhawaki, 5,000 feet, June 18, 1944 (J. Unyal, D). Remarks.—This specimen agrees well with Redtenbacher’s figure and his scanty description. It differs from the species here identified as elvina by being not quite so wide, having an opaque surface, while elvina has the elytra smooth and shiny and having yellowish rather than grayish pubescence. The elytra are compressed at the sides and have a distinct depression near their base. Though the two species are therefore distinct, the evidence as to which is which is much less definite. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 145 27. AFISSA MYSTICOIDES (Sicard) FIGURES 95, 164 S[olanophila] mysticoides S1cArp, 1912b, p. 507. Abdomen.—Abdominal lines subcomplete, evenly rounded, reach- ing to within a third of the hind margin of the first segment. Fifth segment, male, hind margin truncate; female, subtruncate or very slightly convex. Sixth segment, male, subtruncate; female, convex, narrower than that of the male. Male genitalia—Penis seen in profile, lower margin straight until near apex, then curved up; upper margin nearly straight for about half the length, then curved up and sharply bent back to the original direction so that the apical half is considerably wider than the basal half. Seen from below, a tube, about 0.2 mm. wide, split along its entire length and ending in two short, well-separated points. Near the apex the margin of the seam is bent vertically up. The two apical points are joined immediately to the apical margin of-the orifice. Paramera slender, 0.7 mm. long, shorter than the penis, about 0.06 mm. wide, periphery of the apical fourth covered with rather short hairs. Sipho bent through 180° near base and continues nearly straight after that for most of its length. Near apex it is widened and a side branch emerges from it and extends about 0.3 mm. beyond the main part, then is curved inward and thickened at its apex. The orifice is terminal to the main part. Female genitalia—Of the flavicollis type, plates very narrow and dark toward apex. * Length.—4.0 mm. Color and maculation—Upper side brick red; pronotum with a transverse spot which has almost the complete width of the pronotum and is angular behind. Elytra each with five spots, the first elongate oval near the suture, but not touching it; No. 2 crescent-shaped, concave inward, on the outside very close to, but not quite touching, the margin around the humeral angle, partly encircling the callus, the hind tip touching spot No. 1; No. 3 transversely rectangular, No. 4 subquadratic with rounded corners; No. 5 rhomboidal. Pubescence light gray, black on the spots. Under side mostly black, except pro- sternum, parts of mesosternum, and the last abdominal segments, the latter two, however, are not completely light. Appendages and epipleurae light. Type locality.—India. Material examined.—2 specimens: India, near Darjeeling, eastern Himalayas, alt. 7,000-8,000 feet, May 1910 (R. B. Horsfall, AMNH). 146 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Remarks.—This species agrees very well with Sicard’s description except that the head and pronotum have the same color as the elytra and are not yellow as Sicard states. Such color differences occur frequently within one species. As the name indicates, this species re- sembles in its maculation A. mystica (Mulsant) and forms an inter- mediary between the regular 5-spotted species such as flavicollis and mystica. It can easily be distinguished from other 5-spotted species by the form of spot No. 2. 28. AFISSA MYSTICA (Mulsant) FIGURES 100, 169 E[pilachna| mystica MuLSsANrT, 1850, p. 841. Abdomen.—Abdominal lines complete, reaching to within one-third of the hind margin of the first segment, regularly curved into semi- circles. Fifth segment, male, hind margin subtruncate ; female, mildly convex. Sixth segment, male, emarginate; female, subtruncate. Male genitalia.—Penis seen in profile, bent gently upward near the middle and down again just before the tip, which is a sharp point; length 0.9 mm., gradually decreasing in thickness from base to apex. Seen from below, tube split open for basal two-thirds. The last third solid, narrower (0.07 mm. wide) and ending in a single sharp point. Paramera slender, 0.85 mm. long, subparallel, 0.07 mm. wide, rim of apical third with a row of hairs. Sipho bent 180° near base, less strongly in the opposite direction near apex. Orifice rounded, sub- terminal. Female genitalia—Like those of dumerili. Basal ends of plates ending in sharp points, apex folded longitudinally. Length 1.0 mm. Length.—s.0 to 5.8 mm. Body very convex. Tips of elytra broadly margined with heavy punctures. Color and maculation.—Upper side brownish red; pronotum with a triangular discal spot; elytra each with five spots but the pattern considerably modified. Spots 1 and 2 together form practically a complete: circle, 3 laterally and 4 longitudinally widened, the latter usually also extending to the margin. No. 5 when fully developed touching No. 4 near the suture and extending as a narrow curved band practically to the margin. In lighter specimens practically ab- sent or a narrow elongated spot near the suture. Pubescence light, dark on the spots. Under side and appendages ‘light, metasternum and mesosternum and most of abdomen dark. Type locality —East Indies (?). Material examined—s5 specimens: Himalaya (NM); India, Sikkim. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 147 29. AFISSA DECEMGUTTATA (Weise) Ficures 08, 167 Solanophila decemguttata WEISE, 1923b, p. 183. Abdomen.—Abdominal lines not quite complete, forming a regular arc reaching to within one-third of the hind margin of the first seg- ment. Fifth segment, male, subtruncate. Sixth segment, with hind margin evenly convex. Male genitalia—Penis seen from below, a tube split open along the middle; the tube proper narrows gradually toward the apex and ends in a fine straight point. While the tube narrows, a side flange develops on each side so that the total width of the penis remains approximately constant (0.2 mm.). The apex is cut off squarely but the point into which the penis tube ends extends about 0.1 mm. beyond it. Paramera narrow, about 0.7 mm. long, less than 0.1 mm. wide, curved down in their basal third, from then on straight. In their apical half sparsely clothed with long blond hairs. Sipho long, only slightly tapered, about 0.07 mm. thick in middle, curved near the base, from then on straight, ending in a blunt point with the small orifice just before it. Length—4.7 mm. Elytra with very coarse punctures. Color and maculation—Upper side red, pronotum with a small *black spot in the middle nearer the front margin than the base; elytra each with five spots, Nos. 1 and 3 close to the suture but not touching it, No. 4 touching the margin, Nos. 3 and 5 transversely broadened. Pubescence light gray, dark on the spots. Under side light, with the sides and the metasternum black and part of the abdomen dark. Type locality—Formosa, Taihorin. Material examined.—Formosa, Musha, May 3, 1929 (K. Sato, NM). 30. AFISSA ANHWEIANA, new species FIGURE 226 Abdomen.—Abdominal lines subcomplete reaching to within one- fourth of the hind margin of the first segment. Fifth segment, wide in both sexes, slightly emarginate in the male, convex in the female. Sixth segment, not protruding from under the fifth. Male genitalia—Penis 1.5 mm. long; seen from below, a tube 0.3 mm. wide, closed for about two-thirds of its length with the seams open, beyond this an open trough. Apex with two points 0.15 mm. apart, with a semicircular emargination between them. Seen from the side, mildly curved up near its apical third and gradually tapering 148 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 in thickness. Paramera 1.35 mm. long, greatest width 0.2 mm. near the apex. Rim of apical half with long (0.5 mm.) blond hairs. Sipho long and slender, curved 180° near base, a notch on the outside just before the tip carries the oval orifice. The sipho tube laterally com- pressed just before the notch. Female genitalia——Plates very elongate triangle, as in figure 207, but even more elongate. Length 1.2 mm., greatest width 0.27 mm., inner edge distinct and straight. Length—7.5 mm. Elytral epipleurae with traces of cavities for the reception of the hind femora. Mandibles with a broad single apical and one lateral tooth, no dentules (as in fig. 1 F, but less elongate). Color and maculation—Upper side light reddish brown with long whitish-gray pubescence. Pronotum with a median black spot of vary- ing size and shape. It may be transverse about half the width of the pronotum, almost touching the front margin but leaving about a third free at the hind margin. It may be much smaller and diamond- shaped, situated in front of the middle. Elytra each with five rather large black spots. Nos. 1, 3, and 5 close to the suture but not touching it. No. 1 elongately oval, line tangent to the common base passing through the middle of the scutellum. No. 2 behind the callus. No. 3 big, transverse, inner edge straight, outer one semicircular. No. 4. elongately oval, not touching the margin. No. 5 variable in size and* shape. In the darkest specimen the spot almost reaches from suture to margin, and there is a fine dark line parallel to the suture and close to it connecting spots 3 and 5. Under side and appendages light brown, except the tips of the mandibles, which are dark. Type-—Museum of Comparative Zoology, from China, Anhwei Prov., Taipingshien, October 1932 (G. Liu). Material examined.—1 paratype, same data as type, I specimen: Anhwei Prov., Kiuhua Shan, September 1932 (Liu, D). 31. AFISSA LONGISSIMA, new species FIGURES 104, 173, 207 Abdomen.—Abdominal lines subcomplete, reaching to about within one-fifth of the apical margin, broadly rounded. Fifth segment, male, hind margin truncate to slightly concave; female, convex. Sixth segment, male, distinctly emarginate; female, convex. Male genitalia.—Penis a flattened tube 1.8 mm. long; seen in pro- file, practically straight, the apical point turned up, diminishing in thickness from base to apex. Seen from below, tube split open along NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 149 whole length except for apical point, very flattened toward apex. Base of apical point beginning behind the elongate orifice, raised above the rest. Point long, with the sides almost parallel, less than 0.1 mm. wide at base, curving up. Paramera 1.2 mm. long, much shorter than penis, slender, subparallel with rounded apex, greatest width slightly more than 0.1 mm. Sides of apical third covered with a row of long (0.4 mm. max.) light blond hairs. Sipho very long (more than 5 mm.) and slender, width less than 0.05 mm. during most of its length, strongly curved through more than 180° near base, slightly in opposite direction near apex. Orifice terminal, small. Female genitalia—Plates very elongate, 1.03 mm. long, ending at base in sharp point. Emargination on inner margin near apex. Length.—6.5 to 7.0 mm. Apical angles of elytra rounded. Color and maculation——Upper side light brownish red, pronotum with small central spot, elytra with five spots arranged as shown in figure 104. No. 3 very wide and touching or almost touching No. 4, which touches the margin. Pubescence light gray except on the spots. Under side and appendages light except the sides of the metasternum and the tip of the mandibles. Material examined.—Type: U.S.N.M. No. 57146, Formosa, Tai- hoku, September 22, 1927 (T. R. Gardner). I paratype: Same data. 32. AFISSA CHINENSIS (Weise) FIGURE 225 Solanophila chinensis WEISE, 1912, p. 112. Abdomen.—Abdominal lines complete, rounded, reaching to within one-quarter of the hind margin. Fifth segment, male, hind margin truncate or slightly convex ; female, convex. Sixth segment, convex in both sexes. Male genitalia—Penis seen in profile, 1.0 mm. long, lower edge straight for the basal two-thirds, then gently curved up and again curved up at the tip. Seen from below, straight tube, about 0.15 mm. wide, closed with a seam along the middle, orifice subterminal, oval. Paramera 0.9 mm. long, slender, maximum width (near apex) 0.07 mm., rim covered for apical two-thirds sparsely with row of blond hairs (about 0.25 mm. long). Sipho bent near base through an 180° arc, then straight and near apex bent outward again through nearly 180°, but arc smaller, distance between vortices of bends 1.8 mm.; orifice terminal. I50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Female genitalia—Shape as in figure 210. Length of plates 1.0 mm., greatest width 0.3 mm. Length.—5 to 5.5 mm. Mandibles with a tridented apical tooth and two large lateral teeth and one small one. The latter at the base of © the innermost of the two larger teeth. Color and maculation—Upper side brownish red. In the darker specimens with a transverse black spot at the base of the head ; pro- notum with a transverse black spot which occupies about two-thirds the width and one-half the length of the pronotum and is closer to the front than to the hind margin. Elytra with five large black spots, No. I a common spot on the suture reaching or almost reaching the base and enveloping the scutellum, which remains light; No. 2 rounded and practically touching the margin; No. 3 transverse, not reaching the suture; No. 4 a little longer than wide, on the margin; No. 5 rounded, not reaching either suture or side margin. Pubescence light gray, dark on the elytral spots. Under side light, but sides of metasternum dark. Type-——Hamburg Museum. Type locality—Fo-kien. Material examined.—4 specimens: China, Anhwei Prov., Taiping- shien, October 1932 (Liu, MCZ, D). Remarks.—These specimens agree in all particulars with Weise’s description. There are four additional specimens which show no noticeable differences in structure. They have the same general ap- pearance except that the pronotal spot now covers practically the whole pronotum and leaves light only a narrow margin all around. Spot No. 1 does not touch the suture or base and is oval and longer than wide. The other spots show no difference. Unfortunately, all specimens are females, and so in the absence of the male genitalia absolute certainty cannot be obtained on whether this is more than just a color variation. The two forms are sharply differentiated with- out intermediates. These forms are tentatively regarded as a variety of A. chinensis. AFISSA CHINENSIS var. SEPARATA, new variety Like the ground form except that the pronotal spot leaves only a narrow light margin and elytral spot No. 1 does not touch suture or base and is longitudinally oval. Material examined—Type: Museum of Comparative Zoology, from China, Anhwei Prov., Taipingshien, October 1932 (Liu). 3 paratypes, all from China, Anhwei Prov., 1 same data as type, 2 from Kiubua Shan, September 1932 (Liu, MCZ, D). NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE I5I 33. AFISSA PROVISORIA, new species FIGURES 103, 174 Abdomen.—Abdominal plates coming to within about two-fifths of the hind margin of the first segment. Fifth segment, male, mod- erately concave; female, truncate. Sixth segment, male, hind edge convex, with a distinct emargination in the middle, to truncate ; female, convex. Male genitalia—Seen from below, two parallel almost cylindrical bars which unite at apex to form a sharp barely split point. The bars are connected on their upper side by an almost clear membrane. Seen in profile, gently curved up near base, from then on straight to apex. Paramera shorter than penis, about 0.5 mm. long, slender, greatest width about 0.1 mm., covered with hairs on inside of apical half. Sipho curved near base and very slightly in the opposite direction near apex. ; Female genitalia—As in flavicollis. Length—5.5 to 6.0 mm. Mandibles with one lateral tooth, no dentules ; apical tooth trifid. Shape: side margins of elytra nearly straight, parallel. Color and maculation——Upper side brownish yellow, head with a transverse dark spot at the base almost reaching from eye to eye; pronotum with a big discal spot leaving only a very narrow margin in front and back and about a quarter of the width at each side. Elytra with five big dark spots as in figure 103. Spots 1 and 3 almost but not quite reaching the suture. Under side dark, but part of proster- num and apical segments of abdomen slightly lighter. Epipleurae of prothorax and elytra yellow, legs and mouth parts reddish; middle parts of femora usually darker. Material examined.—Type: U.S.N.M. No. 57147, China, Szechwan Prov., Mount Omei, alt. 11,000 feet, August 19, 1934 (D. C. Graham). 5 paratypes, same data as type: Tsu Dien, Mount Omei, alt. 5,000- 7,000 feet, August 7, 1925 (D. C. Graham, NM) (1 specimen) ; Mount Omei, August 6, 1938 (Dean Sage, Jr.. MNH) (2 specimens). Remarks—tThis species agrees nearly with the description of several known species but exactly with that of none and is probably a hitherto undescribed species. 34. AFISSA OCELLATAE-MACULATA (Mader) FIGURES 97, 165 Solanophila ocellatae-maculata MADER, 1930, p. 183. Abdomen.—Abdominal lines variable, not quite complete, reaching to within less than one-fifth of the apical margin of first segment. I52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Fifth segment, male and female, hind margin subtruncate. Sixth segment, male, mildly emarginate; female, entire. Male genitalia—Penis seen from below, a tube split wide open lengthwise. Width 0.23 mm., seams apart 0.1 mm. The penis ends in a straight point. The upper wall of the tube is a semitransparent membrane. Paramera 0.7 mm. long, widened toward apex; greatest width 0.13 mm.; furnished on the rim of more than the apical half with blond hairs. Sipho short, widened near apex, orifice on one of the widened faces just before the tip, small and round. Female gemtalia—Of the flavicollis type. Tergite X subtruncate. Length._—4.8 to 5.4 mm. Color and maculation.—Upper side light reddish brown; pronotum with a transverse black band almost but not quite from side to side. In the lightest specimens resolved into three separate spots, in the darkest continuous and leaving only a narrow light margin on base and apex, usually taking up about the middle third. Elytra with five dark spots arranged 2, 2, I as in flavicollis; Nos. 1 and 3-5 deviating only little from circular shape. No. 2 like an inverted comma, when fully developed touching No. 1. The space between the spots dark, but less so than the spots themselves, and a ring around each spot com- pletely light. Pubescence light gray, black on the spots. Under side almost completely black in the darker specimens with large parts of abdomen and prosternum light in the lighter specimens. Mouth parts, legs, and epipleurae light. Femora with a dark spot in the darker specimens. Type locality—China, Yunnan Prov., mountains near Mengtze. Material examined.—tg specimens (D.C. Graham), all from China, Szechwan Prov., from the following localities: Wei Chow, 65 miles northwest of Chengtu, alt. 9,000-12,500 feet, August 12, 1918; alt. 5,600-8,900 feet, July 26-30, 1933; near O Er, alt. 6,000-15,000 feet, August 6-18, 1933; Beh Luh Din, 30 miles north of Chengtu, alt. 6,000 feet, August-September 1943 ; Hua Yin Shan, 70 miles north of Chungking, alt. 2,500 feet, July 5, 1933; near Fu Liu, alt. 3,000- 8,000 feet, August 18-21, 1926; alt. 5,000-8,200 feet, July 19-21, 1928; near Kuanhsien, alt. 2,000-4,000 feet, August 1933; Ningyu- enfu, alt. 7,800 feet, August 13, 1928. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 153 35. AFISSA PLICATA (Weise) FIGURE 97A Epilachna plicata WEISE, 1889, p. 649. Abdomen.—Abdominal lines nearly complete, curved in regular arc. Fifth segment male, hind margin truncate; female, slightly con- vex. Sixth segment, male, mildly emarginate; female, entire. Male genitalia (see fig. 165).—Penis seen from below, a tube cut in half lengthwise, ending in a straight point, width 0.23 mm. The upper wall of the tube is a semitransparent membrane. Paramera 0.7 mm. long, widened slightly toward apex, greatest width 0.11 mm., furnished on the rim of more than apical half with blond hairs. Sipho short, widened near apex, orifice small, round, on one of the widened faces just before the tip. Genitalia indistinguishable from those of ocellatae-maculata. Female genitalia—Of the flavicollis type, like ocellatae-maculata (Mader). Length—5.0 to 6.0 mm. Elytra near apex with a plica parallel to the side margin. Color and maculation—Upper side yellowish brown; pronotum with a transverse dark band, which in the lighter specimens seems to resolve into six or seven indistinct spots; in the darker specimens a big central spot to which a lateral spot is joined at each side. Elytra with three transverse bands, which can be recognized as a modification of the 5-spotted pattern of the flavicollis type. Spots 1 and 2 are joined together and form a distorted semicircle open in front. Spot 3 is widened laterally, while spot 4 remains nearly circular. In many specimens these two spots are joined together to form a continuous narrow band, reaching from margin to suture but not quite touching either. Spot 5 also widened laterally into a band, which reaches nearly from margin to suture. Under side dark, except the prosternum, most of the middle of the basal seg- ments and the entire apical segments, and often the middle of meso- sternum light ; mouth parts, legs, and epipleurae light. Type locality—China, Kansu Prov. Material examined.—13 specimens (D. C. Graham, NM), all from China, Szechwan Prov.: Near Mupin, alt. 3,000-7,400 feet, alt. 3,000-7,600 feet, July 1-3, 1929; alt. 7,000-13,000 feet, July 6-8, 1929; alt. 5,000-6,500 feet, July 22-24, 1929; Mupin, alt. 4,500 feet, July 24, 1929; alt. 5,000-6,000 feet, July 3, 1929. Remarks.—The very great similarity of the structure of both male and female genitalia shows that A. plicata (Weise) and A. ocellatae- 154 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 maculata (Mader) are very closely related. The presence of the plica in A. plicata and the difference in maculation render it, however, very easy to keep the two species apart. Only a single female is available of each of the following six species, a situation that makes it difficult to establish their affinity. The structure of the female genitalia and, except for lugubris, the spot pattern leave no doubt that they belong to the flavicollis group. They are also sufficiently different in structure from each other and other species to make it relatively certain that they belong to distinct species. 36. AFISSA ANNAMENSIS, new species FIGURE 102 Abdomen.—Abdominal lines subcomplete, rounded, reaching to within about one-fourth of the the hind margin of the first segment. Fifth segment, female, hind margin truncate. Sixth segment, entire. Female genitalia —Of the flavicollis type. Length—8.5 mm. Color and maculation—Upper side red. Pronotum with a small, not very distinct, dark discal spot. Elytra each with five black spots: Nos. 1 and 2 almost touching the base and joined together; No. 3 subquadratic, near the suture; No. 4 almost touching the margin; No. 5 transverse, slightly emarginate behind, a little closer to the margin than to the suture. Pubescence light gray, dark on the spots. Under side and appendages light, except metasternum, the middle part of the first three abdominal segments, and the tips of the mandi- bles, which are dark. Type—vU.S.N.M. No. 57148, Indochina, Annam Prov., Haut Donai, Col de Blao, alt. goo meters, September 30, 1932 (M. Poilane). 37. AFISSA INDOSINENSIS, new species Abdomen.—Abdominal lines complete, wide, reaching to within one-fourth of the hind margin of the first segment. Fifth segment, female, wide, hind margin convex. Sixth segment, convex. Female genitalia—Of the flavicollis type; length of plates 0.95 mm. ; greatest width'0.25 mm. Length.—6.0 mm. Color and maculation.—Upper side dark brownish red; pronotum with two large transverse spots narrowly interrupted in the middle. Elytra each with five black spots: No. 1 on the suture, reaching almost NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 155 to the base but leaving the scutellum light; No. 2 rounded, on the callus; No. 3 slightly transverse; No. 4 touching the margin; No. 5 subapical, rounded closer to the suture than to the margin. None of the spots, because of the relatively dark ground color and their somewhat hazy outlines, are very distinct. Pubescence long, white, brown on the spots. Under side light brown, sides of metasternum and basal part of abdomen darker. Type—uvU.S.N.M. No. 57149, Indochina, Annam Prov., Haut Donai, Agr. Station of Blao, alt. 800 meters, April 10, 1933 (M. Poilane). 38. AFISSA ORTHOFASCIATA, new species FIGURE IOI Abdomen.—Abdominal lines complete, broadly rounded, reaching to within one-third of hind margin of first segment. Fifth segment, female, hind margin convex. Sixth segment, narrow, hind margin convex. Female genitalia—As in flavicollis, plates somewhat wider, 1.05 mim. long. Length—6.5 mm. Color and maculation—Upper side red, pronotum spotless, elytra with two straight transverse black fasciae, the first along the base, leaving only the scutellum red, hind margin nearly straight; the second about the middle of the elytra narrower and slightly com- pressed in the middle of each elytron. In addition a subapical black spot, touching the margin but removed from the suture. Pubescence light red, black on the spots. Under side light, prosternum, meta- sternum, and the middle of the first three abdominal segments black ; appendages light, tip of mandibles dark. Type—vU.S.N.M. No. 57150, Java, Tjibodas, Mount Gede, April 20, 1909 (Bryant and Palmer). Remarks.—This species resembles superficially very closely Epi- lachna parafasciata from the same locality. The two can easily be told apart by the generic characters (structure of claws, sixth ab- dominal segment, and female genitalia). There are other minor dif- ferences, but the similarity of the color pattern is striking. FE. para- fasciata is somewhat more yellowish. 39. AFISSA COMPLETA, new species FIGuRE 108 Abdomen.—Abdominal lines complete, evenly rounded and reaching to within about one-fourth from the hind margin of the first segment. 156 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Fifth segment, female, with moderately convex hind margin. Sixth segment, narrow, strongly convex hind margin. Female genitalia.—Of the flavicollis type. Length.—8.0 mm. Color and maculation—Light reddish brown ; pronotum with a big transverse discal spot that leaves only the margins free. Elytra with four big spots that leave only narrow margins of light color at the suture and between the spots. The first, which is apparently spots Nos. I and 2 of the 5-spotted pattern united, has an emargination at the callus. Surface shining with sparse pubescence (possibly partly rubbed off in the specimen). Under side dark on sides of me- tasternum and the basal part of abdomen, the rest light. Type—vU.S.N.M. No. 57151, China, Szechwan Prov., Lu Ding Chiao, 5,000 feet, July 13, 1930 (D. C. Graham). Remarks.—This species, which, because of the structure of the abdomen and genitalia, must be placed in the flavicollis group and not near convexa which it resembles superficially, seems to be very close to Solanophila lenta Weise (1902, p. 495; type locality, Tonkin). The chief difference seems to be the size and the fact that Jenta is subopaque, but completa shiny. 40. AFISSA INTERMIXTA, new species Abdomen.—Abdominal lines complete, reaching to within about one-third of hind margin of first segment. Fifth segment, female, hind margin convex. Sixth segment, convex, very short. Female genitalia —Of the flavicollis type, as in figure 206. Length—4.8 mm. Color and maculation—Upper side light yellowish brown. Pro- notum with three dark spots, each about one-sixth of the width of the pronotum. The central one a little before the middle, darker than the lateral ones, which are close to the margin. Elytra each with seven dark spots. This can be regarded as the normal 5-spotted regular pattern of the flavicollis group with the following modifi- cations: Spot No. 2, which is crescent-shaped in several species, is completely split in two with the callus between the parts; No. 3 very large, laterally widened; No. 4 circular, not touching the margin; No. 5 about where it is normally in this group. In addition another spot near the margin very slightly farther back. Pubescence light yellowish, dark on the spots. Under side and appendages light to darkish brown. Type—U.S.N.M. No. 57152, India, Assam, Dibrugarh, March 20, 1944 (D. E. Hardy). NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 157 Remarks.—The shape of the female genitalia leaves little doubt that this species belongs to the flavicollis group, although it is the only species of this group known so far having more than five spots. 41. AFISSA LUGUBRIS, new species Ficure 106A Abdomen.—Abdominal lines complete, reaching to within one- fourth of the hind margin of the first abdominal segment and evenly curved. Fifth segment, female, hind margin convex. Sixth segment, hind margin convex, scarcely protruding from under the fifth. Female genitalia—Almost exactly like those of flavicollis (fig. 206). The only noticeable difference is that the inner margins of the plates are more distinct near their base and overlap. Length—6.0 mm. Mandibles with a strongly bifid apical tooth and two lateral teeth of almost equal size without dentules. Color and maculation—Upper side black ; head partly lighter, pro- notum with an indistinct lighter margin. Elytra each with six red- dish-brown spots with indistinct outlines arranged I, 2, 2, 1, the first touching the base. Under side dark with prothorax and mesothorax, epipleurae, abdomen toward apex, and appendages more or less lighter. Type.—vU.S.N.M. No. 57153, China, Kiautschau. Remarks.—This species agrees reasonably well with the descrip- tion of Epilachna pembertoni Crotch (1874, p. 80; type, Indian Museum ; type locality, India, Bhootan). The identity of Jugubris with pembertoni Crotch seems, however, doubtful because of some dif- ferences in coloration and the difference in locality. IV. COMPLICATA GROUP This group is best characterized by the structure of the abdomen and the genitalia. The last visible abdominal segments are greatly modified in most species of this group, particularly in the males, and such modifications have not been observed in any other group of Afissa. These modifications consist of deep emarginations, depres- sions, greater degree of sculpture in general, and processes on some segments, and may affect the third to sixth visible segments. The greatest degree of modification is observed in Afissa complicata. The female genitalia have tergite X with a simple convex to slightly emarginate apical edge, which distinguishes this group sharply from the szechuana and chapini group where the edge is bent double. The plates are suboval, wider than in the flavicollis group and with the II 158 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 greatest width in the apical half. The male genitalia (figs. 175, 176, 179) are rather elaborate in convexa and complicata and in all cases have very wide paramera, wider than in any other Afissa group. The sipho is relatively straight and short. The basic spot pattern in this group, as in the preceding and fol- lowing one, consists of five rather large spots on each elytron which may flow together and cause the upper side to be predominantly black in some species. The group consists of four moderately large (6 to 9 mm. long) species from China, three of them new. 42. AFISSA CONVEXA, new species FIGURES 109, 175, 213 Abdomen.—Abdominal lines terminal, incomplete. The outer mar- gin curved inward and reaching to about the middle of the first seg- ment. Punctation of central part of first segment normal, not consisting of wide circular punctures (as in admurabilis or chapint). Fifth segment, male, in the middle very convex with a depression on each side, hind margin approximately straight ; female, uniformly bent and only slightly convex, hind margin with a broad, shallow emargination. Sixth segment, male, strongly convex with the. hind margin deeply emarginate; female, bilobed with a deep notch be- tween the lobes, but not completely split. Seventh segment appearing through the emargination. Male genitalia—Penis about 2.2 mm. long, a high narrow tube, split lengthwise in the middle on the under side. The apex open, with the upper part reaching farther back, bilobed and ending in two blunt points curved mildly upward. Penis sparsely covered with hairs on its upper middle. Paramera spoon-shaped, about 1.7 mm. long, maximum width 0.5 mm. On the apical part the edge covered densely with long (0.5 mm.) blond hairs. These are much longer and denser on the upper edge than on the lower one. Sipho slightly S-shaped, almost straight; bent sharply just before the apex and ending in a sharp point. Orifice just inside this bend, elongate. Female genitalia —Tergite X, subtruncate with a slight emargina- tion, sides slightly convex, at the base the pigment limited by an al- most semicircular line. Plates 0.75 mm. long, oval, pointed at base, inner edge nearly straight. Length.—7.5 to 8.5 mm. Side of elytra compressed just before apex, so that the reflexed margin appears wider there. Color and maculation—Upper side, color light yellowish brown. NO. 15 LADYBEETLES OF THE GENUS EPILEACH NA—DIEKE 159 Pronotum with a big central spot touching the base and reaching almost or entirely to apex and a smaller spot on each side. Elytra with five large black spots, Nos. 1 and 2 often touching, No. 4 touch- ing the margin. Pubescence blond, dark on the spots. Under side same color as above with metasternum black and the abdomen more or less dark brown to black. In light specimens the last segments are completely light, and the middle of the second to fourth ; in dark specimens only the sixth and the middle of the fifth. In such specimens also the mesosterna, prosterna, and femora are darker. Otherwise appendages light except tips of the mandibles. Material examined.—Type: U.S.N.M. No. 57154, from China, Szechwan Prov., near Mupin, alt. 3,000-7,600 feet, July 1-3, 1929 (D. C. Graham). 7 paratypes, 6 from same locality: alt. 7,000-9,000 feet, July 17, 1929; alt. 7,000 feet, July 18, 1929; 7,000-13,000 feet, July 6-8, 1929. Additional specimens: between Chengtu and Kuanhsien, July 2-5, 1924. Remarks.—tThis species is in appearance very similar to A. sus- tenans, complicata, and subacuta. It can easily be distinguished from them by the structure of the abdomen and the genitalia. The differ- ences in maculation are distinct though slight, but it is not sure whether they would hold in a large series. 43. AFISSA COMPLICATA, new species FIGURES III, 176, 214 Abdomen.—Abdominal lines subterminal, incomplete, the apical part flattened and parallel to the apical edge of the segment, the outer part parallel to the margin of the segment and ending in a small depression not far from the middle of the segment. Male: Third and especially fourth segment much narrower in the middle than at the sides, the fourth with a process beginning at the middle of the apical margin and reaching back over the fifth segment. It is about 0.4 mm. long, ends in a fork, and serves to support the extended genitalia. The fifth segment also shows considerable modifications. It has a wide semi- circular groove reaching from side to side with elevated edges bor- dering it, especially the basal edge near the sides. At the middle of its apical margin it bears an erect emarginate process shorter than the one of the fourth segment. The sixth segment is deeply emarginate, practically to its base, and through this opening the seventh segment emerges. Female: Third and fourth segments are normal, that is, of equal uniform width and with truncate hind margins. The fifth 160 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 segment is strongly emarginate. There is a basal depression in the middle and an apical one on each side of it, but apparently this seg- ment is not very strongly sclerotized and these depressions are smoothed out in some specimens. Sixth segment mildly bilobed, also weakly sclerotized. Male genitalia—tThese are equally as elaborate as the abdominal segments. Figure 176 shows the genitalia of sustenans, which have practically the same structure as those of complicata. The penis tube, straight, 1.0 mm. long, split open on its under side, is under a wide shield and partly fused to it. The shield has parallel smooth edges and ends in a point, which is sharply bent down. On the basal two-thirds of its upper side it carries a sharp median vertical ridge with a few hairs on each side. Paramera spoon-shaped, about 1.0 mm. long, 0.5 mm. in maximum width, with long hairs (about 0.4 mm.) around most of the margin. Sipho very slightly bent near base, the apex as in figure 176. Female genitalia—Tergite X with the apical margin regularly convex. Plates elongately oval, pointed at base with the inner margin emarginate. Length.—7.2 to 9.0 mm. Color and maculation—Upper side light yellowish brown; pro- notum with a median spot closer to base than apex, which shows signs of being due to the confluence of two longitudinal spots close to the middle. In some specimens there are indications of a lateral spot. Elytra each with five large black spots: Nos. 1 and 2 sub- triangular, No. 4 touching the margin. Pubescence light blond, black on the spots. Under side and appendages mostly light on the lighter specimens. Most of metasternum, basal parts of abdomen, and tips of mandibles dark. In the darker specimens the dark color- ation may spread over most of the abdomen, the mesosternum, and part of the prosternum. Also the femora are darker in such specimens. Material examined.—Type: U.S.N.M. No. 57155, from China, Szechwan Prov., near Mupin, July 1-3, 1929, alt. 3,000-7,400 feet, (D.C: Gratiam )). ; 10 paratypes, all China, Szechwan Prov. (Graham, NM): Same data as type (4 specimens) ; between Yachow and Tatsienlu, alt. 2,200- 8,000 feet, July 7-14, 1930; near Washan, alt. 2,000-5,000 feet, July 18, 1925; Mount Omei, Si Gi Pin, alt. 4,000-6,000 feet, August 6-15, 1925. Remarks.—A fissa complicata can easily be distinguished from other species with similar appearance by the remarkable structure of the abdomen. It also is well characterized by the genitalia. NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 161 AFISSA COMPLICATA var. SUSTENANS, new variety FIGURES IIO, 176 This form is so close to A. complicata that although there are a few distinct differences it is probably only a variety. More material would settle this definitely. The differences from the typical com- plicata are as follows: In external appearance sustenans seems to be a form with the black pigment more developed than in complicata s. str. The pronotum is black with a narrow light margin, but there are intermediates between this and the typical complicata form (fig. III). The elytral spots are larger, Nos. 1 and 2 touch (in one specimen both in front and back so that they enclose a small oval light space), Nos. 3 and 4 are joined together, and No. 5 practically extends from suture to margin. The genitalia show some slight differences. The shield of the penis has parallel edges in complicata, but they diverge and are more ragged in sustenans, though the raggedness is somewhat exaggerated in figure 176. The tip of the sipho has the apical process shorter and less distinct and less strongly bent back. Material examined.—Type: U.S.N.M. No. 57156, China, Szech- wan Prov., near Yueh Shi, alt. 4,000-8,000 feet, July 21, 1928 (D. C. Graham). 2 paratypes, both Szechwan Prov. (Graham, NM): Between Yachow and Tatsienlu, 3,000-8,000 feet, July 10-13, 1920; near Tat- sienlu, 5,000-8,500 feet, August 7, 1923. 44. AFISSA HAUSERI (Mader) FIGURES II2, 179 Solanophila Hauseri MAvER, 1930, p. 182. Abdomen.—Abdominal lines subterminal, incomplete, the outer mar- gin parallel to the margin of the segment, at about one-third from the base a sharp depression, which may give the impression that the outer margin curves inward. Male: Hind margin of fourth segment broadly V-shaped; fifth segment deeply emarginate with a strong depression in the middle, sixth segment also deeply emarginate with a process ending in a fork at the middle of apex. Female: Fourth segment in middle with only a very slight broad notch, fifth segment hind margin mildly emarginate, with a semicircular depression less marked than in the male, sixth segment also with hind edge emar- ginate but without trace of a depression. Male genitalia—Penis seen from below, first half open tube with 162 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 approximately parallel side margins, after that widening and flattened out and ending in two widely divergent points; upper side of penis flattened, the flat part widest at apex and narrowing to a sharp point at base; no hairs. Paramera 1.2 mm. long, spoon-shaped, greatest width about 0.4 mm., the apical half dark brown, almost black in some specimens, with long light blond pubescence (about 0.5 mm. long). Sipho short. Female genitalia.—Plates pointed at base, 0.85 mm. long, greatest width 0.35 mm. a little beyond the middle. Tergite X with apical edge gently convex with a trace of an emargination in the middle. Length.—7.0 to 8.2 mm. Color and maculation.—Head light brown, pronotum dark except the front margin and the front angles, to different extent in different specimens. Elytra black with red spots, the first longitudinal half- way between scutellum and callus touching the base; then follows an irregular fascia narrowly interruped at the suture and not reach- ing the sides; in the darkest specimen there is indication that it has resulted from the confluence of two spots: At about three-fifths a similar fascia which in the darkest specimen actually is resolved into two spots on each elytron. Finally there is a red spot at the apex. Pubescence distinct and long, light gray on the black, reddish on the red parts. Under side light brown, darker on metasternum and most of abdomen. Legs light, dark brown in one specimen. Type locality—China, Yunnan Prov., mountains near Mengtze; Kiautschau. Material examined.—5 specimens: China, Szechwan Prov., between Yachow and Mupin, alt. 5,000-10,000 feet, September 1930; near Mupin, alt. 6,000-7,000 feet, July 17, 1929; Mupin, alt. 3,500-5,000 feet, July 1, 1929; Mount Omei, alt. 4,000-6,000 feet, August 19, 1934 (D. C. Graham, NM) ; Shantung Prov., Kiautschau. Remarks.—Mader’s description of hauseri unfortunately is based entirely on color and omits the structure of the abdominal segments, which would have permitted a much more positive identification. None of the five specimens fits Mader’s description exactly, as it was based on darker specimens that showed five light spots on a dark back- ground. However, the coloration of the pubescence, different from that of all other species, leaves little doubt about the identity with hauseri (Mader). NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 163 45. AFISSA DOBZHANSKYI, new species Abdomen.—Abdominal lines terminal, incomplete, the outer edge parallel to the margin of the first abdominal segment. Fifth seg- ment, female, hind margin convex. Sixth segment, truncate. Female genitalia—Very similar to those of A. hauseri; tergite X with apical margin truncate or slightly convex; plates elongate with pointed base, 0.80 mm. long, greatest width 0.27 mm.; inner edge without pigment and straight. The chief difference with the genitalia of hauseri seems to be the slightly slenderer shape and the smaller amount of pigment of the plates. Length—6.0 mm. Epipleurae with a distinct cavity for the re- ception of the tips of the hind femora. Color and maculation—Upper side brownish red; pronotum with the disk dark so that only a relatively narrow light margin remains on all sides. Elytra with a large common scutellar spot, each with an elongate humeral spot. In addition with two dark fasciae, the discal one due to the confluence of spots 3 and 4, somewhat bent back at the suture and narrowly connected with the scutellar spot. The sub- apical fascia due to the widening of the usual subapical spot. Pu- bescence light gray on the dark spots, rubbed off almost everywhere else on the unique specimen. Under side light to dark brown, meta- sternum black. Type—vU.S.N.M. No. 57157, China, Kiautschau, a specimen ob- tained from Prof. Th. Dobzhansky. Remarks.—This species is very close to Afissa hauseri. It differs by the structure of the abdomen and the presence of distinct epi- pleural cavities, which are only slight depressions in hauseri. The elytral pattern also shows differences, and the size is smaller. The evaluation of both of these differences is not safe from a single specimen. «V. SZECHUANA GROUP The species of this and the chapini group are characterized by the fact that tergite X of the female has its apical part folded down and over so that the true apical margin points frontward. It is sepa- rated from the chapini group by the fact that the epipleurae do not have any cavities for the reception of the femora. It consists of two species from China of moderate size (length 6 to 7 mm.), both of them new. The elytra have each five spots, as in all the groups III to VI. The male genitalia are of relatively sim- ple structure and resemble those of the chapini group. 164 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 46. AFISSA SZECHUANA, new species FIGURES 113, 177, 215 Abdomen.—Abdominal lines subterminal, incomplete. The outer part ending near the middle of the width of the first segment, par- allel to the margin, and apex flat or very broadly rounded. Fifth segment, subtruncate in both sexes, with a narrow transverse depres- sion in middle near hind margin in male. Sixth segment, male, with a deep emargination ; female, subtruncate with a very shallow emar- gination. Male genitalia——Penis seen from below, tube about 0.23 mm. wide with nearly parallel sides until it narrows to apical point. A raised seam along practically the whole length, which terminates only when the penis narrows to point. Near the apex the walls of the seam diverge slightly to form the small elongate orifice. Seen in profile, gently curved down to the point, which is slightly bent up. Seen from above, nearly flat. Paramera gently curved down, 1.7 mm. long, slender, gradually narrowing toward apex, width in middle slightly less than 0.2 mm., apex covered with light blond hairs. Sipho almost bent into a semicircle. Orifice subapical on the inside. Female genitalia.—The feature that distinguishes this and related species from the others of the same genus is the structure of tergite X. The apex of this segment is heavily sclerotized and has a deep V-formed emargination. Along this line it is folded downward and over. The true apex of the segment is therefore pointing frontward and is convex toward the front. Plates broadly oval, length 0.65 mm., greatest width 0.47 mm. Length.—6.5 to 7.0 mm. Color and maculation.—Upper side reddish, head with a half cir- cular spot between the eyes, pronotum with a central and a smaller lateral spot. Elytra with five spots, No. 3 constricted in the middle as if it were due to the confluence of two spots. Under side black, side pieces of prothorax and mesothorax light, mouth parts and legs light, including coxae ; femora, however, dark, except tips. Epipleurae dark except a narrow seam on the outer margin. Material examined—Type: U.S.N.M. No. 57158, from China, Szechwan Prov., near Mupin, alt. 3,500-5,000 feet, July 1-2, 1929; (D. C. Graham). 7 paratypes, from the same locality (D. C. Graham, NM): alt. 5,000-6,500 feet, July 22, 1929; alt. 5,000-6,000 feet, July 19, 1924, July 22, 1929. NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 165 47. AFISSA SUBACUTA, new species Ficures 105A, 216 Abdomen.—Abdominal lines subterminal, incomplete; the outer part ending near the middle of the width of the first segment, paral- lel to the margin; apex flat or very broadly rounded. Fifth segment with hind margin convex in both sexes. Sixth segment hardly pro- truding from under the fifth, little sclerotized, with a deep notch in the middle in the male, which practically bisects it entirely, with entire edge in the female. Male genitalia—Penis seen from below as gradually widening tube (0.30 mm. wide near apex) until it suddenly narrows into a fine slender point; tube with a longitudinal raised seam, orifice rounded ; seen from above, with a pronounced ridge on basal half, in profile 0.4 mm. thick near base. Paramera gently curved down, length 1.5 mm., slender, width about 0.2 mm. at apex, with blond hairs. Sipho curved nearly into semicircle, tip compressed, orifice elongate on outside before compression. Very similar to those of szechuana but slightly wider, the orifice more rounded and the apical point slenderer. Female genitaia.—Similar to sgechuana and chapini in the fact that apex of tergite X is strongly sclerotized and folded down and back. The folding line, the apparent apex of the segment, however, is truncate and thin. Plates, length 0.52 mm., greatest width 0.44 mm. Length.—6 to 6.5 mm. Color and maculation—Upper side reddish, head spotless, pro- notum with a big central spot touching the base but not quite the front margin, and two smaller subcircular lateral spots on each side. Elytra with five large spots, No. 3 apparently arising from two con- fluent spots. In two of the three specimens this spot is joined to No. 4, which touches the margin, so that a continuous median fascia results which is narrowly interrupted at the suture. Spot 5 large, diamond-shaped, reaching almost from suture to side margin. Under side mostly black, side pieces of prosternum and mesosternum and part of abdomen light, mouth parts and legs light, except middle parts of femora, which are slightly darker, basal half of epipleura black in middle. Material examined—Type: U.S.N.M. No. 57159, from China, Szechwan Prov., Shin Kai Si, Mount Omei, alt. 4,500 feet, August 4-6, 1929 (D. C. Graham). 2 paratypes: Near Yachow, July 1929 ; between Kiating and Mount Omei, alt. 1,300-4,500 feet, August 21-22, 1925 (Graham, NM). 166 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Remarks.—tThis species is very similar in appearance to A. szech- uana and is closely related to it, although it can be easily separated from it by both differences in the maculation and the structure of the genitalia, especially the female genitalia. It agrees in most details with the description of acuta Weise, syn. acuminata Weise (1889, p. 648). But there is considerable deviation in the exact form of the elytral spots. Besides, acuta is reported to have a deep median notch in the fifth segment which reaches almost to its base. 4. subacuta has such a notch in the sixth segment of the male. My experience with the species in this group makes it appear very doubtful whether Kor- schefsky (1933b, p. 300) was justified in attributing to acuta Weise two specimens from Formosa with different maculation of the same type. VI. CHAPINI GROUP This group consists of the three species chapini, magna, and mili- taris, whose close relationship is indicated by the similarity of their male genitalia (where known) and the fact that they have distinct cavities on the elytral epipleurae for the reception of the tips of the middle and hind femora. By this feature they are separated from the szechuana group, to which they are otherwise closely related. | The female genitalia are very similar to those of the szechuana group with the bent-over apical part of tergite X. Among the Palaearctic and Indomalayan Epilachninae, distinct ely- tral cavities are otherwise found only in Cynegetis, and the presence of these cavities is usually considered a generic character. However, the chapini group is so close to the sgechuana group that it cer- tainly must be considered as belonging to Afissa. Traces of elytral grooves occur in other species. The only differences between Cyn- egetis and Afissa are: Cynegetis has the tarsal claws single with a tooth, whereas those of Afissa are double with both parts nearly the same size, and Cynegetis has the front tibiae grooved. 48. AFISSA CHAPINI, new species FIcuRES I15, 178, 217 Abdomen.—Abdominal lines terminal but incomplete, the outer margin not reaching beyond one-third from the hind margin of the first segment and coming close to the side margin. Fifth segment, male and female, hind margin evenly convex. Sixth segment, not pro- truding, with deep emargination in male, with a slight emargination in female. : NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 167 Male genitalia—Penis seen in profile, first gently bent down, shortly before apex suddenly bent up, becoming gradually thinner from base to apex; no hairs. Seen from below, as in figure 178. The middle opening closed on upper side with a clear membrane. Para- mera slender, curved up, length 1.6 mm., greatest width 0.2 mm., on apical third with long (about 0.4 mm.) light blond hairs. Sipho short, curved nearly in semicircle. Female gemtalia.—Plates, length 0.49 mm., greatest width 0.55 mm., apical edge almost straight. Tergite X large, apex folded double, downward for about 0.3 mm.; both the apparent apex (the folding line) and the real one with a weak emargination. Mandibles with one lateral tooth, no dentules.. Epipleurae with deep cavitites for the reception of the tips of the hind and middle femora. Length.—7 to 7.5 mm. Shape strongly hemispherical. Color and maculation—Upper side light red, pronotum spotless, elytra with five black rounded spots, No. 4 touching margin, 5 about halfway between suture and margin at about one-eighth distance from apex. Pubescence light blond, black on spots. Under side light brown, parts of metasternum and abdomen somewhat darker. Material examined.—Type: U.S.N.M. No. 57160, from Philippine Islands, Luzdn, Mount Maquiling (Baker). 3 paratypes (Baker, NM, D) ; Luzon, Mount Maquiling (2 speci- mens) ; Mindanao, Surigao (1 specimen). Remarks.—This species, though in maculation resembling the flavicollis group, stands out from all other species except magna by the presence of the epipleural cavities. It is also remarkable by the structure of the abdomen and its genitalia, particularly the female ones. 49. AFISSA MAGNA, new species Ficures 114, 180 Abdomen.—Abdominal lines incomplete, subterminal; the outer part reaching to about the middle of the segment and coming to within less than 0.15 mm. of its side margin. Punctation in the middle of the segment with circular points of about 0.07 mm. diameter near base. Fourth segment, male, subtruncate. Fifth segment with convex hind margin. Sixth segment with a deep notch, not protruding from under the fifth. Male genitalia.—Penis a flattened tube, split open on the under side, length 2.4 mm., gently curved down and suddenly bent up at apex 168 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 and ending in a single point. Upper wall a clear membrane. Para- mera 2.1 mm. long, gently curved down, slender, greatest width near apex 0.2 mm.; apical third with long (0.4 mm.) blond hairs. Sipho bent slightly, suddenly becoming narrower before apex and flaring out. Length—8.8 mm. Epipleurae with cavities for the reception of the tips of the middle and hind femora, somewhat shallower than those of chapimi. Metasternum very coarsely punctated. Color and maculation—Upper side brownish red. Pronotum with four rather indistinct dark spots. Elytra each with five large dark spots. Pubescence light gray, dark on the spots. Under side and ap- pendages light brown, sides of metasternum, epipleurae of prothorax, and tips of mandibles darker. Type —U.S.N.M. No. 57161, China, Fukien Prov., near Foochow, 1921-1924 (C. R. Kellogg). Remarks.—This species is very similar to chapini in all morpho- logical details. However, the size, maculation, and structure of the genitalia easily distinguish the two species. A. magna also shows some similarities to szechuana but is easily separated by the presence of the epipleural cavities. 50. AFISSA MILITARIS, new species Abdomen.—Abdominal lines incomplete, terminal, the outer edge parallel to the margin and close to it. Fifth segment, female, hind margin sharply convex. Sixth segment, convex, not protruding from under fifth. Female genitalia——Very similar to those of A. chapini (see fig. 217). The shape of the plates almost exactly the same, length 0.50 mm., greatest width 0.51 mm. Tergite X broad, the apex folded double as in chapint. The apparent apex has a slight emargination. on each side of the middle. The part beyond the bend longer than in chapim and becoming more narrow. Length from bend to real apex, 0.46 mm., width at apex 0.16 mm., apex with a distinct emargination, Length—8.o mm. Epipleurae with distinct cavities for the recep- tion of the tips of the middle and hind femora. These are not quite so deep as those of chapini. Color and maculation—Upper side deep red; pronotum with an almost circular dark spot, which reaches from the base to about one-fifth of the apical margin. Elytra with two basal dark spots, one medial fascia and a subapical spot. The first spot elongate on the suture almost reaching the base; scutellum light. The second spot on the callus leaves a very narrow margin light on the side and NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 169 base. The fascia is not unlike that of figure 105A but is extended to the suture. The subapical spot is rhomboidal, touching the margin and practically reaching the suture but leaving the apex light. Pu- bescence light reddish on the red parts of the elytra, light gray on the black parts. Under side light to dark brown. Type—vU.S.N.M. No. 57162, India, Assam, Dibrugarh, August 1943 (W. L. Jellison). EPIVERTA, new genus Genotype: Solanophila chelonia MAvER. Solanophila chelonia of Mader is so different from Afissa that it is properly regarded as the type of a new genus, of which it is the only known species. The chief differences from Afissa are the following: The elytra are very broadly margined along their whole outer margin. The epipleurae are wide and vertical, whereas they are much narrower and subhorizontal in Afissa. The structure of the spot pattern is also quite different from any found in Afissa, and the mandibles show peculiarities not found anywhere else. The genitalia of the female also are different from those of any oriental Afissa. Those of the male are as yet unknown. Epiverta shows no traces of abdominal lines on the first segment. EPIVERTA CHELONIA (Mader) Ficures 116, 218 Solanophila chelonia MADER, 1933, D. 79. Abdomen.—With no trace of abdominal lines. [Fifth segment, female, broadly emarginate ; sixth segment, convex. Male unknown. Female genitalia—Tergite X with broadly convex hind margin, plates with slightly convex inner margins, overlapping somewhat, length 0.63 mm., greatest half-width 0.33 mm. Greatest width just back of middle. Length—8.o0 mm. Elytra and pronotum with broadly offset and flattened margins. Elytral epipleurae from base to just beyond the end of metasternum with a narrowing horizontal part which is bent outward vertically down. The vertical part considerably wider than the horizontal part, where the latter exists and continuing slightly diminished in width and not quite so vertical to the apex. Mandibles.——Apical tooth trifid, with the two outer components rudimental. Three lateral teeth present, the second about the size of the apical tooth, the third small, both slender and attached to the 170 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 upper side of the rim. The first lateral tooth is considerably larger than any of the others, attached to the lower part of the rim of the mandible between the apical and second lateral and pointing downward. It carries on its lower edge a small side tooth, and beyond that to the apex several small dentules. Color and maculation—Under side black, with some lighter parts on the lighter specimen. Maculation of pronotum and elytra, see figure 116. The other specimen has the light portion much more extended so that the ground color appears to be light reddish brown with irregular black markings. Almost all the light spots of figure 116 connected except that a zigzag-shaped black band separates com- pletely the region of the apical light spots from the others. The dark spots appearing in the centers of the light ones are less dark. Covered above and below with long light-gray pubescence, which is sparser in the dark spots of the elytra. The dark spots on the center of the light patches have, however, the pubescence in full strength, and also their ground color seems to be somewhat lighter than that of the connected black parts. Mouth parts, antennae, and legs reddish. Type locality—Szechwan and Tibet. Material examined.—2 female specimens: China, Szechwan Prov., southwest of Tatsienlu, June 28-July 2, 1923 (D. C. Graham, NM). Remarks.—The form of the epipleurae, together with the absence of abdominal lines, differentiates this species from all other known oriental Epilachninae, and this species is therefore made the type of the new genus Epiverta. Mader, in the original description of Solanophila chelonia, does not mention either of these two characters. There seems, however, little doubt that the species described by him is the same as the two speci- mens before me. He describes the maculation of the elytra as of dark background with 14 light spots arranged 2, 4, 4, I, 2, 1, which may be interpreted as figure 116 with the light color less developed. The following two well-known European species of the Epilach- ninae are included in order to present the structure of abdomen and genitalia of these species in the same manner as for the rest of the subfamily. SUBCOCCINELLA VIGINTIQUATTUORPUNCTATA (Linnaeus) Ficures 1 D, 170 Coccinella 24-punctata LINNAEUS, 1758, p. 366. Abdomen.—Abdominal lines incomplete, reaching to within one- fourth to one-third of the hind margin of the first segment and NO. 15 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE i7I outer part reaching to about one-third of the front margin. Broadly curved: Fifth segment, male, subtruncate or slightly convex; female, with a process in the middle. Sixth segment, male, subtruncate, wide; female, hind margin convex and narrower. Male genitalia—Penis 0.6 mm. long, 0.25 mm. in maximum width, approximately in the form of a wooden shoe, basal part open, upper side a clear membrane; apical part closed, ending in a blunt point. No hairs on the penis. Paramera slender, 0.55 mm. long, clothed with fine pubescence practically on their whole length, at least on the outside part. Sipho rather thick (0.1 mm.), bent about 90° near base and apex. Orifice terminal, on the inner side with a long (0.2 mm.) sclerotized needlelike process; on the outer side with mem- branous flaps. Female genitalia—Tergite X wide, hind margin subtruncate or mildly convex with a very slight emargination in the middle. Only a narrow strip along each side of the apical margin with pigment. Plates 0.3 mm. long, widest (0.23 mm.) near apex. The genitalia are in shape most like figure 218, but the greatest width of the plates is nearer the apex, which is more truncate, while the basal part is rounded. Mandibles (fig. 1 D).—With an apical tooth and two big lateral teeth, the second somewhat smaller than the first. Dentules on all three teeth and the inner margin. CYNEGETIS IMPUNCTATA (Linnaeus) Ficures 1 H, 171 Coccinella impunctata LINNAEUS, 1767, p. 570. ? Abdomen.—Abdominal lines subterminal, subcomplete, outer mar- gin parallel to the side margin of the first segment and close to it. Fifth segment, male, hind margin convex; female, truncate. Sixth segment, male, hind margin emarginate; female, convex. Male genitalia—Penis a very flattened tube almost straight in profile, 0.55 mm. long; seen from below widened from base to apex, widest part 0.3 mm., then suddenly narrowed to rounded tip. Para- mera 0.7 mm. long, slender, gently curved down, 0.1 mm. wide at about four-fifths of their length, covered with hairs on apical part. Sipho curved, orifice with a liplike process bordered by very short bristles, which are clearly visible only at fairly high magnifications (50x or higher). Female genitalia.—These look in general structure much more like those of some Epilachna species than Afissa species. Plates 0.45 mm. 172 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 long, greatest width 0.22 mm. about one-third way from the apex. The general shape of the plates is not unlike that of figure 185, with the apical part more rounded and without the emargination on the inner edge. Tergite X with convex hind margin, the middle a clear, membranous, wide strip, on each side pigmented strips, which do not . quite touch in the middle of the apical margin. Mandibles (fig. 1 H).—With an apical and two lateral teeth. The apical tooth, as in most other Epilachninae, has the middle part much more strongly developed than the side parts. The lateral teeth are relatively short, the second smaller than the first. There are very small dentules on the teeth (clearly visible only with about 50x magnification). The structure of the mandibles conflicts with the description by Ganglbauer (1899, p. 951) who claims that there is only one trifid tooth. He must have examined a specimen with the two lateral teeth broken off. As the material from China and the Philippines that I had a chance to examine was much more extended than anything published pre- viously, I add a list of all the known Epilachninae of these two regions. A few species unknown to me recorded by others have been added, provided there was no serious doubt about the identity of the species. Type localities are printed in italics. CHINA Epilachna sparsa orientalis, new..........+. Fukien, Kiang-su, Shensi, Hong Kong, Szechwan, Anhwei. Var. cinerc@, MEW. oe res cada: ae aes Fukien, Kiang-su, Szechwan. dentiulaias DEW ricco, hace rasa been Fukien, Kiang-su. (Cochin China.) mponscaWewis” 6's.) sce: tae eee othe Chih-li, Kiang-su, Jehol, Manchu- ria, Hopei, Korea. (Japan.) coalescens Maden? . 5 sclswinisrs 521th Szechwan. Libera: Ne6Wirnenain sae cen cede zechwan. ASiGCHLG MAINCHICE, DOW «o's: as spe winys ngs wtp» shales Fukien, Shantung, Tibet (J/ndo- china), Anhwei, Kwangsi. bisquadripunctata Gyllenhal ........... Kwangsi, Hong Kong *. Afissa admirabilis (Crotch).............+. Kiang-si, Kiang-su, Chekiang *, : Anhwei, Hupei. Continentalas,. NEW: :.,.- bercainie sc + sere Szechwan. macularis (Mulsant) ........---.sses Szechwan. (Nepal.) donckiert WeISC .%......-2.-+.0+cee Yunnan, Tibet. aliernans: (CMulsant) oe eines ioe = Szechwan. (Java.) insignis’. (Gorham) od. ce eae ns oe se ee China, Szechwan, Kinkiang *, Anhwei. * Localities originally given by Liu (1935). NO. I5 LADYBEETLES OF THE GENUS EPILACHNA—DIEKE 173 quindecemguttata, NEW ........-+2066%- Szechwan. AIFADILOLACS AMIEW) aie c oicikaltclemialteres Clase sechwan. AUGATACOINS, DEW oes .. | CITY OF WASHINGTON Foe *6, oe0PPeeess (Pusrication 3862) PUBLISHED BY THE SMITHSONIAN INSTITUTION DECEMBER 30, 1946 _ SMITHSONIAN MISCELLANEOUS COLLECTIONS SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 16 NEW BIRDS FROM COLOMBIA BY ALEXANDER WETMORE Secretary, Smithsonian Institution COlAdee ll) “28000000? (PuBLicaTIon 3862) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION DECEMBER 30, 1946 The Lord Battimore Press BALTIMORE, MD., U. & A. , oY ¥ i ‘ én # aye , rn i . 4 F ~ ? P 4 4 % Ah 4 ~ \ . ha | ct i . « ~ . i ‘ ‘ x i , hg ~ d ‘ > tes ‘ i ‘ te Oo v7 NEW BIRDS FROM COLOMBIA By ALEXANDER WETMORE Secretary, Smithsonian Institution The species and subspecies of birds described in the present paper are found in collections made for the Smithsonian Institution from {941 to 1946 in the Departamento de Magdalena and the Comisaria de Guajira in northeastern Colombia. They are based primarily on material in the United States National Museum, with additional comparisons in the American Museum of Natural History and the Chicago Natural History Museum. These previously unknown forms have been determined in connection with the preparation of a report on the complete collection, publication of which will be delayed as the work in the field still continues. Family MomoriDAE MOMOTUS MOMOTA SPATHA, new subspecies Characters——Similar to Momotus momota subrufescens Sclater,* but paler above and below; under wing-coverts slightly paler; loral feathers immediately behind nostril tawny olive, forming a small spot. Description.—Type, U.S.N.M. No. 368,925, male adult, collected at Nazaret, Guajira, Colombia, May 2, 1941, by A. Wetmore and M. A. Carriker, Jr. (orig. No. 11,752). Sides of head, a very narrow line on forehead, and center of crown black, with the auricular feathers nar- rowed and elongated ; loral feathers immediately behind nostrils tawny olive, forming a partly concealed spot; forepart of crown Nile blue, extending back on either side to level of posterior margin of the eye; black feathers on lower margin of ramal area near center edged with Nile blue, producing a narrow line; upper margin of elongated auric- ular feathers edged with Nile blue, producing another narrow line; posterior margin of black pileum, from posterior margin of eye on either side, phenyl blue, forming a broad, well-marked band; sides of neck and hindneck old gold basally, with the tips of the feathers ser- pentine green, giving a greenish cast, most prominent on the hind- neck; back and scapulars serpentine green, with a slight wash of old gold on the central portions of the distal barbs, giving a slight golden 1 Momotus subrufescens Sclater, Rev. Mag. Zool., ser. 2, vol. 5, November 1853, p. 489 (Santa Marta, Depto. Magdalena, Colombia). SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 16 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 wash; rump and upper tail-coverts deep dull yellow green, with the ends of the feathers washed with old gold; wing-coverts, innermost primaries, and secondaries elm green; alula and primary coverts Jouvence blue; outer webs of outer primaries motmot blue, inner webs zinc green, bordered internally with fuscous; basal half of rec- trices dark yellowish green; on central pair shading at center over to dark Chessylite blue, with an overwash of yellowish green when viewed from certain angles, this color extending over the spat- ulate tips except for the distal ends, which are dull black; two ad- jacent pairs of rectrices dark yellowish green at base, and dark Chessylite blue for distal half, the longer pair tipped with dull black; under surface of body lighter than buckthorn brown, with the throat feathers washed with light Paris green; an elongate black spot in center of breast, edged with variscite green; under wing-coverts and inner webs of inner primaries and secondaries pinkish buff; edge of wing isabella color; ends of primaries on under surface dark mouse gray; under surface of tail sooty black, with all except the central rectrices tipped narrowly with mouse gray. Bill black, with a brownish tinge on mandible, especially at the base, tarsus and toes fuscous (from dried skin). Measurements.—Males, 7 specimens, wing 117.1-127.8 (124.5), tail 213-228 (222.3), culmen from base 35.9-39.8 (37.5), tarsus 26.0- 27.8 (26.5) mm. Females, 7 specimens, wing 120.0-129.1 (124.8), tail 213-235 (221.6, average of 6 individuals), culmen from base 33.5-37.5 (36.0Y, tarsus 24.3-27.8 (26.1) mm. Type, male, wing 126.4, tail 218, culmen from base 37.5, tarsus 26.0 mm. Range.—tThe lower levels of the Serrania de Macuire, Guajira, Colombia. Remarks—Motmots were an unexpected find in the lower levels of the Serrania de Macuire, our nearest other record for birds of this group being at Carraipia many miles to the west. The arid desert area of the Guajira Peninsula as a whole is unsuited to these forest inhabitants, which in the Macuire are encountered near the water- bearing arroyos in the foothills of the mountains. The differences of paler coloration on which this subspecies is described are not great, ‘but are sufficient to warrant separation, being very evident when series of the new form are laid out beside equal numbers of subrufescens. The buffy marking on the feathers immedi- ately behind the nostrils is especially interesting, since it is lacking or, rarely, very faintly indicated, in the related form. No. 16 NEW BIRDS FROM COLOMBIA—WETMORE 3 Family TROCHILIDAE METALLURA IRACUNDA, new species Characters—Generally similar to Metallura tyrianthina (Lod- diges)* but decidedly larger; male very much darker throughout; wing-coverts copper color instead of green; female slightly darker above, with the wing-coverts likewise copper color instead of green. Description—Type, U.S.N.M. No. 372,813, male adult, between 9,500 and 10,000 feet elevation above Airoca, Sierra de Perija, Depto. Magdalena, Colombia, collected May 4, 1942, by M. A. Carriker, Jr. (orig. No. 2609). Crown shining cedar green ; sides of neck, hindneck and back sooty black, becoming metallic dark green with occasional coppery reflections near the centers of some of the feathers when viewed at an angle; coppery reflections increasing on rump to cover the upper tail-coverts ; wing-coverts dull copper; primaries and sec- ondaries benzo brown with very faint greenish reflections on outer webs; rectrices very broad, brick red, with a coppery sheen; gorget shining cendre green; under surface chaetura black with faint bronzy reflections that become coppery on the sides; under wing-coverts greenish black; edge of wing ochraceous tawny; extensive fluffy tibial tufts white; under tail-coverts Hessian brown, broadly edged with drab. Bill, tarsus, and toes dull black (from dried skin). Measurements——Males, 14 specimens, wing 60.1-65.9 (63.2), tail 42.0-47.2 (44.5), culmen from base 12.7-14.0 (13.5) mm. Females, 5 specimens, wing 54.8-57.0 (56.0), tail 36.0-38.3 (37.8), culmen from base 13.0-14.0 (13.5) mm. Type, male, wing 65.9, tail 47.2, culmen from base 12.7 mm. Range.—Known from the high paramo region on Cerro Pintado (east of Valledupar), and Las Tres Tetas above Airoca (east of Casacara and southeast of Codazzi), in the Sierra de Perija, Depto. Magdalena, Colombia, crossing into the adjacent sections of Venezuela. | Remarks.—This fine species suggests Metallura tyrianthina but differs definitely from all the component subspecies of that group of forms. It ranges in the Sierra de Perija with Metallura tyrianthina districta Bangs (recorded here for the first time outside the Sierra Nevada de Santa Marta), which, aside from the evident differences in color and size, demonstrates that tyrianthina and iracunda are spe- cifically distinct. In the color of the tail the new bird is more like 2Trochilus tyrianthinus Loddiges, Proc. Comm. Zool. Soc. London, pt. 2, March 29, 1832, p. 6 (Popayan, Colombia). 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 typical tyrianthina than it is like districta, but is even more coppery. The female of iracunda differs from the related forms clearly in de- cidedly larger size, while the coppery color of the wing-coverts, the copper tail, and the average duller, less bright green dorsal surface are also diagnostic. Carriker found Metallura iracunda abundant in the higher open country which is found across the mountain tops along the frontier be- tween Venezuela and Colombia. Though our specimens were collected in Colombia, as the approach to the high country was made from the western side of the range, there is no question but that the species ranges across the border into adjacent Venezuela. Family FoORMICARIIDAE GRALLARIA RUFULA SALTUENSIS, new subspecies Characters Similar to Grallaria rufula spatiator Bangs * but olive, rather than rufescent above; duller grayish brown below; lighter on throat and abdomen; flanks and under tail-coverts lighter. Description—Type, U.S.N.M. No. 373,673, male adult, taken be- tween 9,500 and 10,000 feet elevation, south of the south Teta above Airoca, Sierra de Perija, Depto. Magdalena, Colombia, May 4, 1942, by M. A. Carriker, Jr. (orig. No. 2611). Entire dorsal surface be- tween buffy brown and olive brown; feathers of forehead whitish at base; innermost primaries and secondaries with outer webs snuff brown; outer web of outer primaries avellaneous; tail snuff brown; base of upper loral feathers deep olive buff making an indistinct spot ; narrow indefinite eye circlet whitish ; auricular region like crown, with a wash of wood brown in lower margin; under surface in general light wood brown, paling to whitish on throat, foreneck, and lower breast ; abdomen and under tail-coverts dull white; sides, flanks, and tibiae buffy brown; under wing-coverts avellaneous, becoming olive brown externally; edge of wing whitish. Maxilla clove brown; mandible deep olive buff (tip broken) ; tarsus and toes olive brown (from dried skin). Measurements.—Males, 2 specimens, wing 78.6-80.2 (79.4), tail 36.9-39.9 (38.4), culmen from base 17.8-18.8 (18.3), tarsus 43.3-44.5 (43.9) mm. 1 Females, 2 specimens, 79.3-80.4 (79.8), tail 42.0-42.5 (42.2), cul- men from base 19.1-19.4 (19.2), tarsus 44.8-45.8 (45.3) mm. 3 Grallaria spatiator Bangs, Proc. Biol. Soc. Washington, vol. 12, June 3, 1808, p. 177 (Macotama, Sierra Nevada de Santa Marta, Colombia). No. 16 NEW BIRDS FROM COLOMBIA—-WETMORE 5 Type, male, wing 80.2, tail 36.9, culmen from base 17.8, tarsus 43.3 mm. Range-——Known only from 9,500 to 10,600 feet elevation in the Sierra de Perija (above Airoca and on Cerro Pintado), Depto. Magdalena, Colombia. Remarks.—The form here described is placed as a subspecies of Grallaria rufula with some reservation, since it differs from the nearest races, typical rufula of farther south, and spatiator of the Sierra Nevada de Santa Marta, in being olive in tone, with none of the brighter rufescént tints found in the dullest colored of the popula- tions that have been combined under the specific name in question. This is the more striking since from its geographic location saltuensis would be expected to resemble one or the other of the two races men- tioned rather closely. In addition the new bird appears slighter in bulk, and the tarsi average somewhat more slender. It seems pos- sible that it may be a distinct species, and that a true rufula may be found sometime ranging in the same area. Family TyRANNIDAE CNEMARCHUS ERYTHROPYGIUS ORINOMUS, new subspecies Characters-—Similar to Cnemarchus erythropygius erythropygius (Sclater),* but brown of posterior lower surface and rump paler ; brown area of dorsal surface confined to rump, and not extending so high on lower back; slightly grayer above ; hindneck and back of head slightly darker; dark tips on lateral rectrices shorter; under wing- coverts paler. Description —Type, U.S.N.M. No. 387,513, male adult taken be- tween II,400 and 12,000 feet elevation above Mamancanaca, February 23, 1946, by M. A. Carriker, Jr. (orig. No. 7902). Korepart of crown dull white, with shafts of feathers mouse gray bordered by light mouse gray, producing streaks that are confined to the shaft on the forehead but become broader posteriorly until the white is com- pletely replaced at the center of the crown, the white extending on sides to posterior margin of eye, forming a superciliary streak; pos- terior half of crown and upper hindneck light mouse gray ; posterior half of lores dark mouse gray, the anterior half white merging with the white forehead ; an indistinct streak of dull white beneath eye from rictus to auricular region; auricular area dark mouse gray; lower 4 Taenioptera erythropygia Sclater, Proc. Zool. Soc. London, 1851 (June 20, 1853), Pp. 193, pl. 41 (Ecuador). 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 hindneck and upper back deep mouse gray, becoming darker, almost chaetura drab, on central portion of back; rump and upper tail-coverts ochraceous tawny; wing-coverts chaetura drab; remiges chaetura black ; outer webs of innermost secondaries edged broadly with white near center, forming a prominent elongated spot with indistinct mark- ings of tawny olive on the anterior margin; rectrices ochraceous tawny, tipped with dull black, the central pair mainly black except for base of outer web; throat and foreneck dull white with shaft streak- ings of light mouse gray that become broader posteriorly; breast mouse gray, washed with smoke gray in center; side§ and abdomen ochraceous tawny; under tail-coverts slightly paler; under wing- coverts ochraceous tawny, becoming cinnamon buff on distal part of edge of wing under the alula; inner base of primaries and secondaries pinkish buff for basal half. Bill, tarsus, and feet dull black (from dried skin). Measurements.—Males, 8 specimens, wing 141.7-146.1 (143.7), tail 103.7-112.0 (107.1), culmen from base 18.7-20.2 (19.6), tarsus 28.2-31.6 mm. Females, 7 specimens, wing 141.0-145.4 (143.1), tail 106.5-109.4 (107.6), culmen from base 19.5-20.5 (20.2, average of 6 individuals), tarsus 29.2-30.6 (30.0) mm. Type, male, wing 144.0, tail 108.0, culmen from base 18.7, tarsus 29.8 mm. Range——tThe high paramos of the Sierra Nevada de Santa Marta, between 10,600 and 12,000 feet above sea level (recorded at present from Mamancanaca, including the Cucungaca Valley, and the head of the Rio Guatipuri). Remarks—Cnemarchus erythropygius has not been recorded pre- viously north of the Andes of Ecuador so that Carriker’s discovery of this new race in the Sierra Nevaca de Santa Marta is a notable addition to our knowledge of these birds, and to the avifauna of Colombia. Earlier observers in this area either failed to cover the higher paramos thoroughly, or confused Cuemarchus with Myto- theretes striaticollis striaticollis, common in that area, which is similar in size and general coloration. It is diffcult to understand how so conspicuous a bird could have been overlooked otherwise. In size this northern race is similar to the typical form of Peru and Ecuador. No. 16 NEW BIRDS FROM COLOMBIA—WETMORE 7 OCHTHOECA DIADEMA RUBELLULA, new subspecies Characters —Similar to Ochthoeca diadema diadema (Hartlaub)® but with crown lighter, more olive; back lighter, much more reddish brown; under parts paler yellow. Description—Type, U.S.N.M. No. 373,844, male, between 8,000 and 9,000 feet above Laguna de Junco, slopes of Cerro Pintado, Sierra de Perija, Depto. Magdalena, Colombia, taken July 11, 1942, by M. A. Carriker, Jr. (orig. No. 3245). Crown and hindneck brownish olive; forehead, anterior part of lores and superciliary Strontian yellow, the latter changing above eye to reed yellow for posterior half; back, scapulars, lesser wing-coverts, rump, and upper tail-coverts brighter than snuff brown ; middle and greater wing-coverts and upper surface of remiges, where exposed, dark mouse gray; middle and greater wing-coverts, inner secondaries and tertials edged broadly with sayal brown; rectrices slightly darker than deep mouse gray, with the outer webs edged with citrine drab, especially toward the base; posterior part of lores dusky neutral gray; line behind eye dark grayish olive, with the auricular area below this dark olive buff ; throat deep colonial buff ; upper breast, sides, and flanks light yellowish olive ; lower breast and abdomen between straw yellow and amber yellow; under tail- coverts olive buff; inner under wing-coverts and axillars olive buff, becoming deep colonial buff centrally, and citrine drab externally ; edge of wing dark olive buff; inner webs of innermost primaries and secondaries vinaceous buff on under surface. Under surface of mandible drab; rest of bill and feet fuscous black; tarsus drab (from dried skin). Measurements—Males, 3 specimens, wing 65.1-66.7 (65.9), tail 55-4-55-7 (54.8), culmen from base 11.2-12.1 (11.7), tarsus 19.8-20.1 (19.9) mm. Females, 3 specimens, wing 59.0-60.3 (59.6), tail 50.1-51.9 (50.8), culmen from base 10.8-11.3 (11.1), tarsus 18.5-19.5 (18.9) mm. Type, male, wing 65.1, tail 54.4, culmen from base 11.8, tarsus 19.8 mm. Range——Known only from the type locality, between 8,000 and 9,000 feet elevation above Laguna de Junco on the slopes of Cerro Pintado, near the northern end of the Sierra de Perija, Depto. Magdalena, Colombia. Remarks.—From Ochthoeca diadema jesupi Allen, this new form differs in being much browner on the back, in having brighter brown 5 Myiobius diadema Hartlaub, Rev. Zool., vol. 6, October 1843, p. 289 (Bogota, Colombia). 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 edgings on the wing feathers, the forehead more broadly yellow, and the under surface lighter yellow. In a measure rubellula is inter- mediate between typical diadema and jesupi but is easily distinguished ~ from either. OCTHOECA RUFI-PECTORALIS RUBICUNDULUS, new subspecies Characters.—Similar to Ochthoeca rufi-pectoralis rufopectus (Les- son)® but with back, rump, and scapulars distinctly brown; abdomen grayer ; breast slightly lighter, brighter brown. Description—Type, U.S.N.M. No. 373,864, male adult, taken above Airoca, between 9,500 and 10,000 feet elevation south of the south Teta, Sierra de Perija, Depto. Magdalena, Colombia, May 7, 1942, by M. A. Carriker, Jr. (orig. No. 2648). Crown fuscous black becoming slightly paler on hindneck; prominent superciliary streak dull white, reaching posteriorly to sides of upper hindneck, this ex- tending anteriorly, somewhat narrowed, to the base of the bill; back, scapulars, and rump between sepia and Saccardo’s umber; upper tail-coverts fuscous; lesser and middle wing-coverts fuscous black, faintly margined with bister; greater wing-coverts dull black, tipped broadly with Mikado brown, producing a broad wing band that be- comes narrower as it proceeds inward ; primaries and secondaries dull black, the secondaries and innermost primaries edged narrowly on outer webs with dull white, this edging not extending to the tip; rec- trices dull black, the outer web of the outer rectrix dull white; lores and margin immediately above eye blackish mouse gray ; sides of head dark mouse gray; sides of neck deep mouse gray; throat grayish white ; upper breast and foreneck sayal brown; lower breast and sides pale mouse gray; abdomen and under tail-coverts dull white; under wing-coverts dull white, mixed slightly with fuscous black on edge of wing. Bill dull black, becoming fuscous across base of mandible ; tarsus and toes fuscous black (from dried skin). Measurements.—Males, 3 specimens, wing 71.0-75.4 (72.9), tail 61.3-62.8 (62.0), culmen from base 12.0-13.0 (12.6), tarsus 20.0-21.1 (20.6) mm. Females, 3 specimens, wing 69.7-71.0 (70.4), tail 59.0-62.8 (61.4), culmen from base 12.0-13.0 (12.4), tarsus 19.5-20.2 (19.9) mm. Type, male, adult, wing 75.4, tail 62.0, culmen from base 13.0, tarsus 21.I mm. 6 Tyrannulus rufopectus Lesson, Echo du Monde Savant, vol. 11, No. 10, August 4, 1844, col. 233 (Bogota, Colombia). No. 16 NEW BIRDS FROM COLOMBIA—WETMORE 9 Range.—Known only from the higher elevations (9,500 to 10,600 feet) of the Sierra de Perija (above Airoca, and on Cerro Pintado). Remarks.—It is interesting that two species of Ochthoeca, namely rufipectoralis and diadema, found in the northern Perija range should vary from their representatives in the Andes farther south in exactly the same way, both being definitely brighter brown on the back. The present race is closely similar in size to Ochthoeca rufi-pectoralis rujopectus except that the tarsus is slightly shorter, this measurement in rufopectus being from 18.0 to 18.5 mm. in 3 males and 16.5 mm. in one female. Family TROGLODYTIDAE THRYOPHILUS LEUCOTIS COLLINUS, new subspecies Characters —Similar to Thryophilus-leucotis venezuelanus Lafres- naye,’ but darker above; dark markings on auricular region slightly heavier; flanks and under tail-coverts slightly darker; under wing- coverts duller white; averaging very slightly larger. Description—Type, U.S.N.M. No. 369,737, male adult, collected between 1,500 and 2,000 feet elevation near Nazaret, in the Serrania de Macuire, Guajira, Colombia, May 7, 1941, by A. Wetmore and M. A. Carriker, Jr. (orig. No. 11,842). Crown, hindneck, and upper back snuff brown, changing across lower back to cinnamon brown on rump and upper tail-coverts; tertials and outer webs of secondaries, and inner primaries, slightly darker than sayal brown; outer webs of outermost primaries cinnamon, all barred narrowly with fuscous; inner webs of primaries and secondaries fuscous; rectrices cinnamon brown, with narrow wavy cross bars of dusky neutral gray ; lores and superciliary white; auricular region grayish white, lined indefinitely with neutral gray; a narrow line of bister, edged with dusky neutral gray, behind eye, between the white superciliary and the auricular region ; sides of neck olive gray ; under surface generally white; sides light wood brown, becoming sayal brown on flanks and under tail- coverts ; a faint wash of light wood brown across breast ; under wing- coverts white, with a slight wash of cartridge buff; edge of wing white. Maxilla fuscous black, mandible smoke gray, becoming olive buff at base, tarsus and toes dusky neutral gray, claws fuscous (from dried skin). Measurements.—Males, 4 specimens, wing 64.3-66.5 (65.7), tail 38.6-46.3 (42.6), culmen from base 19.7-21.8 (20.5), tarsus 23.6-24.8 (24.0) mm. 7 Thryothorus venezuelanus Cabanis, Mus. Hein., pt. 1, 1850, p. 78 (Venezuela). IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Females, 3 specimens, wing 58.0-60.9 (59.2), tail 37.8-39.8 (38.9), culmen from base 17.8-19.0 (18.5), tarsus 21.5-24.2 (23.0) mm. Type, male, wing 64.3, tail 42.3, culmen from base 21.8, tarsus 24.8 mm. Range.—Above 1,500 feet elevation in the Serrania de Macuire, Guajira, Colombia. Remarks.—The present race is confined to the green island of the Serrania de Macuire rising in the eastern end of the arid Guajira Peninsula. Its differences in color and size are not great but are suf- ficient to set it off from the paler, slightly smaller Thryophilus leu- cotis venezuelanus that ranges generally through the lowlands of this area. This latter race shows the following measurements in our series from Maicao and Carraipia in the western Guajira, and El Conejo and Distraccion, near Fonseca, in extreme northeastern Magdalena. Males, 8 specimens, wing 60.9-64.3 (62.4), tail 41.0-45.9 (42.5), culmen from base 17.9-19.5 (18.6), tarsus 21.9-23.3 (22.5) mm. Females, 5 specimens, wing 56.8-60.0 (58.2), tail 36.0-38.8 (37.6), culmen from base 17.0-18.5 (17.7, average of 4 specimens), tarsus 21.2-22.0 (21.7) mm. Family TurDIDAE TURDUS LEUCOMELAS CAUTOR, new subspecies Characters —Similar to Turdus leucomelas albiventer Spix® but darker on dorsal surface, with the head grayer, less brownish ; smaller. Description.—Type, U.S.N.M. No. 369,663, male adult, collected at 1,000 feet elevation in the Serrania de Macuire, above Nazaret, Guajira, Colombia, May 5, 1941, by A. Wetmore and M. A. Carriker, Jr. (orig. No. 11,791). Crown and hindneck deep mouse gray ; upper back hair brown; lower back and wing-coverts light sepia; rump light brownish olive; greater wing-coverts edged and tipped with clay color; secondaries and proximal three-quarters of primaries, except the two outermost, sepia, edged on outer web with Saccardo’s umber ; tips of primaries fuscous, the outermost edged narrowly on this distal area with ivory yellow; central rectrices fuscous becoming olive brown on two outermost ; sides of head mouse gray, the auricular feathers with faint, very narrow shaft lines of pallid mouse gray; chin and throat white, the throat streaked prominently with deep mouse gray; breast and sides light mouse gray, with a faint wash of 8 Turdus albiventer Spix, Av. spec. nov. Brasiliam, vol. 1, 1824, p. 70, pl. 60, fig. 2 (Para, Brazil). NO. 16 NEW BIRDS FROM COLOMBIA—WETMORE iT light drab as edging, the breast color shading off to white on the abdomen and under tail-coverts ; tibia mouse gray ; under wing-coverts ochraceous tawny; edge of wing clay color; inner webs of remiges clay color. Bill fuscous, becoming drab at base of mandible; tarsus and toes hair brown (from dried skin). Measurements.—Males, 5 specimens, wing I11.4-114.6 (113.0), tail 86.9-97.1 (91.1), culmen from base 22.2-23.2 (22.7), tarsus 30.2-33.3 (31.7) mm. Females, 2 specimens, wing 108.4-1I1.0 (109.7), tail 86.0-90.1 (88.1), culmen from base 22.4-23.8 (23.1), tarsus 31.4-31.5 (31.5) mm. Type, male, wing 111.8, tail 87.4, culmen from base 22.4, tarsus 33-3 mm. ; Range.—tThe higher forests of the Serrania de Macuire, Guajira, Colombia. Remarks.—In its smaller size and brighter brown of the back this new form suggests typical Turdus leucomelas leucomelas, found from southern Brazil to eastern Pertti and south into Paraguay. In color, however, T. /. cautor is distinctly grayer, especially on the head. The difference in size from T. 1. albiventer, which ranges adjacent to the Guajira home of cautor, is very evident, the wing in 23 males of albiventer measuring from 117.7 to 125.1, and in 18 females from 119.5 to 126.3 mm. The clay-colored spots on the end of the greater wing-coverts in the specimen selected as type are a trace of the juvenile wing mark- ings, which in these thrushes frequently carry over into the succeeding adult plumage stage. This individual bird is fully adult as I deter- mined personally through examination of the ossification of the cranium. The subspecific name given to this new race indicates the shy, wary habit of this forest-dwelling species. Family THRAUPIDAE DUBUSIA CARRIKERI, new species Characters —Similar to Dubusia taeniata (Boissoneau),® but with light brown of breast extending over upper breast and the lower part of the foreneck; blue of forepart of crown and lateral streaks more extensive ; throat streaked with buff; blue of lesser and middle wing- coverts darker ; tibia more bluish; smaller. 9Tanagra (Tachyphonus) taeniata Boissoneau, Rev. Zool., vol. 3, March 1840, p. 67 (Bogota, Colombia). I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Description—Type, U.S.N.M. No. 388,179, male adult, between 8,500 and 9,500 feet on the south side of the main valley of the Rio Guatapuri, Sierra Nevada de Santa Marta, Depto. Magdalena, Colom- bia, taken March 28, 1946, by M. A. Carriker, Jr. (orig. No. 8160). Sides of head and neck, center of crown and hindneck, lores and anterior line of forehead, including area across nostrils, deep black; forepart of crown and a broad streak on either side extending back onto sides of hindneck light forget-me-not blue ; back, scapulars, rump, upper tail-coverts, outer webs of secondaries, and primaries deep slate blue ; lesser and middle wing-coverts light Windsor blue ; greater wing-coverts black basally, tipped with clear Windsor blue; primaries dark neutral gray, edged narrowly on external web with deep slate blue; rectrices dull black, with, the exposed webs washed with deep slate blue on dorsal surface ; throat and upper foreneck black streaked with cartridge buff, the light streaks becoming chamois posteriorly ; upper breast and lower foreneck between cinnamon buff and clay color; lower breast, abdomen, and sides lemon chrome; crissum and under tail-coverts cinnamon buff; tibia dark Payne’s gray. Bill, tarsus, and toes black (from dried skin). Measurements.—Males, 5 specimens, wing 82.1-88.2 (85.4), tail 78.8-82.6 (80.3), culmen from base 14.3-15.1 (14.7), tarsus 26.0-27.9 (26.9) mm. Females, 5 specimens, wing 81.7-86.7 (83.8), tail 77.7-80.7 (79%), culmen from base 14.2-15.4 (14.9, average of 4 specimens), tarsus 25.6-27.7 (26.8) mm. Type, male, wing 84.5, tail 80.8, culmen from base 15.1, tarsus 26.2 mm. Range.—High forests between 8,000 and 10,000 feet elevation in the southeastern section of the Sierra Nevada de Santa Marta (Siminchucua, and head of the Rio Guatapuri), Depto. Magdalena, Colombia. Remarks—While this bird is evidently closely allied to Dubusia taeniata of the Andes to the south, the differences appear so great as to establish carrikeri as a distinct species. It seems to have a defi- nitely restricted range. Male and female appear alike in color, and there is relatively little difference in size between them. An immature female taken at Siminchucua February 7 has the black of the head duller, and the dorsal surface, except for the wing edgings, dusky olive green. The throat and foreneck are cream buff with indefinite markings of dull black, the upper breast olive ocher, and the abdomen and sides rather dull Strontian yellow. NO. 16 NEW BIRDS FROM COLOMBIA—-WET MORE 13 The species is named for the collector, Melbourne A. Carriker, Jr., in recognition of his energy and skill in the investigation of the remote mountain region in which it is found. Family FRINGILLIDAE ARREMON SCHLEGELI FRATRUELIS, new subspecies Characters——Similar to Arremon schlegeli schlegeli Bonaparte 1° but with larger bill; averaging slightly larger in other dimensions. Description—Type, male adult, collected May 6, 1941, between 1,500 and 2,000 feet elevation in the Serrania de Macuire, near Nazaret, Guajira, Colombia, by A. Wetmore and M. A. Carriker, Jr. (orig. No. 11,812). Head, chin, upper parts of hindneck and an oblong patch on sides of neck extending down to upper breast deep black ; upper back and lower hindneck pale neutral gray ; back, scapu- lars, and rump warbler green ; wing-coverts sulphine yellow, becoming empire yellow on bend of wing; edge of wing distal to this white; ‘remiges and alula slate gray; greater wing-coverts slate gray exter- nally and black basally ; rectrices slate color, the shafts black ; throat, breast, abdomen, and under tail-coverts white; sides and flanks light neutral gray; under wing-coverts deep neutral gray, with a slight wash of pyrite yellow toward bend of wing; under side of rectrices blackish mouse gray. Bill colonial buff, fuscous black at base of cul- men ; tarsus and toes fuscous black; claw on hind toe cream buff, and on three anterior toes hair brown (from dried skin). Measurements.—Males, 7 specimens, wing 78.4-82.7 (81.2), tail 58.5-66.7 (62.9), culmen from base 16.5-17.3 (16.9), tarsus 24.7-26.6 (25.9) mm. Females, 5 specimens, wing 70.5-76.3 (73.3), tail 52.0-57.0 (55.5), culmen from base 16.3-16.7 (16.5), tarsus 24.8-26.0 (25.4) mm. Type, male, wing 81.7, tail 65.7, culmen from base 17.0, tarsus 26.0 mm. . Range.—The Serrania de Macuire, Guajira, Colombia, above 1,500 feet elevation. Remarks.—The differences of size characterizing this bird are very evident in series, and are worthy of note when the uniformity of the typical race of schlegeli throughout its extensive range in northern Colombia and northern Venezuela is considered. The size of bill found in fratruelis, while approached closely by a few, is not actually 10 Arremon schlegeli Bonaparte, Consp. Gen. Av., vol. I, pt. 2, 1850, D. 488 (Santa Marta, Colombia). I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 attained by any individual in the extensive series of schlegeli that I have had available for comparison. The Macuire race fratruelis is found only in the higher forests of the mountain range in question and probably consists of only a small population. The nearest groups of schlegeli are found in the eastward extension of the Sierra Nevada de Santa Marta (La Cueva and Los Gorros northwest of Barrancas) and the Sierra Negra (Tierra Nueva, east of Fonseca), separated by the entire extent of the arid Guajira Peninsula. Following are measurements from a series of Arremon schlegeli schlegeli from the Sierra Nevada de Santa Marta, the Sierra Perija, and Norte do Santander, Colombia: Males, 19 specimens, wing 73.1-81.4 (76.3), tail 58.2-65.9 (61.2), culmen from base 14.7-16.4 (15.7), tarsus 22.0-25.6 (24.0) mm. Females, 15 specimens, wing 66.3-73.4 (69.7), tail 52.0-57.7 (54.9), culmen from base 15.0-15.9 (15.4), tarsus 22.4-24.9 (23.8) mm. "SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 17 meena dips ee SOME NEW CAMBRIAN || -BELLEROPHONT GASTROPODS (Winx 2 PLATES) 4 BY: | J. BROOKES KNIGHT i hee Research Associate in Paleontology, U. S. National Museum ( Pusuicarion 3865 ) . _ CITY OF WASHINGTON LMA PUBLISHED BY THE SMITHSONIAN INSTITUTION jis JANUARY 3, 1947 | SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 17 CAalcott Fund SOME NEW CAMBRIAN BELLEROPHONT GASTROPODS (Wirth 2 PLATEs) BY J. BROOKES KNIGHT Research Associate in Paleontology, U. S. National Museum (PuBLicaATION 3865) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JANUARY 3, 1947 The Lord Baftimore Press BALTIMORE, MD., U. 8. A. Galcott Fund SOME NEW CAMBRIAN BELLEROPHONT GASTROPODS By J. BROOKES KNIGHT Research Associate in Paleontology, U. S. National Museum (WitTH Two Ptates) As far as I can learn, only two species and two genera of bellerophonts have been described from Cambrian rocks, and the placement of one of those as a bellerophont is problematical. Owen- ella antiqua (Whitfield) is an undoubted bellerophont assignable to the Sinuitidae. It occurs in beds of the Upper Cambrian, Trem- pealeauan stage in the upper Mississippi Valley, in Texas, and in the Great Basin. Following Whitfield’s faulty original description (Whit- field, 1878, p. 52) it has been described twice, once by Ulrich and Scofield (1897, pp. 847, 1080) and again by Knight (1941, p. 223). The second species that I am regarding very tentatively as a bellero- phont is Coreospira rugosa Saito, 1936. It was described from the Bonnia zone, the uppermost zone of the Upper Redlichia shales of Korea, by Saito (1936, p. 360) and was discussed by Knight (1941, p. 86). Although the Upper Redlichia shales are assigned to the upper part of the Lower Cambrian, the Bonnia zone is referred ques- tionably to the Lower Cambrian and may be lowest Middle Cambrian (Saito, 1936, p. 347). A second species of Coreospira will be de- scribed in this paper, a species collected from the lower Middle Cambrian Ptarmigan formation near Field, British Columbia. Coreospira has the symmetrically coiled shell of a bellerophont but lacks a feature that characterizes all other bellerophonts previously described, an emargination, sinus, or slit, in the supposedly anterior lip. In respect to this, and some other features, Coreospira seems allied to Helcionella Grabau and Shimer, 1909. It may be that Coreo- spira is transitional between Helcionella and the more typical, but still primitive, bellerophonts of the Upper Cambrian, a matter that I intend to discuss again in another paper. For the present I propose to consider it as the oldest known and most primitive bellerophont genus. In addition to the new species of Coreospira, I shall describe four new species of Upper Cambrian bellerophonts, all of which require SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 17 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 106 new genera for their reception. Some of the species described are represented only by steinkerns or fillings. Unless the fillings are of such a nature as to show evidence of the form or outline of the aper- tural lips, it is, in my opinion, generally useless to erect species or genera on them. So important is the form of the lip that a species erected without knowledge of it, or a genus based on such a species, usually is unclassifiable. Even from the utilitarian viewpoint such a species has little value as a stratigraphic guide, for generally it can only be recognized where the beds in which it occurs are identified first on other grounds. This, of course, is stratigraphic paleontology in reverse. The form of the outer lip can be determined (1) when the shell is perfectly preserved, very rare in Paleozoic beds, (2) when the outline of the apertural margins is preserved in the matrix, also relatively rare, and (3) when the growth lines on the outer surface of the shell are preserved in one way or another, for the growth lines are a record of the shell margins of earlier growth stages. If none of the foregoing conditions is fulfilled, there may be still other evidence that is useful if used with caution. Occasionally a filling is found which shows the imprint of rather faint undulations in the inner sur- face of the shell, undulations that record an irregularity of growth at some period in the life history of the individual. Since such an irregularity of growth affects the growing margin of the shell all at once, the undulation in the inner shell surface caused by it will usually give a satisfactory clue to the form of significant parts of the apertural margins. For example, if the reader will examine the figures of Sinuella minuta on plate 2 of this paper he will see that figures 2a-b show a specimen with the shell and its growth lines preserved although the apertural margins themselves are not preserved. These show clearly the form of the sinus in the anterior lip. Figure 2d shows a specimen that is a filling with the shell removed so that the growth lines are not preserved. However this filling shows an undulation recording an irregularity of growth, and this undulation also shows the outlines of the sinus in the outer lip. It is by employing such criteria that the form of the lip was determined for some of the species here described that otherwise would not have warranted description. Acknowledgments and #emarks.—I am deeply indebted to several colleagues, more familiar with the stratigraphy of the Cambrian than I am, for the placement of these species in the Cambrian time-strati- graphic sequence. For this, thanks are due to Virgil E. Barnes, Preston E. Cloud, Jr., G. Arthur Cooper, Norman M. Denson, and Christina Lochman. NO. 17 BELLEROPHONT GASTROPODS—K NIGHT 6) I offer no apologies for the unconventional illumination of the speci- mens photographed. It was necessary to direct the light so as to show objectively certain obscure but significant features on the specimens that would have been lost entirely if conventional lighting from the upper left had been employed. Stratigraphic position—In view of its possible importance for bellerophont phylogeny, as well as for stratigraphic application, the present known occurrence of the two old and four new genera of Cambrian bellerophonts is recapitulated in terms of the American Cambrian succession (Howell, et al, 1944, chart No. 1). Admittedly, the Cambrian succession is not well understood and is subject to re- vision as our understanding of it grows ; nevertheless its main features appear to be well enough understood for our present purpose. Coreospira Saito is now known from two widely separated regions, Korea and the Rocky Mountains of British Columbia. In both regions it occurs at the same, or approximately the same, level; just above (or below?) the boundary of the Lower and Middle Cambrian. Next in order appears Cycloholcus, new, related to Coreospira, near the base of the Upper Cambrian Dresbachian stage. Following these two very primitive and somewhat questionable bellerophonts, the first of the undoubted bellerophonts, Sinuella, new, appears in Upper Cambrian beds of upper Dresbachian or lower Franconian stage. Seemingly somewhat higher in the section, middle Franconian, appears Ancono- chilus, new. Cloudia, new, occurs in the lower Trempealeauan (or upper Franconian?) and thus is contemporaneous with, or possibly precedes, Owenella Ulrich and Scofield which characterizes the lowest zone of the Trempealeau stage and may range higher. Suborder BELLEROPHONTACEA Ulrich and Scofield, 1897 Family CorrosprripAE Knight, new family Genus COREOSPIRA Saito, 1936 Genotype, by original designation: Coreospira rugosa Saito, 1936. Diagnosis —Small, bellerophontiform gastropods with a flat or gently arched outer (dorsal) whorl-face extending laterally as a roundly subangular ridge overhanging the sides. The sides are nearly flat and the umbilici open. There is no emargination (sinus or slit) in the anterior lip and the margins of the lateral lips are approximately tangent to the coil. The ornamentation consists of growth lines and of transverse undulations strong enough to impress themselves on the inner surface of the shell. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Remarks.—Saito, in describing the genus originally, regarded it as coiled “slightly sinistrally (?)” and therefore as not bilaterally symmetrical or bellerophontiform. However, he seems to have been far from certain on this very fundamental point. On studying his description and figures I concluded that the shells he was describing were probably bellerophontiform and redescribed his species tenta- tively as such in 1941 (p. 86). The new species described below appears to confirm my conclusions in this respect. COREOSPIRA WALCOTTI Knight, new species PLATE I, FIGURES 2a-d Description—Small bellerophontiform gastropods seemingly of two or three whorls coiled symmetrically in a plane with widely open umbilici; outer whorl surface (here regarded as dorsal) flatly arched, bounded by rounded angulations on each side, the angulations being the widest part of the shell; sides gently concave sloping gently in- ward to the sharp umbilical shoulders; aperture flaring behind under the coil so that the growth lines are oblique to the radius and tangent to the coil ; anterior lip without an emargination, the growth lines pass- ing directly across the dorsal whorl-face ; ornamentation fine growth lines and broad transverse undulations on the sides of the shell, the latter seemingly more strongly developed inside the shell than outside, for the internal filling shows them much more strongly than does the shell surface. Measurements (approximate).—Holotype, U.S.N.M. No. 111960, length 6 mm., width 2 mm.; figured paratype, U.S.N.M. No. 111961, length 11 mm., width 3 mm. Hypodigm.—to specimens, 8 of them other than the holotype and figured paratype, under U.S.N.M. No. 111962. Remarks.—The specimens are preserved in a dark granular lime- stone and some of them appear to be distorted by pressure. Some patches of shell are preserved. Only one specimen shows the left side and although it is imperfect there is nothing about it to indicate asymmetry. Comparisons.—This species differs from C. rugosa Saito most con- spicuously in that its dorsal surface is gently rounded instead of flat. Occurrence.—Top of the Ptarmigan formation, lower Middle Cam- brian at Walcott’s locality 58k on Mount Stephen, 3 miles east of Field, British Columbia. These beds were originally placed in the Lower Cambrian Mount Whyte formation (Walcott, 1928, p. 317) but are now assigned as given above. NO. 17 BELLEROPHONT GASTROPODS—-KNIGHT 5 Genus CYCLOHOLCUS Knight, new genus Genotype: Cycloholcus nummus Knight, new species. Diagnosis.—Disklike, bellerophontiform gastropods with prominent rounded keels bordering the outer (dorsal) whorl-face and the um- bilici, with a rounded groove between. The lateral margins of the aperture are oblique to the radius and tangent to the coil. There seems to be no emargination in the outer lip. The transverse section of a whorl is roughly dumbbell-like. It differs from Coreospira Saito largely in its coinlike shape and in the relatively prominent keel about the umbilici. It seems to lack transverse undulations of such strength as to be impressed on the inner surface, for, unlike Coreospira, fill- ings are nearly smooth. The genotype is the only known species. CYCLOHOLCUS NUMMUS Knight, new species PLATE I, FIGURES 3a-d Description—Seemingly of about 34 whorls; the keels surrounding the umbilici protruding slightly more than those bordering the outer whorl-face ; the outer whorl-face narrow and rounded ; ornamentation coarse, fasciculated growth lines that show on the inner surface of the shell only sporadically and faintly. Measurements.—Holotype, U.S.N.M. No. 111957, length 10.4 mm., width 2.8 mm.; figured paratype, U.S.N.M. No. 111958, length 11.0 mm. Hypodigm.—7 specimens, 5 of them, other than the holotype and figured paratype, under U.S.N.M. No. 111959. Remarks.—tThe holotype and figured paratype are free fillings ac- companied by the matrix from which they were broken. The other paratypes are still in the dark-gray granular limestone matrix. The block from which the figured paratype was broken retains the shell for the most part, but a few fragments clung to the filling, exposing small areas of the impression of the ornamentation in the matrix. Occurrence——From limestone interbedded with the Nolichucky shales (Dresbachian stage, lower Upper Cambrian) at some point in eastern Tennessee now uncertain; probably from a point on Cub Creek, 1.5 miles southeast of Morristown, Tenn. Family CyrrotitmaE Ulrich and Scofield, 1897 Genus CLOUDIA Knight, new genus Genotype: Cloudia buttsi Knight, new species. Diagnosis.—Bellerophontiform gastropods of moderately large size, 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 with deep, open umbilici surrounded by subangular shoulders, and with roundly subangular dorsum. The whorls are in contact but with- out involution. The anterior lip has a shallow, gently rounded sinus. In many respects the genus reminds one of Cyrtolites Conrad, 1838, but its whorls are not so quadrate in section and there is no dorsal notch-keel; neither does it show the strong transverse undulations that help characterize Cyrtolites. CLOUDIA BUTTSI Knight, new species PLATE 2, FIGURES Ia-e Description—Seemingly of four or more whorls when mature, the whorls nearly twice as wide as high, with wide, steep, very gently arched umbilical slopes; the dorsal surface arched with nearly flat slopes and rounded crest; ornamentation unknown except on the inner part of the umbilical slopes where it consists of fine growth lines ; the sinus in the anterior lip shallow and gently arched. Measurements.—Holotype, U.S.N.M. No. 112054, length 32 mm., width 22 mm.; figured paratype, U.S.N.M. No. 112056, length 21.4 mm., width 15 mm. Hypodigm.—24 specimens, as given below. Remarks.—All the specimens are fillings preserved in rough, dolo- moldic * chert. Only one preserves some silicified shell surface in the middle part of the umbilicus. Several show the form of the outer lip faintly but unmistakably recorded as imprints of slight irregulari- ties of growth. If it had not been for the record of the highly im- portant form of the outer lip I should not have ventured to describe and name either the species or genus. This very interesting species was discovered independently by Dr. Charles Butts, of the United States Geological Survey and by a party headed by Dr. Preston E. Cloud, Jr., of the same organization. Both Dr. Butts and Dr. Cloud called it independently to my attention and both regard it as a useful index fossil for the Copper Ridge dolomite of the southern Appalachians. Occurrence.—All specimens are from the lower part of the middle third of the Copper Ridge dolomite just above a zone of digital, Gymnosolen-like stromatolites. This position seems to be assignable to the Trempealeauan stage of the Upper Cambrian but may be upper Franconian. The collections were made at the following localities in Alabama: The holotype (U.S.N.M. No. 112054) and a paratype 1 See Cloud, Barnes, and Bridge, 1945, p. 135. NO. 17 BELLEROPHONT GASTROPODS—KNIGHT 7 (U.S.N.M. No. 112055) from a point about midlength of the Alabama-Georgia State line, NW4, NW4, fractional sec. 19, T. 11 S., R. 12 E., U.S. G. S. Special Map, Rock Run and vicinity ; a figured paratype (U.S.N.M. No. 112056) and 3 unfigured paratypes (U.S.N.M. No. 112057) from a point at the midlength of the south side NWi, NE, sec. 5, T. 12 S., R. 10 E., Cherokee County, Ala. (rock in place) ; a figured paratype (U.S.N.M. No. 112058) and 4 unfigured paratypes (U.S.N.M. No. 112059) from a point just south of center of SW1, NE}, SW, sec. 12, T. 12 S., R. 9 E., Cherokee County, Ala.; 4 unfigured paratypes (U.S.N.M. No. 112062) from a point near center of E4, NE, NE}, SW3, sec. 22, T. 12S.,R. 9 E., Cherokee County, Ala.; 4 unfigured paratypes (U.S.N.M. No. 112061) from a point about 200 feet south-southwest of a point in the SW1, NE}, SE4, SEI, sec. 15, T. 12 S., R. 9 E.; 6 unfigured paratypes (U.S.N.M. No. 112060) from a point about 2 miles north- west of center of Anniston, Anniston Quadrangle, Calhoun County, Ala. Family StnurripaE Dall, in Zittel-Eastman, 1913 Genus ANCONOCHILUS Knight, new genus Genotype: Anconochilus barnesi Knight, new species. Diagnosis——Rather large, involute, and probably narrowly phan- eromphalous gastropods with a deep, broad, rounded sinus in the anterior lip; posterior lip of the aperture seemingly flaring backward under the coil; whorl section rounded dorsally with the sides of the dorsal slope somewhat flattened; surface characters of the shell entirely unknown. Remarks—When preserved in the form of fillings, Anconochilus is remarkably similar to Sinuites Koken, 1896. Indeed it was only after considerable study of the specimens ,of the genotype species, all of them fillings, that I felt that I could differentiate it from Sinuittes. The basis for the differentiation resides in the form of the lateral and probably the posterior lips as indicated on several of the speci- mens by the impressions of the irregularities of growth. These in- dicate that the margins of the lateral lips did not have the pronounced forward convexity that characterizes Sinuites and led to the name “bilobaius” for its genotype species. On the contrary, the margins of the lateral lips in Anconochilus sweep toward the sides and downward from the apex of the sinus to the circumumbilical shoulders with only a very gentle convexity. Furthermore they appear to have passed tangentially under the whorl in such a way that there must have been 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I06 some backward flaring. If this is the case, as it seems to be, the um- bilicus most probably was open although narrow. To clarify these distinctions the student can compare profitably the illustration show- ing the side view of the holotype of Sinuwites cancellatus (Hall) in Knight, 1941 (pl. 6, fig. 3a) and those of the holotype of the present species. ANCONOCHILUS BARNESI Knight, new species PLATE I, FIGURES Ia-e Description—Since the surface characters of this species are en- tirely unknown, the diagnosis given above for the genus, supplemented by the following measurements, will serve also as the description of the species. Measurements (approximate ).—Holotype, U.S.N.M. No. 112045, length 28 mm., width 24 mm.; figured paratype, U.S.N.M. No. 112046a, length 32.5 mm., width 27.5 mm.; figured paratype, » U.S.N.M. No. 112046b, length 26 mm., width 22 mm. Hypodigm.—tThe holotype, 2 figured paratypes, and 8 unfigured paratypes (U.S.N.M. No. 112047). Occurrence.—Pedernales dolomite member of the Wilberns forma- tion at a level about 50 feet above a stromatolitic bioherm that prob- ably is assignable to the Point Peak shale member of the Wilberns. This suggests that the occurrence is at about the middle of the Franconian stage of the Upper Cambrian. The locality (Univ. Texas 86-T-2-14G) is 2.5 miles airline southeast of Willow City, Gillespie County, Tex., 2,400 feet southeast of the intersection of the Willow City-Sandy Post Office road and the old Fredricksburg-Burnet road ; 1,000 feet east of Willow Creek at a point 1,500 feet south of Willow City-Sandy Post Office road, and 200 feet south of Cretaceous overlap. Genus SINUELLA Knight, new genus Genotype: Sinuella minuta Knight, new species. Diagnosis:—Small, widely phaneromphalous, only slightly involute bellerophontiform gastropods with a gently concave dorsum bordered by low, rounded shoulders. The anterior lip has a moderately deep, rounded sinus. Comparisons.—Sinuella reminds one of Bucanella Meek, 1871, in its broad, rather flat whorls, open umbilici and in the shape of the sinus in the anterior lip. It differs markedly from Bucanella in that its dorsum is slightly concave and faintly bilobate while that of Bucanella is strongly convex and trilobate. In fact, the faintly de- pressed dorsum is unique among bellerophonts. NO. 17 BELLEROPHONT GASTROPODS—KNIGHT 9 SINUELLA MINUTA Knight, new species PLATE 2, FIGURES 2a-g Description With the characters given above for the genus. The 6rnamentation consists of growth lines occasionally expressing them- selves on the dorsum as widely spaced, sharp transverse lirae. There must have been something like 5 or 6 whorls. Measurements (approximate ).—Holotype, U.S.N.M. No. 112049, length 1.27 mm., width 0 mm.; figured paratype, U.S.N.M. No. 112050b, length 1.52 mm., width .g2 mm. ; figured paratype, U.S.N.M. No. 112052, length 3.8 mm. Hypodigm.—About 30 specimens as listed below. Remarks—The minute specimens of this species are mostly im- bedded in a gray, granular, glauconitic limestone. The holotype re- tains the shell. It is broken in two parts, one part free and the other imbedded in matrix. All the other specimens are fillings, either partly exposed in matrix or broken free from it. The shell commonly ad- heres to the outer matrix rather than to the filling. A few of the fillings are in part glauconitic. Occurrence.—The specimens were all collected by the late Dr. C. D. Walcott in 1885, and the precise level of their occurrence in the Upper Cambrian is now undeterminable from the records. Dr. Preston E. Cloud, Jr., with others, has recently restudied in detail the early Paleozoic stratigraphy of the localities from which the specimens came and on the basis of associated fossils and lithology he advises me that these specimens could not have come from beds higher than the Point Peak shale member of the Wilberns formation nor lower than the Cap Mountain limestone member of the Riley formation. He is further of the opinion that they most probably come from either the upper part of the Cap Mountain limestone member or from lime- stone interbeds in the Lion Mountain sandstone member of the Riley formation, or from the Morgan Creek limestone member of the Wil- berns. Thus they are of late Dresbachian or early Franconian age. Other than the Coreospiridae, this is the earliest bellerophont now known. The specimens are represented in three collections from two locali- ties in the Central Mineral Region of Texas. The holotype (U.S.N.M. No. 112049), 2 figured paratypes (U.S.N.M. No. 112050a and U.S.N.M. No. 112050b), and 12 unfigured paratypes (U.S.N.M. No. 112051) in one collection, and 6 unfigured paratypes in another (U.S.N.M. No. 112048) are from Potatotop Hill, 7 miles IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 northwest of Burnet, Tex. One figured paratype (U.S.N.M. No. 112052) and 7 unfigured paratypes (U.S.N.M. No. 112053) are from Packsaddle Mountain, 11 miles southeast of Llano, Tex. REFERENCES Croup, P. E., Jr., Barnes, V. E., and Brincg, J. 1945. Stratigraphy of the Ellenberger group in central Texas, a progress report. Univ. of Texas Publ. 4301. Howe tt, B. F., et al. 1944. Correlation of the Cambrian formations of North America. Bull., Geol. Soc. Amer., vol. 55, pp. 993-1003. KNIGHT JLB.) 4 1941. Paleozoic gastropod genotypes. Geol. Soc. Amer., Spec. Pap. 32, pp. 1-150, pls. 1-96. Sarto, K. 1936. Older Cambrian Brachiopoda, Gastropoda, etc. from northwestern Korea. Imp. Univ. Tokyo, Journ. Fac. Sci., sect. 2, vol. 4, pp. 345-367, pls. 1-3. UtricH, E. O., and Scorietp, W. H. 1897. The Lower Silurian Gastropoda of Minnesota. In Geology of Min- nesota. Final Rep., vol. 3, pt. 2, pp. 813-1081, pls. 61-82. Watcortt, C. D. 1928. Cambrian Geology and paleontology, V. No. 5.—Pre-devonian Pale- ozoic formations of the Cordilleran provinces of Canada. Smith- sonian Misc. Coll., vol. 75, No. 5, pp. 175-377, pls. 26-108. WHuitFIELD, R. P. 1878. Preliminary descriptions of new species of fossils from the~lower geological formations of Wisconsin. Ann. Rep. Wisconsin Geol. Surv. for 1887, pp. 50-86. EXPLANATION OF PLATES PLATE I 1a-e. Anconochilus barnesi Knight, new species. All figures 1. 1a-b, the holotype (U.S.N.M. No. 112045), oblique and side views respectively. Note on ta the irregularities of growth outlining the margin of the outer lip at former stages of growth. Ic, a paratype (U.S.N.M. No. 112046b), side view. 1Id-e, a paratype (U.S.N.M. No. 112046a), front and side views respectively. Irregularities of growth outline the anterior sinus in figure 1d. What appears to be the lateral margin of the lip in figure Ie is actually a line of breakage intersecting the course of the true margin at a high angle. 2a-d. Coreospira walcotti Knight, new species. All figures 4. 2a-b, the holotype (U.S.N.M. No.1r1960), side and oblique views respectively. The specimen is a filling except for the small patch of adhering shell in the lower central part. 2c-d, a paratype (U.S.N.M. No. 111961), side and front views respectively. The specimen is largely a filling except for patches of shell adhering in the umbilical region and on the lt SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 17, PL. 1 CAMBRIAN BELLEROPHONT GASTROPODS (See explanation of plates.) VOLE 106; NOD 17mm SMITHSONIAN MISCELLANEOUS COLLECTIONS CAMBRIAN BELLEROPHONT GASTROPODS ion of plates.) (See explanat ® NOL 7: BELLEROPHONT GASTROPODS—KNIGHT If outer whorl-face. The growth lines and irregularities of growth on the latter show clearly that there is no anterior sinus. 3a-d. Cycloholcus nummus Knight, new species. All figures X 4. 3a-b, the holotype (U.S.N.M. No. 111957), side and marginal views respectively. Note the irregularities of growth tangent to the umbilicus and passing obliquely backward across the sides of the whorl. These indicate positions of former lateral apertural margins. The specimen is a fil- ling, without shell and with the umbilical coil broken through. 3c-d, a paratype (U.S.N.M. No. 111958), side view of filling and corres- ponding inner shell surface adhering to the matrix respectively. There are small patches of shell adhering to the filling and, in the correspond- ing areas of the counterpart, impressions in the matrix of some of the external surface may be seen. These not only give information as to the ornamentation but, together with the irregularities of growth, confirm the course of the lateral apertural margins inferred on the holotype from irregularities of growth alone. PLATE 2 ta-e. Cloudia butisi Knight, new species. All figures X I. Ia, a paratype (U.S.N.M. No. 112058), front view. Note the irregularities of growth near the lower (adapertural) end outlining a former apertural margin and thus showing the shallow, rounded sinus in the anterior lip. 1b, a paratype (U.S.N.M. No. 112056), with the whorl in the upper part sectioned transversely. 1c-e, the holotype (U.S.N.M. No. 112054), three views to show the form. All specimens are fillings. 2a-g. Sinuella minuta Knight, new species. All figures 15. 2a-b, the holo- type (U.S.N.M. No. 112049), two slightly oblique dorsal views show- ing form and growth lines. This specimen retains the shell. 2c-d, a paratype (U.S.N.M. No. 112050b), side and dorsal views respectively. This specimen is a filling and an irregularity of growth shown on figure 2d confirms the form of the anterior lip shown by the growth lines on the holotype. 2e, the holotype, view from below. The specimen is broken across and shows sections of the whorls and the degree of involution. 2f, side view of a paratype (U.S.N.M. No. 112050a). 2g, side view of a paratype (U.S.N.M. No. 112052). This is the largest specimen in the collection and, like many specimens, is poorly preserved as a partial glauconitic filling. . eal st "SMITHSONIAN MISCELLANEOUS COLLECTIONS ARS VOLUME 106, NUMBER 1835 “OF THE PYCNOGONIDA re ‘ewes One Brarey JOEL W.' HEDGPETH) . Marine Biologist, Texas Game, Fish and Oyster Commission | ( PUBLICATION 3866) "CITY OF WASHINGTON "PUBLISHED BY THE SMITHSONIAN INSTITUTION | MARCH 24, 1947 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 18 wi tHE EVOLUTIONARY SIGNIFICANCE OF THE PYCNOGONIDA (WitTH ONE PLATE) BY JOEL W. HEDGPE TH Marine Biologist, Texas Game, Fish and Oyster Commission (Pusrication 3866) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION MARCH 24, 1947 The Lord Baltimore Presse BALTIMORE, MD., U. 8 A. ak ON THE EVOLUTIONARY SIGNIFICANCE OF THE PYCNOGONIDA By JOEL W. HEDGPETH Marine Biologist, Texas Game, Fish and Oyster Commission (WitH ONE PLATE) INTRODUCTORY NOTE The Pycnogonida, or sea spiders, are an anomalous class or sub- phylum of marine arthropods, unknown except by name to most zo- ologists. They are of no economic importance to man, and of little discernible significance in the natural order of things. Yet within the last I0 years more than 50 papers dealing with these creatures have been published, and the complete bibliography now comprises several hundred titles. More than 500 species have been described, but there are relatively few parts of the world in which the pycnogonid fauna is adequately known, and the actual number of extant species may be considerably larger. Also known as Pantopoda, the Pycnogonida have often been con- sidered an “appendix” to the Arachnida in comprehensive treatises, but they have no real relationship to the arachnids, since at no stage in their development do they have a cephalothorax or prominent abdo- men. Their relationship to the Crustacea is even more tenuous, for they do not have biramous appendages, and their own peculiar larval stage, the protonymphon, is distinct from all other arthropod larvae. They are characterized by an extreme reduction of the body, very long legs, which house the sex glands and diverticulae of the gut, and a sub- sidiary pair of egg-bearing legs, or ovigers, which are present in all males but lacking in the females of some genera. In addition to the walking legs, which are usually 8 in number, but may occasionally be IO or 12, and the ovigers, there may be a pair of chelate appendages (chelifores) and sensory palpi. The presence or absence of these accessory appendages constitutes the basis of classification within the group. There is a simple nervous system of ventral ganglia, and a rudimentary circulatory system. There is no respiratory system or any specialized excretory organ, although in the males of many species there is a specialized cement gland which is believed to be of use in attaching the eggs to the ovigers of the male, who carries them around SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 18 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 until they are hatched. Shortly after hatching, the larvae of many species become encysted in hydroids, sea anemones or small medusae, where they live as parasites for a time. Some species are found in holothurians and at least one species spends its early life in the mantle cavity of bivalve mollusks. Pycnogonids are found from the littoral zone to depths of more than 2,000 fathoms, in all oceans. Some live on sargassum at the surface while others appear to be bathypelagic. They vary in size from a span of 3 mm. for small littoral species to about 50 cm. in some of the abyssal forms. In the course of my taxonomic studies of the collections of Pycno- gonida in the United States National Museum, the Museum of Com- parative Zoology, and the Peabody Museum at Yale, I have examined several hundred specimens, including examples of many of the known 10-legged forms, and in this paper I shall attempt to bring together in a coherent manner some of the speculations inspired by this large mass of material. Unfortunately, little is known about these creatures, especially the deep-sea forms, other than that which can be surmised from their pickled corpses. The following discussion, therefore, is not intended to blaze new paths in invertebrate morphology and evolu- tion, but simply to suggest some directions in which future investi- gators might profitably set out. In other words, I have gathered together some speculations on the possible causes of some observed effects, for, as Aristotle said, “Nature does nothing which lacks purpose.” + I wish to acknowledge with thanks the generosity of Dr. Waldo L. Schmitt, Head Curator of Biology of the United States National Museum, in loaning me material and literature, and I also wish to thank Dr. Isabella Gordon, of the British Museum, for her patient answers to my persistent correspondence at a time when the British Museum was in the front line of battle (and suffered accordingly) and for the gift of a specimen of Nymphon hiemale from the Dis- covery collections. I. PHYLOGENY AND PATTERNS OF VARIATION From time to time attempts have been made to divide the Pycno- gonida into orders, but the families are so closely related, and their boundaries so broken down by transitional generic forms, that none of these attempts have been successful. An attempt to separate the families on the basis of the presence of ovigers in both sexes, or in the 1 Gen. Anim. II, v. (741 b, 4-5), Loeb Classics ed., p. 207. ‘No. 18 PYCNOGONIDA—HEDGPETH 3 males only, for example, breaks down in the genus Pallenopsis, which has well-developed ovigers in the male and rudimentary ones in the female. Some workers have assigned this genus to the Phoxichilidi- idae, others consider it a member of the Pallenidae. Diagnoses based on the retention or loss of chelifores in the adult cannot even be applied at the generic rank, particularly in the predominantly achelate genus Achelia, which has several chelate species. Hence, when one comes to draw a family tree, it looks more like a tangled web, or a bush with anastomosed branches, as in figure I. Doaecolopoaa Pentanymphon Decolopoda Pentacolossenaeis yepeesars ers ders CNOGONUIN Mymphon ..2.--7 Colossende paged Phoxichilidium ~orrts---- -- Paranymphon Ainiqma Pigrogromitus Achelia ee iw) 7 RG 4 ‘ ’ iS) S Sa teas. Sx Ny ra S eae ie C. a Ny mphonella Ascorhynchus bet a if ? < CG 1 & RS = fv’ Decognela 0 oS HS) rt? oN 15 Ho ae {i GE C ak aor? Fic. 1.—Hypothetical family tree of the Pycnogonida. (Transitional genera in backhand lettering. ) It will be noted that this family tree—or bush—rises from a hemi- spherical base. This base is the quantitative diagram of the families represented in figure 2, together with diagnostic drawings of the most common types of pycnogonids. Most of the forms illustrated in this diagram represent the type genera of the families concerned. Such attempts as this to erect a family tree naturally bring up the problem of the roots, that is, the phylogenetic relationships of the Pycnogonida with other arthropod groups. This is probably the most difficult problem connected with these animals and may never be solved. Snodgrass, in the most recent and lucid discussion of arthro- pod evolution (1938) is none too certain of the affinities of the Pycnogonida, beyond placing them in the Chelicerata together with the Arachnida and Xiphosurida. But, if we accept his restriction 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 TABLE 1.—Synopsis of the families of Pycnogonida Family Ovigers Chelifores Palpi ag Principal genera NYMPHONIDAE a, @; present, 5 jt. 4-5 | Nymphon Wilson, 1878 |10jt.,with| chelate Heteronymphon compound [Pentanym phon] spines PALLENIDAE op tole present, | absent, or| 4 Callipallene Wilson, 1878 |10jt.,with| chelate | knoblike, Pseudopallene compound or 3-4 jt. Cordylochele spines Austropallene Oropallene Propallene Hannonia Pycnothea Decachela Pigrogromitus Pallenopsis PHOXICHILIDIIDAE oie present, | absent, or} 4 Phoxichilidium G. O. Sars, 5-9 jt.,with} chelate knoblike Anoplodactylus 1891 simple Halosoma spines Hodgsonia Pycnosoma ENDEIDAE fogih absent absent 4 | Endeis Norman, 1908 | 7 jt., with simple spines AMMOTHEIDAE a, 9; present, 6-10 jt. Ammothea Dohrn, 1881 Q-10 jt., usually | (Mympho- Achelia spines achelate, nella, Ammothella simple or | but some- | 17-20 jt.) Nymphopsis compound | times with Ascorhynchus small Nymphonella chelae Eurycyde Cilunculus Boehmia Ephyrogymna Pycnofragilia Heterofragilia Austroraptus Lecythorhynchus Ainigma Paranymphon TANYSTYLIDAE CHa one achelate 4-6 4 Tanystylum Schimkewitsch,| 10 jt., with Clotenia 1913 compound Austrodecus spines Rhynchothorax Discoarachne Dorhynchus Trygaeus Scipiolus NYMPH( oviger second tibia cox / propodus | AL Q\ Achelia spinosa oo ey i= aA pi aX a ee AMMOTHEIDAE Tanystylum orbiculare ff oK (| {] aall A/ / ‘ Awe Ss. tL asst TANYSTYLIDAE SAN NYMPHONIDAE Ts DP Achélia spinosa . PYCNOGONI pPallenopsis, vite S E P caicanea PALLENIDAE Fic. 2.—Diaerz ttictts ait : ; ; 2.—Diagram of qualitative and quantitative relationships of the Pycnogonida. a TANYSTYLIDAE “} no. 18 PYCNOGONIDA—HEDGPETH 5 TABLE 1.—Synopsis of the families of Pycnogonida—Continued Family Ovigers | Chelifores Palpi | jouw] Principal genera COLOSSENDEIDAE o’, 2; | absent, or 8-10 4-6 | Colossendeis Hoek, 1881 10 jt., with | deciduous, Rhophalorhynchus compound | or chelate [Pentacolossendeis spines in extra- Decolopoda legged form Dodecolopoda| ? Pipetta PYCNOGONIDAE cenlt absent absent | 4-5 | Pycnogonum Wilson, 1878 | 6-9 jt., no [Pentapycnon] spines (p. 137) of the Chelicerata to forms in which the genital openings occur always on the eighth postoral somite, the Pycnogonida occupy an anomalous position, since in several forms there are genital openings only on the last pair of legs. If we assign two postoral segments to the proboscis, as some do, this places the genital openings on the ninth postoral segment. This difficulty is recognized by Snodgrass himself (p. 138), and it would seem best not to attempt to include the Pycnogonida with any other group, but to leave it on a limb of its own somewhere between the Annelida and Arachnida. This is essentially the view of those early students of the group, Dohrn and Hoek, who argued in favor of an independent origin for the Pycnogonida (cf. Hoek, 1881, as translated by Morgan, 1891, pp. 26-28). Morgan, however, believed that the Pycnogonida shared a common, albeit remote, origin with the Arachnida, and supported his view by pointing out embryological parallels between marine and terrestrial spiders, as well as such anatomical similarities as the diver- ticulae of the gut and the occurrence of chelae on the first pair of appendages. The most recent statement in favor of an independent position for the Pycnogonida is that of Marcus, (1940b, p. 129) : The Pantopoda do not in any phase possess the crustacean biramous limbs nor the arachnomorphous body composed of cephalothorax (prosoma) with six pairs of appendages and abdomen (opisthosoma). Therefore it seems advisable to consider them as a separate class of the Arthropoda—or the Euarthropoda, if the Malacopoda (Onychophora and Tardigrada) are left aside—and not to include them in Crustacea or Arachnomorpha (Merostomata and Arachnoidea ) and thereby make diagnoses for these classes impossible. The argument in favor of arachnid affinities for the Pycnogonida still has its supporters, however. In the imposing monograph for the Bronn’s Tierreich series, Helfer and Schlottke (1935) consider the Pycnogonida a subclass of the Chelicerata, nearer to the Arachnoidea 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 than to any other group. But they are none too sure on the point, and conclude their discussion of affinities with this escape clause (p. 179) : “Aber dariiber ist das letzte Wort noch nicht gesprochen.”’ Calman and Gordon (1933) are more positive in their view of the arachnid affinities of the Pycnogonida, and advance the theory that discrepan- cies between the Pycnogonida and the Arachnida might be accounted for by metameric instability in the cephalic region of the Pycnogonida. It does not seem to be that this is a tenable view, for although there is Fic. 3.—Some transitional forms of pycnogonids. a, Pigrogromitus timsanus (after Calman, 1927); b, Paranymphon spinosum; c, Rhynchothorax mediter- raneus (after Dohrn, 1881) ; d, Decachela discata; d’', tarsus and propodus; e, Pentacolossendeis reticulata. The line indicates 1 mm. evidence of instability in the Pycnogonida, I am inclined to believe that it is too recent to be of phylogenetic significance, and is of a different character than Calman and Gordon apparently believed. However, I do agree with these authors that certain fossil forms from the Lower Devonian are not pycnogonids, and I further sus- pect that the group is a fairly recent one, and is still undergoing active evolution. Certainly an inspection of the transitional and anomalous genera (see fig. 3) lends support to this view, for they present examples of missing links which would delight and confuse a paleontologist. This occurrence of diversified forms connected by numerous transitional no. 18 PYCNOGONIDA—HEDGPETH 7 types suggests youth rather than age, for we would expect a loss of transitional and experimental forms in an old group. There is, for example, the curious genus Pigrogromitus Calman (1927) from the Suez Canal.2 The body type of this genus is the same as that of Pycnogonum, the type and only genus of the family Pycnogonidae, which is without chelifores and palpi and lacks ovigers in the female. But Pigrogromitus, with its 10-jointed ovigers in both sexes and chelate chelifores, must be placed in the Pallenidae. Nor is it the only transitional genus in this family; it represents but one of the extremes of variation. Most members of the Pallenidae are of the long-legged, extended type, but there are several compact disciform genera, such as Pseudo- pallene, which suggest affiliation with the Tanystylidae. Without its peculiar propodus, which Hilton (1939a) considered to be a family character, Decachela appears to be another transitional form between the Tanystylidae and Pallenidae. The possession of a large spine on the sole, which is opposable to the terminal claw, the two forming a sub- chelate structure, cannot be considered a character entitling this form to family status, although it may be a variation sui generis. There are the usual eight joints in the leg instead of the seven suggested by Hilton’s statement, “legs apparently seven jointed.” The genus Pallenopsis, a transitional form between the Pallenidae and the Phoxichilidiidae, has already been referred to. The phoxi- chilidiid characters of this genus are the possession of femoral cement glands in the male and the overhanging prolongation of the cephalic segment. Because of its 10-jointed ovigers, present in both sexes but reduced in the female, I consider it a pallenid. Thus, in one family alone, there are transitional forms indicating affinities with three diverse groups considered worthy of family rank. If we consider the Tanystylidae, which is related to the Pallenidae through such genera as Pseudopallene and Decachela, we find that it in turn is related to the Pycnogonidae through the genus Rhyncho- thorax, which has a body form approximating that of Pycnogonum. Inasmuch as the Tanystylidae are almost inseparable from the Am- motheidae, observations on the relationships between either of these iamilies and the other families are mutually applicable. The principal differences between the two families are that the proboscis is usually large and bulbous in the Ammotheidae and that the palpus has more 2 Hilton’s (1942c) Pigrogronvitus robustus from Unalaska is actually Pycno- soma strongylocentrott Losina-Losinsky, and constitutes an extension of range for that species from its type locality (48° 58.2’ N., 140° 35.3’ E.) to Alaskan waters. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 joints. The Ammotheidae (and Tanystylidae) comprise a diverse group of forms whose common characters are the reduction of the chelifores (or, when chelate, the chelae are small and lack teeth on the fingers), well-developed palpi, and ovigers in both sexes, usually with compound spines on the terminal segments. Among the interesting genera in the Ammotheidae are Cilunculus Loman (1908), the male of which has femoral cement glands opening through a prominent dorsal tube as in Pallenopsis and some species of Anoplodactylus (Phoxichilidiidae), and Paranymphon Caullery (1896), which has heretofore been referred to the Nymphonidae. In Paranymphon the palpi are seven-jointed, whereas in Nymphon they are always five-jointed, but the principal reason for removing this genus from the Nymphonidae is the discovery of a somewhat similar form, Ainigma Helfer (1938), in which the ammotheid affinities are more clearly marked, and which in turn seems to be more closely re- lated to Paranymphon than to any other genus. These two curious forms are very similar in body form, with high dorsal tubercles on the well-separated lateral processes, and simple tarsal joints. In contrast to the diversity of form in the Ammotheidae, Tany- stylidae, Pallenidae, and Phoxichilidiidae, the families Nymphonidae, Colossendeidae, Pycnogonidae, and Endeidae are remarkably uniform. The Pycnogonidae and Endeidae are monogeneric families, and there are but two indubitable genera in the Nymphonidae and Colossen- deidae, apart from the extra-legged forms, which are a special case. These latter families, however, possess large numbers oi closely re- lated species. The most conspicuous thing about this pattern of variation is the way in which it is correlated with extra-legged forms. It will be noted, from an examination of figure 2, that in families where variation is manifested at generic rank (as indicated by the high proportion of genera to species), 10-legged forms do not occur (so far as we know), whereas in those families in which variation is more active at the species rank, several 10-legged forms have appeared. The only ex- ception to this generalization, is the Endeidae, a small monogeneric family which may actually be an offshoot of the Pycnogonidae. It will be noted that I have not attempted to indicate the compara- tive ages of the various families or branches in my diagram. It is usually contended that the Pycnogonidae are the most specialized family, and the Nymphonidae the most generalized group, retaining more of the primitive attributes than the Pycnogonidae. The Colos- sendeidae are intermediate, according to this view, and the other families branch out from the tree more or less according to individual No. 18 PYCNOGONIDA—HEDGPETH 9 fancy or taste. When t1o-legged forms were first discovered, it was suggested that they were the primitive types, and should be at the base of the tree (Cole, 1905), but this suggestion was made before these forms were well known. It now seems more reasonable to sug- gest that they are recent innovations in form, at least as far as the present pattern of variation is concerned. The occurrence of these Io0- legged forms in three widely divergent branches suggests a common origin for these branches. For that matter, none of the families are actually different enough to enable one to assign any to a higher or lower place on a vertical scale, and the pattern of variation in the Pycnogonida is not amenable to a lineal or two-dimensional inter- pretation, but is three-dimensional, the various families or branches diverging in all directions from a central or nuclear type. How such a structure might be bound into relationship with the other groups of the Arthropoda, is difficult to say. However, it is probably no more difficult to visualize a relationship of three-dimen- sional structures than it is to decide just where, in a simple branching pattern, the Pycnogonida stem out from the remaining Arthropoda. All phylogenetic trees and speculations are influenced by honest errors in evaluating characters, but as Snodgrass said in his concluding remark (1938, p. 149): “Every biologist must have a working creed of phylogeny, but he should not too implicitly believe its tenets.” II. TEN-LEGGED FORMS Forty years ago, most zoologists who interested themselves in the matter believed that the Pycnogonida possessed but four pairs of walking legs, and considered that feature one of the diagnostic charac- ters of the group. Although the first 10-legged form had been col- lected by James Eights on a voyage to the South Shetlands in 1829 and described, with an adequate figure, in 1835 under the name Deco- lopoda australis, it was generally ignored by naturalists? For ex- 3 For the melancholy story of Dr. James Eights (M.D.) and his Antarctic travels, see The Reincarnation of James Eights, Antarctic explorer, by John M. Clark (Sci. Month., vol. 2, No. 2, pp. 189-202, 1916), and James Eights, a pioneer Antarctic naturalist, by W. T. Calman (1937a). The latter paper is based on the earlier one, but a bibliography of Eight’s writings has been added and the discussion of his zoological discoveries is more extended. There is also an interesting diagram of extra-legged pycnogonids, and a facsimile of the original figure of Decolopoda australis. Unfortunately the color of the copy from which the facsimile was made is poor; it should be red instead of slate brown. Further information on Eights and his contemporaries will be found in a brief paper by Lawrence Martin, Early explorations and investigations in southern South America and adjacent Antarctic waters by mariners and scientists from I0 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 ample, Hoek, in his monograph on the Challenger Pycnogonida (1881, p. 6), dismissed Eight’s Decolopoda with these words: “, . . L have not been able to ascertain whether this is a good genus, nor where it has been found.” Apparently he did not see Eights’ paper. When the Rev. T. R. R. Stebbing wrote a series of popular articles on pycnogonids for Knowledge in 1902, he: considered Deco- lopoda an amateurish blunder and denied that there could be such a thing as a 1o-legged pycnogonid. A few years later, Dr. J. C. C. Loman, a well-known Dutch zoologist and author of several papers on the Pycnogonida, published a paper to the effect that Decolopoda must have been a monstrosity. Hardly had the ink dried on these contributions when the South Polar expeditions began to return with not one but two species of 10-legged pycnogonids! The new species was Pentanymphon antarcticum, whose generic name indicates that it differs from the well-established octopodous Nymphon only in the possession of an additional pair of legs. Another species, P. minutum, has recently been described (Gordon, 1944). Shortly after the discovery of Pentanymphon, a third type, related to the octopodous Pycnogonum, was found, one species in the Antarctic and another, strangely enough, from French Guiana, and a second species of Decolopoda was collected in the Antarctic. As if this were not enough, a 12-legged specimen, which has been named Dodeco- lopoda mawsoni, was found by the recent British, Australian, and New Zealand Antarctic Research Expedition on the edge of the Antarctic south of Kerguelen. Finally, among the material belonging to the United States National Museum, I found several specimens of still another form of 10-legged pycnogonid, which I have named Pentacolossendeis reticulata, collected as early as 1872 off the Florida Keys. It is something of a mystery why this species remained unde- scribed so long, since it was first collected by William Stimpson, the United States of America (Nature, vol. 146, pp. 238-230, 1940), and Con- gressional Record, vol. 86, Appendix, pp. 3194-3195, 1940. Now, after more than a hundred years, one of Eights’ original specimens of Decolopoda has been found among the collections in the Museum of Comparative Zoology. Although the only information available concerning this specimen is a cryptic “Parchment No. 952” and a catalog entry, “South Shetlands,” the circumstances indicate that this is one of the long-lost types. It is possible that it was presented to Dr. Amos Binney, together with the manuscript and plate referred to in the letter from Eights to Dr. Binney (included in Calman’s paper), and eventually found its way to the Museum of Comparative Zoology. The specimen is lacking a few of the terminal joints of some of the legs and one of the ovigers is detached, but it is otherwise in good condition. Accordingly, the specimen has been designated “Neoholotype, d, M.C.Z. No. 12271.” NO. 18 PYCNOGONIDA—HEDGPETII II author of several species of pycnogonids, and although he died before finding time to study his material, the species was collected again in 1893 by the State University of Iowa expedition, and three more specimens were collected by the Fish Hawk in 1902.* There are, then, seven, or perhaps eight (there may be a third species of Decolopoda), species of decapodous pycnogonids, and one dodecapodous species. They are so far known only from the Ant- arctic and American tropical regions, and several of them are com- mon, to the extent that every expedition manages to collect several specimens. They are neither isolated freaks nor monstrosities, but relatively stable forms. The most curious thing about these extra-legged pycnogonids is their close resemblance to certain “normal” genera, a resemblance which in some cases extends to a particular species. Pentanymphon is simply a Nymphon with an extra pair of legs, Pentapycnon a I10- legged Pycnogonum, and Pentacolossendeis would be Colossendeis without its extra legs. Decolopoda, however, is somewhat different from the thick-set species of Colossendeis which it resembles in that it possesses chelifores, but since chelifores are occasionally retained through the last moult stage in some individuals of the genus, the difference is not as great as it seems. Dodecolopoda is merely an extra-legged Decolopoda, and is so far known only from a single specimen. The most conspicuous example of resemblance between a decapo- dous and an octopodous form is that of Pentapycnon charcoti Bouvier and Pycnogonum gaini Bouvier. Both of these are Antarctic forms: P. charcoti occurs in the South Shetlands, and P. gaini has been col- lected from the Palmer Archipelago, Ross Sea, and eastward to 54° E. Bouvier, who described both species, commented upon the similarities 4 William Stimpson, M. D. (1832-1872), was “a naturalist of no mean capacity” who gathered a fine collection, wrote largely in Latin and was director of the Chicago Academy of Sciences from 1865 to 1872. He lost all his work and collections (including the Pourtalés collection from the Florida Keys) in the Chicago fire of 1871, and never recovered from the shock. In April 1872 he went to the Keys on the Bache, but even this did not revive him and he died on May 26. Nathaniel Southgate Shaler, in his Autobiography, pp. 128-120, has an interesting little story about Stimpson. It happened in those days when Agassiz pére reigned at Harvard and “that Darwinian hypothesis” was not to be mentioned except in private. According to Shaler, Stimpson “was much puzzled by the transitional varieties between many of the species of molluscs he was studying, especially those occuring among the fresh water gastropods. On one occasion I saw him throw one of these vexatious shapes upon the floor, after he had studied it for a long time, put his heel upon it and grind it to powder, remarking, “That’s the proper way to serve a damned transitional form.’ ” DISTRIBUTION OF TEN-LEGGED PYCNOGONIDS IN THE CARIBBEAN m Pentacolossendeis reticulata A Pentapycnon geayl VPycnogonum Sp. Bathymetric range . A 2 38 +4 452 HH fathoms BY “DISTRIBUTION OF EXTRA-LEGGED PYCNOGONIDS IN THE ANTARCTIC x Pentanymphon amarcticum x1 “Eetorn DO Pentanymphon minutum OS Ee eee laeg ODecolopoda australis “ot S? ee y © Decalopoda antarctica LOR? ra 68S 6: Gear ae BATHYMETRIC RANGE g AS Orkneys xOoe dst BSS Peet eon RT Pentapycnon charcoti Ss ° Sandwich Is 300_FATHOMS Bouvet Id. « Balleny Is.” eAuckl land Id. “eM ecouar 1e Id. eee Us : “4% PLodecolopoda mawsoni: Marion la. *Pr Edward Id Ase iois S sp ; Heard Wve ees - “Crozet Is. Vagnatiee picnic is. Fic. 4.—Distribution maps of polymerous pycnogonids. 12 No. 18 PYCNOGONIDA—-HEDGPETH 13 between them at some length (1913, pp. 157-160), but placed stress on the last dorsal trunk tubercle of P. gaini as a vestigial remnant of the lost fourth somite, in support of his theory that the 10-legged species were the primitive forms. The essential difference between these two forms, according to Bouvier’s figures, is the dilated pro- boscis, adorned with a dorsal tubercle, of Pentapycnon charcott. Ac- cording to Gordon (1944, p. 69), the proboscis of Pycnogonum gaint is sometimes dilated at the tip and may also bear a noticeable tubercle. It would seem, then, that these two forms are closely similar, with essentially the same range of variation, and that it would be impos- sible to refer a specimen lacking the posterior segments to its “genus.” This same parallelism is evident, but not as pronounced, between the tropical American Pentapycnon geayi Bouvier and a West Indian species, Pycnogonum sp.° Here again the most conspicuous difference between the two forms is the shape of the proboscis, which is longer and more tapered in the decapodous form. Also, its do#sal tubercles are taller. The ovigers are almost identical and could not be told apart if separated from the specimens. Unfortunately there is not enough material available of either species to determine the range of variation. Turning to the 10-legged nymphons, we find similar examples of paired species. Although it might be protested that the genus Nymphon is such a large and complex one that it would be easy to find an approximate counterpart of a 10-legged form, the case is strengthened by the existence of a double parallel, in which the two decapodous forms are closely related to a pair of octopodous species, Nymphon hiemale Hodgson and Nymphon gracillimum Calman. The most widely distributed decapodous form, Pentanymphon ant- arcticum,® differs from its cognate “normal” species, Nymphon hie- 5 This is an unpublished species. I regret that this discussion necessitates mention of this species before its formal description in my forthcoming mono- graph of Western Atlantic and Caribbean species. 6 If the future taxonomists act on the suggestion that such decapodous genera as Pentanymphon, Pentapycnon, and Pentacolossendeis cannot stand alone, it should be noted that both antarcticum (Miers, 1879), and minutum (Goodsir, 1842) have been used for Nymphon and that nomina nova might be required for the pentamerous forms, since these names would be unavailable for trinomial designations. As a matter of convenience, the pseudogeneric names should be retained, but in any event, names for new decapodous species should be different from those in the respective octopodous genera. Dr. Hobart M. Smith (Science, vol. 102, No. 2643, pp. 185-189, 1945, Categories of species names in zoology) has proposed an elaborate classification of species names, which does not, how- ever, suggest a solution for this particular problem. It might be feasible, if it 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Fic. 5—a, Pycnogonum gaini; b, Pentapycnon charcoti; c, Pycnogonum sp. }; d, Pentapycnon geayi. (a and b, and oviger of P. geayi after Bouvier, 1913; c and d, original and to same scale.) male, in the following respects: The tarsus of P. antarcticum is somewhat shorter than the same joint in NV. iemale, the third joint of the palpus may be slightly longer, and the compound spines of the turns out these forms actually represent the same species, to adopt some sort of exponential notation, such as Nymphons hiemale [antarcticum], or N. hiemales, etc. It does not seem to me that these pseudogeneric names can be used as subgeneric categories, as the forms represented do not conform with the usual conception of a subgenus. No. 18 PYCNOGONIDA—HEDGPETH 15 oviger are somewhat different (see fig. 6). According to Hodgson’s figures, the differences between these compound spines are conspicu- ous, but in the two specimens I have examined they are not so signifi- Fic. 6.—Nymphon hiemale and Pentanymphon antarcticum. a, dorsal view of trunk; b, terminal joints of oviger; c, chela; d, tarsus and propodus ; e, compound spines of oviger; f, compound spines of oviger (after Hodgson, 1907): 1, P. MECH, 2, N. hiemale. Scale of magnification for each pair of structures is the same. cant. Evidently there is some variation in the conformation of these spines with individual specimens, for the differences are too great to be explained by artistic interpretation of a minute structure whose 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 location renders large-scale camera-lucida drawings impossible. The two species are conspicuously alike in general appearance and con- formation of the chelae. The other decapodous species, P. minutum, is a smaller edition of P. antarcticum, and is very close to N. gracillimum, which in turn appears to be a smaller form of N. hiemale. It is closer to N. gracil- limum than P. antarcticum is to N. hiemale, especially in the structure of the compound spines and proportions of the tarsal joints. Since all four of these forms are highly variable in these and other details, TABLE 2.—Ratios and measurements of Nymphon and Pentanyphon (Compiled principally from Gordon, 1932.) a = iS ae 3 : See © 2 = sm ey = s s ~~ = s So | 8 =. | Seu & ge} & me | : | 8 | ga cS Chen anes s§ oe 5 S “8 s RS ay S z B 2 ae c Length of trunk, mm... 6.-7.2 5.5 4.8 5.8 @ou 2.8 Spinules on chelae..... 24/29-36/40 25/30 | 22/24 Sale 31/33 | 25/29 Palpus, jts. 4F5 Sate e . 81-1 .08 .93 .85 .96* .66 ros Spines on oviger...... 31-43 40 33 40 43 | 28-33 Leg: coxa 2 O0—2.2 I 2a 1 .2—2-0) leiS 25 mseEy 7209s 49 2-2. 7 4 tarsus nranodae sleneyavela Mavelans 83-1 .16 ea 1-33) p03 are 75 93 length; jamimis aoc era eceneye crore 34.95 | 41.90 wees) 275 Omens x00 Leg/trunk ratio....... 7.257 6.4 Shee: |p ALAS 4-5 5.36 * Determined from Gordon’s (1944) figures. + Determined from Hodgson’s (1907) figures. t+ Determined from ‘Calman’s (1915) figures. and all are found in the same general regions of the Antarctic, it seems evident that we are dealing with a common racial stock. These relationships are clearly brought out in table 2, from which it will be noted that the most significant deviation between the octopodous and decapodous forms is the ratio of the length of leg to trunk. This, how- ever, is not surprising, but indicates that there has been no material increase in the length of the leg with the addition of an extra somite, which results in a radical reduction of the ratio. A further interesting fact is brought out by logarithmic plotting of the trunk-leg ratio of these four forms (fig. 7). As is to be expected, the trend is downward, but the much smaller size of the measured no. 18 PYCNOGONIDA—HEDGPETH 17 specimen of Pentanymphon minutum places it outside this trend. However, if it had happened to be twice as large, its position on the graph would be precisely where expected, namely, in the same rela- tion to Pentanymphon antarcticum as Nymphon gracillimum is to N. hiemale. Therefore it can be assumed that the trend of the leg- trunk ratio of Pentanymphon minutum is parallel to that of the other three forms. This close relationship between decapodous and octopodous forms is not so evident for the species of Decolopoda, since the retention of the chelifores sets them apart from Colossendeis at the outset. However, the Colossendeis most closely resembling Decolopoda, C. (see Gl P antarcticum ‘oO ic. (position of ‘4 3 =P minutum if 5 size were doubl led)” SaaleS Sy Sales: BO. aes E Sy (presumed trend of ratio) so SEE oes cy ee “2 minutum 5 log, length of leg ZOmm 30 40 50 60 70 8090 Fic. 7—Logarithmic graph of the ratios of Nymphon and Pentanymphon. wilsoni, shows a close agreement in proportions with D. antarctica, and this resemblance is further emphasized by the fact that C. wilsoni has eight-jointed palpi, instead of the usual nine for the genus. From table 3, giving ratios and comparison of anatomical characters, it can be seen that C. wilsoni agrees more closely with D. antarctica than with D. australis except in the leg-trunk ratio, in which respect it agrees with the specimen of Decolopoda from Heard Island measured by Gordon (1944). It is of interest to note that this Heard Island specimen, which is the nearest record for Decolopoda to the type and only known locality for Dodecolopoda, shows more similarities to Dodecolopoda than to these species of Decolopoda from the American quadrant of the Antarctic. The intermediate character of the ratios of Colossendeis wilsoni, as contrasted with those of Decolopoda antarctica from the Antarctic archipelago and D. sp. from Heard Island, is of 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Fic. 8.—a, Colossendeis wilsoni (after Calman, 1915) ; b, Decolopoda australis ; c, Dodecolopoda mawsoni (after Calman and Gordon, 1932). Dorsal and lateral views of trunk. All drawings to the same scale. NO. 18 PYCNOGONIDA—HEDGPETH 19 further interest in view of the occurrence of C. wilsoni at Cape Adare, about midway between those localities. As Calman and Gordon (1932, p. 110) pointed out, the occurrence of a pycnogonid with six trunk somites “does not really involve any important modification of the problem presented by the ten-legged species.” As table 3 shows, the widest divergence between Dodeco- TABLE 3.—A natomical characters and measurements, Colossendeis, Decolopoda and Dodecolopoda (Compiled principally from Gordon, 1932 and 1944.) Decolopoda Col: dei Decolopod. Decolopod: Dodecolopod si ane | eat am. tena | Poe Trunk: length, mm....... aD Oe Wl | Meter eons ete O=IOR L7G ys oe 18 ARCA eases cae rac eee 2 SM ests ee mle soe 148 207 Age? Vee ea AOS meeb ite ros BLS hl arg Serene 5.28 width, 2d lat. proc. a 94 96-1 .00 aO5 02 97 1.00 Palpus, No. of jts... 8 8-10 9 9 9 Chelifore: length, Ist jt. fe length of trunk’ *" x G7 os =o5-85 746 75 length Ist jt, . a se x 6.00-8.00] 4.0-5.3 30 5.4 ELSIE cs ee x palm long, |palm short,|palm short,|palm short, fingers fingers fingers fingers slightly strongly | strongly | strongly arched arched arched arched Eye tubercle....... more than | more than] less than | less than | less than half width | half width | half width | half width | half width of cephalic | of cephalic | of cephalic | of cephalic | of cephalic seg. seg. seg. seg. seg. Leg: length, mm....... BAS) Fi) nec DOSFOOT Mi ies hs 240.3 tibia 1 6 |1.05-1.08 - 1.1 ea i -O5-I1. I.00-I .04 if G7) -19 mb 122-7 — 2 1.20 ae es 125 . ene OA Tats 1.24 . leg * oe eo 6.5 TO} TO-—Tte 5.40 13.4 * Estimated from photograph of neoholotype. lopoda and Decolopoda is the leg-trunk ratio. Unlike the decapodous nymphons, in which the body length is materially increased by the addition of a fifth somite, Decolopoda and Dodecolopoda appear to incur no noticeable increase in body length with the addition of so- mites over the ratio for the closely related C. wilsoni. Recognizing that their species might just as easily be called a Decolopoda, Calman proposed a new generic name for it simply as a taxonomic conveni- 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 ence, and it is certain that Dodecolopoda represents one more step in the pattern of variation beyond Decolopoda. Its cognate decapodous form is probably Decolopoda australis. Unlike the decapodous forms of the other genera, these polymerous colossendeids are much larger than the related octopodous form. This larger size, however, is not disproportionate, but appears to represent an arithmetic progression between Colossendeis wilsoni, Decolopoda australis and Dodecolopoda mawsoni. Furthermore, when the ratio of trunk length to leg length is plotted against the Dodecolopoda mawsoni-+- trunk-leg ratio -*Colossendeis wilsont 30 40 50 60 70 8090100 120 130 140 150 area of trunk, sg. mm Fic. 9.—Logarithmic graph of trunk-leg ratio as compared to trunk area of polymerous colossendeids. area of the trunk (as determined by projecting the trunk on squared paper), the relationship between the three forms is very similar to the curves for differential growth of a part as compared to the whole of the different stages of the same organism (fig. 9). Many such graphs of heterogonic growth are presented by Huxley in his work on rela- tive growth.” Admittedly, the available data is inadequate, but it appears to be consistent. Plotting the trunk and leg lengths (fig. 10), for example, gives virtually the same curve as the trunk-leg ratio to trunk area graph. There is the same angle of 30° and the location of Decolopoda australis at about one-fourth the distance (or 1:3) be- 7 Huxley, Julian S., Problems of relative growth. 276 pp. New York, 1932. No. 18 PYCNOGONIDA—-HEDGPETH 21 tween C. wilsoni and D. mawsont. A further coincidence is the indica- tion that the curve for leg-trunk ratio of the Nymphon-Pentanymphon group is also at an angle of 30°, although it has a downward rather than an upward trend. More precise and extensive data might reveal some interesting facts about ratios and growth rates, especially if a growth series could be assembled. It would be particularly interesting to verify the Nymphon-Pentanymphon curve, and confirm the apparent trend of Dodecolopoda mawsoni 2 14 15 16 17 18 13 12 Decolopoda australis ob sPentanymphon antarcticurn C wilson Ny mpron hiemale se Vymphon gracillimum Pheu ota ee ee ee agth of trunk mt a S é ie =4-Pentanymphon minutum ow log log. length of leg TR Soimm 60)" 100 130 160 200 230 Fic. 10.—Combined logarithmic graph of trunk-leg ratios of Nymphon, Penta- nymphon, Colossendeis, Decolopoda, and Dodecolopoda. 30° for the polymerous forms in both families. It is not certain that much can be proved by such analysis, other than to demonstrate a common set of numerical values for the different types of polymerous forms. One should be wary of inferring too much from the trends revealed by logarithmic plotting, for they are inherent in the method itself. It is easy to become bemused by these pretty graphs, and they have fascinated several biologists to the extent that their contributions on the subject might be termed a logarithmic analysis of the Adyos. Although Pentacolossendeis reticulata is not a rare species, and may presumably be collected almost at will along the hundred-fathom line south of the Florida Keys, no closely related Colossendeis from 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 the same region has come to light. It is of particular importance, however, in demonstrating that the occurrence of 10-legged pycnog- onids in the tropical American region is not an isolated phenomenon, confined to a single species. It is also of further interest in that in two of the five known specimens the second trunk somites are smaller, and the second pair of legs arising from this somite are slightly shorter than the remaining legs. Since this difference occurs not once, but twice, it would seem that this is not an individual variation, but is in some way correlated with the decapodous condition. It is difficult to ascribe any external environmental cause to the origin of the decapodous (and dodecapodous) forms, for two more diverse sets of conditions than those prevailing in the Antarctic and the American subtropical or Caribbean regions could not be imagined. These regions are widely divergent in both salinity and temperature. For example, the salinity of the Antarctic region of the South Atlantic ranges from 33 to 35 parts per thousand, whereas that of the tropical American region has a range of 36 to 38 parts per thousand. Indeed, the only physical condition these areas appear to have in common is that both overlie tectonic arcs, areas of stress in the earth’s crust where negative anomalies in gravity may occur.* However, this geophysical condition shows little correlation with the occurrence of decapodous pycnogonids, since they are unknown from the East Indies, Japan, and other island are regions. If future collections do bring more decapo- dous pycnogonids to light, however, it is safe to suggest that the most likely area in which they will be found is the East Indies. It may be significant, for reasons still not apparent, that decapodous pycnogonids are most numerous, both in species and numbers, along the Antarctic arc from the Palmer Peninsula to South Georgia. Furthermore, within this critical area the decapodous forms and their corresponding octopodous forms occur only south of the zone of Antarctic-South Atlantic convergence. Although the available data are not extensive, there seems to be a further limitation of decapo- dous forms to the colder waters of the Antarctic, whereas the cor- responding octopodous forms, at least in the genus Nymphon, occur in higher temperatures. This becomes evident when the distribution of the Nymphon-complex is mapped against the temperature distri- bution (fig. 11). Of course it is also true that a similar correlation can be assumed for salinity, but in this case the range does not seem 8 Measurements of isostasy have not yet been made in the Antarctic, but Hess (Proc. Amer. Philos. Soc., vol. 79, No. I, p. 73, 1938) suggests that a negative anomaly strip “will almost be certainly present around the Cape Horn, South Georgia-Antarctica island arc.” NO. 18 PYCNOGONIDA—HEDGPETH 23 to be great enough to justify any generalization, inasmuch as it is in the magnitude of 00.3 parts per thousand. Because of the virtually identical distribution of the two species of Decolopoda, no generalizations as to their distribution can be inferred. As for Pentapycnon charcoti and Pycnogonum gaini, it would seem, in this case, that the decapodous form is the more northern one, for it was found in the South Shetlands, north of the Palmer Peninsula, whereas Pycnogonum gaini occurs near the base of the peninsula, in the Ross Sea, and along the edge of the Antarctica south of New Zealand and Australia. This distribution pattern is tentative, inas- A Nymphon hiemale. @ Pentanymphon antarcticum. Fig. 11.—Distribution of Nymphon-Pentanymphon compared with temperature of 100-meter surface layer. (Isotherms from Deacon, A general account of the hydrology of the South Atlantic Ocean, Discovery Rep., vol. 7, fig. 12, 1933.) much as Pentapycnon charcoti is so far known from a single collection. The distribution of the warm-water forms in the American sub- tropical region is inadequately known, despite the greater accessibility of the area to collectors, and a comprehensive hydrography of the region is yet to be worked out. Hence little can be said about distri- bution in this area that cannot be inferred from an inspection of the map (fig. 4, A) which indicates all the known localities for decapo- dous forms, as well as for the closely related Pycnogonum sp. Once the existence of 10-legged forms had been established, the inevitable discussion as to their phylogenetic significance got under 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 way. Those who participated in the argument immediately divided themselves into opposing camps: the proponents of the theory that the 10-legged condition represented the original, primitive state of the Pycnogonida, and those who believed it to be a secondary phenom- enon arising out of the octopodous condition. The first to suggest the primitive nature of the decapodous condition was Cole (1905), who argued that Decolopoda represented the progenitor of two di- verging series of phylogenetic lines, leading to Pycnogonum on one hand and to Colossendeis on the other. The most persistent advocate of this position was Bouvier, who maintained his belief in the primi- tive origin of the decapodous type in his last paper on Pycnogonida (1937), in spite of the discovery of Dodecolopoda, for he considered the five ventral ganglia of the octopodous pyncogonid an indication of the original number of trunk somites. If that were the case, we might expect a radical change in the anterior ganglia of decapodous forms, but such is not the fact. In Decolopoda there is simply one more ventral ganglion added to the end of the chain, and the enervation of the cephalic region remains unchanged. (See Gordon, 1932, pp. 128- 130, fig. 73.) Bouvier placed particular emphasis upon a larva described by Dogiel (1911) from the Murman Station in the Arctic as evidence in favor of the primitive character of the decapodous condition. This larva (see fig. 12, c) of Nymphon spinosum was fairly well advanced, and possessed a fifth pair of rudimentary legs on the posterior seg- ment. Dogiel believed this to be an atavistic deformity, but it seems more likely that it was simply an isolated example of faulty develop- ment. If it were actually a throw-back, we should expect it to have an indication of the fifth segment, which it does not have, in the illus- tration at least, and we might also expect it to be a more common occurrence. Dogiel’s example is the only one recorded in the litera- ture. For that. matter, anomalies and deformities seem to be rare among the Pycnogonida, aside from those caused by the regeneration of lost parts. The most conspicuous one I have encountered is a specimen of Achelia borealis from the North Pacific, which has but three legs on the right side. (Fig. 13, b.) There is no evidence of traumatic injury in this specimen and it appears to be a congenital deformity. What is apparently the result of regeneration is described by Schimkewitsch and Dogiel (1913) in a specimen of Anoplodactylus petiolatus from Millport, Scotland (fig. 13, ¢). Bouvier (1914) ex- amined a collection of 3,268 specimens of Pycnogonum litorale from Plymouth, England, and found only one abnormal specimen, a female with seven legs, the last pair being replaced by a median one. Bouvier No. 18 PYCNOGONIDA——-HEDGPETH 25 considered this deformity to be the result of an injury at a fairly early stage. Two indubitable examples of congenital abnormalities have recently Fic. 12—a, Protonymphon larva of Achelia echinata (after Dohrn, 1881) ; b, protonymphon larva of Pentanymphon antarcticum; c, larva of Nymphon ea (after Dogiel, 1911) ; d, larva of Nymphonella tapetis (after Ohshima, 1942b). been described by Ohshima (1942a, b). The first of these is a speci- men of Callipallene brevirostris from Sasebo, in which there are but three pairs of walking legs. Otherwise the specimen is a perfectly formed male, bearing eggs. There are four pairs of trunk ganglia, 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Fic. 13.—Ventral view of trunk of six-legged specimen of Callipallene breviros- tris (after Ohshima, 1942a) ; b, ventral view of trunk of seven-legged specimen of Achelia borealis (from Albatross station 5037, 1906) ; c, six-legged Anoplodac- tylus petiolatus with trifid last leg (after Helfer and Schlottke, fig. 142) ; d, six- legged Nymphonella tapetis with trifid leg (after Ohshima, 1942b). NO. 18 PYCNOGONIDA—HEDGPETH 27 the fourth also serving the third pair of legs so that this pair receives a double set of nerves (fig. 13, a). Obviously the fourth pair of legs is missing here. It is interesting to note that the angle of the third pair of lateral processes in this specimen is approximately the mean value of the angles of the third and fourth processes of a normal indi- vidual. The second abnormality described by Ohshima is more com- plicated. In this case it is a six-legged advanced larva of Nymphonella tapetis, a form in which the legs arise by simultaneous budding rather than by addition from the anterior region in successive molts, as in most other pycnogonids in which the larval stages have been observed, On the left side, the second leg receives nerves from the second and third trunk ganglia, whereas on the right side the third and fourth trunk ganglia serve the third leg, which is trifurcate distally (fig. 13, d). This abnormal distal branching of appendages is not rare among arthropods, and there are several reported examples of its occurrence in pycnogonids independent of abnormalities in segmen- tation. (See Gordon, 1932, pp. 130-131.) It appears to have no relation to the problem of polymerism, although the duplication of nerve supply may have induced it in the examples described by Schimkewitsch and Dogiel, and Ohshima. In the seven-legged specimen of Achelia borealis, the odd leg on the right side receives the nerves from the ganglia which serve the second and third legs on the other side (fig. 13, b). This median leg is so located that it balances the second and third legs of the normal side. Like Ohshima’s six-legged specimen of Callipallene brevirostris, this anomalous specimen is an ovigerous male. Ohshima (1942b) sug- gested that the aberrant specimen of Nymphonella tapetis may have been formed by the failure of the limb buds to divide, but as he anticipated, this would not account for a similar abnormality in a form in which the legs were not formed in this manner. Probably the difference in larval development between Nymphonella and other pycnogonids is actually not as great as he seems to believe. From these anomalies it is apparent that the occurrence of four trunk ganglia is very stable. They would also seem to indicate that the loss of ganglia is a rare occurrence, in contradiction to Bouvier’s suggestion that the octopodous forms have lost a trunk ganglion. It seems more likely that it is easier for a pycnogonid to add ganglia than to discard them, although several forms have post-trunk ganglia in the larvae which are coalesced with the last trunk ganglia in the adult forms, and there is a tendency toward fusion of the anterior trunk ganglia and cephalic ganglia in the compact, disciform types. As early as 1905, G. H. Carpenter expressed the opinion that the 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 decapodous condition was a secondary modification, and he was promptly seconded by Calman in 1909, who reaffirmed his position in his Terra Nova report (1915) as a rebuttal of Bouvier’s contentions in behalf of the primitive character of the decapodous condition. Calman also denied Bouvier’s suggestion that the phenomenon could be localized ; it was Bouvier’s belief that the octopodous forms had lost the fourth trunk somite. Although Calman did not believe that the extra segment could be so precisely localized, his suggestion (Cal- man and Gordon, 1932, p. 111) that “the metameric instability which we believe to have affected the trunk somites may possibly have in- fluenced the segmentation of [the palpus of Decolopoda antarctica],” contains a hint that the instability may be primarily effective in the anterior region. This possibility cannot be ignored, although Dr. Cal- man has assured me, in litteris, that he does not believe it can be so localized. Moreover, the fact remains that the major differences among the families and genera involve the varying combinations of anterior appendages. This is further suggested by Nymphonella, in which the essential difference between it and the closely related Asco- rhynchus is the secondary segmentation of the palpus and first pair of legs. Also, the smaller second trunk somites of Pentacolossendets reticulata would seem to indicate that the extra somite, in this decapo- dous form at least, arose in the anterior region. On the other hand, instability in the last trunk segment is suggested by Ohshima’s six- legged specimen, and he (1942a, p. 260) is of the opinion that the pentamerous forms arose through such instability: “Thus either the increase or decrease in the number of body segments, and conse- quently in appendages, takes place at the junction of the trunk and the tail (abdomen), but not as hypertrophy or abortion occurring at the morphological posterior end of body.” The possible localization of this phenomenon in a particular region is not the major problem, however. Even if that could be satisfactorily answered, the question still remains: what, exactly, is the nature, the cause, and significance of the decapodous condition in the Pycno- gonida? It is a phenomenon without counterpart in any other known group of animals, and the various attempts to compare it with the supernumerary pregenital somites of Polyartemia (Calman, 1915) and the additional gill arches of the shark Pliotrema are of little more than academic interest. The uniqueness of the phenomenon can be appreciated when it is remembered that it occurs in three widely divergent family types, yet is at the same time closely correlated with particular species or species complexes. Furthermore, it is apparently correlated with the evolu- No. 18 PYCNOGONIDA——-HEDGPETH 29 tionary force which governs variation within the group as a whole, since it occurs in those groups in which variation is confined to the specific rather than to the generic rank. As a corollary to this, it is interesting to note that the only established example of supernumerary segmentation in other branches occurs in the Ammotheidae (in which, incidentally, the number of joints of the palpus varies, often within the genus), namely, the reduplicated segmentation of the palpus and first pair of legs of Nymphonella.® It should also be pointed out that this phenomenon of reduplicated segmentation, or polymerism, occurs in those branches of the Pycno- gonida which may be considered, because of their large numbers of narrowly separated and numerically abundant species, as the most successful from the evolutionary standpoint. In other words, the Fic. 14.—Ascorhynchus ramipes (after Lou, 1936); b, Nymphonella tapetis (after Ohshima, 1935b). maintenance of generic form throughout a large series of species in certain branches or families is a possible symptom of dynamic tension —Sivayus, as Aristotle would have called it—and when the tension is high, extra-legged forms are the result.1° Conversely, in those groups where the wide divergence of generic pattern and a correspondingly low ratio of species to genus may indicate a low dynamic potential, the basic metameric stability is not upset. It is of further interest to note that, among the Antarctic species at least, the 10-legged forms 9 Bouvier (1910) reported a specimen of Eurycyde with 17 joints in the palpusg, but this appears to have been an individual anomaly. This genus is closely re- lated to Ascorhynchus. 10 For a discussion of the dynamics of evolution, see Lotka, Alfred J., Elements of physical biology (Williams & Wilkins, Baltimore, 1925), chaps. 2-4. Lotka defines evolution as “the history of a system undergoing irreversible changes.” GE 245) 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 are more abundant and more widely distributed than the correspond- ing 8-legged forms. Incidentally, the relatively greater success of the decapodous forms should be another point against the theory of primitive origin, for evolution does not go backward (although it may sometimes stand still) and there would be no conceivable ad- vantage in reverting to a primitive type once the octopodous type _ had proved so successful. The characters of the decapodous pycnogonids, their close resem- blance to particular species, greater success as organisms (as indicated by their wider distribution and abundance vis-a-vis the cognate octo- podous forms), and overlapping but not precisely identical distribu- tion, suggest that they are polymorphic forms of the octopodous species.’ This cannot be proved until studies of the chromosomes are available, but it seems the most plausible explanation in this day and age when chromosomes are quite the fashion. Certainly it is tempting to suggest that decapodous forms are the immediate result of doubled chromosomes and that the dodecapodous form is a possible tetraploid type. There is some support for the suggestion that this is a poly- ploid condition in the fact that the 10-legged forms occur in what are probably the maximum and minimum temperature ranges for pycnogonids. Temperature extremes appear to induce polyploidy, par- ticularly in plants (cf. Huxley, op. cit., p. 337). Unfortunately, live material of the species involved is inaccessible to laboratory workers, and, for that matter, the normal chromosome number of any pycnogonid is yet to be determined, Furthermore, polymorphism is not necessarily a result of polyploidy, complete or partial, and can only be finally determined by the discovery of both 8- and 10-legged forms in a single brood or from successive or alter- nating broods of a single female. Hence, before this problem can be adequately investigated, it will be necessary to determine the chro- mosome number of common species as well as of those involved in the 10-legged problem, and to develop laboratory culture of living material,?? 11 For an extended discussion of polymorphism, see Ford, E.B., Polymorphism and taxonomy, i1 The New Systematics, pp. 493-513, and Huxley, Julian, Evolution: The modern synthesis, especially p. 96 et. seq. 12 Of course, as Goldschmidt maintains, it is possible that chromosome differ- ences may or may not indicate anything, and that the chromosome pattern may change without visible effect on the genotype, but Goldschmidt’s heresies are not well received in the strongholds of the chromosome cartographers. (See Goldschmidt, Richard, The material basis of evolution, Yale University Press, 1940, especially pp. 186 and ror.) As for laboratory culture, it is probable that it will prove to be relatively easy. Dohrn (1881) kept an amputated specimen No. 18 PYCNOGON IDA—-HEDGPETH 31 Such laboratory investigations may demonstrate that metameric re- duplication among the Pycnogonida is a completely different type of variation than heretofore known, but whatever its mechanism, the fact remains that it is too intimately bound up with particular species to be a random coincidence or genetic accident. It may be discovered that the basic chromosome pattern of the three families in which it occurs is identical and possibly different from that of the other families. The success of this variation, as indicated by its relative abundance, indicates that it is in some way advantageous, although we may not be able to perceive wherein the advantage lies. The late Dr. C. Tate Regan, participating in a discussion of this problem at a meeting of the Linnean Society, remarked that Dodecolopoda and the decapodous forms appeared to be an example of “evolution by acci- dent, a phenomenon difficult to understand.” +% Possibly he had in mind the same difficulty which led Aristotle to deny that variation could be accidental (and hence infinite): “Nature, however, avoids what is infinite, because the infinite lacks completion and finality, whereas that is what nature always seeks.” * Undoubtedly William Morton Wheeler would have considered the polymerous Pycnogonida an example of emergence, which he was careful to restrict to its “epigenetic” meaning, as distinct from the all-inclusive sense (with its overtones of spiritual emergence, crea- tive evolution, elan vital and the rest of it) of less realistic biologists and philosophers. Emergence would indeed be a handy name for this phenomenon, but it is little more than a name, and with all deference to the late Dr. Wheeler, a rather dangerous name, because ot its philosophical aura. As a concept, emergence now has little sanction, either in biology or philosophy. alive for 4 weeks while observing regeneration, and Arita (1937) kept a colony alive in flowing sea-water for 10 weeks. Specimens of pycnogonids collected on the shore often live in small jars for a day or more without a change of water. Temperature control, especially for cold water species, may be as im- portant as salinity and oxygen conditions. 13 Cf. Proc. Linn. Soc. London, Sess. 145, 1932-33, pt. 2, pp. 91-93. 14 Gen. Anim. I, 1. (715 b, 15-16), Loeb Classics ed., pp. 6-7: n O€ pilots peviyer TO Grreipoy TO wey Yap Ameipoy aTEdés, 7 SE Plats del (nTEi TEdos. 15 For a statement of Wheeler’s position, see Emergent evolution and the development of societies, in Essays in Philosophical Biology, pp. 143-169 (Har- vard Univ. Press, 1939). Julian Huxley, in his Evolution: The modern syn- thesis, does not even mention emergence, and denies the need for postulating an “elan vital” (p. 568). For the present philosophical status of the concept, see Irwin Edman’s introduction to the Modern Library edition of Bergson’s Creative evolution. 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Nevertheless, this inferentially invites the ghosts of teleology into the discussion, but they implicitly haunt all speculations in theoretical biology, and no one, including the late Dr. Wheeler, despite his unkind remarks about the neo-Thomists, has yet discovered an efficacious formula of exorcism. The whole range of form and variation within the Pycnogonida is a compact, integrated pattern, and patterns are not aimless accidents induced by genes behaving like Mexican jump- ing beans on a warm day. No one has done more to show that growth and form are achieved in conformity with physical laws than that enthusiastic student of Aristotle, D’Arcy Wentworth Thompson, and if any inference can be drawn from his classic monograph, “On Growth and Form,” it is that teleology is far from being a dead con- cept and that at least one purpose of an organism or group of organ- isms is adaptation to and exploitation of its environment to the limit of its capacity to utilize physical laws.1° True, this can be construed simply as a description of the evolutionary process without invoking the Aristotelean réAos, but so clear a process as evolutionary adapta- tion implies a Cause. Dr. Julian Huxley suggests that the purpose, or Cause, of evolution is Progress, and perhaps this is as good a guess as any, although some will protest that it still leaves us within the philosophical circle without a clear way out.** 16 On growth and form (Macmillan Co., New York, new ed., 1942). “Still, all the while, like warp and woof, mechanism and teleology are interwoven together, and we must not cleave to the one nor despise the other, for their union is rooted. in the very nature of totality.” (P. 7.) See also Lotka, op. cit., chap. 9. 17 Huxley, op. cit., chap. 10. It seems to me that “progress” is an unfortunate term for Dr. Huxley’s conception of evolutionary development. Furthermore, while remaining a staunch mechanist up to his last chapter, he inevitably com- mits that anthropomorphic and logical error of the mechanists, i.e., granted that man is the inevitable result of evolutionary progress, he has now attained the power to interfere with that mechanistic process of evolution which produced him and direct its course so as to alter his own evolutionary future. “The future of man, if it is to be progress and not merely a standstill or a degeneration, must be guided by a deliberate purpose.” (P. 577.) This is tantamount to endowing man with the attributes of divinity, of being a First Cause within himself, and while this is not to deny that man is without the power to improve his racial stock by selective breeding, the full implications of this notion would tempt even a liberal clergyman in a university town to resort to St. Thomas Aquinas: “It is possible for an effect to happen outside the order of some particular cause, but not outside the order of the universal cause.” (Summa Theol. I, Q. 103, Art. 7.) Of course, man cannot expect too much from St. Thomas, who denies such power to the angels (ibid., Q. 52, Art. 2), and says also: “For an indi- vidual man cannot be the cause of human nature absolutely, because he would then be the cause of himself; but he is the cause that human nature exists in the man begotten.” (Ibid., Q. 45, Art. 5.) As an exercise in logic, it would be No. 18 PYCNOGONIDA—-HEDGPETH 33 But biologists are not alone in this philosophical dilemma: physi- cists, having pursued matter down to apparently anarchistic particles they call quanta, now find themselves again obliged to become philoso- phers and speculate upon First Causes, after what had seemed for a time a blessed emancipation from philosophy.'* As long as we search for an explanation for the nature of things as we find them in the natu- ral world, so long will we be haunted by teleology, and that will doubt- less be as long as man is on earth. Of course, it is dangerous to argue by analogy from the human mind, but the basic urge of all great intel- lects, be they scientists or philosophers, theologians or poets, to achieve unity out of the multiplicity of things known and perceived, suggests that Nature is up to the same thing in her endless adaptations of diverse yet basically similar forms to the exigencies of the external environment.?® III. CONCERNING DISTRIBUTION AND DISPERSAL Although the observation of Marcus (1940a, p. 197) that “the active and passive means of distribution of the Pycnogonida seem to be less than those in all other marine arthropods” is essentially true for littoral species, there are several noteworthy examples of widespread distribution which are difficult to explain, and future collections, especially of the smaller forms, may prove many apparently local species to be widely distributed. This, however, would not vitiate interesting to know on what grounds Dr. Huxley assumes that the future trend of human evolution is going to be static or even downhill, since he has assumed that it has been “progressive” up to now. One might also infer that Dr. Huxley, like the late Dr. Wheeler, is more of a Lamarckian than he cares to admit in public, although the Chevalier’s august name is mentioned in his book. As for progress, I will have more to say in Bios, March 1947, under the title “The Philosophic Jellyfish.” 18 Cf, Jeans, Sir James, Physics and philosophy (Macmillan Co., New York, 1943). D’arcy Thompson is, naturally, fully aware of this difficulty: “More- over, the naturalist and the physicist will continue to speak of ‘causes,’ just as of old, though it may be with some mental reservations: for, as a French philos- opher said in a kindred difficulty: ‘ce sont 1a des maniéres de s’exprimer, et si elles sont interdites il faut renoncer a parler de ces choses.’” (Op. cit., p. 9.) 19 Perhaps I do not sidestep “the vitalists, teleologists, et hoc genus omne” as adroitly as my former professor Dr. S. J. Holmes does in his paper, The problem of organic form (Sci. Month., vol. 50, pp. 226-232, 253-260, 379-383, 1944). Dr. Holmes discusses form as a result of chemical and physical equilibria and interactions within the organism. His suggestion that life is “a ceaseless striving for a peaceful heterogeneous equilibrium, the attainment of which would result only in death” is not much different, philosophically, from my own state- ment, although I am limited by my material to external morphology. 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Marcus’ suggestion (idem) that the comparatively slight mobility of the Pycnogonida “may in some cases have favoured the development of the great number of minutely separated species,” for the majority of littoral species are probably limited to comparatively small areas. The most puzzling distribution is that of Ammothella bi-ungui- culata, originally described from the Bay of Naples. This littoral species is easily identifiable by the absence of the terminal claws of the legs, and all workers who have identified it from widely scattered parts of the world have been obliging enough to supply adequate figures, hence there can be little doubt that all the records, as widely scattered as they are, represent the same species. Although most of @Ammothella bi-unguiculata Achelia echinata YAchelia spinesa Fic. 15.—Distribution of Ammothella bi-unguiculata, Achelia echinata and A. spinosa. (Based on Goode Base Map No. 205, by permission of the University of Chicago Press.) these records have been described as geographical varieties, there do not seem to be enough anatomical differences among the various speci- mens to merit subspecific names. This species has been found along the shore of southern California, in Japan, Hawaii, and at Rottnest Island near Perth, Western Australia, in addition to the Bay of Naples. Another widely distributed littoral species is Achelia echinata, which has been identified from northern Europe, the Bay of Naples, the Atlantic coast of Morocco, San Francisco, southern Alaska and the Aleutians, Japan, the Siberian coast near Vladivostok, and Kiao- chow, China. There is also a closely related species from northeastern America, Achelia spinosa, which some taxonomists have considered a synonym of A. echinata, although it seems to me to be distinct enough to merit specific rank. Nevertheless, it is probably a member of No. 18 PYCNOGONIDA—-HEDGPETH 35 the same species complex. The range of variation in A. echinata is apparently large enough to justify a number of geographic varieties, and such a range of variation suggests that it is an older species than Ammothella bi-unguiculata. The distribution of Achelia echinata is that of a typical Boreal species, and may represent a dispersal from higher latitudes as a result of the ice age. On the other hand, Ammothella bi-unguiculata is a warm-water form whose distribution cannot be explained on such geological grounds. Furthermore, the uniform character of the specimens from various localities suggests that it is a young species. Unfortunately, we cannot tell whether or not this distribution antedated the sailing ship with its bottom growth of hydroids and crannies in the hull in which such slow-moving organisms as pycnogonids might find refuge, but its pattern of dispersal suggests that sailing vessels had little to do with the distribution of this species.*° Sporadic distribution, such as that of Ammothella bi-unguiculata, is not a rare occurrence among marine invertebrates, including those forms with limited locomotive powers. The most conspicuous ex- ample to come to recent notice is that of a nemertean. Gorgorno- rhynchus, which is represented by closely allied species recently dis- covered in Bermuda and New South Wales. J. F. G. Wheeler, in an extended paper on this form, which differs from all other nemerteans in the possession of a branched proboscis, advanced the suggestion that the Australian and Bermudian forms arose simultaneously within the last few years, possibly by mutation, and that here was an example of evolution caught in the act. This rather extreme hypothesis over- looks,.as Zimmerman pointed out, the accidents of distribution and collecting, and the possibility of fluctuating populations (at:a low cycle of abundance in the past it might easily have been overlooked) .** 20 Concerning the possibility of dispersal on vessel bottoms, this comment (in litteris) by Dr. J. E. Benedict, Government Naturalist for the Falkland Islands, is interesting: “I have taken Caprella in a tow net in, roughly speaking, the middle of the Atlantic. They were dead and could only have come from the fine bottom growth the ship had acquired in harbour in England.” Shipworms are often dispersed on wooden vessel bottoms. See, for example, Edmondson, C. H., Dispersal of shipworms among central Pacific islands, with descriptions of new species, Occ. Pap. Bishop Mus., vol. 18, No. 15, pp. 211-224, 1946. 21 Wheeler, J. F. G., The discovery of the nemertean Gorgornorhynchus and its bearing on evolutionary theory (Amer. Nat., vol. 76, pp. 470-493), and Zim- merman, E. C., On Wheeler’s paper concerning evolution and the nemertean Gorgornorhynchus (ibid., vol. 77, pp. 373-376). Coe, the nemertean authority, considers Wheeler’s idea a “naive assumption.” Cf. Coe, Wesley R., The nemertean Gorgornorhynchus and the fluctuation of populations (ibid., vol. 78, Pp. 94-96). 36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 However, the character of this variation, namely the longitudinal division of the proboscis, suggests that it may be analogous to the decapodous condition in pycnogonids, itself a variation whose distribu- tion is curiously dispersed. When Wheeler suggests that Gorgorno- rhynchus is a simultaneous mutation in two widely separated parts of the world, caused perhaps by ”an internal inevitable disruption of some sort,” he takes two long steps ahead of his data and a hesitant sidestep toward Lamarckianism. However, it is certain that there are conditions prevailing in some areas (such as the American Tropics and the Antarctic insofar as pycnogonids are concerned) which in- duce speciation, or at least give a kind reception to variations. It follows that there must be a tendency within the organism to enable it to respond to those external conditions. If this be Lamarckianism, so be it. One suspects that many biologists have been browbeaten out of their sympathies toward Lamarckianism, for what is paleontology but a long record of organisms which were capable or incapable of responding to changes in their environment, through the inheritance of acquired or induced adaptations? Whatever the explanation for the distribution of Ammothella bi- unguiculata may be (future collections may prove it to be a circum- tropical species), the distribution of many small species in the North and South Atlantic can be explained on the assumption that the Sargassum provides a medium for their dispersal. At least nine small species are found on both sides of the Atlantic, and on the American side six of these are found at Tortugas in the Florida Keys. On the European side of the ocean these species are scattered from Norway to Cape Verde, and the general pattern of distribu- tion suggests a dispersal from the American side of the Atlantic. At least two of these species, Anoplodactylus petiolatus and Endeis spinosa, are permanent members of the sargassum fauna in mid- Atlantic, and I have found Tanystylum orbiculare, a species known from Brazil and the United States, on Sargassum along the coast of Texas.2? The suggestion that the West Indian region may be a center of dispersal for these various species gains some support from the occurrence of identical and similar species on both sides of the Isthmus of Panama. Perhaps more significant than the 22 Another method of dispersal is suggested by Lebour’s (1916) discovery of larvae of Anoplodactylus petiolatus in the medusa stage of hydroids, at Ply- mouth, England. An excellent summary of the sargassum fauna will be found in the paper by G. Timmermann, Biogeographische Untersuchungen tiber die Lebensgemeinschaft des treibenden Golfkrautes, Zeitschr. Morphol. and Oekol. Tiere, vol. 25, pp. 288-335, 1932. NO. 18 PYCNOGONIDA—HEDGPETH 37 identical species, from the standpoint of the distribution pattern, are the pairs of closely related species, for three of the five Atlantic species with closely related Pacific species occur on both sides of the Atlantic. This may indicate that their occurrence in the western Atlantic antedates their distribution to the eastern shores of the ocean. This relationship is best illustrated in tabular form: CARIBBEAN REGION PANAMIC REGION (chiefly ) Callipallene emaciata * (Tortugas) californiensis (Southern California) Ammothella rugulosa (Brazil, Bermuda, Tortugas) heterosetosa (Galapagos) Ascorhynchus armatus * (Hatteras to Cuba) agassizi ~ (Gulf of California) Eurycyde raphiaster * (Tortugas) longisetosa (Colombia) Tanystylum orbiculare (Brazil to Massachusetts ) duospinum (Central California) * On both sides of Atlantic. t Possibly synonymous with armatus. A curious aspect of the distribution of pycnogonids in the Atlantic is the occurrence of several species in Brazilian and European waters and their absence from northeast America and the West Indies. It is possible that this may be more apparent than real, for collec- tions from the northern shore of South America and the West Indies are very inadequate, and several species described from Brazil have turned up in collections from the West Indies.” According to Ekman, the littoral fauna of the North Pacific is six to eight times as rich as that of the North Atlantic.24 While it does not seem that this is altogether true for pycnogonids, this ele- ment of the fauna is more diversified in the North Pacific than it is in the North Atlantic. There is but one endemic genus in the North Atlantic, Paranymphon, and that is a deep-water, not a littoral 23 See Hedgpeth, 1943b. A more exhaustive discussion of western Atlantic and Caribbean species is now in press. ; 24 Ekman, Sven, Tiergeographie des Meeres, p. 231, Leipzig, 1935. An English version of this work is now in preparation, under the direction of Karl P. Schmidt, of the Chicago Natural History Museum. 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 form. Another genus, Pigrogromitus, is known by a single species ; certainly it cannot be called an Atlantic form. There are at least three genera, Lecythorhynchus, Nymphonella, and Decachela, so far known only from the North Pacific. The most striking difference between the pycnogonid faunas of the two oceans is the relatively few species widely distributed along both shores of the Pacific, in contradistinction to the considerable number of -littoral species (in- cluding Boreal-Arctic forms), found on both sides of the Atlantic. As for the Boreal-Arctic species, it should be mentioned that rela- tively few of them are found in the North Pacific: Nymphon gros- sipes, N. longitarse, and Phoxichilidium femoratum are those most D Decachela L Lecythorhynchus N Nymphonella P Paranymphon P Pigrogromitus X Ainigma alae Q Austropallene OAustroraptus @ Boehmia & DiscoarachneW Hannonia VHeteronymphon X Qorhynchus @ Pycnothea Fic. 16.—Distribution of endemic genera of the Northern and Southern Hemispheres. (Because of dubious Northern Hemisphere records, Ammothea, the most abundant and widely distributed endemic genus of the Antarctic, has been omitted.) (Based on Goode Base Map No. 205, by permission of the Uni- versity of Chicago Press.) certainly present. Oddly enough, species of two of the endemic genera are found on both sides of the Pacific: Lecythorhynchus marginatus and Decachela discata. Pycnosoma may also be endemic to the North Pacific, for Helfer’s (1938) reference of a species from Chile to this genus is open to question (cf. Marcus, 1940b, p. 48). The California coast is especially rich in small littoral forms, there being perhaps 30 species in all identified from the coast be- tween Marin County ‘(north of San Francisco) and San Diego. Although the littoral fauna of Japan is still incompletely known it is evidently a rich one.*° In addition to a large number of endemic 25 qn the collections made by the Albatross in 1900 and 1906, there are 18 undescribed species, 6 of which were taken in shallow water. A systematic report on these collections is now awaiting publication. No. 18 PYCNOGONIDA—HEDGPETH 39 forms, it includes a strong element from the East Indies, which is not found north of 35° N. lat., and a somewhat weaker representa- tion of the Pacific Boreal fauna north of 35°. The facies of the combined littoral and shallow-water (less than 100 fathoms) fauna is markedly different from that of the eastern part of the Pacific. One of the most conspicuous differences is the absence of Tany- stylum from Japan and the northwestern Pacific as a whole, although there are several species along the California coast. This divergence between the fauna of the western and eastern shores of the Pacific can be explained in part by the lack of a convenient bridge of floating sargassum such as exists in the North Pacific. ‘The intru- sion of large masses of Arctic water south of the Aleutian chain is probably also an inhibiting factor, and in this connection it is inter- esting to note that the species found both in Japan and California appear to be cold-water forms, with the exception of dAmmothella bi-unguiculata.*® With the exception of the Antarctic, South Africa, and parts of South America, the pycnogonids of the Southern Hemisphere are known only from sporadic records, and much collecting remains to be done before generalizations can be safely drawn. However, enough is known to confirm again that bipolarity, in the sense of identical species occurring in Arctic and Antarctic regions, does not exist except in the case of ubiquitous or cosmopolitan species (particularly the genus Colossendeis) which are found in deep water in all oceans. Fifty or sixty years ago the bipolar hypothesis received much atten- tion, but D’Arcy Thompson gave it a rough handling, pointing out that the theory had been built upon a foundation of inadequate sys- tematics.27 Now the bipolar hypothesis, insofar as Arctic and Ant- arctic faunas is concerned, is no longer accepted, but the name lingers on and has been applied in a different sense than its originators in- tended. In the words of Sverdrup, Johnson, and Fleming, “bipolar animals need not necessarily be bipolar.” ** 26 For a comprehensive comparison between the Japanese and California coasts, see Gislen, T., Physiographical and ecological investigations concerning the littoral of the northern Pacific. Section I, A comparison between the life conditions in the littoral of central Japan and California, Univ. Arsskr. Lund, (2), vol. 39, No. 5, 63 pp., 1943, and Sections II-IV, Regional conditions of the Pacific coast of America and their significance for the development of marine life, ibid., vol. 40, No. 8, 91 pp., 1044. 27 On a supposed resemblance between the marine faunas of the Arctic and the Antarctic regions. (Proc. Roy. Soc. Edinburgh, vol. 22, pp. 311-349, [1898].) 28 Sverdrup, H. U., Johnson, Martin W., and Fleming, Richard H., The oceans: Their physics, chemistry and general biology, p. 849, New York, 1942. 40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 To be sure, there are an extraordinarily large number of species of Nymphon in both Arctic and Antarctic waters, but the genus is far from rare in tropical waters, and all that can be certainly said of this distribution is that this genus flourishes best in the cold waters of the higher latitudes. With one dubious exception (Nymphon longitarse var. antarcticum), there is no bipolar species of Nymphon. Another significant fact is the absence of 10-legged forms from the Arctic, which gives a radically different facies to the Antarctic fauna. There are, however, some examples of distribution which may be construed as support of the revised bipolar—or abipolar—pattern of distribution. The most conspicuous example is that of the genus Rhynchothorax, one species of which is known from the Isle of Capri in the Mediterranean, while the other appears to be a circumpolar Antarctic and Magellanic species. No intervening records are yet known. Several genera, such as Achelia, Tanystylum, and Pallenopsis, prefer the temperate latitudes of both Northern and Southern Hemi- spheres and are poorly represented in the Tropics. The case of the bathypelagic Pallenopsis calcanea is not so clear. This species has been taken at moderately great depths (600 to 800 fathoms) in Davis Strait and the Indian Ocean off South Africa. There is a third record of this species from the vicinity of Bermuda, and it is possible that this may be a widely distributed species which has not often been collected because of its bathypelagic habit. The Antarctic genus Austropallene is apparently the southern counterpart of the Northern Hemisphere Cordylochele, but a South African species described by Flynn (1928) as Pseudopallene gilchristi differs from Cordylochele solely in the possession of a setose fringe around the mouth, and it is probably actually a Cordylochele.*® This would deprive Cordylochele of its status as.a northern genus and weaken the “bipolar” relationship between Cordylochele and Austropallene. Exclusive of extra-legged forms, and of tropical genera which are found on both sides of the Equator, there are perhaps Io genera endemic to the Southern Hemisphere.*® Of these, four are restricted It is unfortunate that the archaic class designation Arachnoida, comprising sea mites, pycnogonids, and Limulus, is sanctioned in this comprehensive treatise. 29 The presence or absence of a setose fringe may be a specific character in this genus as it appears to be in Pallenopsis. Cf. Pallenopsis denticulata Hedg- peth (1044). 30 Ammothea s. str. may be a southern genus, but there are several dubious Northern Hemisphere records which are not yet confirmed. The taxonomic status of other genera is uncertain. No. 18 PYCNOGONIDA—HEDGPETH 4I to the Antarctic (Austropallene, Austrodecus, Austroraptus, and Heteronymphon), and two are known only from the Cape region of South Africa (Boehmia, Aimigma). There are also two other genera which are characteristic members of the Cape fauna (Dztscoarachne and Hannonia), but both of these have been identified from Port Natal. Pycnothea is so far known from one species at Juan Fer- nandez and another at Rottnest Island. The genus Oorhynchus is known only from a deep-water species taken north of New Zealand by the Challenger. It will be noted that four of the Southern Hemi- sphere genera occur along the South African coast. This concentra- tion is not surprising in view of Ekman’s (op. cit., p. 275) summary of endemic forms from this region. Although there are many small genera in scattered parts of the Tropics, there is only one large genus, Anoplodactylus, which might be considered typically tropical. While it is represented by several species in temperate latitudes, it attains its greatest speciation in the Tropics, especially in the West Indies. There is also but one littoral genus which might be said to be cosmopolitan in the sense that its species occur in about the same proportions (usually two or three well-differentiated species in any given region) throughout the oceans. This is Pycnogonum, and its large number of endemic species is pos- sibly due to the heavy body form and sluggish movements which are characteristic of the genus. In general, it appears that the endemic genera of the Southern Hemisphere are more widely distributed than those of the Northern, which is not surprising in view of the more open character of the southern oceans. Each successive Antarctic expedition establishes the circumpolar distribution of more Antarctic species, and littoral col- lections in the South Sea islands will doubtless bring to light many species described from the East Indies and the Indian Ocean. A close relationship between the fauna of South Africa and the East Indies has already been noted by Flynn (1928, p. 3) who suggests that “the great equatorial current is responsible for such phenomena.” SUMMARY 1. The characters and ontogeny of the Pycnogonida entitle them to the stature of a class or subphylum of the Arthropoda, although their relationship to other groups of arthropods still remains uncertain. 2. The Pycnogonida constitute a compact self-contained group of families without ordinal distinctions. 3. The pattern of variation within the Pycnogonida is correlated 42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 with extra-legged or polymerous forms. In families where variation is most active at the generic rank, polymerous forms do not occur, whereas in families composed of a large number of species in one or two genera, several 10-, and in one case, 12-legged forms are pres- ent. The great majority of species, however, are octopodous. 4. Extra-legged forms are closely similar to “normal” octopodous species, and may be polymorphic forms of these species. Certainly the 10-legged forms and their cognate 8-legged species are representa- tives of the same racial stocks. The occurrence of this polymerous condition in the Antarctic and tropical America, at the temperature extremes of the marine environment, is suggestive of polyploidy as it occurs in many plants. These polymerous forms appear to be more numerous and widely distributed than their cognate octopodous forms, suggesting that they are more successful from the evolutionary standpoint. 5. The Sargassum of the North Atlantic is an active agent in the distribution of small, relatively immobile species in that ocean. In the North Pacific such small species tend to remain endemic. There is no evidence in the distribution of the Pycnogonida to sup- port the outworn concept of bipolarity. There are at least twice as many endemic genera in the Southern Hemisphere, as contrasted with northern waters, and most of these are widely distributed, whereas all the endemic genera (except an Atlantic deep-water form and an anomalous genus from Suez Canal) of the Northern Hemi- sphere are restricted to the Pacific, exclusive of tropical forms. BIBLIOGRAPHY This bibliography comprises all the titles on Pycnogonida, insofar as I have been able to ascertain them, which have been published since the comprehensive bibliography in Helfer and Schlottke (1935), together with omissions from that work. References cited in this paper but not included here will be found in the earlier bibliography. I wish to thank Dr. Isabella Gordon and Dr. Ernesto Marcus for assistance in making this compilation. Anorek, M., and Lamy, E. 1938. Pycnogonides parasites de mollusques. Journ. Conchyl., vol 82, pp. 326-331. Arita, K. 1936. Ein wuberzahliges Bein bei einer Pantopoden-Art (Nymphonella tapetis Ohshima). Annot. Zool. Jap., vol. 15, No. 4, pp. 469-477, pl. 32, 3 figs. 1937. Beitrage zur Biologie der Pantopoden. Journ. Dep. Agr., Kyushu Imp. Univ., vol. 5, No. 6, pp. 271-288, 7 figs. No. 18 PYCNOGONIDA—HEDGPETH 43 Bouvier, E. L. 1937. Etude sur les Pycnogonides du “Travailleur’ et du “Talisman” précédée d’observations systématiques sur les Articulés de ce group. Ann. Sci. Nat., Zool. (10), vol. 20, No. 1, pp. 1-42, 13 figs. CALMAN, W. T. 1937a. James Eights, a pioneer Antarctic naturalist. Proc. Linn. Soc. London, 1936-1937, vol. 149, No. 4, pp. 171-184, pl. 4, 3 figs. 1937b. The type specimens of Pallene australiensis Hoek (Pycnogonida). Ann, and Mag. Nat. Hist. (10), vol. 20, pp. 530-534, 6 figs. 1938. Pycnogonida. The John Murray Exp., Sci Rep., vol..5, No. 6, pp. 147-166, 10 figs. CARPENTER, G. H. 1903. On the relationships between the classes of Arthropoda. Proc. Roy. Irish Acad., vol. 24, No. B 4, pp. 320-360. 1905. Notes on segmentation and phylogeny of the Arthropoda. Quart. Journ. Micr. Soc., vol. 49, No. 3, pp. 469-491. Dawson, A. B. 1934. The colored corpuscles of the blood of the purple sea spider, Anoplo- dactylus lentus Wilson. Biol. Bull., vol. 66, pp. 62-68, pl. 1. DERUuUGIN, K. M. (editor). 1935. Pantopoda of the Polar Seas within U.S.S.R. Inst. Arctique U.R.S.S., Materials for the Study of the Arctic, vol. 4, pp. 1-140, 17 figs. Extin_, H. I. 1936. Pycnogonids from Puget Sound. Proc. U. S. Nat. Mus., vol. 83, No. 2001, PP. 413-422, fig. 33. Face, L. 1942. Pycnogonides de la cOte occidentale d’Afrique. Arch. Zool. Exp. et Gén., vol. 82, Notes et revue, pp. 75-90, 7 figs. BnVNNe els 1 1929. Pycnogonida from the Queensland coast. Mem. Queensland Mus., vol. 9, No. 3, pp. 252-260, 9 figs. Gittay, L. 1934a. Pycnogonida from the coast of British Columbia. Can. Field Nat., vol. 48, pp. 49-50, distributional list. 1934b. Rémarques sur le genre Ammothea Leach et description d’une éspéce nouvelle de la mer d’Irlande. Bull. Mus. Roy. Hist. Nat. Belgique, vol. 10, No. 18, pp. 1-6, 3 figs. 1934c. Note sur quelques Pycnogonides de Villefranche S/Mer (Alpes Mari- times). Ibid., No. 35, pp. 1-5, I fig. 1934d. A new Pycnogonid from Bermuda. Ibid., No. 42, pp. 1-3, 6 figs. 1935. Pycnogonides. Rés. Voy. Belgica, Rap. Sci., Zool., pp. 3-16, figs. I-10. 1937. Pycnogonida. Rés. Sci. “Mercator,” 1. Mém. Mus. Roy. Hist. Nat. Belgique (2), vol. 9, pp. 83-80, 6 figs. 1942. New records of Pycnogonida from the Canadian Atlantic coast. Journ. Fish. Res. Board Canada, vol. 5, No. 5, pp. 459-460. Gorpon, I. 1938. Pycnogonida. Australasian Antarctic Expedition, Sci. Rep. (C), Zool. and Bot., vol. 2, No. 8, pp. 1-40, 8 figs. 1944. Pycnogonida. British, Australian, and New Zealand Antarctic Res. Exp., 1929-1931, Rep., ser. B, vol. 5, No. 1, pp. 1-72, 27 figs. 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 HepcretH, J. W. 1939. Some pycnogonids found off the coast of Southern California. Amer. Midl. Nat., vol. 22, No. 2, pp. 458-475, 2 pls. 1940. A new pycnogonid from Pescadero, Calif., and distributional notes on other species. Journ. Washington Acad. Sci., vol 30, No. 2, pp. 84-87, I fig. 1941a. On a new species of Nymphon from the waters of Southern Cali- fornia. Amer. Midl. Nat., vol. 25, No. 2, pp. 447-440, 1 pl. 1941b. A key to the Pycnogonida of the Pacific coast of North America. Trans. San Diego Soc. Nat. Hist., vol. 9, No. 26, pp. 253-264, pls. 9-11. 1942. Sea spiders. Nat. Mag., October, pp. 414-415, figs. (Popular article.) 1943a. Pycnogonida of the Bartlett collections. Journ. Washington Acad. Sci., vol. 33, No. 3, pp. 83-90, 2 figs. 1943b. Pycnogonida from the West Indies and South America collected by the Atlantis and earlier expeditions. Proc. New England Zool. Club, vol. 22, pp. 41-58, pls. 8-10. 1943c. On a species of pycnogonid from the North Pacific. Journ. Wash- ington Acad. Sci., vol. 33, No. 7, pp. 223-224, I fig. 1944. On a new species of Pallenopsis (Pycnogonida) from western Aus- tralia. Proc. New England Zool. Club, vol. 23, pp. 55-58, pls. 11-12. (In press.) Pycnogonida. Encycl. Brit. The Pycnogonida of the western North Atlantic and the Caribbean. Proc. U.S. Nat.)Mus: Report on the Pycnogonida collected by the Albatross in Japanese waters in 1900 and 1906. Ibid. Hetrer, H. 1935. Meerespinnen. Der Naturforscher, October. (Popular article.) 1936a. The fishery grounds near Alexandria. VIII, Pantopoda. Ministry of Commerce and Industry, Egypt, Notes and Mem., No. 16, pp. 1-6, 3 figs. 1936b. Pantopoda. (2 te Nachtrag.) Die Tierwelt Nord und Ostsee, vol. 31, Teil XIa3, pp. I-5, distributional map. 1938. Einige neue Pantopoden aus der Sammlung des Zoologischens Museums in Berlin. Sitzb. Ges. Nat. Fr. Berlin, 1937, pt. 2, pp. 162-185, 11 figs. He rer, H., and ScHLoTTKE, Econ. 1935. Pantopoda. Bronns KI]. u. Ordn. Tierreichs, vol. 5, Abt. IV, Buch 2, Lief. 1, pp. 1-160, 223 figs. Hixton, W. A. 1934. Notes on parasitic pycnognids [sic]. Pomona Journ. Ent. and Zool., vol. 26, No. 4, p. 57. 1939a. A preliminary list of pycnognids [sic] from the shores of California. Ibid., vol. 31, No. 2, pp. 27-35. 1939b. A collection of pycnognids [sic] from Santa Cruz Island. Ibid., No. 4, pp. 72-74, II figs. 1942a. Pantopoda. Pantopoda [sic] chiefly from the Pacific. I. Nympho- nidae. Ibid., vol. 34, No. 1, pp. 3-7. 1942b. Pycnogonids from Allan Hancock Expeditions. Allan Hancock Pa- cific Exp., vol. 5, No. 9, pp. 277-312, pls. 35-48. No. 18 PYCNOGONIDA—HEDGPETH 45 1942c. Pantopoda (continued). II. Family Callipallenidae. Pomona Journ. Ent. and Zool., vol. 34, No. 2, pp. 38-41. 1942d. Pycnogonids from Hawaii. Occ. Pap. Bishop Mus., vol. 17, No. 3, PP. 43-55, 10 figs. 1942e. Pycnogonids from the Pacific. Family Tanystylidae. Pomona Journ. Ent. and Zool., vol. 34, No. 3, pp. 69-70. 1942f. Pycnogonids from the Pacific. Family Phoxichilididae [sic] Sars 1801. Ibid., pp. 71-74. 1943a. Pycnogonids from the Pacific. Family Ammotheidae. Ibid., vol. 34, No. 4, pp. 93-99. 1943b. Pycnogonids from the Pacific. Family Colossendeidae. Ibid., vol. 35, No. I, pp. 2-4. 1943c. Pycnogonids of the Pacific. Family Pycnogonidae. Ibid., No. 2, p. 10. Lesour, M. V. 1945. Notes on the Pycnogonida of Plymouth. Journ. Mar. Biol. Assoc. U. K., vol. 26, pp. 139-165, 7 figs. Loman, J. C. C. 1938. Note préliminaire sur les “Podosomata” (Pycnogonides) du Musée Océanographique de Monaco. Rés. Camp. Sci. Monaco, vol. 97, pp. 277-286, pl. 4, figs. 11, 19-24. (Reprint of Loman, I9g12.) Lou, T1nc-HEN6e. 1936. Sur deux nouvelles varietes des pycnogonides réceuilles a Tsing-Tao, dans la baie de Kiao-Chow, Chine. Contr. Inst. Zool., Nat. Acad. Peiping, vol. 3, No. 1, pp. 1-34, 4 pls., 9 figs. MANE-GaARZzON, F. 1944. Notas sobre pantopodos. I. Colossendeis geoffroyi, nov. sp., de la plataforma continental frente al Rio de la Plata. Comm. Zool. Mus. Hist. Nat. Montevideo, vol. 1, No. 15, pp. 1-7, 4 figs. Marcus, E. 1940a. Pallenopsis fluminensis (Kroyer) e as Pallenopsis sul-atlanticas restantes. Rev. Ent. Rio de Janeiro, vol. 11, Nos. 1-2, pp. 180-199, I fig. 1940b. Os Pantopoda brasileiros e os demais sul-americanos. Bol. Fac. Fil., Cién. e Letr. Univ. Sao Paulo, vol. 19, Zool. 4, pp. 3-179, pl. 1-17. 1940c.Os Pantopoda. Fil. Cién. e Letr., Orgao do Gremio do Fac. Fil. Cién. e Letr. Sao Paulo, vol. 7, pp. 68-73. (Popular summary.) MELto-LerrAo, A. DE. 1945. Uma espécie nova do género Pycnogonum Briinnich, 1764. (Pycno- gonidae, Pantopoda.) Bol. Mus. Nac. Rio de Janeiro, n.s., Zool., No. 42, pp. 1-4, 5 figs. Moorg, H. B. 1933. New faunistic records for the Manx region. Rep. Mar. Biol. Stat. Port Erin, vol. 46, pp. 30-34. 1934. New faunistic records for the Manx region. Ibid., vol 47, pp. 27-28. Mortey, C. 1940. Arachnida of Suffolk: The remaining Arachnida. Trans. Suffolk Nat. Soc., vol. 4, No. 3, pp. 165-174. NEEDLER, A. B. 1943. Pantopoda. Can. Atlantic Fauna, vol. 10. Arthropoda, 1on, Panto- poda, pp. 1-16, 21 figs. 46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 OusuHIMmA, H. 1935a. “On a sea spider inhabiting the Okinawa region.” Dobutsugaku Zasshi (Zool. Soc. Japan), vol. 47, pp. 137-139. (In Japanese.) 1935b. A further note on Nymphonella tapetis: The egg-carrying mature male. Annot. Zool. Jap., vol. 15, No. 1, pp. 95-102, 4 figs. 1936. A list of Pycnogonida recorded from Japanese and adjacent waters. Zool. Mag., vol. 48, Nos. 8, 9, 10, pp. 861-869. 1937. The life-history of “Nymphonella tapetis’” Ohshima. Compt. Rend. XII* Congr. Internat. Zool., Lisbonne, 1935, pp. 1616-1625, pl. 80, 5 figs. 1938. Nymphonellidae, a new family of Pantopoda. Annot. Zool. Jap., vol. 17, Nos. 3, 4, pp. 220-233. 1942a. Six-legged Pantopod, an extraordinary case of hypomery in arthro- pods. Proc. Imp. Acad. Tokyo, vol. 18, No. 5, pp. 257-262, 3 figs. 1942b. A remarkable case of malformed appendages in a pantopod, Nympho- nella tapetis. Ibid., No. 8, pp. 520-523, 2 figs. 1943a. [Résumé of 1942b in Japanese.] Rep. Ann. Meet. Zool. Soc. Japan 1943, 1 p. (No page numbers in reprint.) 1943b. Du maloftaj kazoj de nenormaleco en pantopodoj. Bul. Sci. Fak. Terk. Kyusu Imp. Univ., vol. 10, No. 4, pp. 371-382, 4 figs. (Japa- nese version of 1942a and 1942b, with short Esperanto summary.) PEt, W. G. 1936. Pantopoda. Flora en Fauna der Zuiderzee, suppl., pp. 131-133. Sawaya, M. P. 1941. Sobre uma larva de Pycnogonum pamphorum Marc. Bol. Fac. Fil., Cién. e Letr. Univ. Sao Paulo, vol. 22, Zool. 5, pp. 278-282, 2 figs. 1945. Anoplodactylus stictus Marc. (Pantopoda) em Caioba, Estado do Parana. Arq. Mus. Paranaense, vol. 4, No. 10, pp. 231-236, pl. 23. Scumitt, W. L. 1934. Notes on certain pycnogonids including descriptions of two new species of Pycnogonum. Journ. Washington Acad. Sci., vol 24, No. 1, pp. 61-70, 2 figs. 1945. Miscellaneous zoological material collected by the United States Antarctic Service Expedition, 1939-1941. Proc. Amer. Philos. Soc., vol. 89, No. 1, p. 297. Snoperass, R. E. 1938. Evolution of the Annelida, Onychophora, and Arthropoda. Smith- sonian Misc. Coll., vol. 97, No. 6, pp. 1-159, 54 figs. STEPHENSEN, K. 1936a. Sveriges pycnogonider. Kungl. Vetensk. Vitt.-Sam. Handl. Goteborg (B), vol. 4, No. 14, pp. 1-56, 13 figs. (Meddl. Goteborgs Mus. Zool. Avd., vol. 69.) 1936b. Pycnogonida from Norway and adjacent waters. Bergens Mus. Arb. 1935, No. 7, pp. I-39, I fig. 1937. Pycnogonida. Zool. Iceland, vol. 3, No. 58, pp. I-13. 1938. Amphipoda. Tanaidacea and Pycnogonida. Zool. Ergebn. Reis. Dr. Kohl Larsen... ., vol. 20, Nos. 3-4, pp. 236-264. 1943. Pycnogonida. The Zoology of East Greenland. Meddl. Gronl., vol. 121, No. 8, pp. 1-41, 7 figs. No. 18 PYCNOGONIDA—HEDGPETH 47 TopsEntT, E. 1938a. Les pycnogonides provenant des campagnes du yacht |’Hirondelle, 1886-1887-1888. (Golfe de Gascogne, Terre-neuve, Acores.) Rés. Camp. Sci. Monaco, vol. 97, pp. 272-276. (Reprint of Topsent, 1801.) 1938b. Pycnogonides receuillis par le yacht Princesse Alice. Ibid., pp. 276-277. (Reprint of Topsent, 1897.) WILLIAMS, G. 1933. On Nymphopsis acinacispinatus, a new pycnogonid from Queensland. Ann. and Mag. Nat. Hist. (10), vol. 12, pp. 173-180, 6 figs. 1940. Pycnogonida of Western Australia. Journ. Roy. Soc. Western Aus- tralia, vol. 25 (1938-1939), pp. 197-205, 9 figs. 1941. A revision of the genus Anoplodactylus with a new species from Queensland. Mem. Queensland Mus., vol. 12, No. 1, pp. 33-39, 5 figs. 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(A, about 4; B, nearly natural size; C, slightly enlarged. ) «} ae "SMITHSONIAN MISCELLANEOUS COLLECTIONS © Bethe APA i i" rs Wa? Mate 0 See Bye n 4 4 4 | Peel § 5 { iy eS | fe Suet gh. nN = oe 3.8 4 Sh os 8 ie ; \ als | t Pin “ol iy be | é a 4 { ie eR Os its i ‘ ¢ Bic’: Wye 4 7 ean " when i Pls Dae if ana / See re x 4 , i AI yA 3 iS ¥ |) ARE + TS ene | . f et 77 | Yass Vl hy tee ae . 1) athe . on + ye o) |) eee ) ee ' * a) nM - ; a ; Vt ee ; \ YY my 4 > ae ‘ Gee | © | * : Ee: 2) (a Saruaek ay \ pi vae ako i re r if 2 PAS pete aN Oe er as — oe a / Br aha k | 4 en 7 a ty uh a : hee a At) nance “| Oa 4; ‘- * _ o 7, SM sl (Saas ‘ | hae ~ 7 ‘ | 8 Sareea os f ¢ i ‘9 ohh Gi ult: 4 7 va j 2 ee % Raa PRAY y ted Se ti ,' ye 4 oo Bo "es ND os © ‘ or ‘ ‘ ee Oe Oe Hy 5: RE is oS Se Sa ee ae ae weet woe Ean i _ ge Seer q VOLUME 106, NUMBER £9". THE LAMINA TERMINALIS AND PREOPTIC RECESS a HN AMPHIBIA. . (Wire ey Puates) 3) BY ALBERT M. REESE West Virginia University ( PuBLicaTion 3867) Br Sete at CITY OF WASHINGTON ie “PUBLISHED BY THE SMITHSONIAN INSTITUTION cabana 27, 1947 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 19 THE LAMINA TERMINALIS AN PREOP TIC; RECESS IN AMPHIBIA (WirH Four PLATEs) BY ALBERT M. REESE West Virginia University (PUBLICATION 3867) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JANUARY 27, 1947 «} 4 oT } a : oO te uf i ef i : +S) ee "a i i \ 1 Y y hs fC u uy a i 0 y i] =i : i 7 AS is be ir vy i i Caen Oe Fe \ « we 4) ‘ ‘ aie ee 7 « Ln | The Lord Waltimore (Press BALTIMORE, MD., U. 3. A. ae LAMINA TERMINALIS AND PREOPTIC RECESS IN AMPHIBIA By ALBERT M. REESE West Virginia University (WirH Four PrateEs) Probably every teacher who has a class in the anatomy of the frog has been asked by numberless students the name of the obvious pro- tuberence on the ventral surface of the brain, just anterior to the optic chiasma. For some unknown reason almost all laboratory manuals fail to mention this perfectly obvious structure. The writer’s interest being aroused, he decided to make a comparative study of this struc- ture, the lamina terminalis, and its adjacent cavity, the preoptic recess, in a considerable number of Amphibia. This seems a rather presump- tuous project in view of the numerous papers on the amphibian brain by Dodds, Fish, Hyman, Kingsbury, and many others; and especially by Herrick. The meanings of the lamina terminalis and of the peroptic recess are variously stated by different workers. Dodds (1907) says: “The lamina terminalis in Amphibia is generally considered as extending from the optic chiasma forward and upward to the choroid plexus.” He also says: “Just in front of the chiasma, it (the brain cavity) ex- pands into a narrow pocket, known as the preoptic recess, representing the primitive lumen of the optic vesicle.” Minot (1903) says: “The upper part of the lamina terminalis be- comes much thickened to form a broad band of triangular section uniting the two hemispheres. This band is the anlage of the septum lucidum, the corpus callosum, the fornix and the anterior commissure.” Hyman (1942) says: “Ventrally the anterior boundary of the third ventricle is a thin membrane, the lamina terminalis, believed by many to represent the original anterior end of the embryonic brain ;” again: “From the anterior commissure a delicate membrane, the lamina terminalis, extends ventrally to the optic chiasma.’ Herrick (1910) says: “Accordingly, the lamina terminalis, preoptic recess and adjacent parts, instead of being assigned to the diencephalon, are re- garded as belonging to the telencephalon.’’ According to Johnston (1909): “In all classes the lateral halves of the telencephalon are con- SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 19 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 nected at the rostral end by the lamina terminalis, a membrane formed by the closing of the anterior neuropore of the embryo.” Ranson (1939) says: ‘The lamina terminalis, connecting the two hemispheres in front of the third ventricle, represents the original anterior-boundary of the telencephalon.” Stieda (1875), in a ventral view of the brain of axolotl, published as far back as 1875, shows a very obvious lamina terminalis, but does not name it. As shown in plate I, figure 1, a sagittal section of the brain of a 4- day chick, copied from Patten (1925), it is largely a matter of choice as to whether the lamina terminalis, J, be considered the anterior wall of the diencephalon, d, or of the telencephalon, te. When the dienceph- alon evaginates to form the telencephalon, the median anterior wall of the former becomes, for a time at least, the median anterior wall of the latter. Patten indicates the theoretical boundaries of the cere- bral vesicles by dotted lines, in the figure under discussion, which would make the lamina terminalis, J, the anteroventral wall of the telocoele, te. In the present study, all the ventral views and their corresponding sagittal sections were drawn with a camera lucida and under the same magnification to show relative sizes. In a few cases, where the exact sagittal plane was not cut, dotted boundaries on the camera drawings have been used, especially to show the deepest part of the preoptic recess. From one to six or more brains, depending upon the available ma- terial, were used of each species. Seven species of Caudata, two of Rana, three of Bufo, two of Scaphiopus, three of Hyla, one of Acris, and one of Pseudacris were drawn. Most of the animals were of about average size for the species. Since the cranial nerves, with the excep- tion of the second, are not being considered, they were not dissected © out, thus saving a large amount of time. In a few cases the pituitary was detached from the brain and is not shown in the figures. . While there is considerable variation in the appearance of the ventral aspect of the various brains, including the size, shape, and distinctness of the lamina terminalis, there is great uniformity in the appearance of the preoptic recess and adjacent structures. The first brain to be described here is that of the common toad, Bufo a. americanus, plate 1, figure 2. The most striking characteristic of this brain is its size, which is equaled only by the brain of the giant salamander, plate 4, figure 20. The lamina terminalis, /, is large, with an indistinct V-shaped ventral groove. The infundibulum, 1, and pitui- tary, », are both large; the optic chiasma, c, and optic tract, ¢, are NO. 19 LAMINA TERMINALIS IN AMPHIBIA—-REESE 3 distinct. In the sagittal section, drawn from a smaller brain, the lamina terminalis, J, is thin ventrally, merging into the chiasma, c, caudad, and thickening cephalad as it merges into the other structures of that region. The preoptic recess, 7, is a very distinct, deep cavity, immedi- ately cephalad to the chiasma, c. The pituitary, p, was torn from the tip of the infundibulum, 7, in dissecting the specimen here drawn. The brain of the marine toad, Bufo marinus, plate 1, figure 3, was from a rather small specimen of this very large species. The optic nerves and tract, ¢, are thicker than in B. a. americanus. The infun- dibulum, 7, is of a different shape and the pituitary, p, is smaller. The lamina terminalis, /, is smaller, but has a similar V-shaped ventral groove. The optic nerves and tract, ¢, are thicker than in B. a. americanus. The sagittal section of this species, from which the camera drawing was made, did not pass through the deeper part of the preoptic recess, v, which was added, as a dotted line, from another section. The lamina terminalis thickens and turns dorsad to form what some authors call the anterior commissure. The optic chiasma, c, is very prominent and is distinct from the infundibulun, 7. The pituitary was missing in this section. The brain of the last representative of the genus Bufo to be de- scribed, plate 2, figure 9, is Fowler’s toad B. woodhousii fowleri, which resembles B. a. americanus so closely in appearance that it is some- times difficult to distinguish them. The ventral view of the brain is very similar to that of B. a. americanus, though the lamina terminalis, /, is relatively smaller. The sagittal view is very similar to that of B. a. americanus, the preoptic recess and lamina terminalis being almost identical. Less than half the size of B. a. americanus, is the common spadefoot toad, Scaphiopus h. holbrooku. The ventral view of this brain, plate 1, figure 4, shows the optic tracts, ¢, as quite distinct from the infun- dibulun, 7, by which they are sometimes hidden. The lamina terminalis 1, is distinctly triangular in shape and shows no ventral groove, which is nearly always distinct and is shown by most authors, even when the lamina terminalis is not named. The sagittal section, it will be seen, by comparing it with plate 1, figure 2, and plate 2, figure 9, shows the preoptic recess, p, and the lamina terminalis, J, to be almost identical with those structures in B. a. americanus and B. woodhousu fowleri. The brain of the southwestern spadefoot toad, S. couchii is shown in plate 1, figure 5. This brain is relatively very broad, so that the pituitary, », infundibulum, 7, and lamina terminalis, ], are much wider, 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 from side to side, than their anteroposterior diameters. As in S. h. holbrookii, there is no visible ventral groove in the lamina terminalis. The sagittal section, of which only one series was available, was badly torn anteriorly and the section drawn did not pass through the deeper part of the preoptic recess, 7, which is dotted in from another section on the same slide. The lamina terminalis, 7, at least in this sec- tion, is relatively smaller than in the other Bufonidae. Of the Caudata seven species were examined, varying in size from the small forms, like the spotted newt, Triturus, to the large forms, such as Cryptobranchus, the giant salamander. The variation in sizes of the brains is very much less than the differences in the sizes of the entire animals. While the brain of Cryptobranchus is about twice the length of that of Triturus, the entire body of Cryptobranchus is about four times as long as that of Triturus. Plate 2, figure 6, represents the brain of a small salamander, Gyrino- philus p. porphyriticus, the purple salamander, about 160 mm. in length. The brain is distinctly long and narrow. The lamina termi- nalis, 7, is elongated with a distinct ventral groove. The optic tracts, t, are distinct, and the infundibulum, 7, is strangely narrowed anteriorly. A satisfactory sagittal section was not obtained. In the figure given the preoptic recess, 7, does not open to the brain cavity and the lamina terminalis, J, is seen only on the floor of the recess. The infundibulum, i, and pituitary, p, are both rather shapeless structures. The brain of Plethodon g. glutinosus, the slimy salamander, is shown in plate 2, figure 7. This animal is about 140 mm. in length. The brain is distinctly broad and rectangular in form. The lamina terminalis, J, is somewhat pointed and has a distinct, narrow ventral groove. The optic tract could not be seen. The infundibulum, 7, and pituitary, p, are large and broad. The section drawn did not pass through the deepest part of the preoptic recess, 7, which is dotted in from an adjacent section. The optic chiasma, c, and infundibulum, i, are hardly distinguishable from each other. The lamina terminalis, /, projects dorsad, as in the ma- jority of figures. Plate 2, figure 8, shows the broad, rather rectangular brain of the mud eel, Siren lacertina. This species is said to reach a length of 700 mm., but the specimen from which the brain here figured was taken was only 150 mm. in length. The lamina terminalis, J, is very large and is nearly circular in out- line with an indistinct ventral groove. The optic tracts are not visible and the infundibulum, 7, and pituitary, p, are curiously wide and short. NO. IQ LAMINA TERMINALIS IN AMPHIBIA—REESE 5 The sagittal section is quite typical as far as the lamina terminalis, l, and preoptic recess, 7, are concerned, The infundibulum, 7, and pituitary, ~, are relatively thick and short. The optic chiasma, ¢, is, as in the preceding brain, hardly distinguishable from the infundibulum. The brain of an average-size mud-puppy, Necturus m. maculosus, is shown in plate 4, figure 17. The brain is broad and massive, with a wide, almost circular, infundibulum, 7, and an oval pituitary, p. The optic nerves are relatively narrow and the optic tracts are hidden by the infundibulum. The lamina terminalis, J, is narrow and relatively very small with a distinct median groove. In the sagittal section the preoptic recess, 7, and chiasma, c, are very similar to those structures in the following brain, Tritwrus, and the lamina terminalis, /, extends into a very tall, narrow anterior commis- sure. The infundibulum, 7, as in Triturus, is not sharply distinguish- able from the chiasma and has a thin wall. The pituitary was missing in the section drawn and was not added from other sections. Plate 4, figure 18, shows the brain of the crimson-spotted newt, Triturus v. viridescens, the small salamander mentioned above. The brain is rather broad and shapely. The lamina terminalis, /, is elon- gated, with a distinct ventral groove. The optic tracts, t, are curiously distinct. The infundibulum, i, and pituitary body, p, are of the usual shape. The sagittal section, from which the pituitary is missing, is quite typical except that it is hard to distinguish the chiasma, c, from the rather imperfect infundibulum, 7. The deep preoptic recess, 7, and the lamina terminalis, /, merging into the prominent anterior commissure, are well marked. The brain of the small, red salamander, Pseudotriton ruber ruber, about 100 mm. in length, is shown in plate 4, figure 19. In contrast to the two preceding brains this one is distinctly long and narrow. The optic tracts are obscured by the very large and curiously shaped in- fundibulum, 7; the lamina terminalis, J, is elongated and has a distinct groove: the pituitary is not shown in either view of this brain. The most favorable sagittal section for other structures did not pass through the deepest part of the preoptic recess, which is shown in dotted outline, ry. The lamina terminalis, /, is similar to that seen in plate 4, figure 18. The optic chiasma, ¢c, projects ventrad more distinctly than in the other brains shown. The floor of the infundibulum, 7, is thicker than in the preceding brains. The brain of the last and largest of the tailed Amphibia to be de- scribed is that of the giant salamander or hellbender, Cryptobranchus 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 alleganiensis, plate 4, figure 20. It is, naturally, much larger than the preceding brains and is broad and massive. The cerebral region is relatively short and the olfactory tracts are very wide. The infundi- bulum, 7, and pituitary, p, are wide, heart-shaped structures of about the same size. The optic tract is not visible and the optic nerves are narrow, as might be expected in an animal with such minute eyes. The lamina terminalis, J, is fairly large, with a distinct groove. The sagittal section is very similar to the preceding except that the preoptic recess, 7, is relatively small and the lamina terminalis, J, and floor of infundibulum, 7, are thicker. The chiasma, c, is scarcely distin- guishable from the infundibulum. The pituitary, p, is massive, but this section did not pass through its longest diameter. Five brains of the tree frogs, Hylidae, were studied. The swamp cricket form, Pseudacris nigrita triseriata, is a tiny form, about one-third the length of Fowler’s toad, yet its brain is considerably more than half the length of the latter, as seen by com- parison of plate 2, figures 9 and 10. The brain is distinctly conical in form, with a very large infundibulum, i, and a curiously shaped pitui- tary, p. The optic nerves are relatively thick and the optic tracts, in the only specimen available, were not visible. The lamina terminalis, I, is relatively enormously large with a median groove that seems to split it into equal halves. The sagittal section is rather typical, but with the lamina terminalis, 1, not extending into a tall anterior commissure, as in the preceding sections. The preoptic recess, 7, is deep and the chiasma, c, projects distinctly in front of the infundibulum, 7. The cricket frog, whose brain is shown in plate 2, figure 11, is about the same size as the preceding; the form used was Acris crepitans, which may be a distinct species or may be a variety of A. gryllus. The brain is of about the same size as the preceding but is a some- what different shape. The lamina terminalis, J, is very much smaller, the optic nerves are narrower and the optic tracts are distinct. The infundibulum, 7, is much smaller. The pituitary, p, was dislodged from the specimen drawn and is indicated by a dotted line. The sagittal section is very similar to the preceding and the optic chiasma, c, is distinctly separated from the infundibulum, 7. The preoptic recess, 7, is large and the lamina terminalis, J, here seen, is fused with a solid mass anteriorly and does not show a projecting anterior commissure. The brain of one of the southern tree frogs, Hyla c. cinerea, is shown in plate 3, figure 14. The frog is about 45 mm. long, consider- ably longer than the two preceding tree frogs, and the brain is propor- NO. 19 LAMINA TERMINALIS IN AMPHIBIA—REESE vi tionately longer and is more massive. It is deeply notched anteriorly and shows the optic tracts fairly plainly. The infundibulum, 1, has the usual shape, while the pituitary, p, is very large and is wider than the infundibulum. The lamina terminalis, J, is externally very small and shows the usual ventral groove. The sagittal section is typical, with a very roomy preoptic recess, r. The optic chiasma, c, is separated from the infundibulum, 7, by a deep groove ; and the infundibulum has a thick ventral wall. The common tree frog, Hyla v. versicolor, is 50 mm. in length, about twice the length of Acris and Pseudacris; but it will be noted, by comparing plate 2, figures 10 and 11, and plate 3, figure 15, that the brain, figure 15, while more massive, is no longer than in the two smaller species. The optic nerves extend laterally at right angles to the long axis of the brain and are so closely applied to the very large in- fundibulum that neither the optic tracts nor chiasma are distinguish- able. The pituitary, p, is wide laterally but is very short in the antero- posterior direction. The lamina terminalis, /, is a moderate-size oval, with a distinct median groove. The sagittal section that was drawn did not pass through the deepest part of the preoptic recess, 7, but the dotted line shows the depth of the recess as shown in an adjacent section. As in the preceding species the chiasma, c, is separated from the infundibulum, 7, by a deep fissure. The spring peeper, Hyla c. crucifer, plate 3, figure 16, is the last of the tree frogs to be described. It is slightly smaller then the preceding and the brain is of about the same length but not so broad. The optic nerves extend straight out to the sides, as in H. v. versicolor, and, as in that form, the tracts and chiasma are not visible. The infundibulum, i, is proportionately wider than in H. v. versicolor, and the pituitary, p, is not so wide, but is somewhat longer. The lamina terminalis, J, is approximately like that of the preceding species. The sagittal section is very similar to H. v. versicolor except that the chiasma, c, projects ventrad in a strange way. The last two brains to be described are those of the common genus Rana, R. p. pipiens, the common leopard frog, and FR. pipiens spheno- cephala, the southern leopard frog. Plate 3, figure 12, R. p. pipiens, is about 90 mm. in length and the brain is fairly massive in outline. The optic nerves are thick and may be traced through the chiasma, c, to the very obvious tracts. The in- fundibulum, 7, and pituitary, ~, are of moderate size and of the usual shape. The lamina terminalis, J, is large and broad, with an obvious ventral groove. The sagittal section is quite typical, with a large preoptic recess, 7, 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 and a thin lamina terminalis, /. The optic chiasma, c, is moderately distinct from the infundibulum. Rana pipiens sphenocephala is somewhat smaller than R. p. pipiens. The brain, as seen in plate 3, figure 13, is considerably narrower and more pointed than the preceding. The infundibulum, 7, pituitary, p, and the optic nerves are not very different from those structures in R. p. pipiens. The optic tract and chiasma, c, are distinct. The lamina terminalis, J, is about the same shape as in the preceding brain, but is much smaller. The sagittal section is quite typical, with a very roomy preoptic recess, 7, and a large optic chiasma, c, fairly distinct from the in- fundibulum, 7. The pituitary, p, was accidentally detached and was so drawn. LITERATURE CITED Donps, G. S. 1907. The brain of one of the salamanders (Plethodon glutinosus). Univ. Col. Stud., vol. 4, pp. 97-108. FisH, PIERRE A. 1895. The central nervous system of Desmognathus fusca. Journ. Morph., vol. 10, pp. 231-282. Herrick, C. JUDSON. 1910. The morphology of the forebrain in Amphibia and Reptilia. Journ. Comp. Neur., vol. 20, pp. 413-457. 1924. The amphibian forebrain. 1. Amblystoma, external form. Journ. Comp. Neur., vol. 37, pp. 457-530. Hyman, Lipsy H. 1942. Comparative vertebrate anatomy. Chicago. JoHNSTON, J. B. 1909. The morphology of the forebrain vesicle in vertebrates. Journ. Comp. Neur., vol. 19, pp. 457-539. Kincssury, B. F. 1895. On the brain of Necturus maculatus. Journ. Comp. Neur., vol. 5, Pp. 139-202. Minor, C. S. 1903. Human embryology. Philadelphia. Osporn, H. F. 1888. A contribution to the internal structure of the amphibian brain. Journ. Morph., vol. 2, pp. 51-06. Patten, B. M. 1925. The embryology of the chick. Philadelphia. Ranson, S. W. 1939. The anatomy of the nervous system. 6th ed. Philadelphia. RUBASCHKIN, W. 1903. Zur Morphologie des Vordershirns der Amphibien. Arch. f. Mik. Anat., vol. 62, pp. 207-243. Sterpa, L. 1875. Ueber den Bau des Centralennervensystems des Axolotl. Zeitschr. Wiss. Zool., vol. 25, pp. 285-310. NO. 19 LAMINA TERMINALIS IN AMPHIBIA—REESE 9 EXPLANATION OF PLATES All figures, except the first, were drawn with a camera lucida and with the same magnification. ABBREVIATIONS c, optic chiasma; cb, cerebellum; d, diocoele; e, epiphysis; 7, infundibulum; I, lamina terminalis; m, mesocoele; my, myelocoele; f, pituitary; 7, recessus opticus; ¢, telocoele. Big: 1 oe Fig. 6. Fig. 12. Fig. 17. 19. 20. PLATE I A sagittal section of a typical embryonic brain. A ventral view and sagittal section of the brain of Bufo a. americanus. West Virginia. A ventral view and sagittal section of the brain of B. marinus. British Guiana. A ventral view and sagittal section of the brain of Scaphiopus h. hol- brooku. West Virginia. A ventral view and sagittal section of the brain of S. couchii. Texas. PLATE 2 A ventral view and sagittal section of the brain of Gyrinophilus. p. porphyriticus. West Virginia. A ventral view and sagittal section of the brain of Plethodon g. gluti- nosus. West Virginia. A ventral view and sagittal section of the brain of Siren lacertina. Georgia. A ventral view and sagittal section of the brain of Bufo woodhousii fowleri. West Virginia. A ventral view and sagittal section of the brain of Pseudacris nigrita triseriata. Arkansas. . A ventral view and sagittal section of the brain of Acris crepitans. Arkansas. PLATE 3 A ventral view and sagittal section of the brain of Rana p. pipiens. West Virginia. . A ventral view and sagittal section of the brain of R. pipiens spheno- cephala. Louisiana. . A ventral view and sagittal section of the brain of Hyla c. cinerea. Louisiana. . A ventral view and sagittal section of the brain of H. v. versicolor. West Virginia. A ventral view and sagittal section of the brain of H. c. crucifer. West Virginia. PLATE 4 A ventral view and sagittal section of the brain of Necturus m. macu- losus. West Virginia. . A ventral view and sagittal section of the brain of Triturus v. viride- scens. West Virginia. A ventral view and sagittal section of the brain of Pseudotriton r. ruber. West Virginia. A ventral view and sagittal section of the brain of Cryptobranchus alleganiensis. West Virginia. a SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 19, PL. 1 (For explanation, see p. 9.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 19, PL. 2 (For explanation, see p. 9.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 19, PL. 3 (For explanation, see p. 9.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106, NO. 19, PL. 4 | (For explanation, see p. 9.) ‘SMITHSONTAN MISCELLANEOUS COLLECTIONS sh eX es at Vee 106, NUMBER cat a ATLANTIC MOLLUSKS OF ATE: il tony. ACLIDIDAE : “Winn 6 Praris) \ BY PAUL BARTSCH U.S, National Museum (Pusrication 3868) ~@ ‘CITY OF WASHINGTON isos PUBLISHED BY THE SMITHSONIAN INSTITUTION : “FEBRUARY a, Ig SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 20 PeeMiONOGRAPH OF THE WEST PeaN TIC: MOLLUSKS OF THE Pail vec TDiDAR (WitTH 6 PLates) BY PAUL BARTSCH U. S. National Museum (PuBLicaTION 3868) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION FEBRUARY 24, 1947 i 14 ioe i f ( i) ih te J f $ ei 5 zy ? ' \ 4} hari ————— \ L | | : TBe Lord Baltimore Vress 2 , BALTIMORE, MD., U. 8. A. ay A MONOGRAPH OF THE WEST ATLANTIC MOLLUSKS OF THE FAMILY ACEIDIDAE By PAUL BARTSCH (WITH 6 PLATEs) The mollusks of this family are all small; some are very minute. They vary in shape from elongate-ovate to turreted. They are usually thin and translucent. The nuclear whorls are small and range from one to more than three; they may form a blunt or acute apex ; they are smooth. The postnuclear whorls are rounded and separated by a well- impressed suture; they may be smooth, axially ribbed, or spirally lirate or reticulated, depending upon the particular genus in question. The periphery is usually well rounded; it may be weakly angulated. The base is usually well rounded, and may be umbilicated, rimate, with an umbilical chink, or not umbilicated. The aperture may be ovate or slightly rhomboid. The outer lip is thin and the inner usually concave. The operculum. is thin, corneous, and paucispiral. The animal is said to have thin cylindric tentacles which are approxi- mated basally, bearing the eyes a little lateral, at their base. The mantle cavity is without a gill. A long retractile proboscis is present, also a pair of dentate jaws. The radula is broad and bears numerous very small, needlelike teeth. The sexes are said to be distinct. The West Atlantic members of the family fall into a number of genera which the following key will help to differentiate. KEY TO THE WEST ATLANTIC GENERA OF THE FAMILY ACLIDIDAE Axial and spiral sculpture both present. PoipmicMsOUdereds << .)...saueteccideer oise se vite a ae seas ened ous Bermudaclis RATE HTGAS PL ATICLL ¥ orc ck avs ap sivas Shots, cots ie Neth Giene craic tel OPNS ve teun eae terete ine Graphis Axial and spiral sculpture not both present. EXaa TAO SEO y PLESEME |e -cyisis aise oa eters hee tees saan tae ce eter Costaclis Axial ribs not present. Spiral sculpture present. Wihoms Shomldered: = 122), .575muz). General Electric light meter reading = approximately 2 foot-candles. Intensity in visible estimated to be of order of 3 ergs/mm.2/sec. thesize chlorophyll. Maize seedlings® of the strain Sh 303(4) x were grown at 26-27° C. under exposure to low intensities of violet light for 10 days and the degree of mesocotyl inhibition determined. The plants were then exposed for several days to sunlight so that 5 Seeds kindly furnished by the Maize Genetics Cooperation, Department of Plant Breeding, Cornell University. NO. 2I MESOCOTYL INHIBITION—WEINTRAUB AND PRICE II the chorophyll-deficient plants might be distinguished. This strain is reported to produce plants in the proportions 75 percent normal (containing both chlorophyll and carotenoids in the leaves) to 18.75 percent white (neither chlorophyll nor carotenoids) to 6.25 percent yellow (carotenoids only). Under the conditions of the experiment, however, we were unable to identify all these types with certainty. The green, normal plants could be clearly recognized but the others ranged from very pale yellow through deep yellow to a greenish yellow. In addition a few plants with very nonuniform pigmenta- tion were classified as doubtful. The results of the experiment are given in table 3. The mesocotyls of the dark controls showed no significant differences in length among the various classes of segregates. In view of the small number of TABLE 3.—Inhibition of mesocotyls of normal and chlorophyll-deficient maize seedlings by \4358A. Average mesocotyl length of dark controls of all classes (23 plants) = 73.6 mm. Intensity = 0.00037 Intensity =0.013 ergs/mm.?/sec. ergs/mm.?/sec. fs ‘8.2 838 2 ‘8.8 88 I 25 22 8 S #5 223 . . e = = oy = vo Classification according to o BS. or & a oe 975 =I pigmentation of leaves after 6 5g oss CS) a oss exposure to daylight Z Ay ey a Ay pmo GSCI BRR icrreeis esse eats 20 77 28.0 19 70 44.4 Pale yellow, yellow, and greenish-yellow ...... 5 19 Tee 26 39.0 DOM ERUIN ry eres ,-0/e0ece I 4 15.8 I 4 44.3 seeds available and the uncertainty of classification, relatively little significance can be attached to this experiment. However, the poten- tially chlorophyll-deficient seedlings tend to show less inhibition than the normal. This is the result to be expected if the photoreceptor were also deficient in such plants. Inhibitions at higher intensities—The action curve for oats shown in figure 1 is derived from experiments at low intensities (1. e., those which result in inhibitions less than 75 percent). Data for intensities causing greater inhibition, although available for only a few wave lengths, do not yield any evidence that the action spectrum for this species is markedly influenced by the intensity. Nevertheless there exists the possibility that such an influence might be found in maize. In testing this possibility it was attempted to duplicate closely the technique employed by Flint. “Red” and “blue” light was obtained by filtering the radiation from a Daylight Mazda lamp through red or I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 blue Cellophane, respectively. The intensity in each case was adjusted to give a reading of 3 foot-candles on a Weston Illumination Meter. Seeds of Zea mays var. Funk’s 176A were planted on agar slants and allowed to develop for 21 days at 18+0.5° C., a period sufficient for completion of mesocotyl growth in the illuminated seedlings and in most of the dark controls. The results, presented in table 4, show approximately equal inhibitions by the two treatments. Although this finding, at first glance, might seem in disagreement with the results obtained with monochromatic light it should be emphasized that the two experiments cannot be directly compared. The sample of blue Cellophane employed exhibited a rather high transmission in the red region of the spectrum, while the tungsten filament source is richer in the red than in the blue. An additional complication is introduced by the spectral response of the photocell which extends to about gooo A.; although the cell sensitivity is rela- TaBLe 4.—Inhibition of corn mesocotyls by light transmitted by red and blue Cellophane. Intensity, 3 foot-candles. Temperature, 18 +0.5° C. Dark Red Blue Number of seedlings........... 37 51 55 Average length of mesocotyl.... 169.3 mm. 28.3 mm. 35.3 mm. Inhibifionttess tere see eee —_— 83.37% 79.270 tively low in the infrared, the lamp emission is very high in this region where both colors of Cellophane have high transmission. Consequently the larger portion of the incident radiation in both cases consists of wave lengths which are relatively ineffective for mesocotyl inhibition. From the transmission curves of the Cellophanes (determined with a Beckman spectrophotometer), the energy distribution of the lamp (Taylor, 1943), and the photocell sensitivity (Weston Electric Instrument Corporation spectral response curve for the photronic cell), it was calculated that, in the region most effective for meso- cotyl inhibition (6200-6600 A.), the energy transmitted by the blue Cellophane is somewhat more than one-fourth as great as that trans- mitted by the red Cellophane. Hence, owing to the flatness of the intensity-inhibition curve at higher intensities, the mesocotyls illu- minated through the blue Cellophane would be expected to show nearly as much inhibition as under the red Cellophane, as was indeed observed. NO. 21 MESOCOTYL INHIBITION—WEINTRAUB AND PRICE nS DISCUSSION In contrast to Markton oats, maize seeds of the varieties employed show very marked variability in rate of germination ; at temperatures as low as 18°, at which germination is relatively slow, the nonuni- formity is accentuated. For this reason there is considerable uncer- tainty in average values for mesocotyl length which are derived from small population samples prior to completion of elongation. We believe that this circumstance accounts for the apparent discrepancy between the results of Flint and those of other investigators, as his estimates of inhibition were made by comparison with dark controls which had made only a small fraction of their ultimate growth. The longest average length recorded by Flint for mesocotyls developed in darkness at 18° is 59 mm., and although it is not specified whether growth had ended in these plants, if they are compared with the illuminated mesocotyls in the same experiment, the inhi- bition by red light at 3 foot-candles is 72 percent, and by blue light 84 percent. Hence Flint’s experimental data appear to be in fair agreement with our own (table 4). SUMMARY Comparison has been made of the relative effectiveness of mono- chromatic red (6234 A.) and violet (4358A.) light in suppressing mesocotyl development in 12 species of grasses representing 8 tribes of the Gramineae. The intensity required to effect a given degree of inhibition is, for all these species, of the order of a thousand times as great for violet as for red light. The uniformity of these responses, which are similar to that of Avena sativa, suggests that the action spec- trum determined for the latter is applicable to all the species in- vestigated. LITERATURE. GCLIED ARAKI, T., and Hamapa, H. 1937. Lokalisation der lichtempfindlichen Zonen von Keimorganen bei Avena sativa L. Bot. Mag. (Tokyo), vol. 51, pp. 498-504. Avery, G. S., Jr., BurKHOLpER, P. R., and Crercuton, H. B. 1937. Polarized growth and cell studies in the first internode and coleoptile of Avena in relation to light and darkness. Bot. Gaz., vol. 99, pp. 125-143. 1938. Growth and cell structure in the first internode and coleoptile of Avena as affected by red, green, blue, and violet radiation. Amer. Journ. Bot., vol. 25, p. Ios. BAUMANN, E. 1911. Untersuchungen iiber Ausbildung, Wachstumsweise und mechanische Leistung der Koleoptile der Getreide. Dissertation Munich. 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 BRUNNER, T. 1936. Uber die optischen Eigenschaften phototroper Keimlinge. Planta, vol. 26, pp. 48-63. Bunnine, E. 1937a. Phototropismus und Carotinoide. II. Das Carotin der Reizaufnahme- zonen von Pilobolus, Phycomyces und Avena. Planta, vol. 27, pp. 148-158. 1937b. Phototropismus und Carotinoide. III. Weitere Untersuchungen an Pilzen und hoheren Pflanzen. Planta, vol. 27, pp. 583-610. CLarK, W. G. 1937. Ascorbic acid in the Avena coleoptile. Bot. Gaz., vol. 99, pp. 116-124. puBuy, H. G., and NuERNBERGK, E. 1929. Weitere Untersuchungen iiber den Einfluss des Lichtes auf das Wachstum von Koleoptile und Mesokotyl bei Avena sativa. Proc. Acad. Sci. Amsterdam, vol. 32, pp. 808-817. iii, IL; 18h 1944. Light and the elongation of the mesocotyl in corn. Plant Physiol. vol. 19, pp. 537-543- Goopwin, R. H. 1941. On the inhibition of the first internode of Avena by light. Amer. Journ. Bot., vol. 28, pp. 325-332. Hamapa, H. 1931. Uber die Beeinflussung des Wachstums des Mesokotyls und der Koleoptile von Avena-Keimlingen durch das Licht. Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, vol. 6, pp. 161-238. (Preliminary report in Proc. Imp. Akad. Japan, vol. 5, pp. 438-441, 1920.) Jounston, E. S. 1937. Growth of Avena coleoptile and first internode in different wave- length bands of the visible spectrum. Smithsonian Misc. Coll., vol. 96, No. 6. Kempton, J. H. 1943. Differential effect of nutrient solutions on the size of various parts of maize seedlings grown in the dark. Journ. Agr. Res., vol. 66, pp. 183-228. OcremiA, G. O. 1924. The relation of soil temperature to germination of certain Philippine upland and lowland varieties of rice and infection by the Hel- minthosporium disease. Amer. Journ. Bot., vol. 11, pp. 437-460. SILBERSCHMIDT, K. 1928. Untersuchungen uber die Abhangigkeit des pflanzlichen Wachstums- verlaufes und der erreichten Endlange von konstanten Temperatur- graden. Bibliotheca Botanica, Heft 97, pp. I-95. Taytor, A. H. 1943. Spectral distribution of energy in common illuminants. Gen. Electr. Rev., pp. 410-413. TERAO, H., and Mipusima, U. 1939. Some considerations on the Classification of Orysa sativa L. into two subspecies, so-called “japonica” and “indica.” Jap. Journ. Bot., vol. 10, pp. 213-258. NO. 21 MESOCOTYL INHIBITION—-WEINTRAUB AND PRICE I5 Wa tp, G., and puBuy, H. G. 1936. Pigments of the oat coleoptile. Science, vol. 84, p. 247. WEINTRAUB, R. L., and McAttster, E. D. 1942. Developmental physiology of the grass seedling. I. Inhibition of the mesocotyl of Avena sativa by continuous exposure to light of low intensities. Smithsonian Misc. Coll., vol. 1o1, No. 17. "SMITHSON IAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 22 . (End of Motanieh et | ‘ | Roebling S und | SOLAR CYCLES tare talee aed aL 5 aha ee tk ee CLAYTON (Pusiication 3870) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION MARCH 5, 1947 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 106, NUMBER 22 (End of Volume) Roebling Fund SOLAR CYCERS BY H.-H. CEAY FON (Pustication 3870) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION MARCH 5, 1947 The Lord Baltimore Preas BALTIMORE, MD., U. & A. ’ i cs ‘ ' At ¢ ei ri eres 4 ‘ , ‘ : aD : md Na bye ote } 4 1 oe 1% iN ‘ 1 - Roebling Fund SOLAR CYCLES By H. H. CLAYTON 1 It is a generally accepted fact that certain terrestrial conditions such as auiroras, magnetic character numbers, ionization of the upper atmos- phere, and earth currents vary in close relation to the sunspot period of about II years. There is also a semiannual period in these conditions with maxima at or near the equinoxes. Two suggestions have been made in explana- tion of this semiannual period. One is that the sunspots are more di- rectly pointing toward the earth at the equinoxes. The equator of the sun is inclined about 7 degrees to the plane of the earth’s orbit, so that spots in the southern hemisphere are directed more nearly toward the earth in spring, and spots in the northern hemisphere are directed more nearly toward the earth in autumn. Another view is that the earth produces tides in the solar atmosphere which result in a slight in- crease in sunspots at the time of the equinoxes. Evidence of a semi- annual period in sunspot areas was presented by Henryk Arctowski * in 1916. He found two small maxima in sunspot areas, one near the spring equinox and the other near the autumn equinox. This research was followed by an investigation of Dr. L. A. Bauer*® in 1924. He found a small semiannual oscillation in sunspots of an amplitude of about two spots with maxima in March and October, corresponding with similar oscillations in the atmospheric electrical potential gradient at Ebro, in earth currents, in the annual frequency of auroras, and in magnetic character numbers, all of which are known to be intimately associated with changes in sunspot numbers. Since the first arrangement and publication of sunspot numbers in tables, by R. Wolf in 1859, there has been speculation and research concerning the question as to whether the planets in their revolutions around the sun might not be the origin of sunspot changes.* _1 Published posthumously. The author died October 27, 1946. 2 Mem. Soc. Spettroscopisiti, vol. 5, pp. 98-90, 1916. 3 Terrestrial Magnetism and Atmospheric Electricity, vol. 20, pp. 23-28, 161- 186, 1924. 4 Wolf, R., Compt. Rend. Acad. Sci. Paris, vol. 48, p. 231, 1859; De la Rue, W., Stewart, B., and Loewy B., Proc. Roy. Soc. London, vol. 20, pp. 210- 218, 1872; Birkeland, Kr., Recherches sur les taches du soleil et leur origine, Skrift. Vidensk., vol. 1, Christiania, 1899; Brown, E. W., A possible explanation SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 106, NO. 22 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 The arguments of Prof. Arthur Schuster in regard to the possibility of small tides in the solar atmosphere, published in the Proceedings of the Royal Society of London in 1911, interested me, and I undertook a research on the subject, using more than a hundred years of sunspot observations, namely, from 1837 to 1938. The results of my studies are plotted in figure 1 and compared with the results of Arctowski. EARTH MAR APR MAY JUN JUL AUG SEP OCT NOV DEC SUNSPOT RELATIVE NUMBERS CLAYTON 1837-1938 SUNSPOT AREAS ARCTOWSKI 1874 - 1913 = ~—~ 2 SUNSPOT RELATIVE NUMBERS ‘ CLAYTON a o \ 1837-1940 MS _ Fic. 1.—Variations in the number and area of sunspots during sidereal revolu- tion of the earth and Venus. Small squares show positions when exerting great- est precessional pulls on the sun. of the sunspot period, Monthly Notices, Roy. Astron. Soc., vol. 60, pp. 599-606, 1900; Pocock, R. J., The relative numbers and areas of sunspots east and west of the central meridian during the years 1902-1917, Monthly Notices, Roy. Astron. Soc., vol. 79, p. 54, November 1918; Schuster, A., The influence of the planets on the formation of sunspots, Proc. Roy. Soc. London, vol. 85, pp. 300- 323 1011: NO. 22 SOLAR CYCLES—CLAYTON 3 The dotted curves show the observed numbers and the continuous curve shows the values computed from them by harmonic analysis. The maxima for the earth come very near the points when the earth is at its greatest distance above and below the plane of the sun’s equator, as shown by the small squares in the diagram. In like manner the aver- ages for the planet Venus show maxima near the points of greatest dis- tance above and below the plane of the sun’s equator. Recently I made a renewed study of this subject in which the revolu- tions of Mercury were included. Because of the shorter period of Mer- cury, this study required the use of daily observations of sunspots, for which data from Zurich were available for the years 1917 to 1944. The means gave evidence of a compound period and hence were analyzed by a Fourier series into periods a, and a,, etc. The results for a, are 0° 40 89 120 160 200 240 280 320 360 & Fic. 2—Averages of daily sunspot numbers, 1917-1944, in periods of 87.97 days. The letters N and S indicate the points where the planet was at its greatest distance north and south of the solar equator. shown in figure 2. The plot shows an oscillation in sunspot numbers corresponding with a half-period revolution of Mercury. Maxima oc- curred at the times when Mercury was at its greatest distance south and at the times when Mercury was at its greatest distance north of the sun’s equator. See the letters S and N in figure 2. The results for the three planets are so consistent that accidental coincidence with the re- quirements of the hypothesis seems impossible. Another finding by Arctowski was that, when the sunspot areas given in the publications of the Greenwich Observatory were submit- ted to analysis, the oscillation in sunspot areas in the northern and southern hemispheres of the sun were in opposite phase. When the earth was south of the sun’s equator there was an excess in the sun- spot areas of the southern hemisphere, but when the earth was north of the sun’s equator there was an excess in the areas north of the equator. It seemed worth while to extend this research to recent observations of sunspot areas. Data for this purpose were found in the measurements 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 of sunspot areas made at the United States Naval Observatory and published in the Monthly Weather Review for the years 1927-1945. These data were averaged in periods of 365.25 days, and the means were subjected to harmonic analysis in a Fourier series extending down to harmonics of the annual period of one-fortieth of a year. 120 150 180 210 240 270 330330 360 OC I DEGREES O° 30 a 9 MONTHS S GO N D N S N EARTH, (927-1995 AREAS #20 oO 28 S6 S# 2 140 169 197 225 N § vo, VENUS, (927-1945,.., Me 22 33 44 SS 66 Ss MERCURY, 1927-19 4E 2 9, Fic. 3.—Oscillations in sunspot areas in the northern and southern hemi- spheres of the sun for half the period of revolution of each of three planets, Earth, Venus, and Mercury. The continuous curve, A, shows oscillations in the northern hemisphere of the sun. The dotted curve, B, shows oscillations in the southern hemisphere. These are sine curves computed by harmonic analysis from the observed data. The letters N and S indicate the points where the planets were at their greatest distance north and south of the solar equator. The data on sunspot areas were taken from the reports of the U. S. Naval Observa- tory, Washington, D. C., for the years 1927-1945. The spot areas are measured in millionths of the total area of the sun’s disk. Taking as a starting point the date when the earth was at its greatest distance north of the equator, the position of the maximum spot area in the northern hemisphere for the full annual period was at 300° with an amplitude of 20 spot areas. For the southern hemisphere it was at 345° with an amplitude of 23 spot areas. That is, both max- NO. 22 SOLAR CYCLES—CLAYTON 5 ima occurred when the earth was near perihelion. For the half-year period, however, plotted in figure 3, the oscillations were in opposite phase in the two hemispheres of the sun. DEGREES g° 30 60 90 120150 180 210 240270 300 aa 4 $ | MONTHS $ MA Jk SPOT— = AREAS ; EARTH 3 YEAR ‘ +20 — Fic. 4—Harmonics of the annual period of the earth computed from obser- vations of sunspot areas 1927-1945, averaged in monthly means and analyzed in a Fourier series. The continuous curves, A, show sine curves for the northern hemisphere of the sun. The curves B show sine curves for the southern hemi- sphere. The data were derived from reports of the U. S. Naval Observatory, Washington, D. C. The letters N and S indicate the points when the earth was at its greatest distance north or south of the solar equator. This opposition in phase is shown also by most of the shorter har- monics of the yearly period, in many of which the amplitudes of the oscillations were larger than that of the yearly and half-yearly periods. The oscillations in the period of one-third, one-fourth, one-fifth, and one-eighth of a year are plotted in figure 4. This plot shows that the 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 oscillations for the period of one-fourth and one-fifth years were larger than the amplitude of the annual and semiannual period. They show further that in most cases the oscillations in the southern hemisphere are of a larger amplitude than those in the northern hemisphere. It is difficult to interpret the full meaning of these findings, but the most satisfactory hypothesis is that the annual curve results from the nearer approach of the planet to the sun at perihelion, and that the larger oscillation in the southern hemisphere in the half-year period is due to the fact that the planet is then nearer perihelion where the attraction of the sun is greater. Similar averages of sunspot areas were obtained for the periods of Venus (224.7 days) and of Mercury (87.97 days) and then were sub- jected to harmonic analysis. The full period of Venus shows a maxi- mum near its perihelion, while the half-year period shows opposing os- cillations, as will be seen from figure 3. Opposing oscillations were also found in the periods of one-third, one-fourth, and other harmonics of Venus, but not so strikingly as in the same harmonics of the yearly period of the earth. The inclination of the orbit of Venus to the plane of the sun’s equator is only 3°50’ instead of 7°10’ as in the case of the earth. The orbit of Mercury is inclined 3°23’ to the plane of the sun’s equa- tor. In the half period of Mercury the same opposition is found in the oscillations in the northern and southern hemisphere of the sun as was found for the earth and Venus (see fig. 3). But the harmonics of one- third, one-fourth, and one-fifth of the period of Mercury are found in the same phase in both hemispheres. This result may be due to the fact that the orbit of Mercury is more eccentric than that of the other planets, and the oscillations of sunspots are influenced more by the varying distance of the planet from the sun than by the north and south pull when the planet is above and below the plane of the solar equator. The data for these studies were derived from the table on page xvii of the American Nautical Almanac and from table 36 in “Earth and Sun,” by Ellsworth Huntington.? Also I was aided in their under- standing and proper use by Edgar W. Woolard of the Naval Observatory. It may be noted that the sunspot changes found in the preceding in- vestigation are small, as were also the changes found by preceding in- vestigators. Perhaps it is for this reason that it is generally believed 5 Yale Press, 1923. NO. 22 SOLAR CYCLES—CLAYTON Fi that planetary influence in producing sunspot changes is so small that if it exists it is negligible. However, in the course of my investigations I discovered that when two planets acted simultaneously their combined influence was much greater than the sum of the influences of the two planets acting separ- ately. The periods of Venus and the earth are so related that they come back to almost the same positions after a period of 8 years. Also the periods of Mercury are so related to the annual period of the earth that they reappear in a similar pattern each Io years and come back nearly to the same point after an interval of 20 years. The closest return of Mercury and Venus to the same heliocentric longitude occurs about every II years. The three planets Earth, Venus, and Mercury also ap- proach the same place at an average interval of about II years. When the monthly sunspot numbers from 1749 to 1944 are averaged in periods of 8, 10, and II years, using approximately the dates when the planets were nearest each other, the amplitude of the oscillations were I5 spots for the period of coincidental influence of Venus and Earth, 35 spots for the period of Mercury and Earth, 65 spots for the period of Mercury and Venus, and 68 spots for the average of the near- est approach of the three planets, Mercury, Venus, and Earth. The nearest approach of the three planets since the beginning of con- tinuous observations about 1830, was 1870 when the time of arrival of all three planets at their greatest departure south of the plane of the sun’s equator differed less than 3 days. The monthly sunspot numbers rose from zero in January 1867 to 176 in May 1870, thus showing the largest oscillation in number of sunspots occurring during the past cen- tury. The most logical explanation of the great increase in the number of spots with the coming together of two or more planets is that of res- onance. It is said that a band playing while crossing a bridge may set the entire structure in oscillation if the music includes the keynote of the bridge. Harlan T. Stetson, who suggested the possibility of resonance in his book, “Sunspots and Their Effects,” ® writes on page 183: On the basis of any accepted tidal theory one would expect that each planet in turn would raise tides, however slight, in the solar atmosphere approxi- mately equal and opposite. The raising of such tidal waves would immediately set the whole solar atmosphere into oscillation, sending an atmospheric wave around the sun which would travel at a speed that would depend upon the density and the gravitational attraction. Each planet, in turn, would start its own similar oscillation, and the composite tidal wave at any moment would therefore depend upon the positions of the planets with respect to each other. 6 McGraw Hill, 1937. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 As the sun rotates carrying the atmospheric particles past the point of major attraction, successive pulses would be increasing the amplitude of the waves so long as the period of oscillation of the atmosphere was comparable with the intervals between successive pulses. In this way it is possible that even the slight tide-raising forces of the planets could in the course of time set up a major oscillation in the sun’s atmosphere, very much the way in which syn- chronized footsteps of a regiment may set a steel bridge asway. I have anchored off my summer place in Maine a forty-foot cabin cruiser. It is of sufficient tonnage so that ordinary movements about the boat do not perceptibly set it in motion. Sometimes, as a matter of amusing experiment, I have stood in the cabin astride the fore-and-aft line and allowed my weight to fall alternately first on one foot and then on the other. Now, the weight of 150 pounds pressing in the floor of a fifteen-ton boat, a foot from the fore-and- aft line, produces of itself no perceptible list. Knowing, however, the natural period of roll of the boat and using this as an interval for alternating the pressure from right to left, I can make the boat roll violently in a very few minutes. It can be stopped equally quickly by reversing my movements, thus making the small force which I can exert oppose the natural roll of the boat. It is shown in figure 4 that the annual revolution of the earth pro- duces harmonic oscillations of short period. This is true also for the TABLE I.—Length of the harmonic periods in days in Mercury, Venus, and Earth Solar Earth Venus Mercury rotation 1/5 =73.0 1/3 F74IQ |) Wealisiaceress) Vohra 1/8 =45.6 1/5 AA. 1/2) ==440) | 0) eee 1/10 = 36.5 1/6 B66) ie wee eaices! » (er 1/12 = 30.4 nfs sez 1/4). 2010 P=27 50 1/16 = 22.8 1/ TO 2255 1/4, = 22130 © 2 ehooeeeee 1/2082 1/12 — 18.7 1/5 Ss'T720" |, aelererstonerenceates 1/32/——siteAl 1/20. 01.2 1/8 S10! i)" eee 1/40 — OME 1/24) 193 1/io= 88 1/37 Or 1/48, —= 7.6 1/36—77.5 Tiie—oes 1/4= 168 1/64== 5.6 1/40= 5.6 T/16.—=se5 1/5 = 55 planets Mercury and Venus. Furthermore, many of the harmonics of the three planets agree closely in length of oscillations, as will be noted in table 1. The fact that oscillations in sunspots approximately coincide with north and south positions of the planets above or below the plane of the sun’s equator led William A. Luby to suggest that the oscillations were of the nature of precessional pulls on an oblate sun similar to the pre- cessional pull of the moon on the oblate earth when the moon is north or south of the earth’s equator. If such a force is exerted by the planets on the surface of the equatorial bulge of the sun, there would be a sur- face current moving toward the north when the planet was on the north NO. 22 SOLAR CYCLES—CLAYTON 9 side of the equator and moving toward the south when the planet was on the south side of the equator. On such an hypothesis one could ex- plain the opposing oscillations in the northern and southern hemis- pheres of the sun found by Arctowski and by me. At latitudes 30° to 45° on the sun, matter carried to great heights is falling back toward the surface of the sun. The orbit of all the planets is near the plane of the sun’s equator, and if one could assume a precessional pull toward the equator exerted by the planets on this matter, then one could ex- plain the formations of the horizonal rolls on the surface of the sun such as are demanded by the theory of sunspot formation formulated by Bjerkness, and now widely accepted. Whatever may be the process by which sunspots are generated, the observational data clearly indicate oscillations in the number of sun- spots related to the planetary periods in length. Some of these har- monic periods are shown in figure 4, and my analysis of sunspot and terrestrial data strongly indicate short-period oscillations of 4.5, 5.5, 7; g, and 11 days." These oscillations increase and decrease in intensity, and the united influence of several planets could easily set the sun’s surface into vio- lent agitation. It will be seen from table 1 that periods of 22.5 days, 18.2 days, 11.2 days, and 7.5 days are close to harmonics of the three planets, Earth, Venus, and Mercury, while 9.1 and 5.6 days are close to the harmonics of the solar rotation period and also to harmonics of the planetary periods. In order to investigate this question in more detail, the dates when the planets were nearest together at the critical heliocentric longitudes mentioned above were taken:from British and American Nautical Al- manacs from 1830 to 1946. The tabulation of these data in table 2, shows that still another factor enters into the problem of timing these returns. The earth and Venus, for example, approach and pass each other 5 times in 8 years, that is, at intervals of about 9 months, and at the end of 8 years differ only .083 day from coinciding, 13 periods of Venus being that much shorter than 8 periods of the earth. Table 2 shows that from 1830 to 1886 these planets came closest at intervals of about 8 years when Earth and Venus were south of the solar equator, then in 1886 stepped forward a half year to when Earth and Venus were north of the solar equator. This makes the average lengths of the period somewhat greater than 8 years. 7 See Solar relations to weather, vol. 2, pp. 20-49, 257-260, 268, 368, and 415, 1943. Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 TABLE 2.—Times of arrival of Venus and Earth at their greatest distances above and below the sun’s equator South North Venus Earth eee Venus Earth Hes 1830—Mar. 5 Mar. 5 oO 1895—Sept. II Sept. 7 +4 1838—Mar. 5 Mar. 5 0 1903—Sept. II Sept. 7 +4 1846—Mar. 4 Mar. 5 —I 191I—Sept. 10 Sept. 7 +3 1854—Mar. 3 Mar. 5 —2 I9g19—Sept. 9 Sept. 7 +2 1862—Mar. 2 Mar. 5 —3 1927—Sept. 8 Sept. 7 +1 1870—Mar. I Mar. 5 —4 1935—Sept. 7 Sept. 7 oO 1878—Mar. I Mar. 5 —4 1943—Sept. 7 Sept. 7 O 1886—Feb. 28 Mar. 5 5 1951—Sept. 6 Sept. 7 —I Fic. 5.—Illustrating the revolutions of Venus and Earth. Venus overtakes and passes the earth in about 19 months, as indicated by the letters Fi, Es, Es, and Es. After 8 years they return near to the point E, but about 0.8 day behind Es. This small difference accumulates, so that after about seven periods of 8 years the planet V is more than halfway back to the earth’s position on the opposite side of the sun’s equator. The planets then approach a meeting at that point in their orbit. This occurs in about 12 periods of 8 years so that the average length of the period is somewhat greater than 8 years. See table 2. Figure 5 is intended to illustrate the passing of two planets after intervals of about 19 months and the return of Venus to the point Ve. This difference slowly accumulates after each 8 years and after about 7 periods of 8 years the planet is more than halfway back toward the place where the two planets are on the opposite side of the sun’s equa- tor. That is, assuming that the planets coincide in time when at their nearest approach north of the sun’s equator, after about 12 periods of § NO. 22 , SOLAR CYCLES—CLAYTON Te years they will coincide in time when farthest south of the sun’s equa- tor. This fact is illustrated in table 2, showing the days of nearest ap- proach in time of the two planets when at their greatest distance north and south of the sun’s equator. These data were derived from the U. S. Nautical Almanac. TABLE 3.—Dates of nearest approach in time of the planets Mercury and Venus when at their greatest distance above and below the sun’s equator Planets North Planets South Diff., Diff., Mercury Venus days Mercury Venus days 1834—Mar. 4 Mar. 6 —2 1836—Dec. 6 Dec. 9 —3 1839—Sept. 17 Sept.18 —1I 1842—June 21 June 24 —3 1845—Apr. I Ape 0 1848—Jan. 5 Jany) 27 —2 1850—Oct. 15 Oct. 14 --I 1853—July 20 July 21 —I 1856—Apr. 30 Apr. 27 +3 1859—Feb. 2 Feb: 2 ) 1861—Nov. 13 Nov. 10 +3 1864—Aug. 17 Aug. 16 +1 1867—May 209 May 24 +4 1870—Mar. 2 Mar. 1 +1 1872—Dec. 13 Dec. 6 +7 1875—Sept. 15 Sept. 13 +2 1879—Sept. Io Sept. 13 —3 1881—Mar. 31 Mar. 27 +4 1885—Mar. 25 Mar. 27 —2 1886—Oct. 14 Oct. 10 +4 1890—Oct. 8 Oct. I0 —2 1893—July 12 July 16 —4 1896—Apr. 23 Apr. 23 oO 1899—Jan. 26 Janwe2s —2 190I—Nov. 7 Nov. 6 +1 1904—Aug. II Aug. 13 —2 1907—May 23 May 21 +2 1910—Feb. 24 Feb. 25 —I I912—Dec. 5 Dees 3 +2 1915—Sept. 10 Sept. 9 +1 to18—June 21 June 17 +4 192I—Mar. 26 Mar. 24 +2 pe en 4 Dec. 30 +5 19z6—Oct. 8 Oct. “6 +2 1930—Oct. 3 Oct. 6 —3 1932—Apr. 23 Apr. 20 43 1936—Apr. 17 Apr. 190 —2 1937—Nov. 6 Nov. 2 +4 1941—Oct. 31 Nov. 2 —2 1943—May 22 May 17 +5 1947—May 16 May 17 —I 1950—Feb. 18 Feb. 22 —4 The planets Mercury and Venus return to near the same point in about 11 years, Mercury makes 46 revolutions in 11.07910 years, while Venus makes 18 revolutions in 11.07373 years. The difference is .00537 year or 1.96 days. However, on account of this small differ- ence the cycle after about 5 intervals of 11 years steps forward sud- denly to a length of more than 12 years and returns closely to its orig- inal position after a period of 56.6 years. This makes the average length of the cycle about 11.32 years. This fact is evident in table 3 where the dates are given of closest ap- proach of the planets to each other at their critical points north and south:of the solar equator. Note the alternate intervals.. These data were taken from the British and American Nautical Almanacs and show the computed places from 1834 to 1950. 12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 Another method of approach in studying the movements of the planets is to plot the nearness of approach in time of the planets in pairs when they are at their greatest distance above and below the sun’s equator. Such a plot is shown in figure 6 for Mercury and Venus and for Venus and Earth, for the years 1901 to 1933. The peaks in the curve show near approach of the planets, while bottoms show that the planets were far apart. Mercury and Venus show 8 peaks in 11 years while Venus and Earth show 7, so that the peaks are in the same phase only once during each 11 years. The nearest approaches came in 1909, 1919, and 1930. The greatest activity in sunspots came 2 years earlier in each case when the peaks were approaching each other and plane- tary influence was increasing fastest. TABLE 4.—Pattern of about So years shown by the approximation in time of the three planets, Mercury, Venus, and Earth (within 30 days), when at their critical positions north and south of the sun’s equator North of solar equator Years of nearest approach 1748 ; 1759 1767 1780 1788 1828 1839 1847 1860 1868 1908 1919 1927 1940 1948 South of solar equator Years of nearest approach 1850 1758 1769 1779 1700 1830 1838 1849 18590 1870 1909 1918 1929 1937 1950 Another way of studying the data is to tabulate the dates since the beginning of sunspot observations when the three planets, Mercury, Venus, and Earth, were nearest to their critical points at the same time. Taking the cases where these planets reached their critical points on the same side of the sun, it is surprising to find that the interval be- tween the occurrences is very irregular, when it is considered that the periods of meeting are derived from three periods, each nearly constant in length. There does appear, however, to be a pattern of about 80 years in the data which is illustrated in table 4. The groups north and south tend to come in pairs, as for example 1748-1750, 1758-1759, 1767-1769, etc. It is interesting to note that all these pairs were years of high sunspot activity, while the inter- vals between them were periods of low sunspot activity. This fact will be seen in the plot of sunspot numbers shown in figure 7. In this plot the smoothed sunspot numbers published at Zurich were used and values were plotted at intervals of 6 months. The first two NO. 22 Py) 00 /90/ | (902) /903| /904 (906 N Ss 1905S B= a Ga ES 1914 | /9/5 NY S (91/2 | 1913 1916 19/7 1923 1924| /925 es 1927| 1928 SOLAR CYCLES—CLAYTON 1907 N 1918 N 1929 5 13 1908 | 1909| 1910\ 1941 s s ls N s ; 1919 | 1920 | 1927 | 1922 N s N 5 Ad /930| 1931 | 1932 | 1933 Fic. 6.—Plot of the interval between the dates when the planets Mercury and Venus, and Venus and Earth, are at their greatest distance north and south of the sun’s equator. The letter N indicates the times of their meeting north of the equator, and the letter S the times of their meeting south of the equator. Peaks in the curve show approach to simultaneous time in meeting. The curve A is for Venus and Mercury, and curve B for Venus and Earth. VOL. 106 SMITHSONIAN MISCELLANEOUS COLLECTIONS 14 ‘sjodsuns ur steaf 0g jo poliod oyewixoidde Surmoys ‘YOLINZ ye AtopeATOSGO [e1Ipay SstMG oy} Aq poztodas ‘gP61-0SZ1 ‘ssoquimu jodsuns payjoouls ay} jo jo[g—Z ‘DIY 626/ 696! 6S 6/ NO. 22 SOLAR CYCLES—CLAYTON 15 curves are separated by an interval of 80 years, while the last two are separated by an interval of 79 years. The shortening of the last inter- val is warranted by the last row of figures in table 4. The plot in figure 7 shows that during the first part of the period covered there were large oscillations in sunspots, culminating about the middle of the plot, followed by a long interval of lesser activity. This change may be due to an increase in the intensity of planetary influence as they approach nearer to a simultaneous meeting at their critical points, as indicated in figure 6. The interval separating the three planets was about 12 days in 1788-1790. It was 3 days in 1870 and will be about 9 days in 1951. After 1785 and again after 1870 the intervals of time separating the planets when at their critical peaks decreased and solar activity also decreased. The present period of solar activity is tending to du- plicate the pattern of 1778 to 1790. If this continues there should be a maximum of solar activity in 1947-1948, continuing at a high level through 1950 after which there would follow a long period of relative quietness. It should also be noted that the interval between the maxima is near 10 years when very active and longer when quiet. In closing this consideration of solar activity and planetary relations, I wish to emphasize the fact that these solar periods are irregular in length and amplitude and are not mathematical periods of constant length and amplitude. For this reason they cannot be treated by ordi- nary statistical methods and forecasts based on that assumption. Such forecasts are subject to failure. This is true whether the sunspot period is considered as a single period, or as a combination of periods. That the planet Jupiter is also an influence in the formation of sun- spots seems evident from the fact that during the past 80 years when Jupiter was north of the solar equator at the time of maxima of sun- spots, as in 1854 and 1927, there were an excess of spots in the north- ern hemisphere. When Jupiter was south of the equator as in 1873 to 1895 at sunspot maxima there was an excess of sunspots in the south- ern hemisphere of the sun. This excess of spots in the two hemispheres is shown in figure 8. This diagram of excess of spots in the two hemi- spheres of the sun at successive maxima of spots was published by W. Brunner in the Astronomische Mitteilungen at Zurich, Nr. 144, p. 119, 1944. Figure 9 shows a plot of the closeness of approach of the planets Mercury and Earth when at their critical points north and south of the solar equator. This pattern tends to repeat itself almost exactly at in- tervals of 20 years and approximately at intervals of 10 years. The ex- 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 9 Cecrirg ere [—- AB HE. AIA“ /283 FP SI ILG CIEE, LY peas 99.54 08Fe 49 /ILL,T¢ (ILI, @ CY wes ens epes 10 1386,3 2963, 2 ao | Aes A? Pe 2650 4° PLD - 49049 t ad Carricglec © per Wt 338 ry, 10 (829,80 + ADGQ,D joee. Ars SO fat 3 C6 0 06S [oe Cerrvagles9 487 M8 £0/-I5F 1? 1869.2 +/079,0 Lovrrer & Mars RO MEL EOE bao C. GEC IP GOLD BY AE PODS 1OS6,0 + O65, 2 BO Corring sr LO ROP NET LP L 9 (OFI,9 -GIE,O cd Fic. 8.—Long-period change in excess of spots in northern and southern hemispheres of the sun at time of maxima in sunspots, 1854-1938. (After W. Brunner, Astronomische Mitteilungen Zurich, 1944, Nr. 144, p. 119.) NO. 22 SOLAR CYCLES—CLAYTON 17 901 | 1902\.1903| 1904| 1905|1906 \1907 | 1908 Ss: N $ N rate 19/2 | 1913 | 191% 1915 | 1916 | 1917 | 1918 jee | 1320 | N ‘i s 1934 1935 Fic. 9.—Plot of the intervals between the dates when Mercury and Earth were at their greatest distance north or south of the plane of the sun’s equator. Peaks in the curve indicate the closeness of approach to simultaneous occur- rence. The zero lines indicate no difference. The letters N and S show when the meeting was north or south of the solar equator. Three prominent peaks are found in each 10-year period at intervals of about 40 months. 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 106 act interval appears to be in alternate periods of 10.5 and 9.5 years. However, the 20-year period is not exact. After four intervals the nearest approach is 79 years instead of 80 years, so that the average length is 9.87 years rather than 10 years. This period of about 10 years is believed to be the same as that of the 10-year period found in sunspots, although the conjunction and opposition of Jupiter and Sat- urn is a period of nearly the same length and may contribute to the result. The results arrived at in this discussion were derived chiefly from the researches of Henryk Arctowski and myself. They depend on two apparent facts: 1. That there are small oscillations in the number and size of sun- spots associated with the passage of the planets Earth, Venus, and Mercury through certain critical points in their orbits. 2. These oscillations increase greatly in range when two planets ap- proach their critical points nearly simultaneously. The increase in range follows in the order Venus and Earth, Earth and Mercury, and Mercury and Venus, the last giving the largest range. 3. The range becomes very large when the three planets Mercury, Venus, and Earth reach their critical points nearly simultaneously as in 1870, but more especially if a near approach in the northern hemi- sphere is followed or preceded within a year or two by a near ap- proach in the southern hemisphere (see table 4). I see no way of avoiding the logical conclusion that there is a relation between planetary positions and sunspots, unless it can be shown that the findings are erroneous or else some other explanation of them can be found. The findings are not dependent on any theory but on ob- served facts. I am quite aware that professional astronomers and meteorologists are averse to finding any relations between sunspots and planets, but so were the professional chemists averse to the idea that chemical elements could be changed until the coming of nuclear physics. y | SMITHSONIAN iii LIBRARIES 3 9088 01421 4951